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((Short description|Animals living on land))
((more citations needed|date=February 2014))
[[File:Un okapi à la réserve de faunes à okapis, à Epulu, 2005 (4389219071).jpg|thumb|The [[okapi]] is a species of terrestrial vertebrate.]]
''' Terrestrial animals''' are [[animal]]s that live predominantly or entirely on [[land]] (e.g. [[cat]]s, [[chicken]]s, [[ant]]s, [[spider]]s), as compared with [[aquatic animal]]s, which live predominantly or entirely in the water (e.g. [[fish]], [[lobster]]s, [[octopus]]es), and [[amphibian]]s, which rely on aquatic and terrestrial [[habitat]]s (e.g. [[frog]]s and [[newt]]s). Some groups of [[insects]] are [[wikt:terrestrial|terrestrial]], such as [[ant]]s, [[butterflies]], [[earwigs]], [[cockroaches]], [[grasshopper]]s and many others, while other groups are partially aquatic, such as [[mosquitoes]] and [[dragonflies]], which pass their [[larva]]l stages in water.

Alternatively, terrestrial is used to describe animals that live on the ground, as opposed to [[arboreal]] animals that live in trees.

==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:
*'''[[wiktionary:saxicolous|Saxicolous]]''' creatures are rock dwelling. "Saxicolous" is derived from the Latin word ''saxum'', meaning a rock. 
*'''[[wiktionary:arenicolous|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]].<ref>Shear WA: The early development of terrestrial ecosystems. Nature 1991, 351:283-289.</ref><ref>Vermeij GJ, Dudley R, ''Why are there so few evolutionary transitions between aquatic and terrestrial ecosystems?'' Biol J Linn Soc, 2000, 70:541-554.</ref><ref name="Garwood">((cite journal |last1=Garwood |first1=Russell J. |last2=Edgecombe |first2=Gregory D. |date=September 2011 |title=Early Terrestrial Animals, Evolution, and Uncertainty |journal=Evolution: Education and Outreach |location=New York |publisher=[[Springer Science+Business Media]] |volume=4 |issue=3 |pages=489–501 |doi=10.1007/s12052-011-0357-y |doi-access=free ))</ref> Terrestrial lineages evolved in several<!--refers to "seven" phyla, but see below--> animal [[phylum|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 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.<ref name="Garwood" /> 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 [[parasite]]s are excluded, free living species in terrestrial environments are represented by the following eleven phyla:

*[[Gastrotrich]]s (hairy-backs) live in transient terrestrial water and go dormant during desiccation
*[[Rotifer]]s (wheel animals) live in transient terrestrial water and go dormant during desiccation
*[[Nematoda|Nematodes]] (roundworms) by going dormant during desiccation
*[[Eutardigrada|Eutardigrades]] (water bears) live in transient terrestrial water and go dormant during desiccation
*[[Flatworm]]s ([[land planarian]]s) require moist habitats and have restricted range
*[[Nemertea]]ns (ribbon worms in [[Monostilifera]]) require moist habitats and have restricted range
*[[Onychophora]] (velvet worms) require moist habitats and have restricted range, the only solely terrestrial phylum
*[[Annelid]]s ([[clitellates]]) require moist habitats, highly diverse and derived from their [[Polychaete|marine relatives]]
*[[Arthropod]]s (fully terrestrial members: [[Hexapoda|hexapods]], [[arachnids]], [[myriapods]], [[woodlice]], [[sandhopper]]s, and [[terrestrial crabs]]; semi-terrestrial members include [[Cladocera|water fleas]], [[copepod]]s, and [[Ostracod|seed shrimp]])
*[[Mollusks]] ([[gastropods]]: [[land snail]]s and [[slug]]s)
*[[Chordate]]s ([[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.<ref name=":0">[http://nzetc.victoria.ac.nz/tm/scholarly/tei-Bio11Tuat02-t1-body-d1.html 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===
[[File:Pygoscelis papua -Nagasaki Penguin Aquarium -swimming underwater-8a.jpg|thumb|Animals do not fall neatly into terrestrial or aquatic classification but lie along a continuum: e.g., penguins spend much of their time under water.]]
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]], [[pinniped|seals]], and [[walruses]] sleep on land and feed in the ocean, yet they are all considered terrestrial. Many insects, e.g. [[mosquito]]s, and all [[terrestrial crab]]s, 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 (biology)|nymph]] 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]]. Many hundreds of [[gastropod]] genera and species live in intermediate situations, such as for example, [[Truncatella (gastropod)|''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]]. There is little reason to believe, however, that animals first began living reliably((clarify|date=January 2022)) 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.<ref>MacNaughton, R. B et al. First steps on land: Arthropod trackways in Cambrian-Ordovician eolian sandstone, southeastern Ontario, Canada. Geology, 30, 391 - 394, (2002).</ref>
Three groups of arthropods had independently adapted to land by the end of the Cambrian: [[myriapods]], [[hexapoda|hexapods]] and [[arachnids]].<ref>Lozano-Fernandez, J., Carton, R., Tanner, A. R., Puttick, M. N., Blaxter, M., Vinther, J., Olesen, J., Giribet, G., Edgecombe, G. D., & Pisani, D. (2016). A molecular palaeobiological exploration of arthropod terrestrialization. Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 371(1699), 20150133. https://doi.org/10.1098/rstb.2015.0133</ref> By the late [[Ordovician]], they may have fully terrestrialized. There are other groups of arthropods, all from [[malacostraca]]n crustaceans, which independently became terrestrial at a later date: [[woodlice]], [[talitridae|sandhoppers]], and [[terrestrial crabs]]. Additionally, the sister [[panarthropoda]]n 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]].<ref>Rota-Stabelli, Omar; Daley, Allison C.; Pisani, Davide. "Molecular Timetrees Reveal a Cambrian Colonization of Land and a New Scenario for Ecdysozoan Evolution" Current Biology, Volume 23, Issue 5 (31 January 2013)| https://doi.org/10.1016/j.cub.2013.01.026</ref> 
Among arthropods, many microscopic crustacean groups like [[copepod]]s and [[Amphipoda|amphipods]] and [[Ostracod|seed shrimp]] can go dormant when dry and live in transient bodies of water.((Citation needed|date=September 2022))

