Fossorial
   HOME

TheInfoList



OR:

A fossorial animal () is one that is adapted to digging and which lives primarily (but not solely) underground. Examples of fossorial vertebrates are moles,
badger Badgers are medium-sized short-legged omnivores in the superfamily Musteloidea. Badgers are a polyphyletic rather than a natural taxonomic grouping, being united by their squat bodies and adaptions for fossorial activity rather than by the ...
s,
naked mole-rat The naked mole-rat (''Heterocephalus glaber''), also known as the sand puppy, is a burrowing rodent native to the Horn of Africa and parts of Kenya, notably in Somali regions. It is closely related to the blesmols and is the only species in th ...
s,
meerkat The meerkat (''Suricata suricatta'') or suricate is a small mongoose found in southern Africa. It is characterised by a broad head, large eyes, a pointed snout, long legs, a thin tapering tail, and a brindled coat pattern. The head-and-body ...
s,
armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...
s,
wombat Wombats are short-legged, muscular quadrupedal marsupials of the family Vombatidae that are native to Australia. Living species are about in length with small, stubby tails and weigh between . They are adaptable and habitat tolerant, and are ...
s, and
mole salamander The mole salamanders (genus ''Ambystoma'') are a group of Salamandroidea, advanced salamanders endemic to North America. The group has become famous due to the study of the axolotl (''A. mexicanum'') in research on neoteny, paedomorphosis, and t ...
s. Among invertebrates, many
mollusc 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 ...
s (e.g.,
clam Clam is a common name for several kinds of bivalve mollusc. The word is often applied only to those that are deemed edible and live as infauna, spending most of their lives halfway buried in the sand of the sea floor or riverbeds. Clams h ...
s),
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s (e.g.,
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
s,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s, bees), and
arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s (e.g.
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
s) are fossorial.


Prehistoric evidence

The physical adaptation of fossoriality is widely accepted as being widespread among many
prehistoric Prehistory, also called pre-literary history, is the period of human history between the first known use of stone tools by hominins  million years ago and the beginning of recorded history with the invention of writing systems. The use o ...
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 Phy ...
and
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
, such as
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
and early
eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
s. Furthermore, fossoriality has evolved independently multiple times, even within a single
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
. Fossorial animals appeared simultaneously with the colonization of land by
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s in 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 ...
period (over 440 million years ago). Other notable early burrowers include '' Eocaecilia'' and possibly ''
Dinilysia ''Dinilysia'' (meaning "terrible ilysia") is an extinct genus of snake from the Late Cretaceous (Coniacian) of South America. ''Dinilysia'' was a relatively large ambush predator, measuring approximately long. The skull morphology of ''Dinilysia ...
''. The oldest example of burrowing in
synapsids Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
, the lineage which includes modern mammals and their ancestors, is a
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
, ''
Thrinaxodon liorhinus ''Thrinaxodon'' is an extinct genus (biology), genus of cynodonts which lived in what are now South Africa and Antarctica during the Late Permian - Early Triassic. ''Thrinaxodon'' lived just before, during, and right after the Permian–Triassic ...
'', found in the
Karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe Khoemana (also known as !Orakobab or Korana) word is a semidesert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its extent is ...
of
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
, estimated to be 251 million years old. Evidence shows that this adaptation occurred due to dramatic mass extinctions in the
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
period.


Physical adaptations in vertebrates

There are six major external modifications, as described by H. W. Shimer in 1903, that are shared in all mammalian burrowing species: * Fusiform, a spindle-shaped body tapering at both ends, adapted for the dense subsurface environment. * Lesser developed or missing eyesight, considering subsurface darkness. * Small or missing external ears, to reduce naturally occurring
friction Friction is the force resisting the relative motion of solid surfaces, fluid layers, and material elements sliding against each other. Types of friction include dry, fluid, lubricated, skin, and internal -- an incomplete list. The study of t ...
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. This trait is disputed by Jorge Cubo, who 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. * Short or missing tail, which has little to no locomotor activity or burrowing use to most fossorial mammals.Shimer H.W., 1903, Adaptations to aquatic. Arboreal, fossorial, and cursorial habits in mammals.III. Fossorial Adaptations, The American Naturalist, Vol.XXXVII, No. 444 – December 1903 Other important physical features include a subsurface adjusted skeleton: a triangularly shaped skull, a prenasal
ossicle The ossicles (also called auditory ossicles) are three irregular bones in the middle ear of humans and other mammals, and are among the smallest bones in the human body. Although the term "ossicle" literally means "tiny bone" (from Latin ''ossicu ...
, chisel-shaped teeth, effectively fused and short
lumbar In tetrapod anatomy, lumbar is an adjective that means of or pertaining to the abdominal segment of the torso, between the diaphragm (anatomy), diaphragm and the sacrum. Naming and location The lumbar region is sometimes referred to as the lowe ...
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e, well-developed
sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
, strong forelimb and weaker hind limb bones. Due to the lack of light, one of 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 the air and faster through solid earth, the use of
seismic Seismology (; from Ancient Greek σεισμός (''seismós'') meaning "earthquake" and -λογία (''-logía'') meaning "study of") is the scientific study of earthquakes (or generally, quakes) and the generation and propagation of elastic ...
(
percussive A percussion instrument is a musical instrument that is sounded by being struck or scraped by a beater including attached or enclosed beaters or rattles struck, scraped or rubbed by hand or struck against another similar instrument. Ex ...
) 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 Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...
signaling. The Namib Desert golden mole ( ''Eremitalpa granti namibensis'') can detect
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
colonies and similar prey underground due to the development of a hypertrophied
malleus The ''malleus'', or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibra ...
. This adaptation allows for better detection of low-frequency signals. 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. For animals that burrow by compressing soil, the work required increases exponentially with body diameter. In
amphisbaenia Amphisbaenia (called amphisbaenians or worm lizards) is a group of typically legless lizards, comprising over 200 extant species. Amphisbaenians are characterized by their long bodies, the reduction or loss of the limbs, and rudimentary eyes. A ...
ns, an ancient group of burrowing lizard-like squamates, specializations include the pennation of the
longissimus dorsi The longissimus () is the muscle lateral to the semispinalis muscles. It is the longest subdivision of the erector spinae muscles that extends forward into the transverse processes of the posterior cervical vertebrae. Structure Longissimus th ...
, the main muscle associated with burrowing, to increase muscle cross-sectional area. Constrained to small body diameters by the soil, amphisbaenians can increase muscle mass by increasing body length, not body diameter. In most amphisbaenians, limbs were lost as part of fossorial lifestyle. However the mole lizard '' Bipes'', unlike other amphisbaenians, retains robust digging forelimbs comparable to those of moles and
mole cricket Mole (or Molé) may refer to: Animals * Mole (animal) or "true mole" * Golden mole, southern African mammals * Marsupial mole, Australian mammals Other common meanings * Nevus, a growth on human skin ** Melanocytic nevus, a specific type of ...
s.


