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A mammal () is a
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
animal of the
class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...
Mammalia (). Mammals are characterised by the presence of
milk Milk is a white liquid food produced by the mammary glands of lactating mammals. It is the primary source of nutrition for young mammals (including breastfeeding, breastfed human infants) before they are able to digestion, digest solid food. ...
-producing
mammary gland A mammary gland is an exocrine gland that produces milk in humans and other mammals. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in primates (for example, human ...
s for feeding their young, a broad
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, ...
region of the brain, fur or
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and ...
, and three middle ear bones. These characteristics distinguish them from
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s and
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, from which their ancestors diverged in the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
Period over 300 million years ago. Around 6,640
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
species of mammals have been described and divided into 27 orders. The study of mammals is called mammalogy. The largest orders of mammals, by number of
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
, are the
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s, bats, and eulipotyphlans (including hedgehogs, moles and
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 dif ...
s). The next three are the
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s (including
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s, monkeys and
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
s), the even-toed ungulates (including pigs,
camel A camel (from and () from Ancient Semitic: ''gāmāl'') is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. Camels have long been domesticated and, as livestock, they provid ...
s, and
whale Whales are a widely distributed and diverse group of fully Aquatic animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...
s), and the Carnivora (including
cats The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...
, dogs, and seals). Mammals are the only living members of Synapsida; this
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, together with
Sauropsida Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
(reptiles and birds), constitutes the larger Amniota clade. Early synapsids are referred to as " pelycosaurs." The more advanced
therapsids Therapsida is a clade comprising a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including li ...
became dominant during the
Guadalupian The Guadalupian is the second and middle Series (stratigraphy), series/Epoch (geology), 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 an ...
. Mammals originated from cynodonts, an advanced group of therapsids, during the Late
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
to Early
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
. Mammals achieved their modern diversity in the
Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
and
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
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 mammalian body type is quadrupedal, with most mammals using four limbs for
terrestrial locomotion Terrestrial locomotion has evolution, evolved as animals adapted from ecoregion#Marine, aquatic to ecoregion#Terrestrial, terrestrial environments. Animal locomotion, Locomotion on land raises different problems than that in water, with reduced f ...
; but in some, the limbs are adapted for life
at sea A sea is a large body of salt water. There are particular seas and the sea. The sea commonly refers to the ocean, the interconnected body of seawaters that spans most of Earth. Particular seas are either marginal seas, second-order section ...
,
in the air IN, In or in may refer to: Dans * India (country code IN) * Indiana, United States (postal code IN) * Ingolstadt, Germany (license plate code IN) * In, Russia, a town in the Jewish Autonomous Oblast Businesses and organizations * Independen ...
, in trees or underground. The
bipeds Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) Limb (anatomy), limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from ...
have adapted to move using only the two lower limbs, while the rear limbs of
cetaceans Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
and the sea cows are mere internal vestiges. Mammals range in size from the bumblebee bat to the blue whale—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 lay eggs. The most species-rich group is the viviparous placental mammals, so named for the temporary organ ( placenta) used by offspring to draw nutrition from the mother during
gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregn ...
. Most mammals are intelligent, with some possessing large brains, self-awareness, and tool use. Mammals can communicate and vocalise in several ways, including the production of
ultrasound Ultrasound is sound with frequency, frequencies greater than 20 Hertz, kilohertz. This frequency is the approximate upper audible hearing range, limit of human hearing in healthy young adults. The physical principles of acoustic waves apply ...
, scent marking, alarm signals,
singing Singing is the art of creating music with the voice. It is the oldest form of musical expression, and the human voice can be considered the first musical instrument. The definition of singing varies across sources. Some sources define singi ...
, echolocation; and, in the case of humans, complex
language Language is a structured system of communication that consists of grammar and vocabulary. It is the primary means by which humans convey meaning, both in spoken and signed language, signed forms, and may also be conveyed through writing syste ...
. Mammals can organise themselves into fission–fusion societies, harems, and hierarchies—but can also be solitary and territorial. Most mammals are polygynous, but some can be monogamous or polyandrous.
Domestication Domestication is a multi-generational Mutualism (biology), mutualistic relationship in which an animal species, such as humans or leafcutter ants, takes over control and care of another species, such as sheep or fungi, to obtain from them a st ...
of many types of mammals by humans played a major role in the Neolithic Revolution, and resulted in
farming Agriculture encompasses crop and livestock production, aquaculture, and forestry for food and non-food products. Agriculture was a key factor in the rise of sedentary human civilization, whereby farming of domesticated species created ...
replacing hunting and gathering as the primary source of food for humans. This led to a major restructuring of human societies from nomadic to sedentary, with more co-operation among larger and larger groups, and ultimately the development of the first civilisations. Domesticated mammals provided, and continue to provide, power for transport and agriculture, as well as food (
meat Meat is animal Tissue (biology), tissue, often muscle, that is eaten as food. Humans have hunted and farmed other animals for meat since prehistory. The Neolithic Revolution allowed the domestication of vertebrates, including chickens, sheep, ...
and
dairy product Dairy products or milk products are food products made from (or containing) milk. The most common dairy animals are cow, water buffalo, goat, nanny goat, and Sheep, ewe. Dairy products include common grocery store food around the world such as y ...
s), fur, and
leather Leather is a strong, flexible and durable material obtained from the tanning (leather), tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffal ...
. Mammals are also hunted and raced for sport, kept as pets and working animals of various types, and are used as
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
s in science. Mammals have been depicted in art since
Paleolithic The Paleolithic or Palaeolithic ( years ago) ( ), also called the Old Stone Age (), is a period in human prehistory that is distinguished by the original development of stone tools, and which represents almost the entire period of human prehist ...
times, and appear in literature, film, mythology, and religion. Decline in numbers and
extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
of many mammals is primarily driven by human
poaching Poaching is 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 against the huntin ...
and habitat destruction, primarily
deforestation Deforestation or forest clearance is the removal and destruction 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. Ab ...
.


