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Mammals () are a group of vertebrate animals constituting the
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
Mammalia (), characterized by the presence of mammary glands which in
females Female ( symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction. A female has larger gametes than a male. Femal ...
produce
milk Milk is a white liquid food produced by the mammary glands of mammals. It is the primary source of nutrition for young mammals (including breastfed human infants) before they are able to digest solid food. Immune factors and immune-modulati ...
for feeding (nursing) their young, a
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, ...
(a region of the brain),
fur Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
or
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal an ...
, and three middle ear bones. These characteristics distinguish them from
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchoceph ...
s (including birds) from which they diverged in the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
, over 300 million years ago. Around 6,400 extant species of mammals have been described divided into 29
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
. The largest
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
, in terms of number of species, are the
rodents Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
,
bats Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
, and
Eulipotyphla Eulipotyphla (, which means "truly fat and blind") is an order of mammals suggested by molecular methods of phylogenetic reconstruction, which includes the laurasiatherian members of the now-invalid polyphyletic order Lipotyphla, but not th ...
(
hedgehog A hedgehog is a spiny mammal of the subfamily Erinaceinae, in the eulipotyphlan family Erinaceidae. There are seventeen species of hedgehog in five genera found throughout parts of Europe, Asia, and Africa, and in New Zealand by introductio ...
s,
mole Mole (or Molé) may refer to: Animals * Mole (animal) or "true mole", mammals in the family Talpidae, found in Eurasia and North America * Golden moles, southern African mammals in the family Chrysochloridae, similar to but unrelated to Talpida ...
s,
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to differ ...
s, and others). The next three are the Primates (including humans,
ape Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister ...
s,
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incom ...
s, and others), the
Artiodactyla The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
(
cetacean Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...
s and
even-toed ungulate The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poste ...
s), and the
Carnivora Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
(
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s,
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. ...
s, seals, and others). In terms of
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived ch ...
, which reflects evolutionary history, mammals are the only living members of the Synapsida (synapsids); this clade, together with
Sauropsida Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early sy ...
(reptiles and birds), constitutes the larger
Amniota Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
clade. The early synapsids were sphenacodonts, a group that included the famous ''
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodont ...
''. The synapsids split into several diverse groups of non-mammalian synapsids — traditionally and incorrectly referred to as mammal-like reptiles or by the term pelycosaurs, and now known as stem mammals or protomammals — before giving rise to
therapsids Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
during the beginning of the
Middle Permian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± ...
period. Mammals originated from
cynodont The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide var ...
s, an advanced group of therapsids, during the Late
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
-Early
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
. The modern mammalian orders arose in the
Paleogene The Paleogene ( ; also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period million years ago ( Mya) to the beginning o ...
and
Neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
periods of the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configur ...
era, after the extinction of non-avian dinosaurs, and have been the dominant terrestrial animal group from 66 million years ago to the present. The basic body type is
quadruped Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor ...
, and most mammals use their four extremities for
terrestrial locomotion Terrestrial locomotion has evolved as animals adapted from aquatic to terrestrial environments. Locomotion on land raises different problems than that in water, with reduced friction being replaced by the increased effects of gravity. As view ...
; but in some, the extremities are adapted for life at sea, in the air, in trees, underground, or on two legs. Mammals range in size from the bumblebee bat to the
blue whale The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can b ...
—possibly the largest animal to have ever lived. Maximum lifespan varies from two years for the shrew to 211 years for the
bowhead whale The bowhead whale (''Balaena mysticetus'') is a species of baleen whale belonging to the family Balaenidae and the only living representative of the genus '' Balaena''. They are the only baleen whale endemic to the Arctic and subarctic waters ...
. All modern mammals give birth to live young, except the five species of
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals ( Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their bra ...
s, which are egg-laying mammals. The most species-rich group of mammals, the
cohort Cohort or cohortes may refer to: * Cohort (educational group), a group of students working together through the same academic curriculum * Cohort (floating point), a set of different encodings of the same numerical value * Cohort (military unit ...
called
placentals Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
, have a
placenta The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate matern ...
, which enables the feeding of the fetus during
gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pr ...
. Most mammals are intelligent, with some possessing large brains,
self-awareness In philosophy of self, self-awareness is the experience of one's own personality or individuality. It is not to be confused with consciousness in the sense of qualia. While consciousness is being aware of one's environment and body and lifest ...
, and
tool use Tool use by animals is a phenomenon in which an animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, defence, communication, recreation or construction. Originally thought to be a skill possessed o ...
. Mammals can communicate and vocalize in several ways, including the production of
ultrasound Ultrasound is sound waves with frequencies higher than the upper audible limit of human hearing. Ultrasound is not different from "normal" (audible) sound in its physical properties, except that humans cannot hear it. This limit varies ...
, scent-marking,
alarm signal In animal communication, an alarm signal is an antipredator adaptation in the form of signals emitted by social animals in response to danger. Many primates and birds have elaborate alarm calls for warning conspecifics of approaching predato ...
s, singing, and echolocation. Mammals can organize themselves into fission-fusion societies,
harems Harem ( Persian: حرمسرا ''haramsarā'', ar, حَرِيمٌ ''ḥarīm'', "a sacred inviolable place; harem; female members of the family") refers to domestic spaces that are reserved for the women of the house in a Muslim family. A har ...
, and
hierarchies A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
—but can also be solitary and
territorial A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
. Most mammals are
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
, but some can be
monogamous Monogamy ( ) is a form of dyadic relationship in which an individual has only one partner during their lifetime. Alternately, only one partner at any one time ( serial monogamy) — as compared to the various forms of non-monogamy (e.g., po ...
or
polyandrous Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...
.
Domestication Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
of many types of mammals by humans played a major role in the
Neolithic Revolution The Neolithic Revolution, or the (First) Agricultural Revolution, was the wide-scale transition of many human cultures during the Neolithic period from a lifestyle of hunting and gathering to one of agriculture and settlement, making an in ...
, and resulted in
farming Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people ...
replacing hunting and gathering as the primary source of food for humans. This led to a major restructuring of human societies from nomadic to sedentary, with more co-operation among larger and larger groups, and ultimately the development of the first
civilization A civilization (or civilisation) is any complex society characterized by the development of a state, social stratification, urbanization, and symbolic systems of communication beyond natural spoken language (namely, a writing system). ...
s. Domesticated mammals provided, and continue to provide, power for transport and agriculture, as well as food (
meat Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chick ...
and dairy products),
fur Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
, and
leather Leather is a strong, flexible and durable material obtained from the tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffalo, pigs and ho ...
. Mammals are also hunted and raced for sport, and are used as
model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the working ...
s in science. Mammals have been depicted in
art Art is a diverse range of human activity, and resulting product, that involves creative or imaginative talent expressive of technical proficiency, beauty, emotional power, or conceptual ideas. There is no generally agreed definition of wh ...
since
Paleolithic The Paleolithic or Palaeolithic (), also called the Old Stone Age (from Greek: παλαιός '' palaios'', "old" and λίθος '' lithos'', "stone"), is a period in human prehistory that is distinguished by the original development of stone t ...
times, and appear in literature, film, mythology, and religion. Decline in numbers and
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
of many mammals is primarily driven by human
poaching Poaching has been defined as the illegal hunting or capturing of wild animals, usually associated with land use rights. Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set a ...
and
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, primarily
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
.


Classification

Mammal classification has been through several revisions since Carl Linnaeus initially defined the class, and at present, no classification system is universally accepted. McKenna & Bell (1997) and Wilson & Reeder (2005) provide useful recent compendiums.
Simpson Simpson most often refers to: * Simpson (name), a British surname *''The Simpsons'', an animated American sitcom **The Simpson family, central characters of the series ''The Simpsons'' Simpson may also refer to: Organizations Schools *Simps ...
(1945) provides
systematics Biological systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: cladograms, phylogenetic tr ...
of mammal origins and relationships that had been taught universally until the end of the 20th century. However, since 1945, a large amount of new and more detailed information has gradually been found: The paleontological record has been recalibrated, and the intervening years have seen much debate and progress concerning the theoretical underpinnings of systematization itself, partly through the new concept of
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived ch ...
. Though fieldwork and lab work progressively outdated Simpson's classification, it remains the closest thing to an official classification of mammals, despite its known issues. Most mammals, including the six most species-rich
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
, belong to the placental group. The three largest orders in numbers of species are Rodentia:
mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus'' ...
,
rat Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' ( pack rats), '' Bandicota'' (bandicoo ...
s,
porcupine Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethizo ...
s,
beaver Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers ...
s,
capybara The capybaraAlso called capivara (in Brazil), capiguara (in Bolivia), chigüire, chigüiro, or fercho (in Colombia and Venezuela), carpincho (in Argentina, Paraguay and Uruguay) and ronsoco (in Peru). or greater capybara (''Hydrochoerus hydro ...
s, and other gnawing mammals;
Chiroptera Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
: bats; and
Soricomorpha Soricomorpha (from Greek "shrew-form") is a formerly used taxon within the class of mammals. In the past it formed a significant group within the former order Insectivora. However, Insectivora was shown to be polyphyletic and various new orders ...
:
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to differ ...
s, moles, and
solenodon Solenodons (from el, τέλειος , 'channel' or 'pipe' and el, ὀδούς , 'tooth') are venomous, nocturnal, burrowing, insectivorous mammals belonging to the family Solenodontidae . The two living solenodon species are the Cuban sole ...
s. The next three biggest orders, depending on the
biological classification In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
scheme used, are the Primates:
ape Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister ...
s,
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incom ...
s, and
lemur Lemurs ( ) (from Latin ''lemures'' – ghosts or spirits) are wet-nosed primates of the superfamily Lemuroidea (), divided into 8 families and consisting of 15 genera and around 100 existing species. They are endemic to the island of Madaga ...
s; the
Cetartiodactyla The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
:
whale Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s and even-toed ungulates; and the
Carnivora Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
which includes
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s,
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. ...
s,
weasel Weasels are mammals of the genus ''Mustela'' of the family Mustelidae. The genus ''Mustela'' includes the least weasels, polecats, stoats, ferrets and European mink. Members of this genus are small, active predators, with long and slender bod ...
s,
bear Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the Nort ...
s, seals, and allies. According to ''
Mammal Species of the World ''Mammal Species of the World: A Taxonomic and Geographic Reference'' is a standard reference work in mammalogy giving descriptions and bibliographic data for the known species of mammal Mammals () are a group of vertebrate Verte ...
'', 5,416 species were identified in 2006. These were grouped into 1,229 
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenc ...
, 153 
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
and 29 orders. In 2008, the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of nat ...
(IUCN) completed a five-year Global Mammal Assessment for its IUCN Red List, which counted 5,488 species. According to research published in the ''
Journal of Mammalogy The ''Journal of Mammalogy'' is a bimonthly peer-reviewed scientific journal published by Oxford University Press on behalf of the American Society of Mammalogists. Both the society and the journal were established in 1919. The journal covers rese ...
'' in 2018, the number of recognized mammal species is 6,495, including 96 recently extinct.


Definitions

The word "
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
" is modern, from the scientific name ''Mammalia'' coined by Carl Linnaeus in 1758, derived from the Latin ''
mamma Mama(s) or Mamma or Momma may refer to: Roles * Mother, a female parent * Mama-san, in Japan and East Asia, a woman in a position of authority *Mamas, a name for female associates of the Hells Angels Places * Mama, Russia, an urban-type settlem ...
'' ("teat, pap"). In an influential 1988 paper, Timothy Rowe defined Mammalia
phylogenetically In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
as the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
of mammals, the clade consisting of the
most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. Th ...
of living
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals ( Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their bra ...
s (
echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and th ...
s and
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or monotyp ...
es) and
Theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...
n mammals (
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a p ...
s and
placental Placental mammals ( infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distingui ...
s) and all descendants of that ancestor. Since this ancestor lived in the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
period, Rowe's definition excludes all animals from the earlier
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, despite the fact that Triassic fossils in the
Haramiyida Haramiyida ("thief" from Arabic الحرامية (al ḥarāmiyah), "thief, bandit") is a possibly polyphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. Their teeth, which are by far the most common remain ...
have been referred to the Mammalia since the mid-19th century. If Mammalia is considered as the crown group, its origin can be roughly dated as the first known appearance of animals more closely related to some extant mammals than to others. '' Ambondro'' is more closely related to monotremes than to therian mammals while ''
Amphilestes ''Amphilestes'' is a genus of extinct eutriconodont mammal from the Middle Jurassic of the United Kingdom. It was one of the first Mesozoic mammals discovered and described. Discovery The first specimen of ''Amphilestes'' was discovered along ...
'' and ''
Amphitherium ''Amphitherium'' is an extinct genus of stem cladotherian mammal that lived during the Middle Jurassic of England. It was one of the first Mesozoic mammals ever described. A recent phylogenetic study found it to be the sister taxon of '' Palaeox ...
'' are more closely related to the therians; as fossils of all three genera are dated about in the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formation ...
, this is a reasonable estimate for the appearance of the crown group. T.S. Kemp has provided a more traditional definition: " Synapsids that possess a
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bo ...
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral c ...
jaw articulation and occlusion between upper and lower molars with a transverse component to the movement" or, equivalently in Kemp's view, the clade originating with the last common ancestor of ''
Sinoconodon ''Sinoconodon'' is an extinct genus of mammaliamorphs that appears in the fossil record of the Lufeng Formation of China in the Sinemurian stage of the Early Jurassic period, about 193 million years ago. While sharing many plesiomorphic traits w ...
'' and living mammals. The earliest known synapsid satisfying Kemp's definitions is '' Tikitherium'', dated , so the appearance of mammals in this broader sense can be given this
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch ...
date.


