Mammals described in 1780
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Mammals () are a group of
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
animals constituting the class Mammalia (), characterized by the presence of
mammary gland A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in pri ...
s 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. Females ...
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-modula ...
for feeding (nursing) their young, a neocortex (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, and three middle ear bones. These characteristics distinguish them from reptiles (including
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s) from which they diverged in the Carboniferous, over 300 million years ago. Around 6,400 extant
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of mammals have been described divided into 29 orders. The largest orders, in terms of number of species, are the rodents,
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 bir ...
, 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 the ...
(
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
Primate 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 including ...
s (including
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s,
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 g ...
s, monkeys, 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 ...
( cetaceans 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 poster ...
s), and the Carnivora (
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. Do ...
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 cha ...
, which reflects evolutionary history, mammals are the only living members of the
Synapsid 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 reptil ...
a (synapsids); this clade, together with Sauropsida (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 dis ...
clade. The early synapsids were sphenacodonts, a group that included the famous '' Dimetrodon''. 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 mo ...
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 cynodonts, an advanced group of therapsids, during the Late
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
-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 and Neogene periods of the Cenozoic 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 viewe ...
; but in some, the extremities are adapted for
life at sea Life is a quality that distinguishes matter that has biological processes, such as Cell signaling, signaling and self-sustaining processes, from that which does not, and is defined by the capacity for Cell growth, growth, reaction to Stimu ...
, in the air, in trees, underground, or on two legs. Mammals range in size from the
bumblebee bat Kitti's hog-nosed bat (''Craseonycteris thonglongyai''), also known as the bumblebee bat, is a near-threatened species of bat and the only extant member of the family Craseonycteridae. It occurs in western Thailand and southeast Myanmar, where i ...
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 ...
—possibly the largest animal to have ever lived. Maximum lifespan varies from two years for the shrew to 211 years for the bowhead whale. All modern mammals give birth to live young, except the five species of monotremes, 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, 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 mate ...
, which enables the feeding of the fetus during gestation. Most mammals are intelligent, with some possessing large brains, self-awareness, 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 ...
. 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 In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species) using agonistic behaviors or (less commonly) real physical aggression. A ...
,
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 Singing is the act of creating musical sounds with the voice. A person who sings is called a singer, artist or vocalist (in jazz and/or popular music). Singers perform music (arias, recitatives, songs, etc.) that can be sung with or with ...
, 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. Most mammals are polygynous, but some can be monogamous 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 wive ...
.
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. ...
of many types of mammals by humans played a major role in the Neolithic Revolution, 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 peopl ...
replacing
hunting and gathering 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, ...
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 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 hog ...
. Mammals are also hunted and raced for sport, and are used as model organisms 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 wha ...
since Paleolithic 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 ...
.


Classification

Mammal classification has been through several revisions since
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
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 *Simpso ...
(1945) provides systematics 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 cha ...
. 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, belong to the placental group. The three largest orders in numbers of species are
Rodent 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 ...
ia: mice, rats, porcupines, beavers, capybaras, 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 bir ...
: bats; and Soricomorpha:
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 Moles can refer to: * Moles de Xert, a mountain range in the Baix Maestrat comarca, Valencian Community, Spain * The Moles (Australian band) *The Moles, alter ego of Scottish band Simon Dupree and the Big Sound People *Abraham Moles, French engin ...
, 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 solen ...
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 give ...
scheme used, are the
Primate 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 including ...
s:
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 g ...
s, monkeys, 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 Madagas ...
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 poste ...
:
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 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 ...
; and the Carnivora 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. Do ...
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 b ...
s, bears, 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 mammals. It is now in its third edition, published in late 2005, ...
'', 5,416 species were identified in 2006. These were grouped into 1,229  genera, 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 The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
, 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 r ...
'' in 2018, the number of recognized mammal species is 6,495, including 96 recently extinct.