Classification

Mammal classification has been through several revisions since
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
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 (1945) provides
systematics Systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: phylogenetic trees, phylogenies). Phy ...
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 systematisation itself, partly through the new concept of
cladistics Cladistics ( ; from Ancient Greek 'branch') is an approach to Taxonomy (biology), biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesiz ...
. 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 (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
ia:
mice A mouse (: mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
, rats, porcupines, beavers, capybaras, and other gnawing mammals; Chiroptera: bats; and Eulipotyphla:
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 dif ...
s, moles, and solenodons. 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 is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s: apes, monkeys, and
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
s; Cetartiodactyla:
whale Whales are a widely distributed and diverse group of fully Aquatic animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...
s and even-toed ungulates; and Carnivora which includes cats, dogs, weasels,
bear Bears are carnivoran mammals of the family (biology), family Ursidae (). They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats ...
s, seals, and allies. According to '' Mammal Species of the World'', 5,416 species were identified in 2006. These were grouped into 1,229 
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
, 153  families and 29 orders. In 2008, the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the stat ...
(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 an inventory of the global conservation status and extinction risk of biological ...
, which counted 5,488 species. According to research published in the '' Journal of Mammalogy'' in 2018, the number of recognised mammal species is 6,495, including 96 recently extinct.


Definitions

The word "
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
" is modern, from the scientific name ''Mammalia'' coined by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in 1758, derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
'' mamma'' ("teat, pap"). In an influential 1988 paper, Timothy Rowe defined Mammalia phylogenetically as the crown group of mammals, the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
consisting of the most recent common ancestor of living monotremes ( echidnas and platypuses) and
theria Theria ( or ; ) is a scientific classification, subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the Placentalia, placental mammals) and the metatherians (including the marsupials) but excludes the egg-lay ...
ns (
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s and placentals) and all descendants of that ancestor. Since this ancestor lived in the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
period, Rowe's definition excludes all animals from the earlier
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
, 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 The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
, this is a reasonable estimate for the appearance of the crown group. T. S. Kemp has provided a more traditional definition: " Synapsids that possess a dentarysquamosal jaw articulation and occlusion between upper and lower molars with a transverse component to the movement" or, equivalently in Kemp's view, the clade originating with the last common ancestor of '' Sinoconodon'' 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 (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
date. However, this animal may have actually evolved during the Neogene.


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 presence/absence data. Classification systems based on molecular studies reveal three major groups or lineages of placentals— Afrotheria,
Xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a superorder and major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and ...
and Boreoeutheria—which diverged in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
. 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 (basal Boreoeutheria), Epitheria (basal Xenarthra) and Exafroplacentalia (basal Afrotheria). Boreoeutheria in turn contains two major lineages— Euarchontoglires and Laurasiatheria. 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. Mammal phylogeny according to Álvarez-Carretero ''et al.'', 2022:


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� ...
. The cladogram takes Mammalia to be the crown group.


Evolution from older amniotes

The first fully terrestrial
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s were
amniote Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s. Like their amphibious early
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
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
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s generally need to lay their eggs in water. The first amniotes apparently arose in the Pennsylvanian subperiod of the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
. They descended from earlier reptiliomorph amphibious tetrapods, which lived on land that was already inhabited by
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s and other invertebrates as well as
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s,
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
es and other plants. Within a few million years, two important amniote lineages became distinct: the synapsids, which would later include the common ancestor of the mammals; and the sauropsids, which now include
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s,
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s,
snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
s,
crocodilian Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s and
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s (including
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s). Synapsids have a single hole ( temporal fenestra) 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 251.902 Mya. It is the s ...
such as '' Dimetrodon''. Nonmammalian synapsids were traditionally—and incorrectly—called "mammal-like reptiles" or pelycosaurs; we now know they were neither reptiles nor part of reptile lineage. Therapsids, a group of synapsids, evolved in the Middle Permian, 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 incisors which are equal in size in therapsids, but not for eupelycosaurs. The therapsid lineage leading to mammals went through a series of stages, beginning with animals that were very similar to their early synapsid ancestors and ending with
probainognathia Probainognathia is one of the two major subgroups of the clade Eucynodontia, the other being Cynognathia. The earliest forms were carnivorous and insectivorous, though some groups eventually also evolved herbivorous diets. The earliest and most b ...
n
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
s, some of which could easily be mistaken for mammals. Those stages were characterised by: * The gradual development of a bony secondary palate. * Abrupt acquisition of endothermy among Mammaliamorpha, 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. 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 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 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 pterosaurs 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 file:Common brown robberfly with prey.jpg, A Asilidae, robber fly eating a hoverfly An insectivore is a carnivore, carnivorous animal or plant which eats insects. An alternative term is entomophage, which can also refer to the Entomophagy ...
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
spalacotheriid Spalacotheriidae is a family of extinct mammals belonging to the paraphyletic group 'Symmetrodonta'. They lasted from the Early Cretaceous to the Campanian in North America, Europe, Asia and North Africa. Spalacotheriids are characterised by hav ...
s. The earliest-known fossil of the Metatheria ("changed beasts") is '' Sinodelphys'', found in 125-million-year-old
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
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'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...
("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 bones extend forwards from the pelvis. These are not found in any modern placental, but they are found in marsupials, monotremes, other nontherian mammals and '' Ukhaatherium'', an early Cretaceous animal in the eutherian order Asioryctitheria. 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 (; : fetuses, foetuses, rarely feti or foeti) is the unborn offspring of a viviparous animal that develops from an embryo. Following the embryonic development, embryonic stage, the fetal stage of development takes place. Pren ...
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 gestation, gestates inside a woman's uterus. A multiple birth, multiple pregnancy involves more than one offspring, such as with twins. Conception (biology), Conception usually occurs ...
was short, as in modern marsupials. This suggests that the placenta was a later development. One of the earliest-known monotremes was '' Teinolophos'', which lived about 120 million years ago in Australia. Monotremes have some features which may be inherited from the original amniotes such as the same orifice to urinate, defecate and reproduce (
cloaca A cloaca ( ), : cloacae ( or ), or vent, is the rear orifice that serves as the only opening for the digestive (rectum), reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles, birds, cartilagin ...
)—as reptiles and birds also do— and they lay eggs 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 Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
, 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 maxillae and premaxillae (bones in the front of the upper jaw) of cynodonts were channels which supplied blood vessels and nerves to vibrissae ( whiskers) 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'' has foramina that are almost identical to those found in the nonmammalian cynodont '' Thrinaxodon''. Popular sources, nevertheless, continue to attribute whiskers to ''Thrinaxodon''. Studies on Permian coprolites suggest that non-mammalian
synapsids Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
of the epoch already had fur, setting the evolution of hairs possibly as far back as dicynodonts. When endothermy 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 clade comprising a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including li ...
. 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 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 Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
between them and Mammaliaformes. They are omnipresent in non-placental Mammaliaformes, though ''
Megazostrodon ''Megazostrodon'' is an extinct genus of basal mammaliaforms belonging to the order Morganucodonta. It is approximately 200 million years old.
'' and '' Erythrotherium'' 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 process ...
(
milk Milk is a white liquid food produced by the mammary glands of lactating mammals. It is the primary source of nutrition for young mammals (including breastfeeding, breastfed human infants) before they are able to digestion, digest solid food. ...
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

