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A fossorial (from Latin fossor, "digger") animal is one that is adapted to digging and lives underground. Some examples are the badger, the naked mole-rat, or the mole salamander. Most bees and wasps are called "fossorial Hymenoptera", and many rodents are considered fossorial despite their physical adaptations to living underground being minimal.[clarification needed] The term fossorial has been used for rodent species that live in burrows most of the day but are surface-dwelling during other parts of the day. While species that live all or almost all of their life underground, the term subterranean has been applied. Some organisms are fossorial to aid in temperature regulation, while others use the underground habitat for protection from predators or for food storage. An animal is said to be sub-fossorial if it shows limited adaptations to a fossorial lifestyle.[1]

Contents

1 Prehistoric evidence 2 Physical adaptations 3 Physiological modifications 4 Geological and ecological implications 5 See also 6 References

Prehistoric evidence[edit] The physical adaption of fossoriality is widely accepted as being widespread among many prehistoric phyla and taxa, such as bacteria and early eukaryotes. Fossorial
Fossorial
animals appeared simultaneously with the colonization of land by arthropods in the late Ordovician (over 440 million years ago).[2] Other notable early burrowers include Eocaecilia
Eocaecilia
and possibly Dinilysia.[3] The oldest example of burrowing in synapsids,the lineage which lincludes modern mammals and their ancestors, is a Cynodont, Thrinaxodon liorhinus, found in the Karoo
Karoo
of South Africa, estimated to be 251 million years old. Evidence shows that this adaption occurred due to dramatic mass extinctions in the Permian
Permian
period.[1] Physical adaptations[edit]

European Mole - Note the strong and short forelimbs

There are six major external modifications, as described by H.W. Shimer in 1903,[4] that are shared in all mammalian burrowing species:

Fusiform, as an adaption to the dense subsurface environment below earth. Lesser developed or missing eyesight, considering subsurface darkness. Small or missing external ears, to reduce natural occurring friction during burrowing. Short and stout limbs, since swiftness or speed of movement is less important than the strength to dig. Broad and stout forelimbs (Manus), including long claws, designed to loosen the burrowing material for the hind feet to disperse in the back. Short or missing tail, which also has little to no locomotive or burrowing use to most fossorial Mammals.[4] This trait is also argued by Jorge Cubo, as he states that the skull is the main tool during excavation, but that the most active parts are the forelimbs for digging and that the hind-limbs are used for stability.[5]