McKenna/Bell classification

In 1997, the mammals were comprehensively revised by Malcolm C. McKenna and Susan K. Bell, which has resulted in the McKenna/Bell classification. The authors worked together as
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fos ...
s at the
American Museum of Natural History The American Museum of Natural History (abbreviated as AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. In Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 26 inte ...
. McKenna inherited the project from Simpson and, with Bell, constructed a completely updated hierarchical system, covering living and extinct taxa, that reflects the historical genealogy of Mammalia. Their 1997 book, ''Classification of Mammals above the Species Level'', is a comprehensive work on the systematics, relationships and occurrences of all mammal taxa, living and extinct, down through the rank of genus, though molecular genetic data challenge several of the groupings. In the following list, extinct groups are labelled with a
dagger A dagger is a fighting knife with a very sharp point and usually two sharp edges, typically designed or capable of being used as a thrusting or stabbing weapon.State v. Martin, 633 S.W.2d 80 (Mo. 1982): This is the dictionary or popular-use de ...
(†). Class Mammalia * Subclass
Prototheria Prototheria (; from Greek πρώτος, ''prōtos'', first, + θήρ, ''thēr'', wild animal) is a subclass to which the orders Monotremata, Morganucodonta, Docodonta, Triconodonta and Multituberculata have been assigned, although the validi ...
: monotremes:
echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and th ...
s and the
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or monotyp ...
* Subclass
Theriiformes Theriiformes is a clade of mammals. The term was coined by Timothy B. Rowe in his doctoral dissertation, and is defined as the clade formed by the most recent common ancestor of multituberculates and theria Theria (; Greek: , wild beast) ...
: live-bearing mammals and their prehistoric relatives ** Infraclass †
Allotheria Allotheria (meaning "other beasts", from the Greek , '–other and , '–wild animal) is an extinct branch of successful Mesozoic mammals. The most important characteristic was the presence of lower molariform teeth equipped with two longitudi ...
: multituberculates ** Infraclass †
Eutriconodonta Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack ''et al.'' in 1973 as a replacement name for the ...
: eutriconodonts ** Infraclass
Holotheria Mammalia is a class of animal within the phylum Chordata. Mammal classification has been through several iterations since Carl Linnaeus initially defined the class. No classification system is universally accepted; McKenna & Bell (1997) and Wilson ...
: modern live-bearing mammals and their prehistoric relatives *** Superlegion †
Kuehneotheria Kuehneotheriidae is an extinct family of mammaliaforms traditionally placed within ' Symmetrodonta', though now generally considered more basal than true symmetrodonts. All members of Kuehneotheriidae which have been found so far are represented ...
*** Supercohort
Theria Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes. ...
: live-bearing mammals **** Cohort
Marsupialia Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a p ...
: marsupials ***** Magnorder
Australidelphia Australidelphia is the superorder that contains roughly three-quarters of all marsupials, including all those native to Australasia and a single species — the monito del monte — from South America. All other American marsupials are members o ...
: Australian marsupials and the
monito del monte The monito del monte or colocolo opossum, ''Dromiciops gliroides'', also called ''chumaihuén'' in Mapudungun, is a diminutive marsupial native only to southwestern South America (Argentina and Chile). It is the only extant species in the anci ...
***** Magnorder
Ameridelphia Ameridelphia is traditionally a superorder that includes all marsupials living in the Americas except for the Monito del monte (''Dromiciops''). It is now regarded as a paraphyletic group. Orders The orders within this group are listed below: ...
: New World marsupials. Now considered paraphyletic, with shrew opossums being closer to australidelphians. **** Cohort
Placentalia Placental mammals ( infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distingui ...
: placentals ***** Magnorder Xenarthra: xenarthrans ***** Magnorder
Epitheria Epitherians comprise all the placental mammals except the Xenarthra. They are primarily characterized by having a stirrup-shaped stapes in the middle ear, which allows for passage of a blood vessel. This is in contrast to the column-shaped sta ...
: epitheres ****** Superorder †
Leptictida Leptictida (''leptos iktis'' "small/slender weasel") is a possibly paraphyletic extinct order of eutherian mammals. Their classification is contentious: according to cladistic studies, they may be (distantly) related to Euarchontoglires (rodent ...
****** Superorder
Preptotheria Preptotheria is a superorder of placental mammals proposed by McKenna & Bell in their classification of mammals. Classification The Linnean taxonomy of Preptotheria according to the scheme of McKenna & Bell (1997): Cohort Placentalia Magnord ...
******* Grandorder
Anagalida Anagaloidea is an extinct order of placental mammals that first appeared during the Paleocene epoch. Taxonomy According to the traditional (morphological) view, Anagaloidea is part of the superorder Anagalida, along with the elephant shrews, rod ...
:
lagomorphs The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (hares and rabbits) and the Ochotonidae (pikas). The name of the order is derived from the Ancient Greek ''lagos'' (λαγ ...
, rodents and
elephant shrew Elephant shrews, also called jumping shrews or sengis, are small insectivorous mammals native to Africa, belonging to the family Macroscelididae, in the order Macroscelidea. Their traditional common English name "elephant shrew" comes from a pe ...
s ******* Grandorder
Ferae Ferae ( , , "wild beasts") is a mirorder of placental mammalsMalcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'' in Columbia University Press, New York (1997), 631 Seiten. that groups together clades Pan- ...
:
carnivora Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
ns,
pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smu ...
s, †
creodont Creodonta ("meat teeth") is a former order of extinct carnivorous placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ...
s and relatives ******* Grandorder
Lipotyphla Lipotyphla is a formerly used order of mammals, including the members of the order Eulipotyphla (i.e. the solenodons, family Solenodontidae; hedgehogs and gymnures, family Erinaceidae; desmans, moles, and shrew-like moles, family Talpidae; and tru ...
:
insectivora The order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order Eulipotyphla, ...
ns ******* Grandorder
Archonta The Archonta are a now-abandoned group of mammals, considered a superorder in some classifications, which consists of these orders: *Primates *Plesiadapiformes (extinct primate-like archontans) * Scandentia (treeshrews) * Dermoptera (colugos) Whi ...
:
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most b ...
s, primates,
colugo Colugos () are arboreal gliding mammals that are native to Southeast Asia. Their closest evolutionary relatives are primates. There are just two living species of colugos: the Sunda flying lemur (''Galeopterus variegatus'') and the Philippine fl ...
s and treeshrews (now considered paraphyletic, with bats being closer to other groups) ******* Grandorder Ungulata: ungulates ******** Order
Tubulidentata Orycteropodidae is a family of afrotherian mammals. Although there are many fossil species, the only species surviving today is the aardvark, ''Orycteropus afer''. Orycteropodidae is recognized as the only family within the order Tubulidentata ...
''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertaint ...
'':
aardvark The aardvark ( ; ''Orycteropus afer'') is a medium-sized, burrowing, nocturnal mammal native to Africa. It is the only living species of the order Tubulidentata, although other prehistoric species and genera of Tubulidentata are known. Unlike ...
******** Mirorder
Eparctocyona Eparctocyona is a clade of placental mammals comprising the artiodactyls (even-toed ungulates), cetacea (whales and related), and the extinct condylarths Condylarthra is an informal group – previously considered an order – of extinct plac ...
: †
condylarth Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wa ...
s,
whale Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s and
artiodactyls The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
(even-toed ungulates) ******** Mirorder †
Meridiungulata South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals of controversial affinities that were indigenous to South America prior to the Great American Biotic Interchange. They comprise five major groups c ...
: South American ungulates ******** Mirorder
Altungulata Altungulata or Pantomesaxonia (''sensu'' and later authors) is an invalid clade ( mirorder) of ungulate mammals comprising the perissodactyls, hyracoids, and tethytheres ( sirenians, proboscideans, and related extinct taxa). The name "Pantom ...
:
perissodactyls Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) ...
(odd-toed ungulates),
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantid ...
s,
manatee Manatees (family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing three of the four living specie ...
s and
hyrax Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simil ...
es


Molecular classification of placentals

As of the early 21st century, molecular studies based on DNA analysis have suggested new relationships among mammal families. Most of these findings have been independently validated by
retrotransposon Retrotransposons (also called Class I transposable elements or transposons via RNA intermediates) are a type of genetic component that copy and paste themselves into different genomic locations ( transposon) by converting RNA back into DNA throug ...
presence/absence data. Classification systems based on molecular studies reveal three major groups or lineages of placental mammals—
Afrotheria Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a clade of mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephant shrews (also kn ...
, Xenarthra and
Boreoeutheria Boreoeutheria (, "northern true beasts") is a magnorder of placental mammals that groups together superorders Euarchontoglires and Laurasiatheria. With a few exceptionsExceptional clades whose males lack the usual boreoeutherian scrotum are mo ...
—which diverged in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period o ...
. The relationships between these three lineages is contentious, and all three possible hypotheses have been proposed with respect to which group is
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is ne ...
. These hypotheses are
Atlantogenata Atlantogenata is a proposed clade of mammals containing the cohorts or superorders Xenarthra and Afrotheria. These groups originated and radiated in the South American and African continents, respectively, presumably in the Cretaceous. Together ...
(basal Boreoeutheria),
Epitheria Epitherians comprise all the placental mammals except the Xenarthra. They are primarily characterized by having a stirrup-shaped stapes in the middle ear, which allows for passage of a blood vessel. This is in contrast to the column-shaped sta ...
(basal Xenarthra) and
Exafroplacentalia Exafroplacentalia or Notolegia is a clade of placental mammals proposed in 2001 on the basis of molecular research. Exafroplacentalia places Xenarthra as a sister group to the Boreoeutheria (comprising Laurasiatheria and Euarchontoglires), thus ...
(basal Afrotheria). Boreoeutheria in turn contains two major lineages—
Euarchontoglires Euarchontoglires (synonymous with Supraprimates) is a clade and a superorder of mammals, the living members of which belong to one of the five following groups: rodents, lagomorphs, treeshrews, colugos, and primates. Evolutionary affinities ...
and
Laurasiatheria Laurasiatheria ("laurasian beasts") is a superorder of placental mammals that groups together true insectivores ( eulipotyphlans), bats ( chiropterans), carnivorans, pangolins ( pholidotes), even-toed ungulates (artiodactyls), odd-toed ungulates ...
. Estimates for the divergence times between these three placental groups range from 105 to 120 million years ago, depending on the type of DNA used (such as
nuclear Nuclear may refer to: Physics Relating to the nucleus of the atom: * Nuclear engineering *Nuclear physics *Nuclear power *Nuclear reactor *Nuclear weapon *Nuclear medicine *Radiation therapy *Nuclear warfare Mathematics * Nuclear space * Nucle ...
or
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used t ...
) and varying interpretations of paleogeographic data. The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related t ...
above is based on Tarver ''et al.''. (2016)


Evolution


Origins

Synapsida, a clade that contains mammals and their extinct relatives, originated during the Pennsylvanian subperiod (~323 million to ~300 million years ago), when they split from the reptile lineage. Crown group mammals evolved from earlier
mammaliaforms Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent comm ...
during the
Early Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma&nbs ...
. The cladogram takes Mammalia to be the crown group.


Evolution from older amniotes

The first fully terrestrial vertebrates were
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are dist ...
s. Like their amphibious early
tetrapod Tetrapods (; ) are four- limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
predecessors, they had lungs and limbs. Amniotic eggs, however, have internal membranes that allow the developing
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm c ...
to breathe but keep water in. Hence, amniotes can lay eggs on dry land, while amphibians generally need to lay their eggs in water. The first amniotes apparently arose in the Pennsylvanian subperiod of the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
. They descended from earlier
reptiliomorph Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians (lissamphibians). It was defi ...
amphibious tetrapods, which lived on land that was already inhabited by insects and other invertebrates as well as ferns,
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta ('' sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and ...
es and other plants. Within a few million years, two important amniote lineages became distinct: the synapsids, which would later include the common ancestor of the mammals; and the
sauropsid Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early ...
s, which now include
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necke ...
s,
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s,
snake Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more joints ...
s,
crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest livin ...
s and dinosaurs (including birds). Synapsids have a single hole (
temporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets ...
) low on each side of the skull. Primitive synapsids included the largest and fiercest animals of the early
Permian The Permian ( ) is a geologic period and 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 251.9 Mya. It is the last period of the Paleozo ...
such as
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodont ...
. Nonmammalian synapsids were traditionally – and incorrectly – called "mammal-like reptiles" or
pelycosaurs Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an order, but this is now ...
; we now know they were neither reptiles nor part of reptile lineage.
Therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s, a group of synapsids, evolved in the
Middle Permian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± ...
, about 265 million years ago, and became the dominant land vertebrates. They differ from basal eupelycosaurs in several features of the skull and jaws, including: larger skulls and
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, w ...
s which are equal in size in therapsids, but not for eupelycosaurs. The therapsid lineage leading to mammals went through a series of stages, beginning with animals that were very similar to their early synapsid ancestors and ending with
probainognathia Probainognathia is one of the two major subgroups of the clade Eucynodontia, the other being Cynognathia. The earliest forms were carnivorous and insectivorous, though some groups eventually also evolved herbivorous diets. The earliest and m ...
n
cynodont The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide var ...
s, some of which could easily be mistaken for mammals. Those stages were characterized by: * The gradual development of a bony secondary
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
. * Abrupt acquisition of
endothermy An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inst ...
among
Mammaliamorpha Mammaliamorpha is a clade of cynodonts. It contains the clades Tritylodontidae and Mammaliaformes, as well as a few genera that do not belong to either of these groups. The family Tritheledontidae has also been placed in Mammaliamorpha by som ...
, thus prior to the origin of mammals by 30-50 millions of years '. * Progression towards an erect limb posture, which would increase the animals' stamina by avoiding
Carrier's constraint Carrier's constraint is the observation that air-breathing vertebrates which have two lungs and flex their bodies sideways during locomotion find it very difficult to move and breathe at the same time, because the sideways flexing expands one lun ...
. But this process was slow and erratic: for example, all herbivorous nonmammaliaform therapsids retained sprawling limbs (some late forms may have had semierect hind limbs); Permian carnivorous therapsids had sprawling forelimbs, and some late Permian ones also had semisprawling hindlimbs. In fact, modern monotremes still have semisprawling limbs. * The
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bo ...
gradually became the main bone of the lower jaw which, by the Triassic, progressed towards the fully mammalian jaw (the lower consisting only of the dentary) and middle ear (which is constructed by the bones that were previously used to construct the jaws of reptiles).


First mammals

The
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, a ...
about 252 million years ago, which was a prolonged event due to the accumulation of several extinction pulses, ended the dominance of carnivorous therapsids. In the early Triassic, most medium to large land carnivore niches were taken over by
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s which, over an extended period (35 million years), came to include the
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
, the
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
s and the dinosaurs; however, large cynodonts like ''
Trucidocynodon ''Trucidocynodon'' is an extinct genus of ecteniniid cynodonts from Upper Triassic of Brazil. It contains a single species, ''Trucidocynodon riograndensis''. Fossils of ''Trucidocynodon'' were discovered in Santa Maria Formation outcrops in ...
'' and traversodontids still occupied large sized carnivorous and herbivorous niches respectively. By the Jurassic, the dinosaurs had come to dominate the large terrestrial herbivore niches as well. The first mammals (in Kemp's sense) appeared in the Late Triassic epoch (about 225 million years ago), 40 million years after the first therapsids. They expanded out of their nocturnal
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
niche from the mid-Jurassic onwards; The Jurassic ''
Castorocauda ''Castorocauda'' is an extinct, semi-aquatic, superficially otter-like genus of docodont mammaliaforms with one species, ''C. lutrasimilis''. It is part of the Yanliao Biota, found in the Daohugou Beds of Inner Mongolia, China dating to the Mi ...
'', for example, was a close relative of true mammals that had adaptations for swimming, digging and catching fish. Most, if not all, are thought to have remained nocturnal (the nocturnal bottleneck), accounting for much of the typical mammalian traits. The majority of the mammal species that existed in the
Mesozoic Era The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
were multituberculates, eutriconodonts and spalacotheriids. The earliest known
metatherian Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as wel ...
is ''
Sinodelphys ''Sinodelphys'' is an extinct eutherian from the Early Cretaceous, estimated to be 125 million years old. It was discovered and described in 2003 in rocks of the Yixian Formation in Liaoning Province, China, by a team of scientists including Zhe ...
'', found in 125 million-year-old
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous ( chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pr ...
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments ( silt-sized particles) of other minerals, especi ...
in China's northeastern
Liaoning Province Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
. The fossil is nearly complete and includes tufts of fur and imprints of soft tissues. The oldest known fossil among the
Eutheria Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials. Eutherians are distinguished from noneutherians by various phenotypic tr ...
("true beasts") is the small shrewlike '' Juramaia sinensis'', or "Jurassic mother from China", dated to 160 million years ago in the late Jurassic. A later eutherian relative, ''
Eomaia ''Eomaia'' ("dawn mother") is a genus of extinct fossil mammals containing the single species ''Eomaia scansoria'', discovered in rocks that were found in the Yixian Formation, Liaoning Province, China, and dated to the Barremian Age of the Lowe ...
'', dated to 125 million years ago in the early Cretaceous, possessed some features in common with the marsupials but not with the placentals, evidence that these features were present in the last common ancestor of the two groups but were later lost in the placental lineage. In particular, the epipubic bones extend forwards from the pelvis. These are not found in any modern placental, but they are found in marsupials, monotremes, other nontherian mammals and '' Ukhaatherium'', an early Cretaceous animal in the eutherian order
Asioryctitheria Asioryctitheria ("Asian digging beasts") is an extinct order of early eutherians. Skull structure With the exception of '' Prokennalestes'', these advanced forms lacked a Meckelian groove. Furthermore, they were equipped with double-rooted ca ...
. This also applies to the multituberculates. They are apparently an ancestral feature, which subsequently disappeared in the placental lineage. These epipubic bones seem to function by stiffening the muscles during locomotion, reducing the amount of space being presented, which placentals require to contain their
fetus A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal devel ...
during gestation periods. A narrow pelvic outlet indicates that the young were very small at birth and therefore pregnancy was short, as in modern marsupials. This suggests that the placenta was a later development. One of the earliest known monotremes was '' Teinolophos'', which lived about 120 million years ago in Australia. Monotremes have some features which may be inherited from the original amniotes such as the same orifice to urinate, defecate and reproduce (
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and bir ...
)—as lizards and birds also do— and they lay
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
which are leathery and uncalcified.