==Vertebrate terrestrialization==
((main|Vertebrate land invasion))
By approximately 375 million years ago<ref name="Garwood" /> 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),<ref>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.</ref> 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 to being aquatic and conquer the air also.

== Terrestrial gastropods ==
((main|land snail|semi-slug|slug))
[[Gastropod]] mollusks are one of the most successful animals that have diversified in the fully terrestrial habitat.<ref name="Kameda 2011"/> They have evolved terrestrial taxa in more than nine lineages.<ref name="Kameda 2011"/> They are commonly referred to as [[land snail]]s and [[slug]]s.

Terrestrial invasion of gastropod mollusks has occurred in [[Neritopsina]], [[Cyclophoroidea]], [[Littorinoidea]], [[Rissooidea]], [[Ellobioidea]], [[Onchidioidea]], [[Veronicelloidea]]<!--listed as Rathouisioidea that consist of the same two families-->, [[Succineoidea]], and [[Stylommatophora]], and in particular, each of Neritopsina, Rissooidea and Ellobioidea has likely achieved land invasion more than once.<ref name="Kameda 2011"/>

Most terrestrialization events have occurred during the [[Paleozoic]] or [[Mesozoic]].<ref name="Kameda 2011"/> Gastropods are especially unique due to several fully terrestrial and epifaunal lineages that evolved during the [[Cenozoic]].<ref name="Kameda 2011"/> Some members of rissooidean families [[Truncatellidae]], [[Assimineidae]], and [[Pomatiopsidae]] are considered to have colonized to land during the Cenozoic.<ref name="Kameda 2011"/> Most truncatellid and assimineid snails amphibiously live in [[intertidal]] and [[supratidal]] zones from [[brackish water]] to pelagic areas.<ref name="Kameda 2011"/> Terrestrial lineages likely evolved from such ancestors.<ref name="Kameda 2011"/> The [[rissooidea]]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.<ref name="Kameda 2011"/> 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]].<ref name="Kameda 2011"/>

About one-third of gastropod species are terrestrial.<ref name="Raz 2012"/> In terrestrial habitats they are subjected to daily and seasonal variation in temperature and water availability.<ref name="Raz 2012"/> Their success in colonizing different habitats is due to physiological, behavioral, and morphological adaptations to water availability, as well as ionic and thermal balance.<ref name="Raz 2012"/> They are adapted to most of the habitats on Earth.<ref name="Raz 2012"/> The shell of a snail is constructed of [[calcium carbonate]], but even in acidic soils one can find various species of shell-less slugs.<ref name="Raz 2012"/> Land-snails, such as [[Xerocrassa seetzeni]] and [[Sphincterochila boissieri]], also live in deserts, where they must contend with heat and aridity.<ref name="Raz 2012"/> Terrestrial gastropods are primarily herbivores and only a few groups are carnivorous.<ref name="Siriboon 2013"/> Carnivorous gastropods usually feed on other gastropod species or on weak individuals of the same species; some feed on insect larvae or earthworms.<ref name="Siriboon 2013"/>