Therians 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 K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
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). The increase in mammalian diversity was not, however, solely because of expansion into large-bodied niches. Mammals diversified very quickly, displaying 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 ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
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 come from the early
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
, after the extinction of the non-avian dinosaurs. (Scientists identified an early Paleocene animal named '' Protungulatum donnae'' as one of the first placental mammals, but it has since 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 more recent 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 ''cre ...
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 specialised 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 (the lower jaw bone, which carries the teeth) and the squamosal (a small cranial bone) meet to form the joint. In most gnathostomes, including early
therapsids Therapsida is a clade comprising a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including li ...
, the joint consists of the articular (a small bone at the back of the lower jaw) and quadrate (a small bone at the back of the upper jaw). * Middle ear – In crown-group mammals, sound is carried from the eardrum by a chain of three bones, the malleus, the
incus The ''incus'' (: incudes) or anvil in the ear is one of three small bones (ossicles) in the middle ear. The incus receives vibrations from the malleus, to which it is connected laterally, and transmits these to the stapes medially. The incus i ...
and the stapes. Ancestrally, the malleus and the incus are derived from the articular and the quadrate bones that constituted the jaw joint of early therapsids. * Tooth replacement – Teeth can be replaced once ( diphyodonty) or (as in toothed whales and murid rodents) not at all ( monophyodonty). Elephants, manatees, and kangaroos continually grow new teeth throughout their life (
polyphyodont A polyphyodont is any animal whose tooth (animal), teeth are continually replaced. In contrast, diphyodonts are characterized by having only two successive sets of teeth. Polyphyodonts include most toothed fishes (most notably sharks), many repti ...
y). * Prismatic enamel – The enamel coating on the surface of a tooth consists of prisms, solid, rod-like structures extending from the
dentin Dentin ( ) (American English) or dentine ( or ) (British English) () is a calcified tissue (biology), tissue of the body and, along with tooth enamel, enamel, cementum, and pulp (tooth), pulp, is one of the four major components of teeth. It i ...
to the tooth's surface. * Occipital condyles – Two knobs at the base of the skull fit into the topmost neck vertebra; most other
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
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 sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, although some orders do not show any bias or are significantly female-biased ( Lagomorpha). 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 In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
(bones in the neck). The exceptions are the
manatee Manatees (, family (biology), family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivory, herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing t ...
and the two-toed sloth, which have six, and the three-toed sloth which has nine. All mammalian brains possess a
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, ...
, a brain region unique to mammals. Placental brains have a corpus callosum, unlike monotremes and marsupials.


Circulatory systems

The mammalian
heart The heart is a muscular Organ (biology), organ found in humans and other animals. This organ pumps blood through the blood vessels. The heart and blood vessels together make the circulatory system. The pumped blood carries oxygen and nutrie ...
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 in the pulmonary capillaries (blood vessels in the lungs), oxygen-rich blood returns to the left atrium via one of the four pulmonary veins. 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 arteries, arterial blood vessels of coronary circulation, which transport oxygenated blood to the Cardiac muscle, heart muscle. The heart requires a continuous supply of oxygen to function and survive, much like any ...
.