Definitions

The word " mammal" is modern, from the scientific name ''Mammalia'' coined by Carl Linnaeus in 1758, derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
'' mamma'' ("teat, pap"). In an influential 1988 paper, Timothy Rowe defined Mammalia phylogenetically as the crown group 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. The ...
of living 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 the ...
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 mono ...
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 ...
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 distinguishe ...
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 (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
, despite the fact that Triassic fossils in the Haramiyida 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'' and '' Amphitherium'' are more closely related to the therians; as fossils of all three genera are dated about in the Middle Jurassic, this is a reasonable estimate for the appearance of the crown group. T.S. Kemp has provided a more traditional definition: "
Synapsid 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 reptil ...
s 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 bone ...
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 co ...
jaw articulation and
occlusion Occlusion may refer to: Health and fitness * Occlusion (dentistry), the manner in which the upper and lower teeth come together when the mouth is closed * Occlusion miliaria, a skin condition * Occlusive dressing, an air- and water-tight trauma ...
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 tra ...
'' 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 fossi ...
s at the American Museum of Natural History. 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 validity ...
: 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 the ...
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 mono ...
* 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 longitudin ...
: multituberculates ** Infraclass † Eutriconodonta: 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 Wilso ...
: modern live-bearing mammals and their prehistoric relatives *** Superlegion † Kuehneotheria *** 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 po ...
: marsupials ***** Magnorder Australidelphia: 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 ancient ...
***** 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 opossum The family Caenolestidae contains the seven surviving species of shrew opossum: small, shrew-like marsupials that are confined to the Andes mountains of South America. The order is thought to have diverged from the ancestral marsupial line very e ...
s being closer to australidelphians. **** Cohort Placentalia: placentals ***** Magnorder
Xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. ...
: 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 stape ...
: 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 (rodents ...
****** Superorder Preptotheria ******* Grandorder Anagalida: lagomorphs,
rodent 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 ...
s 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 perc ...
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-C ...
: carnivorans, pangolins, †
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 ance ...
s and relatives ******* Grandorder
Lipotyphla Lipotyphla is a formerly used order (biology), 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 T ...
:
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, wi ...
ns ******* Grandorder Archonta:
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 ...
s,
primate 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 including ...
s,
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 fly ...
s and
treeshrew The treeshrews (or tree shrews or banxrings) are small mammals native to the tropical forests of South and Southeast Asia. They make up the entire order Scandentia, which split into two families: the Tupaiidae (19 species, "ordinary" treeshrew ...
s (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, uncertain ...
'':
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. Unlik ...
******** Mirorder Eparctocyona: †
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 waste ...
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 co ...
: South American ungulates ******** Mirorder Altungulata:
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) o ...
(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 Elephantidae ...
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 speci ...
s and hyraxes


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 through ...
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 k ...
,
Xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. ...
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 of ...
. The relationships between these three lineages is contentious, and all three possible hypotheses have been proposed with respect to which group is basal. 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 stape ...
(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 wi ...
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 ungulat ...
. 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 or mitochondrial) 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 to ...
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 during the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
. The cladogram takes Mammalia to be the crown group.