== 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: [[Geoplanidae|land planarians]], [[Monostilifera|land ribbon worms]], [[Nematode|roundworms]] (nematodes), and [[Clitellata|land annelids]] (clitellates) who are very primitive and [[cutaneous respiration|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 [[Polychaeta|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 [[Amphipoda|amphipods]] and [[Ostracod|seed shrimps]] are known to go dormant when dry and live in transient bodies of water too.<ref name=":0" />

== See also ==
((Portal|Wetlands))
((col div|colwidth=40em))
* [[Aquatic animal]]
* [[Aquatic ecosystem]]
* [[Aquatic locomotion]]
* [[Aquatic mammal]]
* [[Aquatic plant]]
* [[Marine invertebrates]]
* [[Marine mammal]]
* [[Terrestrial (disambiguation)|Terrestrial]] 
* [[Terrestrial ecosystem]]
* [[Terrestrial locomotion]]
* [[Terrestrial plant]]
* ((annotated link|Wetland))
* [[Wetland indicator status]]
* [[Zoology]]
((colend))

== Further reading ==
* [[Jennifer A. Clack|Clack J. A.]] (2002). ''Gaining ground: the origin and evolution of tetrapods''. [[Indiana University Press]], 369 pp., ((ISBN|978-0-253-34054-2)).
* Cloudsley-Thompson J. L. (1988). ''Evolution and adaptation of terrestrial arthropods''. [[Springer Science+Business Media|Springer]], 141 pp., ((ISBN|978-3-540-18188-0)).
* Dejours P. et al. (1987). ''Comparative physiology: life in water and on land''. Liviana Editrice, Italy, 556 pp., ((ISBN|978-0-387-96515-4)).
* Gordon M. S. & Olson E. C. (1995). ''Invasions of the land: the transitions of organisms from aquatic to terrestrial life''. [[Columbia University Press]], 312 pp., ((ISBN|978-0-231-06876-5)).
* Little C. (1983). ''The colonisation of land: Origins and adaptations of terrestrial animals''. [[Cambridge University Press]], Cambridge. 290 pp., ((ISBN|978-0-521-25218-8)).
* Little C. (1990). ''The terrestrial invasion. An ecophysiological approach to the origin of land animals''. Cambridge University Press, Cambridge. 304 pp. ((ISBN|978-0-521-33669-7)).
*((cite book|last=Zimmer|first=Carl|title=At the Water's Edge : Fish with Fingers, Whales with Legs, and How Life Came Ashore but Then Went Back to Sea|date=1999|publisher=Touchstone|location=New York|isbn=0684856239))

== References ==
This article incorporates CC-BY-2.0 text from the reference<ref name="Kameda 2011">Kameda Y. & Kato M. (2011). "Terrestrial invasion of pomatiopsid gastropods in the heavy-snow region of the Japanese Archipelago". ''[[BMC Evolutionary Biology]]'' '''11''': 118. ((doi|10.1186/1471-2148-11-118)).</ref> and CC-BY-2.5 text from the reference<ref name="Raz 2012">Raz 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. ((doi|10.1371/journal.pone.0041840)).</ref> and CC-BY-3.0 text from the reference<ref name="Siriboon 2013">((Cite journal | last1 = Siriboon | first1 = T. | last2 = Sutcharit | first2 = C. | last3 = Naggs | first3 = F. | last4 = Panha | first4 = S. | title = Three new species of the carnivorous snail genus ''Perrottetia'' Kobelt, 1905 from Thailand (Pulmonata, Streptaxidae) | doi = 10.3897/zookeys.287.4572 | journal = ZooKeys | pages = 41–57 | year = 2013 | issue = 287 | pmid = 23794847| pmc = 3677355| bibcode = 2013ZooK..287...41S | doi-access = free ))</ref>
((Reflist))

((DEFAULTSORT:Terrestrial Animal))
[[Category:Zoology]]
[[Category:Animals by adaptation|Terrestrial]]

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