Respiratory systems

The
lung The lungs are the primary Organ (biology), organs of the respiratory system in many animals, including humans. In mammals and most other tetrapods, two lungs are located near the Vertebral column, backbone on either side of the heart. Their ...
s 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, 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 A contract is an agreement that specifies certain legally enforceable rights and obligations pertaining to two or more parties. A contract typically involves consent to transfer of goods, services, money, or promise to transfer any of thos ...
, increasing pressure on the diaphragm, which forces air out quicker and more forcefully. The rib cage 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.


Integumentary systems

The integumentary system (skin) is made up of three layers: the outermost
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
, the
dermis The dermis or corium is a layer of skin between the epidermis (skin), epidermis (with which it makes up the cutis (anatomy), cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from s ...
and the hypodermis. 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 (also known as body fat or simply fat) is a loose connective tissue composed mostly of adipocytes. It also contains the stromal vascular fraction (SVF) of cells including preadipocytes, fibroblasts, Blood vessel, vascular endothel ...
, which stores lipids and provides cushioning and insulation. The thickness of this layer varies widely from species to species;
marine mammal Marine mammals are mammals that rely on marine ecosystems for their existence. They include animals such as cetaceans, pinnipeds, sirenians, sea otters and polar bears. They are an informal group, unified only by their reliance on marine enviro ...
s require a thick hypodermis (
blubber Blubber is a thick layer of Blood vessel, vascularized adipose tissue under the skin of all cetaceans, pinnipeds, penguins, and sirenians. It was present in many marine reptiles, such as Ichthyosauria, ichthyosaurs and plesiosaurs. Description ...
) for insulation, and right whales have the thickest blubber at . Although other animals have features such as whiskers,
feather 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 an exa ...
s, setae, or
cilia The cilium (: cilia; ; in Medieval Latin and in anatomy, ''cilium'') is a short hair-like membrane protrusion from many types of eukaryotic cell. (Cilia are absent in bacteria and archaea.) The cilium has the shape of a slender threadlike proj ...
that superficially resemble it, no animals other than mammals have
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and ...
. It is a definitive characteristic of the class, though some mammals have very little.


Digestive systems

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 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 specialised for grinding foliage and
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s.
Grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to free range (roam around) and consume wild vegetations in order to feed conversion ratio, convert the otherwise indigestible (by human diges ...
animals that tend to eat hard,
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , commonly found in nature as quartz. In many parts of the world, silica is the major constituent of sand. Silica is one of the most complex and abundant f ...
-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 ...
teeth (of varying length depending on diet), long canines and similar tooth replacement patterns. The stomach of even-toed ungulates (Artiodactyla) is divided into four sections: the
rumen The rumen, also known as a paunch, is the largest stomach compartment in ruminants. The rumen and the reticulum make up the reticulorumen in ruminant animals. The diverse microbial communities in the rumen allows it to serve as the primary si ...
, the reticulum, the omasum and the abomasum (only
ruminant Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microb ...
s have a rumen). After the plant material is consumed, it is mixed with saliva in the rumen and reticulum and separates into solid and liquid material. The solids lump together to form a bolus (or cud), and is regurgitated. When the bolus enters the mouth, the fluid is squeezed out with the tongue and swallowed again. Ingested food passes to the rumen and reticulum where cellulolytic microbes (
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
, protozoa and
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
) produce cellulase, which is needed to break down the
cellulose Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
in plants. Perissodactyls, in contrast to the ruminants, store digested food that has left the stomach in an enlarged
cecum The cecum ( caecum, ; plural ceca or caeca, ) 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 (a ...
, 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 fibres. The cecum is either absent or short and simple, and the large intestine is not sacculated or much wider than the small intestine.


Excretory and genitourinary systems

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 excret ...
involves many components. Like most other land animals, mammals are ureotelic, and convert
ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
into
urea Urea, also called carbamide (because it is a diamide of carbonic acid), is an organic compound with chemical formula . This amide has two Amine, amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest am ...
, which is done by the
liver The liver is a major metabolic organ (anatomy), organ exclusively found in vertebrates, which performs many essential biological Function (biology), functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of var ...
as part of the urea cycle.
Bilirubin Bilirubin (BR) (adopted from German, originally bili—bile—plus ruber—red—from Latin) is a red-orange compound that occurs in the normcomponent of the straw-yellow color in urine. Another breakdown product, stercobilin, causes the brown ...
, a waste product derived from blood cells, is passed through
bile Bile (from Latin ''bilis''), also known as gall, is a yellow-green/misty green fluid produced by the liver of most vertebrates that aids the digestion of lipids in the small intestine. In humans, bile is primarily composed of water, is pro ...
and
urine Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...
with the help of enzymes excreted by the liver. The passing of bilirubin via bile through the intestinal tract gives mammalian
feces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
a distinctive brown coloration. Distinctive features of the mammalian kidney include the presence of the renal pelvis and renal pyramids, 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, 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 placentals have no remaining trace of the
cloaca A cloaca ( ), : cloacae ( or ), or vent, is the rear orifice that serves as the only opening for the digestive (rectum), reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles, birds, cartilagin ...
. 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 or
urogenital sinus The urogenital sinus is a body part of a human or other Placentalia, placental only present in the development of the urinary system, development of the urinary and development of the reproductive organs, reproductive organs. It is the ventral p ...
that receives the urethra and
vagina In mammals and other animals, the vagina (: vaginas or vaginae) is the elastic, muscular sex organ, reproductive organ of the female genital tract. In humans, it extends from the vulval vestibule to the cervix (neck of the uterus). The #Vag ...
, while in males it forms the entirety of the penile urethra. However, the afrosoricids 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 dif ...
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 into "single hole", have a true cloaca. Urine flows from the ureters into the cloaca in monotremes and into the bladder in placentals.