Earliest appearances of features

''
Hadrocodium ''Hadrocodium wui'' is an extinct mammaliaform that lived during the Sinemurian stage of the Early Jurassic approximately in the Lufeng Formation of the Lufeng Basin in what is now the Yunnan province in south-western China (, paleocoordinat ...
'', whose fossils date from approximately 195 million years ago, in the early
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
, provides the first clear evidence of a jaw joint formed solely by the squamosal and dentary bones; there is no space in the jaw for the articular, a bone involved in the jaws of all early synapsids. The earliest clear evidence of hair or fur is in fossils of ''
Castorocauda ''Castorocauda'' is an extinct, semi-aquatic, superficially otter-like genus of docodont mammaliaforms with one species, ''C. lutrasimilis''. It is part of the Yanliao Biota, found in the Daohugou Beds of Inner Mongolia, China dating to the Mi ...
'' and ''
Megaconus ''Megaconus'' is an extinct genus of allotherian mammal from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. The type and only species, ''Megaconus mammaliaformis'' was first described in the journal '' Nature'' in 2013. ''Me ...
'', from 164 million years ago in the mid-Jurassic. In the 1950s, it was suggested that the foramina (passages) in the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...
e and
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
e (bones in the front of the upper jaw) of cynodonts were channels which supplied blood vessels and nerves to vibrissae (
whiskers Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are coarser ...
) and so were evidence of hair or fur; it was soon pointed out, however, that foramina do not necessarily show that an animal had vibrissae, as the modern lizard ''
Tupinambis ''Tupinambis'' is a lizard genus which belongs to the family Teiidae and contains eight described species. These large lizards are commonly referred to as tegus (''teiús'' in Portuguese). '' T. merianae'' (Argentine black and white tegu), '' ...
'' has foramina that are almost identical to those found in the nonmammalian cynodont ''
Thrinaxodon ''Thrinaxodon'' is an extinct genus of cynodonts, most commonly regarded by its species ''T. liorhinus'' which lived in what are now South Africa and Antarctica during the Early Triassic. ''Thrinaxodon'' lived just after the Permian–Triassic ...
''. Popular sources, nevertheless, continue to attribute whiskers to ''Thrinaxodon''. Studies on Permian
coprolites A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is der ...
suggest that non-mammalian
synapsids Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes rept ...
of the epoch already had fur, setting the evolution of hairs possibly as far back as
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typic ...
s. When
endothermy An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inst ...
first appeared in the evolution of mammals is uncertain, though it is generally agreed to have first evolved in non-mammalian
therapsids Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
. Modern monotremes have lower body temperatures and more variable metabolic rates than marsupials and placentals, but there is evidence that some of their ancestors, perhaps including ancestors of the therians, may have had body temperatures like those of modern therians. Likewise, some modern therians like afrotheres and xenarthrans have secondarily developed lower body temperatures. The evolution of erect limbs in mammals is incomplete—living and fossil monotremes have sprawling limbs. The parasagittal (nonsprawling) limb posture appeared sometime in the late Jurassic or early Cretaceous; it is found in the eutherian ''Eomaia'' and the metatherian ''Sinodelphys'', both dated to 125 million years ago.
Epipubic Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them). They first o ...
bones, a feature that strongly influenced the reproduction of most mammal clades, are first found in
Tritylodontidae Tritylodontidae ("three-knob teeth", named after the shape of their cheek teeth) is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, bearing several mammalian traits like erect limbs, endothermy and details ...
, suggesting that it is a
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
between them and
mammaliformes Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent comm ...
. They are omnipresent in non-placental mammaliformes, though ''
Megazostrodon ''Megazostrodon'' is an extinct genus of basal mammaliaforms belonging to the order Morganucodonta. It is approximately 200 million years old.
'' and ''
Erythrotherium ''Erythrotherium'' (meaning "red beast") is an extinct genus of basal mammaliaforms from the Late Triassic to Lower Jurassic. It is related to '' Morganucodon''. Only one species is recorded, ''Erythrotherium parringtoni'', from Red Beds, Stor ...
'' appear to have lacked them. It has been suggested that the original function of
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proces ...
(
milk Milk is a white liquid food produced by the mammary glands of mammals. It is the primary source of nutrition for young mammals (including breastfed human infants) before they are able to digest solid food. Immune factors and immune-modulati ...
production) was to keep eggs moist. Much of the argument is based on monotremes, the egg-laying mammals. In human females, mammary glands become fully developed during puberty, regardless of pregnancy.


Rise of the mammals

Therian mammals took over the medium- to large-sized ecological niches in the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configur ...
, after the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With ...
approximately 66 million years ago emptied ecological space once filled by non-avian dinosaurs and other groups of reptiles, as well as various other mammal groups, and underwent an exponential increase in body size (
megafauna In terrestrial zoology, the megafauna (from Greek μέγας ''megas'' "large" and New Latin ''fauna'' "animal life") comprises the large or giant animals of an area, habitat, or geological period, extinct and/or extant. The most common threshol ...
). Then mammals diversified very quickly; both birds and mammals show an exponential rise in diversity. For example, the earliest known bat dates from about 50 million years ago, only 16 million years after the extinction of the non-avian dinosaurs. Molecular phylogenetic studies initially suggested that most placental orders diverged about 100 to 85 million years ago and that modern families appeared in the period from the late
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "dawn ...
through the Miocene. However, no placental fossils have been found from before the end of the Cretaceous. The earliest undisputed fossils of placentals comes from the early
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''palai ...
, after the extinction of the non-avian dinosaurs. In particular, scientists have identified an early Paleocene animal named '' Protungulatum donnae'' as one of the first placental mammals. however it has been reclassified as a non-placental eutherian. Recalibrations of genetic and morphological diversity rates have suggested a
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'' ...
origin for placentals, and a Paleocene origin for most modern clades. The earliest known ancestor of primates is '' Archicebus achilles'' from around 55 million years ago. This tiny primate weighed 20–30 grams (0.7–1.1 ounce) and could fit within a human palm.


Anatomy


Distinguishing features

Living mammal species can be identified by the presence of
sweat gland Sweat glands, also known as sudoriferous or sudoriparous glands, , are small tubular structures of the skin that produce sweat. Sweat glands are a type of exocrine gland, which are glands that produce and secrete substances onto an epithelial su ...
s, including those that are specialized to produce milk to nourish their young. In classifying fossils, however, other features must be used, since soft tissue glands and many other features are not visible in fossils. Many traits shared by all living mammals appeared among the earliest members of the group: * Jaw joint – The
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bo ...
(the lower jaw bone, which carries the teeth) and the
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral c ...
(a small
cranial Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pr ...
bone) meet to form the joint. In most
gnathostomes Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, livin ...
, including early
therapsids Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
, the joint consists of the
articular The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals. Anatomy In most vertebrates, the articular bone is connected to two oth ...
(a small bone at the back of the lower jaw) and
quadrate Quadrate may refer to: * Quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squa ...
(a small bone at the back of the upper jaw). *
Middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
– In crown-group mammals, sound is carried from the
eardrum In the anatomy of humans and various other tetrapods, the eardrum, also called the tympanic membrane or myringa, is a thin, cone-shaped membrane that separates the external ear from the middle ear. Its function is to transmit sound from the ...
by a chain of three bones, the
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 vibrations ...
, the
incus The ''incus'' (plural incudes) or anvil is a bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store ...
and the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the f ...
. Ancestrally, the malleus and the incus are derived from the articular and the quadrate bones that constituted the jaw joint of early therapsids. * Tooth replacement – Teeth can be replaced once ( diphyodonty) or (as in toothed whales and
murid In Sufism, a ''murīd'' (Arabic مُرِيد 'one who seeks') is a novice committed to spiritual enlightenment by ''sulūk'' (traversing a path) under a spiritual guide, who may take the title murshid, '' pir'' or '' shaykh''. A '' sālik'' or S ...
rodents) not at all (
monophyodont A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
y). Elephants, manatees, and kangaroos continually grow new teeth throughout their life (
polyphyodont A polyphyodont is any animal whose teeth are continually replaced. In contrast, diphyodonts are characterized by having only two successive sets of teeth. Polyphyodonts include most toothed fishes, many reptiles such as crocodiles and geckos, ...
y). * Prismatic enamel – The enamel coating on the surface of a tooth consists of prisms, solid, rod-like structures extending from the
dentin Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by e ...
to the tooth's surface. * Occipital condyles – Two knobs at the base of the skull fit into the topmost neck vertebra; most other
tetrapod Tetrapods (; ) are four- limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s, in contrast, have only one such knob. For the most part, these characteristics were not present in the Triassic ancestors of the mammals. Nearly all mammaliaforms possess an epipubic bone, the exception being modern placentals.


Sexual dimorphism

On average, male mammals are larger than females, with males being at least 10% larger than females in over 45% of investigated species. Most mammalian orders also exhibit male-biased
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, although some orders do not show any bias or are significantly female-biased (
Lagomorpha The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (hares and rabbits) and the Ochotonidae ( pikas). The name of the order is derived from the Ancient Greek ''lagos'' (λα ...
). Sexual size dimorphism increases with body size across mammals (
Rensch's rule Rensch's rule is a biological rule on allometrics, concerning the relationship between the extent of sexual size dimorphism and which sex is larger. Across species within a lineage, size dimorphism increases with increasing body size when the m ...
), suggesting that there are parallel selection pressures on both male and female size. Male-biased dimorphism relates to sexual selection on males through male–male competition for females, as there is a positive correlation between the degree of sexual selection, as indicated by mating systems, and the degree of male-biased size dimorphism. The degree of sexual selection is also positively correlated with male and female size across mammals. Further, parallel selection pressure on female mass is identified in that age at weaning is significantly higher in more
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
species, even when correcting for body mass. Also, the reproductive rate is lower for larger females, indicating that fecundity selection selects for smaller females in mammals. Although these patterns hold across mammals as a whole, there is considerable variation across orders.


