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Terrestrial animals are animals that live predominantly or entirely on land (e.g. cats, chickens, ants, most spiders), as compared with
aquatic animal An aquatic animal is any animal, whether vertebrate or invertebrate, that lives in a body of water for all or most of its lifetime. Aquatic animals generally conduct gas exchange in water by extracting dissolved oxygen via specialised respirato ...
s, which live predominantly or entirely in the water (e.g. fish, lobsters, octopuses), and semiaquatic animals, which rely on both aquatic and terrestrial habitats (e.g. platypus, most amphibians). Some groups of insects are terrestrial, such as ants, butterflies, earwigs, cockroaches, grasshoppers and many others, while other groups are partially aquatic, such as mosquitoes and dragonflies, which pass their larval stages in water. Alternatively, terrestrial is used to describe animals that live on the ground, as opposed to arboreal animals that live in trees.


Ecological subgroups

The term "terrestrial" is typically applied to species that live primarily on or in the ground, in contrast to arboreal species, who live primarily in trees, even though the latter are actually a specialized subgroup of the terrestrial fauna. There are other less common terms that apply to specific subgroups 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, 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 are a polyphyletic group that 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, whereas few animals became fully terrestrial during the Cenozoic. If internal parasites are excluded, eleven phyla include free living species in terrestrial environments. These can be grouped as follows: Three phyla contain species that have adapted totally to dry terrestrial environments, and which have no aquatic phase in their life cycles: * Arthropods — fully terrestrial members: hexapods ( crown group being insects), arachnids, myriapods, and some land-evolved crustaceans such as woodlice, sandhoppers, coconut crab and terrestrial crabs; semi-terrestrial members include most species of crabs, crayfish, water fleas, copepods, and seed shrimp * Molluscs — mainly terrestrial 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 molluscs, terrestrial gastropod mollusks that have gastropod shell, shel ...
s and slugs; one species of coleoid cephalopod, the algae octopus, is also known to be routinely terrestrial * Chordates — specifically tetrapod vertebrates (especially amniotes); semiterrestrial members: non- caecilian amphibians ( frogs and salamanders) and amphibious fish Four phyla include species that depend on more or less moist habitats: * Annelids — mainly oligochaete clitellates such as earthworms, pot worms and some leeches, but require moist soil habitats, highly diverse and derived from their marine relatives * Onychophores (velvet worms) — the only solely terrestrial phylum, though require moist habitats and have restricted range * Platyhelminthes (flatworms) — specifically land planarians, require moist habitats and have restricted range * Nemerteans (ribbon worms) — specifically the 12 terrestrial species from the suborder Monostilifera, require moist habitats and have restricted range Species in four more phyla, as well as 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> * Gastrotrichs (hairy-backs) — live in transient terrestrial water and go dormant during desiccation * Rotifers (wheel animals) — live in transient terrestrial water and go dormant during desiccation * Nematodes (roundworms) — mostly parasitic but some (e.g. '' Caenorhabditis'') are free-living detritivores, live in topsoil and go dormant during desiccation * Tardigrades (water bears) — live in transient terrestrial water and go dormant during desiccation


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.
mosquito Mosquitoes, the Culicidae, are a Family (biology), family of small Diptera, flies consisting of 3,600 species. The word ''mosquito'' (formed by ''Musca (fly), mosca'' and diminutive ''-ito'') is Spanish and Portuguese for ''little fly''. Mos ...
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 (; ; sometimes spelled nymphe) is a minor female nature deity in ancient Greek folklore. Distinct from other Greek goddesses, nymphs are generally regarded as personifications of nature; they are typically tied to a specific place, land ...
or larva. 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 a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
. 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 ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
. There is little reason to believe, however, that animals first began living permanently 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 The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
, they may have fully terrestrialized. There are other groups of arthropods, all from
malacostraca Malacostraca is the second largest of the six classes of pancrustaceans behind insects, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crab ...
n crustaceans, which independently became terrestrial at a later date: woodlice, sandhoppers, and terrestrial crabs. Additionally, the sister
panarthropoda Panarthropoda is a clade comprising the greatest diversity of animal groups. It contains the extant phyla Arthropoda (Euarthropoda), Tardigrada (water bears) and Onychophora (velvet worms), although the precise relationships among these remained ...
n groups
Onychophora Onychophora (from , , "claws"; and , , "to carry"), commonly known as velvet worms (for their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, ''Peripatus''), is a phylum of el ...
(velvet worms) are also terrestrial, while the
Eutardigrada Eutardigrada are a class of tardigrades (Tardigrada) without lateral appendages. Primarily freshwater bound, some species have secondarily gained the ability to live in marine environments ('' Halobiotus''). By cryptobiosis many species are able ...
are also adapted for land to some degree; both groups probably becoming so during the
Early Devonian The Early Devonian is the first of three Epoch (geology), epochs comprising the Devonian period, corresponding to the Lower Devonian Series (stratigraphy), series. It lasted from and began with the Lochkovian Stage , which was followed by the Pr ...
. Among arthropods, many microscopic crustacean groups like copepods and
amphipods Amphipoda () is an order (biology), 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 10,700 amphip ...
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 The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
, tetrapods (losing their gills) became fully terrestrialized, allowing their expansion into most terrestrial niches, though later on some will return to being aquatic and conquer the air also.