Sound production

As in all other tetrapods, mammals have a
larynx The larynx (), commonly called the voice box, is an organ (anatomy), organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal ...
that can quickly open and close to produce sounds, and a supralaryngeal
vocal tract The vocal tract is the cavity in human bodies and in animals where the sound produced at the sound source (larynx in mammals; syrinx in birds) is filtered. In birds, it consists of the trachea, the syrinx, the oral cavity, the upper part of t ...
which filters this sound. The lungs and surrounding musculature provide the air stream and pressure required to phonate. The larynx controls the pitch and
volume Volume is a measure of regions in three-dimensional space. It is often quantified numerically using SI derived units (such as the cubic metre and litre) or by various imperial or US customary units (such as the gallon, quart, cubic inch) ...
of sound, but the strength the lungs exert to exhale also contributes to volume. More primitive mammals, such as the echidna, can only hiss, as sound is achieved solely through exhaling through a partially closed larynx. Other mammals phonate using vocal folds. The movement or tenseness of the vocal folds can result in many sounds such as
purr A purr or whirr is a tonal fluttering sound made by some species of felids, including both larger, wild cats and the domestic cat (''Felis catus''), as well as two species of genets. It varies in loudness and tone among species and in the same ...
ing and
screaming A scream is a loud/hard speech production, vocalization in which air is passed through the vocal cords with greater force than is used in regular or close-distance vocalisation. This can be performed by any creature possessing lungs, including h ...
. 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 while pushing the lungs, heart, and trachea into the
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
. 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. All cats descend from the ''Felidae'' family, sharing similar musculature, c ...
s. The production of infrasound is possible in some mammals such as the African elephant (''Loxodonta'' spp.) and
baleen whale Baleen whales (), also known as whalebone whales, are marine mammals of the order (biology), parvorder Mysticeti in the infraorder Cetacea (whales, dolphins and porpoises), which use baleen plates (or "whalebone") in their mouths to sieve plankt ...
s. Small mammals with small larynxes have the ability to produce
ultrasound Ultrasound is sound with frequency, frequencies greater than 20 Hertz, kilohertz. This frequency is the approximate upper audible hearing range, limit of human hearing in healthy young adults. The physical principles of acoustic waves apply ...
, which can be detected by modifications to the middle ear 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 (cochlea), modiolus. A core component of the cochlea is the organ of Cort ...
. 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 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 in the brain, and supplied by the recurrent laryngeal nerve and the superior laryngeal nerve, branches of the
vagus nerve The vagus nerve, also known as the tenth cranial nerve (CN X), plays a crucial role in the autonomic nervous system, which is responsible for regulating involuntary functions within the human body. This nerve carries both sensory and motor fibe ...
. The vocal tract is supplied by the hypoglossal nerve 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 ta ...
s. Electrical stimulation of the periaqueductal grey (PEG) region of the mammalian midbrain elicit vocalisations. The ability to learn new vocalisations 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 involved in the planning, motor control, 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 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 ...
. Others include protection, sensory purposes, waterproofing, and camouflage. Different types of fur serve different purposes: * Definitive – which may be
shed A shed is typically a simple, single-storey (though some sheds may have two or more stories and or a loft) roofed structure, often used for storage, for hobby, hobbies, or as a workshop, and typically serving as outbuilding, such as in a bac ...
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 defence (such as in porcupines) * Bristles – 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 fiber 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 some properties similar to animal w ...
– 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 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 mammals, conversely, trap air in their fur to conserve heat by keeping the skin dry.


Coloration

Mammalian coats are coloured 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 mechanism of evolution in which members of one sex mate choice, choose mates of the other sex to mating, 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; eumelanins for brown and black colours and pheomelanin for a range of yellowish to reddish colours, giving mammals an earth tone. Some mammals have more vibrant colours; certain monkeys such
mandrill The mandrill (''Mandrillus sphinx'') is a large Old World monkey native to west central Africa. It is one of the most colorful mammals in the world, with red and blue skin on its face and posterior. The species is Sexual dimorphism, sexually ...
s and vervet monkeys, and opossums such as the Mexican mouse opossums and Derby's woolly opossums, have blue skin due to light diffraction in
collagen Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
fibres. Many sloths appear green because their fur hosts green
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
; 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 (''Alopex lagopus''), collared lemming (''Dicrostonyx groenlandicus''), stoat (''Mustela erminea''), and snowshoe hare (''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 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 comm ...
.
Aposematism Aposematism is the Advertising in biology, advertising by an animal, whether terrestrial or marine, to potential predation, predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the pr ...
, 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. Coat color is sometimes
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, 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 coloured 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 zebras appear to provide some protection from biting flies.