Biological systems

The majority of mammals have seven
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In sa ...
(bones in the neck). The exceptions are the
manatee Manatees (family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing three of the four living specie ...
and the
two-toed sloth ''Choloepus'' is a genus of xenarthran mammals of Central and South America within the monotypic family Choloepodidae, consisting of two-toed sloths, sometimes also called two-fingered sloths. The two species of ''Choloepus'' (which means "lame ...
, which have six, and the
three-toed sloth The three-toed or three-fingered sloths are arboreal neotropical mammals . They are the only members of the genus ''Bradypus'' and the family Bradypodidae. The four living species of three-toed sloths are the brown-throated sloth, the maned slo ...
which has nine. All mammalian brains possess a
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, ...
, a brain region unique to mammals. Placental brains have a
corpus callosum The corpus callosum (Latin for "tough body"), also callosal commissure, is a wide, thick nerve tract, consisting of a flat bundle of commissural fibers, beneath the cerebral cortex in the brain. The corpus callosum is only found in placenta ...
, unlike monotremes and marsupials. The lungs of mammals are spongy and honeycombed. Breathing is mainly achieved with the diaphragm, which divides the thorax from the abdominal cavity, forming a dome convex to the thorax. Contraction of the diaphragm flattens the dome, increasing the volume of the lung cavity. Air enters through the oral and nasal cavities, and travels through the larynx, trachea and
bronchi A bronchus is a passage or airway in the lower respiratory tract that conducts air into the lungs. The first or primary bronchi pronounced (BRAN-KAI) to branch from the trachea at the carina are the right main bronchus and the left main bronch ...
, and expands the
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * ...
. Relaxing the diaphragm has the opposite effect, decreasing the volume of the lung cavity, causing air to be pushed out of the lungs. During exercise, the abdominal wall
contracts A contract is a legally enforceable agreement between two or more parties that creates, defines, and governs mutual rights and obligations between them. A contract typically involves the transfer of goods, services, money, or a promise to ...
, increasing pressure on the diaphragm, which forces air out quicker and more forcefully. The
rib cage The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a se ...
is able to expand and contract the chest cavity through the action of other respiratory muscles. Consequently, air is sucked into or expelled out of the lungs, always moving down its pressure gradient. This type of lung is known as a bellows lung due to its resemblance to blacksmith
bellows A bellows or pair of bellows is a device constructed to furnish a strong blast of air. The simplest type consists of a flexible bag comprising a pair of rigid boards with handles joined by flexible leather sides enclosing an approximately airtig ...
. The mammalian
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide ...
has four chambers, two upper atria, the receiving chambers, and two lower ventricles, the discharging chambers. The heart has four valves, which separate its chambers and ensures blood flows in the correct direction through the heart (preventing backflow). After
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a b ...
in the pulmonary capillaries (blood vessels in the lungs), oxygen-rich blood returns to the left atrium via one of the four
pulmonary vein The pulmonary veins are the veins that transfer oxygenated blood from the lungs to the heart. The largest pulmonary veins are the four ''main pulmonary veins'', two from each lung that drain into the left atrium of the heart. The pulmonary ve ...
s. Blood flows nearly continuously back into the atrium, which acts as the receiving chamber, and from here through an opening into the left ventricle. Most blood flows passively into the heart while both the atria and ventricles are relaxed, but toward the end of the ventricular relaxation period, the left atrium will contract, pumping blood into the ventricle. The heart also requires nutrients and oxygen found in blood like other muscles, and is supplied via
coronary arteries The coronary arteries are the arterial blood vessels of coronary circulation, which transport oxygenated blood to the heart muscle. The heart requires a continuous supply of oxygen to function and survive, much like any other tissue or organ o ...
. The integumentary system (skin) is made up of three layers: the outermost
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
, the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided ...
and the
hypodermis The subcutaneous tissue (), also called the hypodermis, hypoderm (), subcutis, superficial fascia, is the lowermost layer of the integumentary system in vertebrates. The types of cells found in the layer are fibroblasts, adipose cells, and macr ...
. The epidermis is typically 10 to 30 cells thick; its main function is to provide a waterproof layer. Its outermost cells are constantly lost; its bottommost cells are constantly dividing and pushing upward. The middle layer, the dermis, is 15 to 40 times thicker than the epidermis. The dermis is made up of many components, such as bony structures and blood vessels. The hypodermis is made up of
adipose tissue Adipose tissue, body fat, or simply fat is a loose connective tissue composed mostly of adipocytes. In addition to adipocytes, adipose tissue contains the stromal vascular fraction (SVF) of cells including preadipocytes, fibroblasts, vascular ...
, which stores lipids and provides cushioning and insulation. The thickness of this layer varies widely from species to species;
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their ...
s require a thick hypodermis (
blubber Blubber is a thick layer of vascularized adipose tissue under the skin of all cetaceans, pinnipeds, penguins, and sirenians. Description Lipid-rich, collagen fiber-laced blubber comprises the hypodermis and covers the whole body, except for ...
) for insulation, and
right whale Right whales are three species of large baleen whales of the genus ''Eubalaena'': the North Atlantic right whale (''E. glacialis''), the North Pacific right whale (''E. japonica'') and the Southern right whale (''E. australis''). They are cla ...
s have the thickest blubber at . Although other animals have features such as whiskers,
feathers Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premie ...
,
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
, or
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike proj ...
that superficially resemble it, no animals other than mammals have
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal an ...
. It is a definitive characteristic of the class, though some mammals have very little. Herbivores have developed a diverse range of physical structures to facilitate the consumption of plant material. To break up intact plant tissues, mammals have developed
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tea ...
structures that reflect their feeding preferences. For instance,
frugivore A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...
s (animals that feed primarily on fruit) and herbivores that feed on soft foliage have low-crowned teeth specialized for grinding foliage and
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm p ...
s.
Grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and other ...
animals that tend to eat hard,
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
-rich grasses, have high-crowned teeth, which are capable of grinding tough plant tissues and do not wear down as quickly as low-crowned teeth. Most carnivorous mammals have
carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified ...
iforme teeth (of varying length depending on diet), long canines and similar tooth replacement patterns. The stomach of
even-toed ungulate The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poste ...
s (Artiodactyla) is divided into four sections: the
rumen The rumen, also known as a paunch, is the largest stomach compartment in ruminants and the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. The rumen's microbial favoring environment all ...
, the
reticulum Reticulum is a small, faint constellation in the southern sky. Its name is Latin for a small net, or reticle—a net of crosshairs at the focus of a telescope eyepiece that is used to measure star positions. The constellation is best viewed ...
, the
omasum The omasum, also known as the bible, the fardel, the manyplies and the psalterium, is the third compartment of the stomach in ruminants. The omasum comes after the rumen and reticulum and before the abomasum. Different ruminants have different o ...
and the
abomasum The abomasum, also known as the maw,The Cham ...
(only
ruminant Ruminants (suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The ...
s have a rumen). After the plant material is consumed, it is mixed with saliva in the rumen and reticulum and separates into solid and liquid material. The solids lump together to form a bolus (or cud), and is regurgitated. When the bolus enters the mouth, the fluid is squeezed out with the tongue and swallowed again. Ingested food passes to the rumen and reticulum where cellulolytic
microbe A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s (
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were amo ...
,
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
and
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fro ...
) produce
cellulase Cellulase (EC 3.2.1.4; systematic name 4-β-D-glucan 4-glucanohydrolase) is any of several enzymes produced chiefly by fungi, bacteria, and protozoans that catalyze cellulolysis, the decomposition of cellulose and of some related polysacchari ...
, which is needed to break down the
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
in plants.
Perissodactyls Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) ...
, in contrast to the ruminants, store digested food that has left the stomach in an enlarged
cecum The cecum or caecum is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix, to which it is joined). The w ...
, where it is fermented by bacteria. Carnivora have a simple stomach adapted to digest primarily meat, as compared to the elaborate digestive systems of herbivorous animals, which are necessary to break down tough, complex plant fibers. The
caecum The cecum or caecum is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix, to which it is joined). The w ...
is either absent or short and simple, and the large intestine is not sacculated or much wider than the small intestine. The mammalian
excretory system The excretory system is a passive biological system that removes excess, unnecessary materials from the body fluids of an organism, so as to help maintain internal chemical homeostasis and prevent damage to the body. The dual function of excreto ...
involves many components. Like most other land animals, mammals are
ureotelic Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds, ...
, and convert
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
into
urea Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid. Urea serves an important ...
, which is done by the
liver The liver is a major organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the synthesis of proteins and biochemicals necessary for digestion and growth. In humans, it i ...
as part of the
urea cycle The urea cycle (also known as the ornithine cycle) is a cycle of biochemical reactions that produces urea (NH2)2CO from ammonia (NH3). Animals that use this cycle, mainly amphibians and mammals, are called ureotelic. The urea cycle converts hig ...
.
Bilirubin Bilirubin (BR) (Latin for "red bile") is a red-orange compound that occurs in the normal catabolic pathway that breaks down heme in vertebrates. This catabolism is a necessary process in the body's clearance of waste products that arise from th ...
, a waste product derived from
blood cell A blood cell, also called a hematopoietic cell, hemocyte, or hematocyte, is a cell produced through hematopoiesis and found mainly in the blood. Major types of blood cells include red blood cells (erythrocytes), white blood cells (leukocytes), ...
s, is passed through
bile Bile (from Latin ''bilis''), or gall, is a dark-green-to-yellowish-brown fluid produced by the liver of most vertebrates that aids the digestion of lipids in the small intestine. In humans, bile is produced continuously by the liver (liver bil ...
and
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excreted from the body through the urethra. Cellu ...
with the help of enzymes excreted by the liver. The passing of bilirubin via bile through the intestinal tract gives mammalian
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a rela ...
a distinctive brown coloration. Distinctive features of the mammalian kidney include the presence of the
renal pelvis The renal pelvis or pelvis of the kidney is the funnel-like dilated part of the ureter in the kidney. It is formed by the covnvergence of the major calyces, acting as a funnel for urine flowing from the major calyces to the ureter. It has a mucous ...
and
renal pyramid The renal medulla is the innermost part of the kidney. The renal medulla is split up into a number of sections, known as the renal pyramids. Blood enters into the kidney via the renal artery, which then splits up to form the segmental arteries whi ...
s, and of a clearly distinguishable cortex and
medulla Medulla or Medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ovary * Medulla of t ...
, which is due to the presence of elongated loops of Henle. Only the mammalian kidney has a bean shape, although there are some exceptions, such as the multilobed reniculate kidneys of pinnipeds,
cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel t ...
ns and bears. Most adult placental mammals have no remaining trace of the
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and bir ...
. In the embryo, the embryonic cloaca divides into a posterior region that becomes part of the anus, and an anterior region that has different fates depending on the sex of the individual: in females, it develops into the vestibule that receives the
urethra The urethra (from Greek οὐρήθρα – ''ourḗthrā'') is a tube that connects the urinary bladder to the urinary meatus for the removal of urine from the body of both females and males. In human females and other primates, the urethra ...
and
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hyme ...
, while in males it forms the entirety of the penile urethra. However, the
tenrec A tenrec is any species of mammal within the afrotherian family Tenrecidae endemic to Madagascar. Tenrecs are wildly diverse; as a result of convergent evolution some resemble hedgehogs, shrews, opossums, rats, and mice. They occupy aquatic ...
s,
golden mole Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae and as such they are taxonomically distinct from the true moles, family Talpidae, and other mole-like families, al ...
s, and some
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to differ ...
s retain a cloaca as adults. In marsupials, the genital tract is separate from the anus, but a trace of the original cloaca does remain externally. Monotremes, which translates from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
into "single hole", have a true cloaca.


Sound production

As in all other tetrapods, mammals have a
larynx The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is ab ...
that can quickly open and close to produce sounds, and a supralaryngeal
vocal tract The vocal tract is the cavity in human bodies and in animals where the sound produced at the sound source ( larynx in mammals; syrinx in birds) is filtered. In birds it consists of the trachea, the syrinx, the oral cavity, the upper part of t ...
which filters this sound. The lungs and surrounding musculature provide the air stream and pressure required to phonate. The larynx controls the pitch and
volume Volume is a measure of occupied three-dimensional space. It is often quantified numerically using SI derived units (such as the cubic metre and litre) or by various imperial or US customary units (such as the gallon, quart, cubic inch). ...
of sound, but the strength the lungs exert to
exhale Exhalation (or expiration) is the flow of the breath out of an organism. In animals, it is the movement of air from the lungs out of the airways, to the external environment during breathing. This happens due to elastic properties of the lung ...
also contributes to volume. More primitive mammals, such as the echidna, can only hiss, as sound is achieved solely through exhaling through a partially closed larynx. Other mammals phonate using vocal folds. The movement or tenseness of the vocal folds can result in many sounds such as
purr A purr is a tonal fluttering sound made by some species of felids and two species of genets. It varies in loudness and tone among species and in the same animal. Felids are a family of mammals that belong to the order Carnivora and are inf ...
ing and
screaming A scream is a loud vocalization in which air is passed through the vocal cords with greater force than is used in regular or close-distance vocalisation. This can be performed by any creature possessing lungs, including humans. A scream is of ...
. Mammals can change the position of the larynx, allowing them to breathe through the nose while swallowing through the mouth, and to form both oral and nasal sounds; nasal sounds, such as a dog whine, are generally soft sounds, and oral sounds, such as a dog bark, are generally loud. Some mammals have a large larynx and thus a low-pitched voice, namely the hammer-headed bat (''Hypsignathus monstrosus'') where the larynx can take up the entirety of the
thoracic cavity The thoracic cavity (or chest cavity) is the chamber of the body of vertebrates that is protected by the thoracic wall ( rib cage and associated skin, muscle, and fascia). The central compartment of the thoracic cavity is the mediastinum. Ther ...
while pushing the lungs, heart, and trachea into the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
. Large vocal pads can also lower the pitch, as in the low-pitched roars of
big cat The term "big cat" is typically used to refer to any of the five living members of the genus ''Panthera'', namely the tiger, lion, jaguar, leopard, and snow leopard. Despite enormous differences in size, various cat species are quite simil ...
s. The production of
infrasound Infrasound, sometimes referred to as low status sound, describes sound waves with a frequency below the lower limit of human audibility (generally 20 Hz). Hearing becomes gradually less sensitive as frequency decreases, so for humans to perce ...
is possible in some mammals such as the African elephant (''Loxodonta'' spp.) and
baleen whale Baleen whales ( systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea ( whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") i ...
s. Small mammals with small larynxes have the ability to produce
ultrasound Ultrasound is sound waves with frequencies higher than the upper audible limit of human hearing. Ultrasound is not different from "normal" (audible) sound in its physical properties, except that humans cannot hear it. This limit varies ...
, which can be detected by modifications to the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
and
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory ...
. Ultrasound is inaudible to birds and reptiles, which might have been important during the Mesozoic, when birds and reptiles were the dominant predators. This private channel is used by some rodents in, for example, mother-to-pup communication, and by bats when echolocating. Toothed whales also use echolocation, but, as opposed to the vocal membrane that extends upward from the vocal folds, they have a
melon A melon is any of various plants of the family Cucurbitaceae with sweet, edible, and fleshy fruit. The word "melon" can refer to either the plant or specifically to the fruit. Botanically, a melon is a kind of berry, specifically a " pepo". T ...
to manipulate sounds. Some mammals, namely the primates, have air sacs attached to the larynx, which may function to lower the resonances or increase the volume of sound. The vocal production system is controlled by the
cranial nerve nuclei A cranial nerve nucleus is a collection of neurons ( gray matter) in the brain stem that is associated with one or more of the cranial nerves. Axons carrying information to and from the cranial nerves form a synapse first at these nuclei. Lesions ...
in the brain, and supplied by the
recurrent laryngeal nerve The recurrent laryngeal nerve (RLN) is a branch of the vagus nerve ( cranial nerve X) that supplies all the intrinsic muscles of the larynx, with the exception of the cricothyroid muscles. There are two recurrent laryngeal nerves, right and ...
and the superior laryngeal nerve, branches of the
vagus nerve The vagus nerve, also known as the tenth cranial nerve, cranial nerve X, or simply CN X, is a cranial nerve that interfaces with the parasympathetic control of the heart, lungs, and digestive tract. It comprises two nerves—the left and righ ...
. The vocal tract is supplied by the
hypoglossal nerve The hypoglossal nerve, also known as the twelfth cranial nerve, cranial nerve XII, or simply CN XII, is a cranial nerve that innervates all the extrinsic and intrinsic muscles of the tongue except for the palatoglossus, which is innervated b ...
and
facial nerve The facial nerve, also known as the seventh cranial nerve, cranial nerve VII, or simply CN VII, is a cranial nerve that emerges from the pons of the brainstem, controls the muscles of facial expression, and functions in the conveyance of t ...
s. Electrical stimulation of the
periaqueductal gray The periaqueductal gray (PAG, also known as the central gray) is a brain region that plays a critical role in autonomic function, motivated behavior and behavioural responses to threatening stimuli. PAG is also the primary control center for d ...
(PEG) region of the mammalian
midbrain The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal ( alertness), and temperature regulation. The name comes from the Greek ''mesos'', " ...
elicit vocalizations. The ability to learn new vocalizations is only exemplified in humans, seals, cetaceans, elephants and possibly bats; in humans, this is the result of a direct connection between the
motor cortex The motor cortex is the region of the cerebral cortex believed to be involved in the planning, control, and execution of voluntary movements. The motor cortex is an area of the frontal lobe located in the posterior precentral gyrus immediate ...
, which controls movement, and the
motor neuron A motor neuron (or motoneuron or efferent neuron) is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly or indirectl ...
s in the spinal cord.


Fur

The primary function of the fur of mammals is thermoregulation. Others include protection, sensory purposes, waterproofing, and camouflage. Different types of fur serve different purposes: * Definitive – which may be
shed A shed is typically a simple, single-story roofed structure that is used for hobbies, or as a workshop in a back garden or on an allotment. Sheds vary considerably in their size and complexity of construction, from simple open-sided ones ...
after reaching a certain length * Vibrissae – sensory hairs, most commonly
whisker Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are coarser ...
s * Pelage – guard hairs, under-fur, and awn hair * Spines – stiff guard hair used for defense (such as in
porcupine Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethizo ...
s) *
Bristle A bristle is a stiff hair or feather (natural or artificial), either on an animal, such as a pig, a plant, or on a tool such as a brush or broom. Synthetic types Synthetic materials such as nylon Nylon is a generic designation for a family ...
s – long hairs usually used in visual signals. (such as a lion's
mane Mane may refer to: *Mane (horse), the line of hair along the spine of the neck * Mane (lion), the hair found around the male mammal's neck In arts and entertainment * ''Mane'' (film) is a 1990 Kannada language film directed by Girish Kasaravalli ...
) * Velli – often called "down fur" which insulates newborn mammals *
Wool Wool is the textile fibre obtained from sheep and other mammals, especially goats, rabbits, and camelids. The term may also refer to inorganic materials, such as mineral wool and glass wool, that have properties similar to animal wool. As ...
– long, soft and often curly


Thermoregulation

Hair length is not a factor in thermoregulation: for example, some tropical mammals such as sloths have the same length of fur length as some arctic mammals but with less insulation; and, conversely, other tropical mammals with short hair have the same insulating value as arctic mammals. The denseness of fur can increase an animal's insulation value, and arctic mammals especially have dense fur; for example, the
musk ox Musk ( Persian: مشک, ''Mushk'') is a class of aromatic substances commonly used as base notes in perfumery. They include glandular secretions from animals such as the musk deer, numerous plants emitting similar fragrances, and artificial sub ...
has guard hairs measuring as well as a dense underfur, which forms an airtight coat, allowing them to survive in temperatures of . Some desert mammals, such as camels, use dense fur to prevent solar heat from reaching their skin, allowing the animal to stay cool; a camel's fur may reach in the summer, but the skin stays at .
Aquatic mammal Aquatic and semiaquatic mammals are a diverse group of mammals that dwell partly or entirely in bodies of water. They include the various marine mammals who dwell in oceans, as well as various freshwater species, such as the European otter. The ...
s, conversely, trap air in their fur to conserve heat by keeping the skin dry.