Terrestrial gastropods

Gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...
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 molluscs, terrestrial gastropod mollusks that have gastropod shell, shel ...
s and slugs. Terrestrial invasion of gastropod mollusks has occurred in Neritopsina, Cyclophoroidea,
Littorinoidea Littorinoidea are a Taxonomic rank, superfamily of both sea snails and land snails which have a gill and an operculum (gastropod), operculum, Terrestrial animal, terrestrial and marine (ocean), marine gastropod Mollusc, mollusks in the clade Litt ...
, Rissooidea,
Ellobioidea Ellobiidae, common name the hollow-shelled snails, is a family (biology), family of small air-breathing Land snail, land, Sea snail, marine, and brackish snails; pulmonate gastropod mollusks in the clade Eupulmonata.Bouchet, P. (2012). Ellobiida ...
, Onchidioidea, Veronicelloidea, Succineoidea, and Stylommatophora, 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. Gastropods are especially unique due to several fully terrestrial and epifaunal lineages that evolved during the Cenozoic. Some members of rissooidean families Truncatellidae, Assimineidae, 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 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 estuary ...
to pelagic areas. Terrestrial lineages likely evolved from such ancestors. The rissooidean gastropod family Pomatiopsidae is one of the few groups that have evolved fully terrestrial taxa during the late Cenozoic in the
Japanese Archipelago The is an archipelago of list of islands of Japan, 14,125 islands that form the country of Japan. It extends over from the Sea of Okhotsk in the northeast to the East China Sea, East China and Philippine Sea, Philippine seas in the southwest al ...
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 Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
, 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 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 The rotifers (, from Latin 'wheel' and 'bearing'), sometimes 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 Harris ...
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 Amphipoda () is an order (biology), 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 10,700 amphip ...
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 vertebrate or invertebrate, that lives in a body of water for all or most of its lifetime. Aquatic animals generally conduct gas exchange in water by extracting dissolved oxygen via specialised respirato ...
*
Aquatic ecosystem An aquatic ecosystem is an ecosystem found in and around a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms—aquatic life—that are dependent on each other and on their environ ...
*
Aquatic locomotion Aquatic locomotion or swimming is biologically propulsion , 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 arth ...
*
Aquatic mammal Aquatic mammals and semiaquatic mammals are a diverse group of mammals that dwell partly or entirely in body of water, bodies of water. They include the various ''marine mammals'' who dwell in oceans, as well as various freshwater species, such a ...
*
Aquatic plant Aquatic plants, also referred to as hydrophytes, are vascular plants and Non-vascular plant, non-vascular plants that have adapted to live in aquatic ecosystem, aquatic environments (marine ecosystem, saltwater or freshwater ecosystem, freshwater ...
*
Marine invertebrates Marine invertebrates are invertebrate animals that live in marine habitats, and make up most of the macroscopic life in the oceans. It is a polyphyletic blanket term that contains all marine animals except the marine vertebrates, including the ...
*
Marine mammal Marine mammals are mammals that rely on marine ecosystems for their existence. They include animals such as cetaceans, pinnipeds, sirenians, sea otters and polar bears. They are an informal group, unified only by their reliance on marine enviro ...
* Terrestrial *
Terrestrial ecosystem Terrestrial ecosystems are ecosystems that 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 presen ...
*
Terrestrial locomotion Terrestrial locomotion has evolution, evolved as animals adapted from ecoregion#Marine, aquatic to ecoregion#Terrestrial, terrestrial environments. Animal locomotion, Locomotion on land raises different problems than that in water, with reduced f ...
*
Terrestrial plant A terrestrial plant is a plant that grows on, in or from land. Other types of plants are aquatic plant, aquatic (living in or on water), semiaquatic (living at edge or seasonally in water), epiphyte, epiphytic (living on other plants), and litho ...
* *
Wetland indicator status Wetland indicator status denotes the probability of individual species of vascular plants occurring in freshwater, brackish and Salt marsh, saltwater wetlands in the United States. The wetland status of 7,000 plants is determined upon information ...
*
Zoology Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...


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 ...
, 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 New York, often called New York City (NYC), is the most populous city in the United States, located at the southern tip of New York State on one of the world's la ...
, 312 pp., . * Little C. (1983). ''The colonisation of land: Origins and adaptations of terrestrial animals''.
Cambridge University Press Cambridge University Press was the university press of the University of Cambridge. Granted a letters patent by King Henry VIII in 1534, it was the oldest university press in the world. Cambridge University Press merged with Cambridge Assessme ...
, 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