Reproductive system

Mammals reproduce by internal fertilisation and are solely gonochoric (an animal is born with either male or female genitalia, as opposed to
hermaphrodite A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic. The individuals of many ...
s where there is no such schism). Male mammals ejaculate
semen Semen, also known as seminal fluid, is a bodily fluid that contains spermatozoon, spermatozoa which is secreted by the male gonads (sexual glands) and other sexual organs of male or hermaphrodite, hermaphroditic animals. In humans and placen ...
during copulation through a
penis A penis (; : penises or penes) is a sex organ through which male and hermaphrodite animals expel semen during copulation (zoology), copulation, and through which male placental mammals and marsupials also Urination, urinate. The term ''pen ...
, which may be contained in a prepuce when not erect. Male placentals also urinate through a penis, and some placentals also have a penis bone (
baculum The baculum (: bacula), also known as the penis bone, penile bone, ''os penis'', ''os genitale'', or ''os priapi'', is a bone in the penis of many placental mammals. It is not present in humans, but is present in the penises of some primates, ...
). Marsupials typically have forked penises, while the echidna penis generally has four heads with only two functioning. Depending on the species, an
erection An erection (clinically: penile erection or penile tumescence) is a Physiology, physiological phenomenon in which the penis becomes firm, engorged, and enlarged. Penile erection is the result of a complex interaction of psychological, neural, ...
may be fuelled by blood flow into vascular, spongy tissue or by muscular action. The testicles of most mammals descend into the scrotum which is typically posterior to the penis but is often anterior in marsupials. Female mammals generally have a
vulva In mammals, the vulva (: vulvas or vulvae) comprises mostly external, visible structures of the female sex organ, genitalia leading into the interior of the female reproductive tract. For humans, it includes the mons pubis, labia majora, lab ...
(
clitoris In amniotes, the clitoris ( or ; : clitorises or clitorides) is a female sex organ. In humans, it is the vulva's most erogenous zone, erogenous area and generally the primary anatomical source of female Human sexuality, sexual pleasure. Th ...
and
labia The labia are the major externally visible structures of the vulva. In humans and other primates, there are two pairs of labia: the ''labia majora'' (outer lips) are large and thick folds of skin that cover the vulva's other parts, while the ''la ...
) on the outside, while the internal system contains paired oviducts, one or two uteri, one or two cervices and a
vagina In mammals and other animals, the vagina (: vaginas or vaginae) is the elastic, muscular sex organ, reproductive organ of the female genital tract. In humans, it extends from the vulval vestibule to the cervix (neck of the uterus). The #Vag ...
. Marsupials have two lateral vaginas and a medial vagina. The "vagina" of monotremes is better understood as a "urogenital sinus". The uterine systems of placentals can vary between a duplex, where there are two uteri and cervices which open into the vagina, a bipartite, where two uterine horns have a single cervix that connects to the vagina, a bicornuate, which consists where two uterine horns that are connected distally but separate medially creating a Y-shape, and a simplex, which has a single uterus. The ancestral condition for mammal reproduction is the birthing of relatively undeveloped young, either through direct vivipary 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'', which produced undeveloped perinates, but at much higher litter sizes than any modern mammal, 38 specimens. Most modern mammals are viviparous, giving birth to live young. However, the five species of monotreme, the platypus and the four species of echidna, lay eggs. The monotremes have a
sex-determination system A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. Most organisms that create their offspring using sexual reproduction have two common sexes, males and females, and in ...
different from that of most other mammals. In particular, the
sex chromosome Sex chromosomes (also referred to as allosomes, heterotypical chromosome, gonosomes, heterochromosomes, or idiochromosomes) are chromosomes that carry the genes that determine the sex of an individual. The human sex chromosomes are a typical pair ...
s of a platypus are more like those of a chicken than those of a therian mammal. Viviparous mammals are in the subclass Theria; those living today are in the marsupial and placental infraclasses. Marsupials have a short
gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregn ...
period, typically shorter than its estrous cycle 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 gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
. This is the plesiomorphic condition among viviparous mammals; the presence of epipubic bones in all non-placentals 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, which connects the developing fetus to the uterine wall to allow nutrient uptake. In placentals, 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 that produces milk in humans and other mammals. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in primates (for example, human ...
s of mammals are specialised to produce milk, the primary source of nutrition for newborns. The monotremes branched early from other mammals and do not have the teats 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 milk while monotreme and marsupial milk is dominated by
oligosaccharide An oligosaccharide (; ) is a carbohydrate, saccharide polymer containing a small number (typically three to ten) of monosaccharides (simple sugars). Oligosaccharides can have many functions including Cell–cell recognition, cell recognition and ce ...
s. Weaning is the process in which a mammal becomes less dependent on their mother's milk and more on solid food.


Endothermy

Nearly all mammals are endothermic ("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 ectothermic ("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 produces little metabolic heat, so it is considered an operational poikilotherm. 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 dif ...
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 recognise DNA double-strand breaks as well as the level of the DNA repair protein Ku80. 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 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. 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 mi ...
s are unguligrade, 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 Insectivore, insectivorous mammal native to Central America, Central and South America. It is the largest of the four living species of anteaters, which are classified with sloths in the or ...
s and platypuses are also knuckle-walkers. Some mammals are
bipeds Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) Limb (anatomy), limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from ...
, 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 Vision, Visions, or The Vision may refer to: Perception Optical perception * Visual perception, the sense of sight * Visual system, the physical mechanism of eyesight * Computer vision, a field dealing with how computers can be made to gain und ...
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 rats. 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, then there are three faster gaits which, from slowest to fastest, are the trot, the canter and the gallop. 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 as an " inverted pendulum" gait in which the body vaults o ...
and
running Running is a method of terrestrial locomotion by which humans and other animals move quickly on foot. Running is a gait with an aerial phase in which all feet are above the ground (though there are exceptions). This is in contrast to walkin ...
, but they employ many other gaits occasionally, including a four-legged crawl in tight spaces. Mammals show a vast range of
gait Gait is the pattern of Motion (physics), movement of the limb (anatomy), limbs of animals, including Gait (human), humans, during Animal locomotion, locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on s ...
s, the order that they place and lift their appendages in locomotion. Gaits can be grouped into categories according to their patterns of support sequence. For quadrupeds, there are three main categories: walking gaits, running gaits and leaping gaits. Walking is the most common gait, where some feet are on the ground at any given time, and found in almost all legged animals. Running is considered to occur when at some points in the stride all feet are off the ground in a moment of suspension.