Evolution from older amniotes

The first fully terrestrial
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
s were amniotes. 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 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. 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
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s and other invertebrates as well as
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s, mosses and other plants. Within a few million years, two important amniote lineages became distinct: the
synapsid 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 reptil ...
s, which would later include the common ancestor of the mammals; and the sauropsids, 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 necked t ...
s, lizards,
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 j ...
s, crocodilians and
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s (including
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s). 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 System (stratigraphy), stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last ...
such as Dimetrodon. 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, wher ...
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 probainognathian cynodonts, 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 separ ...
. * 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 inste ...
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 some ph ...
, 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 lung ...
. 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 bone ...
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, as ...
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 archosaurs which, over an extended period (35 million years), came to include the crocodylomorphs, 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'' 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 niche from the mid-Jurassic onwards; The Jurassic '' Castorocauda'', 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 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 well ...
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 shale in China's northeastern Liaoning Province. 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 tra ...
("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'', 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 bone 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 ...
s 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 ''Ukhaatherium'' is a now extinct species of mammal that lived during the upper Cretaceous about 84 to 72 million years ago in today's East Asia. It is known above all from the fossil locality Ukhaa Tolgod, Mongolia. An adult ''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 Canin ...
. 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 dev ...
during gestation periods. A narrow pelvic outlet indicates that the young were very small at birth and therefore
pregnancy Pregnancy is the time during which one or more offspring develops (gestation, gestates) inside a woman, woman's uterus (womb). A multiple birth, multiple pregnancy involves more than one offspring, such as with twins. Pregnancy usually occur ...
was short, as in modern marsupials. This suggests that the placenta was a later development. One of the earliest known monotremes was ''
Teinolophos ''Teinolophos'' is a prehistoric species of monotreme, or egg-laying mammal, from the Teinolophidae. It is known from four specimens, each consisting of a partial lower jawbone collected from the Wonthaggi Formation at Flat Rocks, Victoria, Aus ...
'', 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)—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'', 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'' and '' Megaconus'', 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. T ...
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 coarse ...
) 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), '' T ...
'' 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 m ...
''. 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 d ...
suggest that non-mammalian synapsids 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 inste ...
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 mo ...
. 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, 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 hav ...
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 recen ...
. 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, Stormber ...
'' 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-modula ...
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, 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 the ...
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). 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'', " ...
through the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
. 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 ''pal ...
, 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 glands, 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 bone ...
(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 co ...
(a small cranial bone) meet to form the joint. In most gnathostomes, 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 mo ...
, 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 (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 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 f ...
, the incus 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 foo ...
. 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 A diphyodont is any animal with two ss of teeth, initially the ''deciduous'' set and consecutively the '' permanent'' set. Most mammals are diphyodonts—as to chew their food they need a strong, durable and complete set of teeth. Diphyodonts contr ...
) 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 Su ...
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 to the tooth's surface. *
Occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
s – 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 an ...
, 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'' (λαγ ...
). Sexual size dimorphism increases with body size across mammals ( Rensch's rule), 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 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 (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 speci ...
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 "la ...
, 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 slot ...
which has nine. All mammalian brains possess a neocortex, 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 placental m ...
, 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 bronchus. ...
, and expands the alveoli. 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, 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 sem ...
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 to t ...
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 ...
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 vei ...
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 dermis 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 macro ...
. 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) for insulation, and right whales 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 premier ...
,
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. ...
, or cilia that superficially resemble it, no animals other than mammals have hair. 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, t ...
structures that reflect their feeding preferences. For instance, frugivores (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 angiospe ...
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 ...
-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 f ...
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 poster ...
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 al ...
, 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 b ...
, 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 oma ...
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 Cud is a portion of food that returns from a ruminant's stomach to the mouth to be chewed for the second time. More precisely, it is a bolus of semi-degraded food regurgitated from the reticulorumen of a ruminant. Cud is produced during the phy ...
), 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 microbes (
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
, protozoa 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 from ...
) 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 polysacchar ...
, 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 w ...
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) o ...
, 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 wo ...
, 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 wo ...
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 wa ...
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 ...
as part of the
urea cycle The urea cycle (also known as the ornithine cycle) is a cycle of Biochemistry, 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 ...
. Bilirubin, 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 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. Cellular ...
with the help of enzymes excreted by the liver. The passing of bilirubin via bile through the intestinal tract gives mammalian feces a distinctive brown coloration. Distinctive features of the mammalian kidney include the presence of the renal pelvis 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 Cortex or cortical may refer to: Biology * Cortex (anatomy), the outermost layer of an organ ** Cerebral cortex, the outer layer of the vertebrate cerebrum, part of which is the ''forebrain'' *** Motor cortex, the regions of the cerebral cortex i ...
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, cetaceans and bears. Most adult placental mammals have no remaining trace of the cloaca. 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 c ...