Coloration

Mammalian coats are colored for a variety of reasons, the major selective pressures including
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
,
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ( ...
, communication, and thermoregulation. Coloration in both the hair and skin of mammals is mainly determined by the type and amount of
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amin ...
;
eumelanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amin ...
s for brown and black colors and
pheomelanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amin ...
for a range of yellowish to reddish colors, giving mammals an earth tone. Some mammals have more vibrant colors; certain monkeys such
mandrill The mandrill (''Mandrillus sphinx'') is a large Old World monkey native to west central Africa. It is one of the most colorful mammals in the world, with red and blue skin on its face and posterior. The species is sexually dimorphic, as male ...
s and
vervet monkey The vervet monkey (''Chlorocebus pygerythrus''), or simply vervet, is an Old World monkey of the family Cercopithecidae native to Africa. The term "vervet" is also used to refer to all the members of the genus '' Chlorocebus''. The five distinct ...
s, and opossums such as the Mexican mouse opossums and Derby's woolly opossums, have blue skin due to light diffraction in
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
fibers. Many sloths appear green because their fur hosts green
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular ...
; this may be a
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or paras ...
relation that affords
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
to the sloths. Camouflage is a powerful influence in a large number of mammals, as it helps to conceal individuals from predators or prey. In arctic and subarctic mammals such as the
arctic fox The Arctic fox (''Vulpes lagopus''), also known as the white fox, polar fox, or snow fox, is a small fox native to the Arctic regions of the Northern Hemisphere and common throughout the Arctic tundra biome. It is well adapted to living i ...
(''Alopex lagopus''),
collared lemming ''Dicrostonyx'' is a genus of rodent Rodents (from Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower T ...
(''Dicrostonyx groenlandicus''),
stoat The stoat (''Mustela erminea''), also known as the Eurasian ermine, Beringian ermine and ermine, is a mustelid native to Eurasia and the northern portions of North America. Because of its wide circumpolar distribution, it is listed as Least Co ...
(''Mustela erminea''), and
snowshoe hare The snowshoe hare (''Lepus americanus''), also called the varying hare or snowshoe rabbit, is a species of hare found in North America. It has the name "snowshoe" because of the large size of its hind feet. The animal's feet prevent it from sin ...
(''Lepus americanus''), seasonal color change between brown in summer and white in winter is driven largely by camouflage. Some arboreal mammals, notably primates and marsupials, have shades of violet, green, or blue skin on parts of their bodies, indicating some distinct advantage in their largely
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, but others are exclusively arboreal. The habitats pose nu ...
habitat due to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.
Aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste o ...
, warning off possible predators, is the most likely explanation of the black-and-white pelage of many mammals which are able to defend themselves, such as in the foul-smelling
skunk Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or gi ...
and the powerful and aggressive
honey badger The honey badger (''Mellivora capensis''), also known as the ratel ( or ), is a mammal widely distributed in Africa, Southwest Asia, and the Indian subcontinent. Because of its wide range and occurrence in a variety of habitats, it is listed ...
. Coat color is sometimes
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most anim ...
, as in many primate species. Differences in female and male coat color may indicate nutrition and hormone levels, important in mate selection. Coat color may influence the ability to retain heat, depending on how much light is reflected. Mammals with a darker colored coat can absorb more heat from solar radiation, and stay warmer, and some smaller mammals, such as
vole Voles are small rodents that are relatives of lemmings and hamsters, but with a stouter body; a longer, hairy tail; a slightly rounder head; smaller eyes and ears; and differently formed molars (high-crowned with angular cusps instead of low ...
s, have darker fur in the winter. The white, pigmentless fur of arctic mammals, such as the polar bear, may reflect more solar radiation directly onto the skin. The dazzling black-and-white striping of zebras appear to provide some protection from biting flies.


Reproductive system

Mammals are solely
gonochoric In biology, gonochorism is a sexual system where there are only two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric. Gonochorism ...
(an animal is born with either male or female genitalia, as opposed to
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have s ...
s where there is no such schism). In male placentals, the
penis A penis (plural ''penises'' or ''penes'' () is the primary sexual organ that male animals use to inseminate females (or hermaphrodites) during copulation. Such organs occur in many animals, both vertebrate and invertebrate, but males d ...
is used both for urination and copulation. Depending on the species, an
erection An erection (clinically: penile erection or penile tumescence) is a physiological phenomenon in which the penis becomes firm, engorged, and enlarged. Penile erection is the result of a complex interaction of psychological, neural, vascular, ...
may be fueled by blood flow into vascular, spongy tissue or by muscular action. A penis may be contained in a prepuce when not erect, and some placentals also have a penis bone (
baculum The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a bone found in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as th ...
). Marsupials typically have forked penises, while the
echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and th ...
penis generally has four heads with only two functioning. The
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoste ...
of most mammals descend into the
scrotum The scrotum or scrotal sac is an anatomical male reproductive structure located at the base of the penis that consists of a suspended dual-chambered sac of skin and smooth muscle. It is present in most terrestrial male mammals. The scrotum con ...
which is typically posterior to the penis but is often anterior in marsupials. Female mammals generally have a
clitoris The clitoris ( or ) is a female sex organ present in mammals, ostriches and a limited number of other animals. In humans, the visible portion – the glans – is at the front junction of the labia minora (inner lips), above the op ...
,
labia majora The labia majora (singular: ''labium majus'') are two prominent longitudinal cutaneous folds that extend downward and backward from the mons pubis to the perineum. Together with the labia minora they form the labia of the vulva. The labia ma ...
and
labia minora The labia minora (Latin for 'smaller lips', singular: ''labium minus'', 'smaller lip'), also known as the inner labia, inner lips, vaginal lips or nymphae are two flaps of skin on either side of the human vaginal opening in the vulva, situated be ...
on the outside, while the internal system contains paired
oviduct The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, o ...
s, 1–2
uteri The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The u ...
, 1–2
cervices The cervix or cervix uteri (Latin, 'neck of the uterus') is the lower part of the uterus (womb) in the human female reproductive system. The cervix is usually 2 to 3 cm long (~1 inch) and roughly cylindrical in shape, which changes during ...
and a
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hyme ...
. Marsupials have two lateral vaginas and a medial vagina. The "vagina" of monotremes is better understood as a "urogenital sinus". The uterine systems of placental mammals can vary between a duplex, where there are two uteri and cervices which open into the vagina, a bipartite, where two uterine horns have a single cervix that connects to the vagina, a bicornuate, which consists where two uterine horns that are connected distally but separate medially creating a Y-shape, and a simplex, which has a single uterus. The ancestral condition for mammal reproduction is the birthing of relatively undeveloped, either through direct
vivipary In plants, vivipary occurs when seeds or embryos begin to develop before they detach from the parent. Plants such as some Iridaceae and Agavoideae grow cormlets in the axils of their inflorescences. These fall and in favourable circumstances the ...
or a short period as soft-shelled eggs. This is likely due to the fact that the torso could not expand due to the presence of
epipubic bones Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them). They first occur ...
. The oldest demonstration of this reproductive style is with ''
Kayentatherium ''Kayentatherium'' is an extinct genus of tritylodontid cynodonts that lived during the Early Jurassic. It is one of two tritylodonts from the Kayenta Formation of northern Arizona, United States. ''Kayentatherium'' means "Kayenta Beast", and i ...
'', which produced undeveloped
perinate A perinate is a member of a viviparous species from approximately one month before birth to one month after it. See also * Fetus A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an an ...
s, but at much higher litter sizes than any modern mammal, 38 specimens. Most modern mammals are
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
, giving birth to live young. However, the five species of monotreme, the platypus and the four species of echidna, lay eggs. The monotremes have a
sex determination system A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. Most organisms that create their offspring using sexual reproduction have two sexes. In some species there are hermaphr ...
different from that of most other mammals. In particular, the
sex chromosome A sex chromosome (also referred to as an allosome, heterotypical chromosome, gonosome, heterochromosome, or idiochromosome) is a chromosome that differs from an ordinary autosome in form, size, and behavior. The human sex chromosomes, a typica ...
s of a platypus are more like those of a chicken than those of a therian mammal. Viviparous mammals are in the subclass Theria; those living today are in the marsupial and placental infraclasses. Marsupials have a short
gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pr ...
period, typically shorter than its
estrous cycle The estrous cycle (, originally ) is the set of recurring physiological changes that are induced by reproductive hormones in most mammalian therian females. Estrous cycles start after sexual maturity in females and are interrupted by anestrous ...
and generally giving birth to a number of undeveloped newborns that then undergo further development; in many species, this takes place within a pouch-like sac, the marsupium, located in the front of the mother's
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
. This is the
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, an ...
condition among viviparous mammals; the presence of epipubic bones in all non-placental mammals prevents the expansion of the torso needed for full pregnancy. Even non-placental eutherians probably reproduced this way. The placentals give birth to relatively complete and developed young, usually after long gestation periods. They get their name from the
placenta The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate matern ...
, which connects the developing fetus to the uterine wall to allow nutrient uptake. In placental mammals, the epipubic is either completely lost or converted into the baculum; allowing the torso to be able to expand and thus birth developed offspring. The mammary glands of mammals are specialized to produce milk, the primary source of nutrition for newborns. The monotremes branched early from other mammals and do not have the nipples seen in most mammals, but they do have mammary glands. The young lick the milk from a mammary patch on the mother's belly. Compared to placental mammals, the milk of marsupials changes greatly in both production rate and in nutrient composition, due to the underdeveloped young. In addition, the mammary glands have more autonomy allowing them to supply separate milks to young at different development stages.
Lactose Lactose is a disaccharide sugar synthesized by galactose and glucose subunits and has the molecular formula C12H22O11. Lactose makes up around 2–8% of milk (by mass). The name comes from ' (gen. '), the Latin word for milk, plus the suffix ' ...
is the main sugar in placental mammal milk while monotreme and marsupial milk is dominated by
oligosaccharide An oligosaccharide (/ˌɑlɪgoʊˈsækəˌɹaɪd/; from the Greek ὀλίγος ''olígos'', "a few", and σάκχαρ ''sácchar'', "sugar") is a saccharide polymer containing a small number (typically two to ten) of monosaccharides (simple suga ...
s. Weaning is the process in which a mammal becomes less dependent on their mother's milk and more on solid food.


Endothermy

Nearly all mammals are
endothermic In thermochemistry, an endothermic process () is any thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, H.P., Butler, L. J. (2015).''Principle of Modern Chemistry'', Brooks Cole. p. ...
("warm-blooded"). Most mammals also have hair to help keep them warm. Like birds, mammals can forage or hunt in weather and climates too cold for
ectotherm An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life ...
ic ("cold-blooded") reptiles and insects. Endothermy requires plenty of food energy, so mammals eat more food per unit of body weight than most reptiles. Small insectivorous mammals eat prodigious amounts for their size. A rare exception, the
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 ...
produces little metabolic heat, so it is considered an operational
poikilotherm A poikilotherm () is an animal whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important stressors is temperature change, which can lead to alterations in membrane ...
. Birds are also endothermic, so endothermy is not unique to mammals.


Species lifespan

Among mammals, species maximum lifespan varies significantly (for example the
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to differ ...
has a lifespan of two years, whereas the oldest
bowhead whale The bowhead whale (''Balaena mysticetus'') is a species of baleen whale belonging to the family Balaenidae and the only living representative of the genus '' Balaena''. They are the only baleen whale endemic to the Arctic and subarctic waters ...
is recorded to be 211 years). Although the underlying basis for these lifespan differences is still uncertain, numerous studies indicate that the ability to repair DNA damage is an important determinant of mammalian lifespan. In a 1974 study by Hart and Setlow, it was found that DNA excision repair capability increased systematically with species lifespan among seven mammalian species. Species lifespan was observed to be robustly correlated with the capacity to recognize DNA double-strand breaks as well as the level of the DNA repair protein
Ku80 Ku80 is a protein that, in humans, is encoded by the ''XRCC5'' gene. Together, Ku70 and Ku80 make up the Ku heterodimer, which binds to DNA double-strand break ends and is required for the non-homologous end joining (NHEJ) pathway of DNA repa ...
. In a study of the cells from sixteen mammalian species, genes employed in DNA repair were found to be up-regulated in the longer-lived species. The cellular level of the DNA repair enzyme
poly ADP ribose polymerase Poly (ADP-ribose) polymerase (PARP) is a family of proteins involved in a number of cellular processes such as DNA repair, genomic stability, and programmed cell death. Members of PARP family The PARP family comprises 17 members (10 putative ...
was found to correlate with species lifespan in a study of 13 mammalian species. Three additional studies of a variety of mammalian species also reported a correlation between species lifespan and DNA repair capability.


Locomotion


Terrestrial

Most vertebrates—the amphibians, the reptiles and some mammals such as humans and bears—are
plantigrade 151px, Portion of a human skeleton, showing plantigrade habit In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. T ...
, walking on the whole of the underside of the foot. Many mammals, such as cats and dogs, are
digitigrade In terrestrial vertebrates, digitigrade () locomotion is walking or running on the toes (from the Latin ''digitus'', 'finger', and ''gradior'', 'walk'). A digitigrade animal is one that stands or walks with its toes (metatarsals) touching the groun ...
, walking on their toes, the greater stride length allowing more speed. Digitigrade mammals are also often adept at quiet movement. Some animals such as
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 millio ...
s are
unguligrade Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
, walking on the tips of their toes. This even further increases their stride length and thus their speed. A few mammals, namely the great apes, are also known to walk on their knuckles, at least for their front legs.
Giant anteater The giant anteater (''Myrmecophaga tridactyla'') is an insectivorous mammal native to Central and South America. It is one of four living species of anteaters, of which it is the largest member. The only extant member of the genus ''Myrmecoph ...
s and platypuses are also knuckle-walkers. Some mammals are
bipeds Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
, using only two limbs for locomotion, which can be seen in, for example, humans and the great apes. Bipedal species have a larger field of vision than quadrupeds, conserve more energy and have the ability to manipulate objects with their hands, which aids in foraging. Instead of walking, some bipeds hop, such as kangaroos and
kangaroo rat Kangaroo rats, small mostly nocturnal rodents of genus ''Dipodomys'', are native to arid areas of western North America. The common name derives from their bipedal form. They hop in a manner similar to the much larger kangaroo, but developed th ...
s. Animals will use different gaits for different speeds, terrain and situations. For example, horses show four natural gaits, the slowest
horse gait Horses can use various gaits (patterns of leg movement) during locomotion across solid ground, either naturally or as a result of specialized training by humans.Ensminger, M. E. ''Horses and Horsemanship'' 6th edition USA: Interstate Publisher ...
is the
walk Walking (also known as ambulation) is one of the main gaits of terrestrial locomotion among legged animals. Walking is typically slower than running and other gaits. Walking is defined by an ' inverted pendulum' gait in which the body vaults ...
, then there are three faster gaits which, from slowest to fastest, are the
trot The trot is a ten-beat diagonal horse gait where the diagonal pairs of legs move forward at the same time with a moment of suspension between each beat. It has a wide variation in possible speeds, but averages about . A very slow trot is somet ...
, the
canter The canter and gallop are variations on the fastest gait that can be performed by a horse or other equine. The canter is a controlled three-beat gait, while the gallop is a faster, four-beat variation of the same gait. It is a natural gait pos ...
and the
gallop The canter and gallop are variations on the fastest gait that can be performed by a horse or other equine. The canter is a controlled three-beat gait, while the gallop is a faster, four-beat variation of the same gait. It is a natural gait pos ...
. Animals may also have unusual gaits that are used occasionally, such as for moving sideways or backwards. For example, the main human gaits are bipedal
walking Walking (also known as ambulation) is one of the main gaits of terrestrial locomotion among legged animals. Walking is typically slower than running and other gaits. Walking is defined by an ' inverted pendulum' gait in which the body vaults ...
and
running Running is a method of terrestrial locomotion allowing humans and other animals to move rapidly on foot. Running is a type of gait characterized by an aerial phase in which all feet are above the ground (though there are exceptions). This is ...
, but they employ many other gaits occasionally, including a four-legged
crawl Crawl, The Crawl, or crawling may refer to: Biology * Crawling (human), any of several types of human quadrupedal gait * Limbless locomotion, the movement of limbless animals over the ground * Undulatory locomotion, a type of motion characteriz ...
in tight spaces. Mammals show a vast range of
gait Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficien ...
s, the order that they place and lift their appendages in locomotion. Gaits can be grouped into categories according to their patterns of support sequence. For quadrupeds, there are three main categories: walking gaits, running gaits and leaping gaits. Walking is the most common gait, where some feet are on the ground at any given time, and found in almost all legged animals. Running is considered to occur when at some points in the stride all feet are off the ground in a moment of suspension.