Physiological modifications

Many fossorial and sub-fossorial mammals that live in temperate zones with partially frozen grounds tend to hibernate due to the seasonal lack of soft, succulent herbage and other sources of nutrition. W. H. Shimer concluded that, in general, species that adopted fossorial lifestyles likely did so because they failed, aboveground, to find food and protection from predators. Additionally, some, such as E. Nevo, propose that fossorial lifestyles could have occurred because aboveground climates were harsh. Shifts towards an underground lifestyle also entail changes in
metabolism Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the co ...
and energetics, often in a weight-dependent manner. Sub-fossorial species weighing more than have comparably lower
basal rate Basal metabolic rate (BMR) is the rate of energy expenditure per unit time by endothermic animals at rest.. In other words it is the energy required by body organs to perform normal It is reported in energy units per unit time ranging from watt ( ...
s than those weighing lower than . The average fossorial animal has a basal rate between 60% and 90%. Further observations conclude that larger burrowing animals, such as
hedgehog A hedgehog is a spiny mammal of the subfamily Erinaceinae, in the eulipotyphlan family Erinaceidae. There are 17 species of hedgehog in five genera found throughout parts of Europe, Asia, and Africa, and in New Zealand by introduction. The ...
s or
armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...
s, have lower thermal conductance than smaller animals, most likely to reduce heat storage in their burrows.


Geological and ecological implications

One important impact on the environment caused by fossorial animals is
bioturbation Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a ...
, defined by Marshall Wilkinson as the alteration of fundamental properties of the soil, including surface geomorphic processes.Wilkinson, M.T, Richards, P.J., Humphreys, G.S., 2009, Breaking ground: Pedological, geological, and ecological implications of soil bioturbation, Earth-Science Reviewss, Vol. 97, Issues 1-4, pp. 257–272 It is measured that small fossorials, such as
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s,
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s, and
earthworm An earthworm is a soil-dwelling terrestrial invertebrate that belongs to the phylum Annelida. The term is the common name for the largest members of the class (or subclass, depending on the author) Oligochaeta. In classical systems, they we ...
s displace a massive amount of soil. The total global rates displaced by these animals are equivalent to the total global rates of
tectonic uplift Tectonic uplift is the orogeny, geologic uplift of Earth#Surface, Earth's surface that is attributed to plate tectonics. While Isostasy, isostatic response is important, an increase in the mean elevation of a region can only occur in response to ...
. The presence of burrowing animals also has a direct impact on the soil's 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 A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
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 Homogeneity and heterogeneity are concepts relating to the Uniformity (chemistry), uniformity of a Chemical substance, substance, process or image. A homogeneous feature is uniform in composition or character (i.e., color, shape, size, weight, ...
of the crops is affected. Burrowing also impacts the
nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmosphere, atmospheric, terrestrial ecosystem, terrestrial, and marine ecosystems. The conversion of nitrogen can ...
in the affected soil. Mounds and bare soils that contain burrowing animals have considerably higher amounts of and as well as greater
nitrification ''Nitrification'' is the biological oxidation of ammonia to nitrate via the intermediary nitrite. Nitrification is an important step in the nitrogen cycle in soil. The process of complete nitrification may occur through separate organisms or ent ...
potential and microbial consumption than in vegetated soils. The primary mechanism for this occurrence is caused by the removal of the covering grassland. Burrowing snakes may be more vulnerable to changing environments than non-burrowing snakes, although this may not be the case for other fossorial groups such as lizards. This may form an evolutionary dead end for snakes.


See also

*
Arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
*
Burrow file:Chipmunk-burrow (exits).jpg, An eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of Animal lo ...
*
Cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
* Fossa


References

* * {{Cite web , title = Fossorial Legs , url = http://bugs.bio.usyd.edu.au/learning/resources/Entomology/externalMorphology/imagePages/legs_fossorial.html , publisher = University of Sydney , access-date = 1 September 2012 , archive-date = 15 September 2012 , archive-url = https://web.archive.org/web/20120915031224/http://bugs.bio.usyd.edu.au/learning/resources/Entomology/externalMorphology/imagePages/legs_fossorial.html , url-status = dead Habitats Animal physiology Animal locomotion