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 hymen ...
, 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, a ...
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, all ...
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 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 th ...
which filters this sound. The lungs and surrounding musculature provide the air stream and pressure required to
phonate The term phonation has slightly different meanings depending on the subfield of phonetics. Among some phoneticians, ''phonation'' is the process by which the vocal folds produce certain sounds through quasi-periodic vibration. This is the defin ...
. 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). Th ...
of sound, but the strength the lungs exert to exhale 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 fold The human voice consists of sound made by a human being using the vocal tract, including talking, singing, laughing, crying, screaming, shouting, humming or yelling. The human voice frequency is specifically a part of human sound product ...
s. 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 info ...
ing and screaming. 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. There ...
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 to ...
. 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 similar ...
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 whales. 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 or ...
. 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". Th ...
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 The superior laryngeal nerve is a branch of the vagus nerve. It arises from the middle of the inferior ganglion of vagus nerve and in its course receives a branch from the superior cervical ganglion of the sympathetic nervous system. The superi ...
, 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 by ...
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 taste ...
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 immediately ...
, which controls movement, and the motor neurons 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 after reaching a certain length * Vibrissae – sensory hairs, most commonly whiskers * Pelage – guard hairs, under-fur, and awn hair * Spines – stiff guard hair used for defense (such as in porcupines) *
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 are also used to make bristles in items such as ...
s – long hairs usually used in visual signals. (such as a lion's mane) * 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. ...
– 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 amino ...
; eumelanins 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 a ...
for a range of yellowish to reddish colors, giving mammals an
earth tone Earth tone is a color scheme with multiple meanings. In its narrowest sense, it refers to "any color containing some brown" – the color of ground or soil (earth). It can also refer to "natural colors" (colors found in nature) such as brown soil, ...
. Some mammals have more vibrant colors; certain monkeys such mandrills 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 opossum Derby's woolly opossum (''Caluromys derbianus''), or the Central American woolly opossum, is an opossum found in deciduous and moist evergreen forests of Central America, from southern Mexico to western Ecuador and Colombia. It was first describ ...
s, have blue skin due to light diffraction in collagen fibers. Many sloths appear green because their fur hosts green algae; this may be a symbiotic 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 in ...
(''Alopex lagopus''),
collared lemming ''Dicrostonyx'' is a genus of rodent in the family Cricetidae The Cricetidae are a family of rodents in the large and complex superfamily Muroidea. It includes true hamsters, voles, lemmings, muskrats, and New World rats and mice. At almo ...
(''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 Conc ...
(''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, 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 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 liste ...
. 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 ani ...
, 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 lo ...
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
zebra Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: the Grévy's zebra (''Equus grevyi''), plains zebra (''E. quagga''), and the mountain zebra (''E. zebr ...
s 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 c ...
(an animal is born with either male or female genitalia, as opposed to hermaphrodites 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 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). 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 the ...
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 co ...
which is typically posterior to the penis but is often anterior in marsupials. Female mammals generally have a clitoris,
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 majo ...
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 b ...
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, 1–2 cervices 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 hymen ...
. 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 horn The uterine horns (cornua of uterus) are the points in the upper uterus where the fallopian tubes exit to meet the ovaries. They are one of the points of attachment for the round ligament of uterus (the other being the mons pubis). They also provi ...
s 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. 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 is ...
'', which produced undeveloped perinates, 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 ...
, 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 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 typical ...
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 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 anestro ...
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 to ...
. This is the plesiomorphic 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 mate ...
, 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 gland A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in pri ...
s 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
nipple The nipple is a raised region of tissue on the surface of the breast from which, in females, milk leaves the breast through the lactiferous ducts to feed an infant. The milk can flow through the nipple passively or it can be ejected by smooth m ...
s 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 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 sug ...
s.
Weaning Weaning is the process of gradually introducing an infant human or another mammal to what will be its adult diet while withdrawing the supply of its mother's milk. The process takes place only in mammals, as only mammals produce milk. The infan ...
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. ...
("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 the ...
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 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 repair ...
. 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, walking on the whole of the underside of the foot. Many mammals, such as cats and dogs, are digitigrade, 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 million yea ...
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, ca ...
, 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 ''Myrmecophag ...
s and platypuses are also knuckle-walkers. Some mammals are bipeds, 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 t ...
s. Animals will use different gaits for different speeds, terrain and situations. For example, horses show four natural gaits, the slowest horse gait 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 someti ...
, 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 po ...
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 gaits, 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 anteater The silky anteater, also known as the pygmy anteater, has traditionally been considered a single species of anteater, ''Cyclopes didactylus'', in the genus ''Cyclopes'', the only living genus in the family Cyclopedidae. Found in southern Mexico ...
s, 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 a ...
, 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 squirrels 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 l ...
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 cell Merkel cells, also known as Merkel-Ranvier cells or tactile epithelial cells, are oval-shaped mechanoreceptors essential for light touch sensation and found in the skin of vertebrates. They are abundant in highly sensitive skin like that of the f ...
s, 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 the ...
s. Many
rodent 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 ...
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 Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
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.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 are ...
s, for example, are only semi-fossorial and have very small yet functional eyes, in the fully fossorial
marsupial mole Marsupial moles, the Notoryctidae , are highly specialized marsupial mammals, known from two species found at the Australian interior. * ''Notoryctes typhlops'' (southern marsupial mole, known as the ''itjaritjari'' by the Pitjantjatjara and Yan ...
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 of the family Chrysochloridae, the golden mole Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family ...
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 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, wi ...
.