Arboreal

Arboreal animals frequently have elongated limbs that help them cross gaps, reach fruit or other resources, test the firmness of support ahead and, in some cases, to
brachiate Brachiation (from "brachium", Latin for "arm"), or arm swinging, is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb ...
(swing between trees). Many arboreal species, such as tree porcupines, silky anteaters, spider monkeys, and
possums Possum may refer to: Animals * Phalangeriformes, or possums, any of a number of arboreal marsupial species native to Australia, New Guinea, and Sulawesi ** Common brushtail possum (''Trichosurus vulpecula''), a common possum in Australian urban ...
, use
prehensile tail A prehensile tail is the tail of an animal that has adapted to grasp or hold objects. Fully prehensile tails can be used to hold and manipulate objects, and in particular to aid arboreal creatures in finding and eating food in the trees. If the t ...
s to grasp branches. In the spider monkey, the tip of the tail has either a bare patch or adhesive pad, which provides increased friction. Claws can be used to interact with rough substrates and reorient the direction of forces the animal applies. This is what allows
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
s to climb tree trunks that are so large to be essentially flat from the perspective of such a small animal. However, claws can interfere with an animal's ability to grasp very small branches, as they may wrap too far around and prick the animal's own paw. Frictional gripping is used by primates, relying upon hairless fingertips. Squeezing the branch between the fingertips generates frictional force that holds the animal's hand to the branch. However, this type of grip depends upon the angle of the frictional force, thus upon the diameter of the branch, with larger branches resulting in reduced gripping ability. To control descent, especially down large diameter branches, some arboreal animals such as squirrels have evolved highly mobile ankle joints that permit rotating the foot into a 'reversed' posture. This allows the claws to hook into the rough surface of the bark, opposing the force of gravity. Small size provides many advantages to arboreal species: such as increasing the relative size of branches to the animal, lower center of mass, increased stability, lower mass (allowing movement on smaller branches) and the ability to move through more cluttered habitat. Size relating to weight affects gliding animals such as the
sugar glider The sugar glider (''Petaurus breviceps'') is a small, omnivorous, arboreal, and nocturnal gliding possum belonging to the marsupial infraclass. The common name refers to its predilection for sugary foods such as sap and nectar and its abili ...
. Some species of primate, bat and all species of
sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their ...
achieve passive stability by hanging beneath the branch. Both pitching and tipping become irrelevant, as the only method of failure would be losing their grip.


Aerial

Bats are the only mammals that can truly fly. They fly through the air at a constant speed by moving their wings up and down (usually with some fore-aft movement as well). Because the animal is in motion, there is some airflow relative to its body which, combined with the velocity of the wings, generates a faster airflow moving over the wing. This generates a lift force vector pointing forwards and upwards, and a drag force vector pointing rearwards and upwards. The upwards components of these counteract gravity, keeping the body in the air, while the forward component provides thrust to counteract both the drag from the wing and from the body as a whole. The wings of bats are much thinner and consist of more bones than those of birds, allowing bats to maneuver more accurately and fly with more lift and less drag. By folding the wings inwards towards their body on the upstroke, they use 35% less energy during flight than birds. The membranes are delicate, ripping easily; however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly. The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cells, also found on human fingertips. These sensitive areas are different in bats, as each bump has a tiny hair in the center, making it even more sensitive and allowing the bat to detect and collect information about the air flowing over its wings, and to fly more efficiently by changing the shape of its wings in response.


Fossorial and subterranean

A fossorial (from Latin ''fossor'', meaning "digger") is an animal adapted to digging which lives primarily, but not solely, underground. Some examples are
badger Badgers are short-legged omnivores in the family Mustelidae (which also includes the otters, wolverines, martens, minks, polecats, weasels, and ferrets). Badgers are a polyphyletic rather than a natural taxonomic grouping, being united by ...
s, and
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. Many rodent species are also considered fossorial because they live in burrows for most but not all of the day. Species that live exclusively underground are subterranean, and those with limited adaptations to a fossorial lifestyle sub-fossorial. Some organisms are fossorial to aid in temperature regulation while others use the underground habitat for protection from
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s or for
food storage Food storage is a way of decreasing the variability of the food supply in the face of natural, inevitable variability. p.507 It allows food to be eaten for some time (typically weeks to months) after harvest rather than solely immediately. I ...
.Damiani, R, 2003, Earliest evidence of cynodont burrowing, The Royal Society Publishing, Volume 270, Issue 1525 Fossorial mammals have a fusiform body, thickest at the shoulders and tapering off at the tail and nose. Unable to see in the dark burrows, most have degenerated eyes, but degeneration varies between species;
pocket gopher Pocket gophers, commonly referred to simply as gophers, are burrowing rodents of the family Geomyidae. The roughly 41 speciesSearch results for "Geomyidae" on thASM Mammal Diversity Database are all endemic to North and Central America. They a ...
s, for example, are only semi-fossorial and have very small yet functional eyes, in the fully fossorial marsupial mole the eyes are degenerated and useless, talpa moles have
vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
eyes and the
cape golden mole The Cape golden mole (''Chrysochloris asiatica'') is a small, insectivorous mammal Mammals () are a group of vertebrate Vertebrates () comprise all animal Animals are multicellular, eukaryotic organisms in the biologi ...
has a layer of skin covering the eyes. External ears flaps are also very small or absent. Truly fossorial mammals have short, stout legs as strength is more important than speed to a burrowing mammal, but semi-fossorial mammals have
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 u ...
legs. The front paws are broad and have strong claws to help in loosening dirt while excavating burrows, and the back paws have webbing, as well as claws, which aids in throwing loosened dirt backwards. Most have large incisors to prevent dirt from flying into their mouth. Many fossorial mammals such as shrews, hedgehogs, and moles were classified under the now obsolete order
Insectivora The order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order Eulipotyphla, ...
.


Aquatic

Fully aquatic mammals, the cetaceans and
sirenia The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct ...
ns, have lost their legs and have a tail fin to propel themselves through the water. Flipper movement is continuous. Whales swim by moving their tail fin and lower body up and down, propelling themselves through vertical movement, while their flippers are mainly used for steering. Their skeletal anatomy allows them to be fast swimmers. Most species have a
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through c ...
to prevent themselves from turning upside-down in the water. The flukes of sirenians are raised up and down in long strokes to move the animal forward, and can be twisted to turn. The forelimbs are paddle-like flippers which aid in turning and slowing.
Semi-aquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semi aquatic animals include: * Ver ...
mammals, like pinnipeds, have two pairs of flippers on the front and back, the fore-flippers and hind-flippers. The elbows and ankles are enclosed within the body. Pinnipeds have several adaptions for reducing
drag Drag or The Drag may refer to: Places * Drag, Norway, a village in Tysfjord municipality, Nordland, Norway * ''Drág'', the Hungarian name for Dragu Commune in Sălaj County, Romania * Drag (Austin, Texas), the portion of Guadalupe Street a ...
. In addition to their streamlined bodies, they have smooth networks of muscle bundles in their skin that may increase
laminar flow In fluid dynamics, laminar flow is characterized by fluid particles following smooth paths in layers, with each layer moving smoothly past the adjacent layers with little or no mixing. At low velocities, the fluid tends to flow without lateral mi ...
and make it easier for them to slip through water. They also lack
arrector pili The arrector pili muscles, also known as hair erector muscles, are small muscles attached to hair follicles in mammals. Contraction of these muscles causes the hairs to stand on end, known colloquially as goose bumps (piloerection). Structure ...
, so their fur can be streamlined as they swim. They rely on their fore-flippers for locomotion in a wing-like manner similar to
penguin Penguins (order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adapt ...
s and
sea turtles Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhe ...
. Fore-flipper movement is not continuous, and the animal glides between each stroke. Compared to terrestrial carnivorans, the fore-limbs are reduced in length, which gives the locomotor muscles at the shoulder and elbow joints greater mechanical advantage; the hind-flippers serve as stabilizers. Other semi-aquatic mammals include beavers,
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
es,
otter Otters are carnivorous mammals in the subfamily Lutrinae. The 13 extant otter species are all semiaquatic, aquatic, or marine, with diets based on fish and invertebrates. Lutrinae is a branch of the Mustelidae family, which also includes wea ...
s and platypuses. Hippos are very large semi-aquatic mammals, and their barrel-shaped bodies have graviportal skeletal structures, adapted to carrying their enormous weight, and their
specific gravity Relative density, or specific gravity, is the ratio of the density (mass of a unit volume) of a substance to the density of a given reference material. Specific gravity for liquids is nearly always measured with respect to water at its densest ...
allows them to sink and move along the bottom of a river.


Behavior


Communication and vocalization

Many mammals communicate by vocalizing. Vocal communication serves many purposes, including in mating rituals, as warning calls, to indicate food sources, and for social purposes. Males often call during mating rituals to ward off other males and to attract females, as in the
roaring A roar is a type of animal vocalization that is deep and resonating. Many mammals have evolved to produce roars and other roar-like vocals for purposes such as long-distance communication and intimidation. These include various species of big ca ...
of
lion The lion (''Panthera leo'') is a large cat of the genus ''Panthera'' native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphic; adult ...
s and
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or hart, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of we ...
. The
songs A song is a musical composition intended to be performed by the human voice. This is often done at distinct and fixed pitches (melodies) using patterns of sound and silence. Songs contain various forms, such as those including the repetit ...
of the humpback whale may be signals to females; they have different dialects in different regions of the ocean. Social vocalizations include the
territorial A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
calls of gibbons, and the use of frequency in greater spear-nosed bats to distinguish between groups. The
vervet monkey The vervet monkey (''Chlorocebus pygerythrus''), or simply vervet, is an Old World monkey of the family Cercopithecidae native to Africa. The term "vervet" is also used to refer to all the members of the genus '' Chlorocebus''. The five distinct ...
gives a distinct alarm call for each of at least four different predators, and the reactions of other monkeys vary according to the call. For example, if an alarm call signals a python, the monkeys climb into the trees, whereas the eagle alarm causes monkeys to seek a hiding place on the ground.
Prairie dogs Prairie dogs (genus ''Cynomys'') are herbivorous burrowing ground squirrels native to the grasslands of North America. Within the genus are five species: black-tailed, white-tailed, Gunnison's, Utah, and Mexican prairie dogs. In Mexico, ...
similarly have complex calls that signal the type, size, and speed of an approaching predator. Elephants communicate socially with a variety of sounds including snorting, screaming, trumpeting, roaring and rumbling. Some of the rumbling calls are
infrasonic Infrasound, sometimes referred to as low status sound, describes sound waves with a frequency below the lower limit of human audibility (generally 20 Hz). Hearing becomes gradually less sensitive as frequency decreases, so for humans to perce ...
, below the hearing range of humans, and can be heard by other elephants up to away at still times near sunrise and sunset. left, Orca calling including occasional echolocation clicks Mammals signal by a variety of means. Many give visual anti-predator signals, as when deer and
gazelle A gazelle is one of many antelope species in the genus ''Gazella'' . This article also deals with the seven species included in two further genera, ''Eudorcas'' and ''Nanger'', which were formerly considered subgenera of ''Gazella''. A third f ...
stot, honestly indicating their fit condition and their ability to escape, or when
white-tailed deer The white-tailed deer (''Odocoileus virginianus''), also known as the whitetail or Virginia deer, is a medium-sized deer native to North America, Central America, and South America as far south as Peru and Bolivia. It has also been introduced ...
and other prey mammals flag with conspicuous tail markings when alarmed, informing the predator that it has been detected. Many mammals make use of scent-marking, sometimes possibly to help defend territory, but probably with a range of functions both within and between species.
Microbat Microbats constitute the suborder Microchiroptera within the order Chiroptera (bats). Bats have long been differentiated into Megachiroptera (megabats) and Microchiroptera, based on their size, the use of echolocation by the Microchiroptera an ...
s and
toothed whale The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales possessing teeth, such as the beaked whales and sperm whales. Seventy-three species o ...
s including
oceanic dolphin Oceanic dolphins or Delphinidae are a widely distributed family of dolphins that live in the sea. Close to forty extant species are recognised. They include several big species whose common names contain "whale" rather than "dolphin", such as t ...
s vocalize both socially and in echolocation.