Other important physical features include a subsurface adjusted skeleton: a triangular shaped skull, a prenasal ossicle, chisel shaped teeth, more or less fused and short lumbar vertebrae, well-developed sternum and strong forelimb and weaker hind limb bones.[4] Due to the lack of light, one the most important features of fossorial animals are the development of physical sensory traits that allow them to communicate and navigate in the dark subsurface environment. Considering that sound travels slower in air and faster through solid earth, the use of seismic (percussive) waves on a small scale is more advantageous in these environments. Several different uses are well documented. The cape mole rat Georychus capensis uses drumming behavior to send messages to its kin through conspecific signaling. The Namib desert golden mole Eremitalpa granti namibensis can detect termite colonies and similar prey underground due to the development of a hypertrophied malleus. This adaptation allows for better detection of low-frequency signals.[6] The most likely explanation of the actual transmission of these seismic inputs, captured by the auditory system, is the use of bone conduction; whenever vibrations are applied to the skull, the signals travel through many routes to the inner ear.[7] Physiological modifications[edit] Many fossorial and semi-fossorial mammals that live in temperate zones with partially frozen grounds tend to hibernate. Hibernation occurs due to the seasonal lack of soft succulent herbage and other sources of nutrition; therefore leading mammals to hibernate.[4] A conclusion made by W.H. Shimer is that, in general, a species that chose, voluntarily or not, adaptions to the fossorial lifestyle, likely originated as primitive and defenseless rodents, insectivores or edentates that failed to abundantly find food and protection from predators.[4] The life underground in these subsurface environments also have direct links to the animal’s metabolism and energetics. The weight of the individual specimen has direct implications here. Animals weighing more than 80 grams have comparably lower basal rates and animals weighing lower than 60 grams have comparably high basal rates, considering species that spend only part of their time burrowing. The average fossorial animal has a basal rate between 60% and 90%. Further observations conclude that larger burrowing animals, such as hedgehogs or armadillos, have lower thermal conductances than smaller animals, most likely to reduce heat storage in their burrows.[8] Many fossorial and semi-fossorial mammals that live in temperate zones with partially frozen grounds tend to hibernate. Hibernation occurs due to the seasonal lack of soft succulent herbage and other sources of nutrition; therefore leading mammals to hibernate. Geological and ecological implications[edit] One important impact on the environment caused by fossorial animals is bioturbation, defined by Marshall Wilkinson as the alteration of fundamental properties of the soil, including surface geomorphic processes.[9] It is measured that small fossorials, such as ants, termites, and earthworms displace a massive amount of soil. The total global rates displaced by these animals are equivalent to the total global rates of tectonic uplift.[9] The presence of burrowing animals has also a direct impact on the soils composition, structure, and growing vegetation. The impact these animals have can range from feeding, harvesting, caching and soil disturbances, but can differ considering the large diversity of fossorial species - especially herbivorous species. The net effect is usually composed of an alteration of the composition of plant species and increased plant diversity, which can cause issues with standing crops, as the homogeneity of the crops are affected.[10] Burrowing also impacts the nitrogen cycle in the affected soil. Mounds and bare soils, that contain burrowing animals, have considerably higher amounts of NH4 and NO3 as well as the nitrification potential and the microbial NO3 consumption than in vegetated soils. The primary mechanism for this occurrence is caused by the removal of the covering grassland.[11] See also[edit]

Arboreal Burrow Cursorial Fossa

References[edit]

^ a b Damiani, R, 2003, Earliest evidence of cynodont burrowing, The Royal Society Publishing, Volume 270, Issue 1525 ^ http://science.sciencemag.org/content/235/4784/61 ^ http://advances.sciencemag.org/content/1/10/e1500743 ^ a b c d e Shimer H.W., 1903, Adaptations to aquatic. arboreal, fossorial, and cursorial habits in mammals.III. Fossorial
Fossorial
Adaptations, The American Naturalist, Vol.XXXVII, No. 444 - December 1903 ^ Cubo, J, 2005, A heterochronic interpretation of the origin of digging adaptions in the northern water vole, Arvicola terrestris (Rodentia: Arvicolidae), Biological Journal of Linnean Society, Volume 87, pp. 381–391 ^ Narins, P.M, 1997, Use of seismic signals by fossorial south African mammals: a neurological goldmine, Brain research bulletin, Vol. 44, Issue 5, pp. 641–646 ^ Mason, M.J., 2001, Middle ear structures in fossorial mammals: a comparison with non-fossorial species, Journal of Zoology, Vol. 255, Issue 4, pp. 467–486 ^ McNab, B, 1979, The Influence of body size on the Energetics
Energetics
and Distribution of Fossorial
Fossorial
and Burrowing Mammals, Ecology, Volume 60, pages 1010-1021 ^ a b Wilkinson, M.T, Richards, P.J., Humphreys, G.S., 2009, Breaking ground: Pedological, geological, and ecological implications of soil bioturbation, Earth Science Reviews, Vol. 97, Issues 1-4, pp. 257–272 ^ Huntly, N, Reichman, O.J., 1994, Effects of Subterranean Mammalian Herbivores on Vegetation, Journal of Mammalogy, Volume 75, pp. 852–859 ^ Canals, H, 2003, How Disturbance by Fossorial
Fossorial
Mammals Alters N Cycling in a California Annual Grassland. Ecology, Volume 84, pp. 875–881

" Fossorial
Fossorial
- Definition of Fossorial". Amateur Entomologists' Society. Retrieved 1 September 2012.  " Fossorial
Fossorial
Legs". University of Sydney. Retrieved 1 September