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 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. In addition to their streamlined bodies, they have smooth networks of muscle bundles in their skin that may increase laminar flow and make it easier for them to slip through water. They also lack arrector pili, so their fur can be streamlined as they swim. They rely on their fore-flippers for locomotion in a wing-like manner similar to penguins and sea turtles. 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, otters 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 of lions 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 repetition ...
of the humpback whale may be signals to females; they have different dialects in different regions of the ocean. Social vocalizations include the territorial 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 Audio frequency, frequency below the lower limit of human audibility (generally 20 Hertz, Hz). Hearing becomes gradually less sensitive as frequency decreases, ...
, 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 ...
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 t ...
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 In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species) using agonistic behaviors or (less commonly) real physical aggression. A ...
, 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 of ...
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 the ...
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 diet (and includes insectivorous diets). Other mammals, called herbivores, 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. 132 ...
(seed eating), folivory (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 (nectar eating),
gummivory A gummivore is an omnivorous animal whose diet consists primarily of the gums and saps of trees (about 90%) and bugs for protein. Notable gummivores include arboreal, terrestrial primates like certain marmosets and lemurs. These animals that live o ...
(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 stomach i ...
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 to absorb the nutrients not digested when the food was first ingested. An omnivore 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 and ...
because the
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s,
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 includ ...
s and
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...
s found in meat require little in the way of specialized digestion. Exceptions to this include baleen whales who also house gut flora 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 that ...
). 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 ce ...
. 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 of ...
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 Blatto ...
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 (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 hyenas, allowing them to consume bones; some extinct groups, notably the Machairodontinae, 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 People with the surn ...
. 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 o ...
. 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 inclu ...
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. In the period preceding hibernation, larger mammals, such as bears, become
polyphagic Polyphagia or hyperphagia is an abnormally strong, incessant sensation of hunger or desire to eat often leading to overeating. In contrast to an increase in appetite following exercise, polyphagia does not subside after eating and often leads to ...
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 squirrels can drop to , however the head and neck always stay above . A few mammals in hot environments aestivate in times of drought or extreme heat, for example the
fat-tailed dwarf lemur The fat-tailed dwarf lemur (''Cheirogaleus medius''), also known as the lesser dwarf lemur, western fat-tailed dwarf lemur, or spiny forest dwarf lemur, is endemic to Madagascar. Hibernation Recent research has shown that ''C. medius'' hibernat ...
(''Cheirogaleus medius'').