Feeding

To maintain a high constant body temperature is energy expensive—mammals therefore need a nutritious and plentiful diet. While the earliest mammals were probably predators, different species have since adapted to meet their dietary requirements in a variety of ways. Some eat other animals—this is a
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other so ...
diet (and includes insectivorous diets). Other mammals, called
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthp ...
s, eat plants, which contain complex carbohydrates such as cellulose. An herbivorous diet includes subtypes such as
granivory Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed predators) feed on the seeds of plants as a main or exclusive food source,Hulme, P.E. and Benkman, C.W. (2002) "Granivory", pp. 1 ...
(seed eating),
folivory In zoology, a folivore is a herbivore that specializes in eating leaves. Mature leaves contain a high proportion of hard-to-digest cellulose, less energy than other types of foods, and often toxic compounds.Jones, S., Martin, R., & Pilbeam, D. (1 ...
(leaf eating),
frugivory A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...
(fruit eating),
nectarivory In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants. Nectar as a food source presents a number of benefits ...
(nectar eating), gummivory (gum eating) and
mycophagy Fungivory or mycophagy is the process of organisms consuming fungi. Many different organisms have been recorded to gain their energy from consuming fungi, including birds, mammals, insects, plants, amoebas, gastropods, nematodes, bacteria and othe ...
(fungus eating). The digestive tract of an herbivore is host to bacteria that ferment these complex substances, and make them available for digestion, which are either housed in the multichambered
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomac ...
or in a large cecum. Some mammals are
coprophagous Coprophagia () or coprophagy () is the consumption of feces. The word is derived from the grc, κόπρος , "feces" and , "to eat". Coprophagy refers to many kinds of feces-eating, including eating feces of other species (heterospecifics), of ...
, consuming
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a rela ...
to absorb the nutrients not digested when the food was first ingested. An
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the n ...
eats both prey and plants. Carnivorous mammals have a simple
digestive tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans an ...
because the proteins,
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
s and minerals found in meat require little in the way of specialized digestion. Exceptions to this include
baleen whale Baleen whales ( systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea ( whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") i ...
s who also house
gut flora Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gu ...
in a multi-chambered stomach, like terrestrial herbivores. The size of an animal is also a factor in determining diet type (
Allen's rule Allen's rule is an ecogeographical rule formulated by Joel Asaph Allen in 1877, broadly stating that animals adapted to cold climates have thicker limbs and bodily appendages than animals adapted to warm climates. More specifically, it states tha ...
). Since small mammals have a high ratio of heat-losing surface area to heat-generating volume, they tend to have high energy requirements and a high
metabolic rate Metabolism (, from el, μεταβολή ''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 cel ...
. Mammals that weigh less than about are mostly insectivorous because they cannot tolerate the slow, complex digestive process of an herbivore. Larger animals, on the other hand, generate more heat and less of this heat is lost. They can therefore tolerate either a slower collection process (carnivores that feed on larger vertebrates) or a slower digestive process (herbivores). Furthermore, mammals that weigh more than usually cannot collect enough insects during their waking hours to sustain themselves. The only large insectivorous mammals are those that feed on huge colonies of insects (
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total o ...
s or
termite Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blat ...
s). Some mammals are omnivores and display varying degrees of carnivory and herbivory, generally leaning in favor of one more than the other. Since plants and meat are digested differently, there is a preference for one over the other, as in bears where some species may be mostly carnivorous and others mostly herbivorous. They are grouped into three categories: mesocarnivory (50–70% meat),
hypercarnivory A hypercarnivore is an animal which has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
(70% and greater of meat), and hypocarnivory (50% or less of meat). The dentition of hypocarnivores consists of dull, triangular carnassial teeth meant for grinding food. Hypercarnivores, however, have conical teeth and sharp carnassials meant for slashing, and in some cases strong jaws for bone-crushing, as in the case of
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the c ...
s, allowing them to consume bones; some extinct groups, notably the
Machairodontinae Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million unt ...
, had saber-shaped
canines Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth In mammalian oral anatomy, the canine teet ...
. Some physiological carnivores consume plant matter and some physiological herbivores consume meat. From a behavioral aspect, this would make them omnivores, but from the physiological standpoint, this may be due to
zoopharmacognosy Zoopharmacognosy is a behaviour in which non-human animals self-medicate by selecting and ingesting or topically applying plants, soils, insects, and psychoactive drugs to prevent or reduce the harmful effects of pathogens, toxins, and even ...
. Physiologically, animals must be able to obtain both energy and nutrients from plant and animal materials to be considered omnivorous. Thus, such animals are still able to be classified as carnivores and herbivores when they are just obtaining nutrients from materials originating from sources that do not seemingly complement their classification. For example, it is well documented that some ungulates such as giraffes, camels, and cattle, will gnaw on bones to consume particular minerals and nutrients. Also, cats, which are generally regarded as obligate carnivores, occasionally eat grass to regurgitate indigestible material (such as
hairball A hairball is a small collection of hair or fur formed in the stomach of animals, and uncommonly in humans, that is occasionally vomited up when it becomes too big. Hairballs are primarily a tight elongated cylinder of packed fur, but may incl ...
s), aid with hemoglobin production, and as a laxative. Many mammals, in the absence of sufficient food requirements in an environment, suppress their metabolism and conserve energy in a process known as
hibernation Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It mo ...
. In the period preceding hibernation, larger mammals, such as bears, become polyphagic to increase fat stores, whereas smaller mammals prefer to collect and stash food. The slowing of the metabolism is accompanied by a decreased heart and respiratory rate, as well as a drop in internal temperatures, which can be around ambient temperature in some cases. For example, the internal temperatures of hibernating
arctic ground squirrel The Arctic ground squirrel (''Urocitellus parryii'') ( Inuktitut: ''ᓯᒃᓯᒃ, siksik'') is a species of ground squirrel native to the Arctic and Subarctic of North America and Asia. People in Alaska, particularly around the Aleutians, refer ...
s can drop to , however the head and neck always stay above . A few mammals in hot environments
aestivate Aestivation ( la, aestas (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a ...
in times of drought or extreme heat, for example the fat-tailed dwarf lemur (''Cheirogaleus medius'').


Intelligence

In intelligent mammals, such as primates, the
cerebrum The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb ...
is larger relative to the rest of the brain.
Intelligence Intelligence has been defined in many ways: the capacity for abstraction, logic, understanding, self-awareness, learning, emotional knowledge, reasoning, planning, creativity, critical thinking, and problem-solving. More generally, it can be des ...
itself is not easy to define, but indications of intelligence include the ability to learn, matched with behavioral flexibility.
Rats Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' ( pack rats), '' Bandicota'' ( bandico ...
, for example, are considered to be highly intelligent, as they can learn and perform new tasks, an ability that may be important when they first colonize a fresh habitat. In some mammals, food gathering appears to be related to intelligence: a deer feeding on plants has a brain smaller than a cat, which must think to outwit its prey.
Tool use by animals Tool use by animals is a phenomenon in which an animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, defence, communication, recreation or construction. Originally thought to be a skill possessed ...
may indicate different levels of
learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, values, attitudes, and preferences. The ability to learn is possessed by humans, animals, and some machines; there is also evidence for some kind of l ...
and
cognition Cognition refers to "the mental action or process of acquiring knowledge and understanding through thought, experience, and the senses". It encompasses all aspects of intellectual functions and processes such as: perception, attention, thou ...
. The
sea otter The sea otter (''Enhydra lutris'') is a marine mammal native to the coasts of the northern and eastern North Pacific Ocean. Adult sea otters typically weigh between , making them the heaviest members of the weasel family, but among the sma ...
uses rocks as essential and regular parts of its foraging behaviour (smashing
abalone Abalone ( or ; via Spanish , from Rumsen ''aulón'') is a common name for any of a group of small to very large marine gastropod molluscs in the family Haliotidae. Other common names are ear shells, sea ears, and, rarely, muttonfish or mut ...
from rocks or breaking open shells), with some populations spending 21% of their time making tools. Other tool use, such as
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s using twigs to "fish" for termites, may be developed by watching others use tools and may even be a true example of animal teaching. Tools may even be used in solving puzzles in which the animal appears to experience a "Eureka moment". Other mammals that do not use tools, such as dogs, can also experience a Eureka moment.
Brain size The size of the brain is a frequent topic of study within the fields of anatomy, biological anthropology, animal science and evolution. Brain size is sometimes measured by weight and sometimes by volume (via MRI scans or by skull volume). Neu ...
was previously considered a major indicator of the intelligence of an animal. Since most of the brain is used for maintaining bodily functions, greater ratios of brain to body mass may increase the amount of brain mass available for more complex cognitive tasks.
Allometric Allometry is the study of the relationship of body size to shape, anatomy, physiology and finally behaviour, first outlined by Otto Snell in 1892, by D'Arcy Thompson in 1917 in '' On Growth and Form'' and by Julian Huxley in 1932. Overview Allo ...
analysis indicates that mammalian brain size scales at approximately the or exponent of the body mass. Comparison of a particular animal's brain size with the expected brain size based on such allometric analysis provides an encephalisation quotient that can be used as another indication of animal intelligence.
Sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the genus '' Physeter'' and one of three extant species in the sperm whale fam ...
s have the largest brain mass of any animal on earth, averaging and in mature males.
Self-awareness In philosophy of self, self-awareness is the experience of one's own personality or individuality. It is not to be confused with consciousness in the sense of qualia. While consciousness is being aware of one's environment and body and lifest ...
appears to be a sign of abstract thinking. Self-awareness, although not well-defined, is believed to be a precursor to more advanced processes such as metacognitive reasoning. The traditional method for measuring this is the
mirror test The mirror test—sometimes called the mark test, mirror self-recognition (MSR) test, red spot technique, or rouge test—is a behavioral technique developed in 1970 by American psychologist Gordon Gallup Jr. as an attempt to determine whether a ...
, which determines if an animal possesses the ability of self-recognition. Mammals that have passed the mirror test include Asian elephants (some pass, some do not); chimpanzees;
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and less often the dwarf chimpanzee or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus '' Pan,'' the other being the co ...
s;
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the gen ...
s; humans, from 18 months (
mirror stage The mirror stage (french: stade du miroir) is a concept in the psychoanalytic theory of Jacques Lacan. The mirror stage is based on the belief that infants recognize themselves in a mirror (literal) or other symbolic contraption which induces apper ...
); bottlenose dolphins killer whales; and false killer whales.


Social structure

Eusociality Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping gen ...
is the highest level of social organization. These societies have an overlap of adult generations, the division of reproductive labor and cooperative caring of young. Usually insects, such as
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfa ...
s, ants and termites, have eusocial behavior, but it is demonstrated in two rodent species: the naked mole-rat and the Damaraland mole-rat. Presociality is when animals exhibit more than just sexual interactions with members of the same species, but fall short of qualifying as eusocial. That is, presocial animals can display communal living, cooperative care of young, or primitive division of reproductive labor, but they do not display all of the three essential traits of eusocial animals. Humans and some species of
Callitrichidae The Callitrichidae (also called Arctopitheci or Hapalidae) are a family of New World monkeys, including marmosets, tamarins, and lion tamarins. At times, this group of animals has been regarded as a subfamily, called the Callitrichinae, of th ...
(
marmoset The marmosets (), also known as zaris or sagoin, are 22 New World monkey species of the genera '' Callithrix'', '' Cebuella'', '' Callibella'', and '' Mico''. All four genera are part of the biological family Callitrichidae. The term "marmoset ...
s and
tamarin The tamarins are squirrel-sized New World monkeys from the family Callitrichidae in the genus ''Saguinus''. They are the first offshoot in the Callitrichidae tree, and therefore are the sister group of a clade formed by the lion tamarins, Goe ...
s) are unique among primates in their degree of cooperative care of young. Harry Harlow set up an experiment with
rhesus monkey The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies that are split between two groups, the Chinese-derived and the Indian-derived. Generally b ...
s, presocial primates, in 1958; the results from this study showed that social encounters are necessary in order for the young monkeys to develop both mentally and sexually. A fission-fusion society is a society that changes frequently in its size and composition, making up a permanent social group called the "parent group". Permanent social networks consist of all individual members of a community and often varies to track changes in their environment. In a fission–fusion society, the main parent group can fracture (fission) into smaller stable subgroups or individuals to adapt to
environmental A biophysical environment is a biotic and abiotic surrounding of an organism or population, and consequently includes the factors that have an influence in their survival, development, and evolution. A biophysical environment can vary in scale f ...
or social circumstances. For example, a number of males may break off from the main group in order to hunt or forage for food during the day, but at night they may return to join (fusion) the primary group to share food and partake in other activities. Many mammals exhibit this, such as primates (for example orangutans and
spider monkey Spider monkeys are New World monkeys belonging to the genus ''Ateles'', part of the subfamily Atelinae, family Atelidae. Like other atelines, they are found in tropical forests of Central and South America, from southern Mexico to Brazil. T ...
s), elephants, spotted hyenas, lions, and dolphins. Solitary animals defend a territory and avoid social interactions with the members of its species, except during breeding season. This is to avoid resource competition, as two individuals of the same species would occupy the same niche, and to prevent depletion of food. A solitary animal, while foraging, can also be less conspicuous to predators or prey. In a
hierarchy A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
, individuals are either dominant or submissive. A despotic hierarchy is where one individual is dominant while the others are submissive, as in wolves and lemurs, and a
pecking order In biology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social groups interact, creating a ranking system. A dominant higher-ranking individual is som ...
is a linear ranking of individuals where there is a top individual and a bottom individual. Pecking orders may also be ranked by sex, where the lowest individual of a sex has a higher ranking than the top individual of the other sex, as in hyenas. Dominant individuals, or alphas, have a high chance of reproductive success, especially in
harems Harem ( Persian: حرمسرا ''haramsarā'', ar, حَرِيمٌ ''ḥarīm'', "a sacred inviolable place; harem; female members of the family") refers to domestic spaces that are reserved for the women of the house in a Muslim family. A har ...
where one or a few males (resident males) have exclusive breeding rights to females in a group. Non-resident males can also be accepted in harems, but some species, such as the
common vampire bat The common vampire bat (''Desmodus rotundus'') is a small, leaf-nosed bat native to Latin America. It is one of three extant species of vampire bat, the other two being the hairy-legged and the white-winged vampire bats. The common vampire ba ...
(''Desmodus rotundus''), may be more strict. Some mammals are perfectly
monogamous Monogamy ( ) is a form of dyadic relationship in which an individual has only one partner during their lifetime. Alternately, only one partner at any one time ( serial monogamy) — as compared to the various forms of non-monogamy (e.g., po ...
, meaning that they mate for life and take no other partners (even after the original mate's death), as with wolves,
Eurasian beaver The Eurasian beaver (''Castor fiber'') or European beaver is a beaver species that was once widespread in Eurasia, but was hunted to near-extinction for both its fur and castoreum. At the turn of the 20th century, only about 1,200 beavers surv ...
s, and otters. There are three types of polygamy: either one or multiple dominant males have breeding rights (
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
), multiple males that females mate with (polyandry), or multiple males have exclusive relations with multiple females (
polygynandry Polygynandry is a mating system in which both males and females have multiple mating partners during a breeding season. In sexually reproducing diploid animals, different mating strategies are employed by males and females, because the cost of ga ...
). It is much more common for polygynous mating to happen, which, excluding
leks A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an avail ...
, are estimated to occur in up to 90% of mammals. Lek mating occurs when males congregate around females and try to attract them with various
courtship display A courtship display is a set of display behaviors in which an animal, usually a male, attempts to attract a mate; the mate exercises choice, so sexual selection acts on the display. These behaviors often include ritualized movement ("dances"), ...
s and vocalizations, as in harbor seals. All higher mammals (excluding monotremes) share two major adaptations for care of the young: live birth and lactation. These imply a group-wide choice of a degree of
parental care Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal k ...
. They may build nests and dig burrows to raise their young in, or feed and guard them often for a prolonged period of time. Many mammals are K-selected, and invest more time and energy into their young than do
r-selected In ecology, ''r''/''K'' selection theory relates to the selection of combinations of traits in an organism that trade off between quantity and quality of offspring. The focus on either an increased quantity of offspring at the expense of indivi ...
animals. When two animals mate, they both share an interest in the success of the offspring, though often to different extremes. Mammalian females exhibit some degree of maternal aggression, another example of parental care, which may be targeted against other females of the species or the young of other females; however, some mammals may "aunt" the infants of other females, and care for them. Mammalian males may play a role in child rearing, as with tenrecs, however this varies species to species, even within the same genus. For example, the males of the
southern pig-tailed macaque The southern pig-tailed macaque (''Macaca nemestrina''), also known as the Sundaland pig-tailed macaque and Sunda pig-tailed macaque, is a medium-sized macaque that lives in southern Thailand, Malaysia, and Indonesia. It is known locally as ber ...
(''Macaca nemestrina'') do not participate in child care, whereas the males of the Japanese macaque (''M. fuscata'') do.


Humans and other mammals


In human culture

Non-human mammals play a wide variety of roles in human culture. They are the most popular of
pet A pet, or companion animal, is an animal kept primarily for a person's company or entertainment rather than as a working animal, livestock, or a laboratory animal. Popular pets are often considered to have attractive appearances, intelligence ...
s, with tens of millions of dogs, cats and other animals including
rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit ...
s and mice kept by families around the world. Mammals such as
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks and, ...
s, horses and deer are among the earliest subjects of art, being found in
Upper Paleolithic The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories coi ...
cave painting In archaeology, Cave paintings are a type of parietal art (which category also includes petroglyphs, or engravings), found on the wall or ceilings of caves. The term usually implies prehistoric origin, and the oldest known are more than 40,000 ...
s such as at
Lascaux Lascaux ( , ; french: Grotte de Lascaux , "Lascaux Cave") is a network of caves near the village of Montignac, in the department of Dordogne in southwestern France. Over 600 parietal wall paintings cover the interior walls and ceilings of ...
. Major artists such as
Albrecht Dürer Albrecht Dürer (; ; hu, Ajtósi Adalbert; 21 May 1471 – 6 April 1528),Müller, Peter O. (1993) ''Substantiv-Derivation in Den Schriften Albrecht Dürers'', Walter de Gruyter. . sometimes spelled in English as Durer (without an umlaut) or Due ...
,
George Stubbs George Stubbs (25 August 1724 – 10 July 1806) was an English painter, best known for his paintings of horses. Self-trained, Stubbs learnt his skills independently from other great artists of the 18th century such as Reynolds or Gainsborou ...
and Edwin Landseer are known for their portraits of mammals. Many species of mammals have been hunted for sport and for food; deer and
wild boar The wild boar (''Sus scrofa''), also known as the wild swine, common wild pig, Eurasian wild pig, or simply wild pig, is a suid native to much of Eurasia and North Africa, and has been introduced to the Americas and Oceania. The species is ...
are especially popular as game animals. Mammals such as
horses The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million y ...
and
dogs The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. ...
are widely raced for sport, often combined with betting on the outcome. There is a tension between the role of animals as companions to humans, and their existence as individuals with rights of their own. Mammals further play a wide variety of roles in literature, film, mythology, and religion.


