|This article needs additional citations for verification. (February 2014)|
Terrestrial animals are animals that live predominantly or entirely on land (e.g., cats, ants, snails), as compared with aquatic animals, which live predominantly or entirely in the water (e.g., fish, lobsters, octopuses), or amphibians, which rely on a combination of aquatic and terrestrial habitats (e.g., frogs). The term terrestrial is also frequently used for species that live primarily on the ground, in contrast to arboreal species, which live primarily in trees.
Terrestrial invasion is one of the most important events in the history of life., Terrestrial lineages evolved in several animal phyla, among which vertebrates, arthropods, and mollusks are representatives of more successful groups of the epifaunal terrestrial life.
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 has evolved independently and successfully many times by various groups of animals. Most terrestrial lineages originated under a mild or tropical climate during the Paleozoic and Mesozoic, whereas few animals became fully terrestrial during the Cenozoic.
When excluding internal parasites, free living species in terrestrial environments are represented by the following ten phyla: Flatworms (Planarians), Nemertea (ribbon worms), Nematoda (roundworms), Rotifers, Tardigrada (water bears), Onychophora (velvet worms), Arthropods, mollusks (gastropods: land snails and slugs), Annelida and Chordata (tetrapods). Roundworms, tardigrades, and rotifers are microscopic animals that require a film of water to live in, and are not considered truly terrestrial. Flatworms, ribbon worms, velvet worms and annelids all depend on more or less moist habitats, as do the Arthropods centipedes and millipedes. The three remaining phyla, Arthropods, Gastropods and Chordata, all contain species that have adapted totally to dry terrestrial environments, and contain species that have no aquatic phase in their life cycles.
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.
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. 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.
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 snails and slugs.
Terrestrial invasion of gastropod mollusks has occurred in Neritopsina, Cyclophoroidea, Littorinoidea, Rissooidea, Ellobioidea, 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 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 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.
About one-third of gastropod species is 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. Interestingly, land-snails also live in deserts, where they must contend with heat and aridity.
- Shear WA: The early development of terrestrial ecosystems. Nature 1991, 351:283-289.
- Vermeij GJ, Dudley R, Why are there so few evolutionary transitions between aquatic and terrestrial ecosystems? Biol J Linn Soc, 2000, 70:541-554.
- 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.
- 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.
- Clark 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, 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.
- Zimmer, Carl (1999). At the Water's Edge : Fish with Fingers, Whales with Legs, and How Life Came Ashore but Then Went Back to Sea. New York: Touchstone. ISBN 0684856239.