Arboreal

Arboreal animals frequently have elongated limbs that help them cross gaps, reach fruit or other resources, test the firmness of support ahead and, in some cases, to brachiate (swing between trees). Many arboreal species, such as tree porcupines, silky anteaters, spider monkeys, and
possums Possum may refer to: Animals * Didelphimorphia, or (o)possums, an order of marsupials native to the Americas ** Didelphis, a genus of marsupials within Didelphimorphia *** Common opossum, native to Central and South America *** Virginia opossum, ...
, use prehensile tails 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. Some species of primate, bat and all species of
sloth Sloths are a Neotropical realm, Neotropical group of xenarthran mammals constituting the suborder Folivora, including the extant Arboreal locomotion, arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of move ...
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 manoeuvre more accurately and fly with more lift and less drag. By folding the wings inwards towards their body on the upstroke, they use 35% less energy during flight than birds. The membranes are delicate, ripping easily; however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly. The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cells, also found on human fingertips. These sensitive areas are different in bats, as each bump has a tiny hair in the center, making it even more sensitive and allowing the bat to detect and collect information about the air flowing over its wings, and to fly more efficiently by changing the shape of its wings in response.


Fossorial and subterranean

A fossorial (from Latin ''fossor'', meaning "digger") is an animal adapted to digging which lives primarily, but not solely, underground. Some examples are
badger Badgers are medium-sized short-legged omnivores in the superfamily Musteloidea. Badgers are a polyphyletic rather than a natural taxonomic grouping, being united by their squat bodies and adaptions for fossorial activity rather than by the ...
s, and naked mole-rats. Many
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
species are also considered fossorial because they live in burrows for most but not all of the day. Species that live exclusively underground are subterranean, and those with limited adaptations to a fossorial lifestyle sub-fossorial. Some organisms are fossorial to aid in temperature regulation while others use the underground habitat for protection from
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
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 ar ...
s, for example, are only semi-fossorial and have very small yet functional eyes, in the fully fossorial marsupial mole, the eyes are degenerated and useless, '' Talpa'' moles have vestigial eyes and the Cape golden mole 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 (biology), 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 ...
.


Aquatic

Fully aquatic mammals, the cetaceans and sirenians, 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 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, hippopotamuses,
otter Otters are carnivorous mammals in the subfamily Lutrinae. The 13 extant otter species are all semiaquatic, aquatic, or marine. Lutrinae is a branch of the Mustelidae family, which includes weasels, badgers, mink, and wolverines, among ...
s and platypuses. Hippos are very large semi-aquatic mammals, and their barrel-shaped bodies have graviportal skeletal structures, adapted to carrying their enormous weight, and their specific gravity allows them to sink and move along the bottom of a river.


Behavior


Communication and vocalisation

Many mammals communicate by vocalising. 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
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body (biology), body; a short, rounded head; round ears; and a dark, hairy tuft at the ...
s and
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or Hart (deer), hart, and a female is called a doe or hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Ir ...
. The songs of the humpback whale may be signals to females; they have different dialects in different regions of the ocean. Social vocalisations include the territorial calls of gibbons, and the use of frequency in greater spear-nosed bats to distinguish between groups. The vervet monkey 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 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, 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 stot, honestly indicating their fit condition and their ability to escape, or when
white-tailed deer The white-tailed deer (''Odocoileus virginianus''), also known Common name, commonly as the whitetail and the Virginia deer, is a medium-sized species of deer native to North America, North, Central America, Central and South America. It is the ...
and other prey mammals flag with conspicuous tail markings when alarmed, informing the predator that it has been detected. Many mammals make use of scent-marking, sometimes possibly to help defend territory, but probably with a range of functions both within and between species. Microbats and toothed whales including oceanic dolphins vocalise 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
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s, eat plants, which contain complex carbohydrates such as cellulose. An herbivorous diet includes subtypes such as granivory (seed eating), folivory (leaf eating), frugivory (fruit eating), nectarivory (nectar eating), gummivory (gum eating) and mycophagy (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 upper gastrointestinal tract of Human, humans and many other animals, including several invertebrates. The Ancient Greek name for the stomach is ''gaster'' which is used as ''gastric'' in medical t ...
or in a large cecum. Some mammals are coprophagous, consuming
feces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
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 Digestion, digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascula ...
because the
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
s,
lipid Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include storing ...
s and
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid substance with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2011): Mi ...
s found in meat require little in the way of specialised digestion. Exceptions to this include
baleen whale Baleen whales (), also known as whalebone whales, are marine mammals of the order (biology), parvorder Mysticeti in the infraorder Cetacea (whales, dolphins and porpoises), which use baleen plates (or "whalebone") in their mouths to sieve plankt ...
s 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). 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. 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 Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s or
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s). Some mammals are omnivores and display varying degrees of carnivory and herbivory, generally leaning in favour 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 sabre-shaped canines. Some physiological carnivores consume plant matter and some physiological herbivores consume meat. From a behavioural aspect, this would make them omnivores, but from the physiological standpoint, this may be due to zoopharmacognosy. 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 hairballs), aid with haemoglobin 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 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 (''Cheirogaleus medius'').