Uses and importance

The domestication of mammals was instrumental in the Neolithic development of agriculture and of
civilization A civilization (or civilisation) is any complex society characterized by the development of a state, social stratification, urbanization, and symbolic systems of communication beyond natural spoken language (namely, a writing system). ...
, causing farmers to replace
hunter-gatherer A traditional hunter-gatherer or forager is a human living an ancestrally derived lifestyle in which most or all food is obtained by foraging, that is, by gathering food from local sources, especially edible wild plants but also insects, fungi ...
s around the world. This transition from hunting and gathering to herding flocks and growing crops was a major step in human history. The new agricultural economies, based on domesticated mammals, caused "radical restructuring of human societies, worldwide alterations in biodiversity, and significant changes in the Earth's landforms and its atmosphere... momentous outcomes".
Domestic Domestic may refer to: In the home * Anything relating to the human home or family ** A domestic animal, one that has undergone domestication ** A domestic appliance, or home appliance ** A domestic partnership ** Domestic science, sometimes ca ...
mammals form a large part of the
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animals w ...
raised for
meat Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chick ...
across the world. They include (2009) around 1.4 billion
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...
, 1 billion sheep, 1 billion
domestic pig The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus s ...
s, and (1985) over 700 million rabbits. Working domestic animals including cattle and horses have been used for work and
transport Transport (in British English), or transportation (in American English), is the intentional movement of humans, animals, and goods from one location to another. Modes of transport include air, land ( rail and road), water, cable, pipe ...
from the origins of agriculture, their numbers declining with the arrival of mechanised transport and
agricultural machinery Agricultural machinery relates to the mechanical structures and devices used in farming or other agriculture. There are many types of such equipment, from hand tools and power tools to tractors and the countless kinds of farm implements that ...
. In 2004 they still provided some 80% of the power for the mainly small farms in the third world, and some 20% of the world's transport, again mainly in rural areas. In mountainous regions unsuitable for wheeled vehicles,
pack animal A pack animal, also known as a sumpter animal or beast of burden, is an individual or type of working animal used by humans as means of transporting materials by attaching them so their weight bears on the animal's back, in contrast to draft an ...
s continue to transport goods. Mammal skins provide
leather Leather is a strong, flexible and durable material obtained from the tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffalo, pigs and ho ...
for
shoe A shoe is an item of footwear intended to protect and comfort the human foot. They are often worn with a sock. Shoes are also used as an item of decoration and fashion. The design of shoes has varied enormously through time and from culture to ...
s,
clothing Clothing (also known as clothes, apparel, and attire) are items worn on the human body, body. Typically, clothing is made of fabrics or textiles, but over time it has included garments made from animal skin and other thin sheets of materials ...
and
upholstery Upholstery is the work of providing furniture, especially seats, with padding, springs, webbing, and fabric or leather covers. The word also refers to the materials used to upholster something. ''Upholstery'' comes from the Middle English ...
.
Wool Wool is the textile fibre obtained from sheep and other mammals, especially goats, rabbits, and camelids. The term may also refer to inorganic materials, such as mineral wool and glass wool, that have properties similar to animal wool. As ...
from mammals including sheep, goats and
alpaca The alpaca (''Lama pacos'') is a species of South American camelid mammal. It is similar to, and often confused with, the llama. However, alpacas are often noticeably smaller than llamas. The two animals are closely related and can successf ...
s has been used for centuries for clothing. Mammals serve a major role in science as experimental animals, both in fundamental biological research, such as in genetics, and in the development of new medicines, which must be tested exhaustively to demonstrate their
safety Safety is the state of being "safe", the condition of being protected from harm or other danger. Safety can also refer to the control of recognized hazards in order to achieve an acceptable level of risk. Meanings There are two slightly dif ...
. Millions of mammals, especially mice and rats, are used in
experiments An experiment is a procedure carried out to support or refute a hypothesis, or determine the efficacy or likelihood of something previously untried. Experiments provide insight into cause-and-effect by demonstrating what outcome occurs when ...
each year. A knockout mouse is a genetically modified mouse with an inactivated gene, replaced or disrupted with an artificial piece of DNA. They enable the study of
sequenced In genetics and biochemistry, sequencing means to determine the primary structure (sometimes incorrectly called the primary sequence) of an unbranched biopolymer. Sequencing results in a symbolic linear depiction known as a sequence which succ ...
genes whose functions are unknown. A small percentage of the mammals are non-human primates, used in research for their similarity to humans. Despite the benefits domesticated mammals had for human development, humans have an increasingly detrimental effect on wild mammals across the world. It has been estimated that the mass of all ''wild'' mammals has declined to only 4% of all mammals, with 96% of mammals being humans and their livestock now (see figure). In fact, terrestrial wild mammals make up only 2% of all mammals.


Hybrids

Hybrids are offspring resulting from the breeding of two genetically distinct individuals, which usually will result in a high degree of heterozygosity, though hybrid and heterozygous are not synonymous. The deliberate or accidental hybridizing of two or more species of closely related animals through captive breeding is a human activity which has been in existence for millennia and has grown for economic purposes. Hybrids between different subspecies within a species (such as between the Bengal tiger and Siberian tiger) are known as intra-specific hybrids. Hybrids between different species within the same genus (such as between lions and tigers) are known as interspecific hybrids or crosses. Hybrids between different genera (such as between sheep and goats) are known as intergeneric hybrids. Natural hybrids will occur in
hybrid zone A hybrid zone exists where the ranges of two interbreeding species or diverged intraspecific lineages meet and cross-fertilize. Hybrid zones can form ''in situ'' due to the evolution of a new lineage but generally they result from secondary conta ...
s, where two populations of species within the same genera or species living in the same or adjacent areas will interbreed with each other. Some hybrids have been recognized as species, such as the red wolf (though this is controversial).
Artificial selection Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant m ...
, the deliberate selective breeding of domestic animals, is being used to breed back recently extinct animals in an attempt to achieve an animal breed with a
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological prope ...
that resembles that extinct
wildtype The wild type (WT) is the phenotype of the typical form of a species as it occurs in nature. Originally, the wild type was conceptualized as a product of the standard "normal" allele at a locus, in contrast to that produced by a non-standard, ...
ancestor. A breeding-back (intraspecific) hybrid may be very similar to the extinct wildtype in appearance, ecological niche and to some extent genetics, but the initial
gene pool The gene pool is the set of all genes, or genetic information, in any population, usually of a particular species. Description A large gene pool indicates extensive genetic diversity, which is associated with robust populations that can surv ...
of that wild type is lost forever with its
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
. As a result, bred-back breeds are at best vague look-alikes of extinct wildtypes, as
Heck cattle Heck cattle are a hardy breed of domestic cattle. These cattle are the result of an attempt by Heinz and Lutz Heck to breed back the extinct aurochs (''Bos primigenius'') from modern aurochs-derived cattle in the 1920s and 1930s. Controve ...
are of the
aurochs The aurochs (''Bos primigenius'') ( or ) is an extinct cattle species, considered to be the wild ancestor of modern domestic cattle. With a shoulder height of up to in bulls and in cows, it was one of the largest herbivores in the Holocene ...
.
Purebred Purebreds are " cultivated varieties" of an animal species achieved through the process of selective breeding. When the lineage of a purebred animal is recorded, that animal is said to be " pedigreed". Purebreds breed true-to-type which means ...
wild species evolved to a specific ecology can be threatened with extinction through the process of
genetic pollution Genetic pollution is a controversial term for uncontrolled gene flow into wild populations. It is defined as "the dispersal of contaminated altered genes from genetically engineered organisms to natural organisms, esp. by cross-pollination", but ...
, the uncontrolled hybridization,
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. In ...
genetic swamping which leads to homogenization or out-competition from the heterosic hybrid species. When new populations are imported or selectively bred by people, or when habitat modification brings previously isolated species into contact, extinction in some species, especially rare varieties, is possible.
Interbreeding In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Hybrids are not always intermediates between their parents (such as in ...
can swamp the rarer gene pool and create hybrids, depleting the purebred gene pool. For example, the endangered
wild water buffalo The wild water buffalo (''Bubalus arnee''), also called Asian buffalo, Asiatic buffalo and wild buffalo, is a large bovine native to the Indian subcontinent and Southeast Asia. It has been listed as '' Endangered'' in the IUCN Red List since 1 ...
is most threatened with extinction by genetic pollution from the domestic water buffalo. Such extinctions are not always apparent from a morphological standpoint. Some degree of
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
is a normal evolutionary process, nevertheless, hybridization threatens the existence of rare species.


Threats

The loss of species from ecological communities,
defaunation Defaunation is the global, local or functional extinction of animal populations or species from ecological communities. The growth of the human population, combined with advances in harvesting technologies, has led to more intense and efficient e ...
, is primarily driven by human activity. This has resulted in
empty forest Empty forest is a term coined by Kent H. Redford's article "The Empty Forest" (1992), which was published in ''BioScience''. An "empty forest" refers to an ecosystem that is void of large mammals. Empty forests are characterized by an otherwise e ...
s, ecological communities depleted of large vertebrates. In the
Quaternary extinction event The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecolog ...
, the mass die-off of
megafauna In terrestrial zoology, the megafauna (from Greek μέγας ''megas'' "large" and New Latin ''fauna'' "animal life") comprises the large or giant animals of an area, habitat, or geological period, extinct and/or extant. The most common threshol ...
l variety coincided with the appearance of humans, suggesting a human influence. One hypothesis is that humans hunted large mammals, such as the
woolly mammoth The woolly mammoth (''Mammuthus primigenius'') is an extinct species of mammoth that lived during the Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with '' Mammuthus subp ...
, into extinction. The 2019 '' Global Assessment Report on Biodiversity and Ecosystem Services'' by IPBES states that the total
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms b ...
of wild mammals has declined by 82 percent since the beginning of human civilization. Wild animals make up just 4% of mammalian
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms b ...
on earth, while humans and their domesticated animals make up 96%. Various species are predicted to become extinct in the near future, among them the
rhinoceros A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species ...
,
giraffes The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, '' Giraffa camelopardali ...
, and species of
primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter includ ...
and
pangolins Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smuts ...
. According to the WWF's 2020 ''
Living Planet Report The ''Living Planet Report'' is published every two years by the World Wide Fund for Nature since 1998. It is based on the Living Planet Index and ecological footprint calculations. The ''Living Planet Report'' is the world's leading, science- ...
'', vertebrate
wildlife Wildlife refers to undomesticated animal species, but has come to include all organisms that grow or live wild in an area without being introduced by humans. Wildlife was also synonymous to game: those birds and mammals that were hunted ...
populations have declined by 68% since 1970 as a result of human activities, particularly
overconsumption Overconsumption describes a situation where a consumer overuses their available goods and services to where they can't, or don't want to, replenish or reuse them. In microeconomics, this may be described as the point where the marginal cost ...
,
population growth Population growth is the increase in the number of people in a population or dispersed group. Actual global human population growth amounts to around 83 million annually, or 1.1% per year. The global population has grown from 1 billion in 1800 to ...
and
intensive farming Intensive agriculture, also known as intensive farming (as opposed to extensive farming), conventional, or industrial agriculture, is a type of agriculture, both of crop plants and of animals, with higher levels of input and output per unit of ...
, which is evidence that humans have triggered a sixth mass extinction event. Hunting alone threatens hundreds of mammalian species around the world. Scientists claim that the growing demand for
meat Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chick ...
is contributing to
biodiversity loss Biodiversity loss includes the worldwide extinction of different species, as well as the local reduction or loss of species in a certain habitat, resulting in a loss of biological diversity. The latter phenomenon can be temporary or permanent, d ...
as this is a significant driver of
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
and
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
; species-rich habitats, such as significant portions of the
Amazon rainforest The Amazon rainforest, Amazon jungle or ; es, Selva amazónica, , or usually ; french: Forêt amazonienne; nl, Amazoneregenwoud. In English, the names are sometimes capitalized further, as Amazon Rainforest, Amazon Forest, or Amazon Jungle. ...
, are being converted to agricultural land for meat production. Another influence is over-hunting and
poaching Poaching has been defined as the illegal hunting or capturing of wild animals, usually associated with land use rights. Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set a ...
, which can reduce the overall population of game animals, especially those located near villages, as in the case of
peccaries A peccary (also javelina or skunk pig) is a medium-sized, pig-like hoofed mammal of the family Tayassuidae (New World pigs). They are found throughout Central and South America, Trinidad in the Caribbean, and in the southwestern area of Nort ...
. The effects of poaching can especially be seen in the
ivory trade The ivory trade is the commercial, often illegal trade in the ivory tusks of the hippopotamus, walrus, narwhal, mammoth, and most commonly, African and Asian elephants. Ivory has been traded for hundreds of years by people in Africa and Asia ...
with African elephants. Marine mammals are at risk from entanglement from fishing gear, notably
cetaceans Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel the ...
, with discard mortalities ranging from 65,000 to 86,000 individuals annually. Attention is being given to endangered species globally, notably through the
Convention on Biological Diversity The Convention on Biological Diversity (CBD), known informally as the Biodiversity Convention, is a multilateral treaty. The Convention has three main goals: the conservation of biological diversity (or biodiversity); the sustainable use of its ...
, otherwise known as the Rio Accord, which includes 189 signatory countries that are focused on identifying endangered species and habitats. Another notable conservation organization is the IUCN, which has a membership of over 1,200 governmental and
non-governmental A non-governmental organization (NGO) or non-governmental organisation (see spelling differences) is an organization that generally is formed independent from government. They are typically nonprofit entities, and many of them are active in h ...
organizations. List of recently extinct mammals, Recent extinctions can be directly attributed to human influences. The IUCN characterizes 'recent' extinction as those that have occurred past the cut-off point of 1500, and around 80 mammal species have gone extinct since that time and 2015. Some species, such as the Père David's deer are extinct in the wild, and survive solely in captive populations. Other species, such as the Florida panther, are Ecological extinction, ecologically extinct, surviving in such low numbers that they essentially have no impact on the ecosystem. Other populations are only Local extinction, locally extinct (extirpated), still existing elsewhere, but reduced in distribution, as with the extinction of gray whales in the Atlantic Ocean, Atlantic.


Notes


See also

* List of mammal genera – living mammals * List of mammalogists * List of monotremes and marsupials * List of placental mammals * List of prehistoric mammals * List of threatened mammals of the United States * Lists of mammals by population, Lists of mammals by population size * Lists of mammals by region * Mammals described in the 2000s * Mammals in culture * Small mammal


References


Further reading

* * * * * * * * *


External links


Biodiversitymapping.org – All mammal orders in the world with distribution maps

Paleocene Mammals
a site covering the rise of the mammals, paleocene-mammals.de
Evolution of Mammals
a brief introduction to early mammals, enchantedlearning.com
European Mammal Atlas EMMA
from Societas Europaea Mammalogica, European-mammals.org
Marine Mammals of the World
– An overview of all marine mammals, including descriptions, both fully aquatic and semi-aquatic, noaa.gov
Mammalogy.org
The American Society of Mammalogists was established in 1919 for the purpose of promoting the study of mammals, and this website includes a mammal image library {{Authority control Mammals, Bathonian first appearances Extant Middle Jurassic first appearances Taxa named by Carl Linnaeus