Intelligence

In intelligent mammals, such as
primate 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 including ...
s, 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'' (bandicoot ...
, 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 ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
. 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 and cognition. The sea otter uses rocks as essential and regular parts of its foraging behaviour (smashing abalone from rocks or breaking open shells), with some populations spending 21% of their time making tools. Other tool use, such as chimpanzees 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). Neur ...
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 Allome ...
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 whales have the largest brain mass of any animal on earth, averaging and in mature males. Self-awareness 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 an ...
, 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; bonobos;
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 genu ...
s;
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s, 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 appe ...
); 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 generat ...
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 bees, ants and termites, have eusocial behavior, but it is demonstrated in two rodent species: the naked mole-rat and the
Damaraland mole-rat The Damaraland mole-rat (''Fukomys damarensis''), Damara mole rat or Damaraland blesmol, is a burrowing rodent found in southern Africa. Along with the smaller, less hairy, naked mole rat, it is a species of eusocial mammal. Description Like ot ...
. 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 the ...
(
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 Harry Frederick Harlow (October 31, 1905 – December 6, 1981) was an American psychologist best known for his maternal-separation, dependency needs, and social isolation experiments on rhesus monkeys, which manifested the importance of caregi ...
set up an experiment with rhesus monkeys, 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 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. The ...
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, 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 so ...
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 bat ...
(''Desmodus rotundus''), may be more strict. Some mammals are perfectly monogamous, meaning that they
mate for life In biology, a pair bond is the strong affinity that develops in some species between a mating pair, often leading to the production and rearing of offspring and potentially a lifelong bond. Pair-bonding is a term coined in the 1940s that is freque ...
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 survi ...
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). 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 ki ...
. 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 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 ...
, 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 individ ...
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 The Japanese macaque (''Macaca fuscata''), also known as the snow monkey, is a terrestrial Old World monkey species that is native to Japan. Colloquially, they are referred to as "snow monkeys" because some live in areas where snow covers the gr ...
(''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 rabbits 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 an ...
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 coin ...
cave paintings 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,
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 Joshua Reynolds, Reynold ...
and
Edwin Landseer Sir Edwin Henry Landseer (7 March 1802 – 1 October 1873) was an English painter and sculptor, well known for his paintings of animals – particularly horses, dogs, and stags. However, his best-known works are the lion sculptures at the bas ...
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 and dogs 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-gatherers 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 c ...
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 animal ...
raised for meat 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 ma ...
, 1 billion
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to domesticated ...
, 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, 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, pipelin ...
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 ani ...
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 hog ...
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 ...
s,
clothing Clothing (also known as clothes, apparel, and attire) are items worn on the 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 natural ...
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 w ...
.
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. ...
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 success ...
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 A knockout mouse, or knock-out mouse, is a genetically modified mouse (''Mus musculus'') in which researchers have inactivated, or " knocked out", an existing gene by replacing it or disrupting it with an artificial piece of DNA. They are importa ...
is a genetically modified mouse with an inactivated
gene In biology, the word gene (from , ; "... Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
, 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 suc ...
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 The Bengal tiger is a population of the '' Panthera tigris tigris'' subspecies. It ranks among the biggest wild cats alive today. It is considered to belong to the world's charismatic megafauna. The tiger is estimated to have been present in ...
and
Siberian tiger The Siberian tiger or Amur tiger is a population of the tiger subspecies '' Panthera tigris tigris'' native to the Russian Far East, Northeast China and possibly North Korea. It once ranged throughout the Korean Peninsula, but currently inh ...
) 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 contact ...
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, the deliberate
selective breeding 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 ...
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 pr ...
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 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 are of the aurochs.
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 the ...
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 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 Bovinae, bovine native to the Indian subcontinent and Southeast Asia. It has been listed as ''Endangered species, Endangered'' i ...
is most threatened with extinction by genetic pollution from the
domestic water buffalo The water buffalo (''Bubalus bubalis''), also called the domestic water buffalo or Asian water buffalo, is a large bovid originating in the Indian subcontinent and Southeast Asia. Today, it is also found in Europe, Australia, North America, So ...
. Such extinctions are not always apparent from a morphological standpoint. Some degree of gene flow is a normal evolutionary process, nevertheless, hybridization threatens the existence of rare species.


Threats

The loss of species from ecological communities, defaunation, is primarily driven by human activity. This has resulted in empty forests, 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 megafaunal 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 s ...
, into extinction. The 2019 ''
Global Assessment Report on Biodiversity and Ecosystem Services Global means of or referring to a globe and may also refer to: Entertainment * ''Global'' (Paul van Dyk album), 2003 * ''Global'' (Bunji Garlin album), 2007 * ''Global'' (Humanoid album), 1989 * ''Global'' (Todd Rundgren album), 2015 * Bruno ...
'' by
IPBES The Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES) is an intergovernmental organization established to improve the interface between science and policy on issues of biodiversity and ecosystem services. It ...
states that the total biomass of wild mammals has declined by 82 percent since the beginning of human civilization. Wild animals make up just 4% of mammalian biomass 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 camelopardal ...
, and species of primates 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 ''Smutsia'' ...
. 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 of ...
, population growth 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 The Holocene extinction, or Anthropocene extinction, is the ongoing extinction event during the Holocene epoch. The extinctions span numerous families of bacteria, fungi, plants, and animals, including mammals, birds, reptiles, amphibians, fish, ...
event. Hunting alone threatens hundreds of mammalian species around the world. Scientists claim that the growing demand for meat 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, de ...
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 ...
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, 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 North ...
. The effects of poaching can especially be seen in the ivory trade with African elephants. Marine mammals are at risk from entanglement from fishing gear, notably cetaceans, 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 ...
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