Drinking


Intelligence

In intelligent mammals, such as
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s, the
cerebrum The cerebrum (: cerebra), 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 olfac ...
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. It can be described as t ...
itself is not easy to define, but indications of intelligence include the ability to learn, matched with behavioural flexibility. Rats, 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 colonise a fresh
habitat In ecology, habitat refers to the array of resources, 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 manifestation of its ...
. 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 non-humans is a phenomenon in which a non-human animal uses any kind of tool in order to achieve a goal such as acquiring food and water, Personal grooming, grooming, combat, defence, Animal communication, communication, Play (activ ...
may indicate different levels of
learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, value (personal and cultural), values, Attitude (psychology), attitudes, and preferences. The ability to learn is possessed by humans, non-human animals, and ...
and
cognition Cognition is the "mental action or process of acquiring knowledge and understanding through thought, experience, and the senses". It encompasses all aspects of intellectual functions and processes such as: perception, attention, thought, ...
. 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
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s using twigs to "fish" for termites, may be developed by watching others use tools and may even be a true example of animal teaching. Tools may even be used in solving puzzles in which the animal appears to experience a "Eureka moment". Other mammals that do not use tools, such as dogs, can also experience a Eureka moment. Brain size 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 analysis indicates that mammalian brain size scales at approximately the or exponent of the body mass. Comparison of a particular animal's brain size with the expected brain size based on such allometric analysis provides an encephalisation quotient that can be used as another indication of animal intelligence.
Sperm whale The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the Genus (biology), genus ''Physeter'' and one of three extant species in the s ...
s 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, which determines if an animal possesses the ability of self-recognition. Mammals that have passed the mirror test include
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living ''Elephas'' species. It is the largest living land animal in Asia and the second largest living Elephantidae, elephantid in the world. It is char ...
s (some pass, some do not); chimpanzees;
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
s; orangutans; humans, from 18 months ( mirror stage); common bottlenose dolphins; orcas; and false killer whales.


Social structure

Eusociality Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
is the highest level of social organisation. These societies have an overlap of adult generations, the division of reproductive labour and cooperative caring of young. Usually insects, such as bees, ants and termites, have eusocial behaviour, but it is demonstrated in two rodent species: the naked mole-rat and the Damaraland mole-rat. Presociality is when animals exhibit more than just sexual interactions with members of the same species, but fall short of qualifying as eusocial. That is, presocial animals can display communal living, cooperative care of young, or primitive division of reproductive labour, but they do not display all of the three essential traits of eusocial animals. Humans and some species of Callitrichidae ( marmosets and tamarins) are unique among primates in their degree of cooperative care of young. Harry Harlow 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 monkeys), elephants, spotted hyenas, lions, and dolphins. Solitary animals defend a territory and avoid social interactions with the members of its species, except during breeding season. This is to avoid resource competition, as two individuals of the same species would occupy the same niche, and to prevent depletion of food. A solitary animal, while foraging, can also be less conspicuous to predators or prey. In a
hierarchy A hierarchy (from Ancient Greek, 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 ...
, 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 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 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 (''Desmodus rotundus''), may be more strict. Some mammals are perfectly monogamous, meaning that they mate for life and take no other partners (even after the original mate's death), as with wolves, Eurasian beavers, and otters. There are three types of polygamy: either one or multiple dominant males have breeding rights ( polygyny), 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, 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 displays and vocalisations, as in harbour 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. They may build nests and dig burrows to raise their young in, or feed and guard them often for a prolonged period of time. Many mammals are K-selected, and invest more time and energy into their young than do r-selected 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 (''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 g ...
(''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 pets, 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.'' They lived from the late Miocene epoch (from around 6.2 million years ago) into the Holocene until about 4,000 years ago, with mammoth species at various times inhabi ...
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 ...
cave paintings such as at
Lascaux Lascaux ( , ; , "Lascaux Cave") is a network of caves near the village of Montignac, Dordogne, Montignac, in the Departments of France, department of Dordogne in southwestern France. Over 600 Parietal art, parietal cave painting, wall paintin ...
. Major artists such as
Albrecht Dürer Albrecht Dürer ( , ;; 21 May 1471 – 6 April 1528),Müller, Peter O. (1993) ''Substantiv-Derivation in Den Schriften Albrecht Dürers'', Walter de Gruyter. . sometimes spelled in English as Durer or Duerer, was a German painter, Old master prin ...
, George Stubbs and Edwin Landseer are known for their portraits of mammals. Many species of mammals have been hunted for sport and for food; deer and wild boar 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 civilisation, causing farmers to replace
hunter-gatherer A hunter-gatherer or forager is a human living in a community, or according to an ancestrally derived Lifestyle, lifestyle, in which most or all food is obtained by foraging, that is, by gathering food from local naturally occurring sources, esp ...
s around the world. This transition from hunting and gathering to herding flocks and growing crops was a major step in human history. The new agricultural economies, based on domesticated mammals, caused "radical restructuring of human societies, worldwide alterations in biodiversity, and significant changes in the Earth's landforms and its atmosphere... momentous outcomes". Domestic mammals form a large part of the
livestock Livestock are the Domestication, domesticated animals that are raised in an Agriculture, agricultural setting to provide labour and produce diversified products for consumption such as meat, Egg as food, eggs, milk, fur, leather, and wool. The t ...
raised for
meat Meat is animal Tissue (biology), tissue, often muscle, that is eaten as food. Humans have hunted and farmed other animals for meat since prehistory. The Neolithic Revolution allowed the domestication of vertebrates, including chickens, sheep, ...
across the world. They include (2009) around 1.4 billion
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
, 1 billion sheep, 1 billion domestic pigs, 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 Motion, movement of humans, animals, and cargo, goods from one location to another. Mode of transport, Modes of transport include aviation, air, land tr ...
from the origins of agriculture, their numbers declining with the arrival of mechanised transport and
agricultural machinery Agricultural machinery relates to the machine (mechanical), mechanical structures and devices used in farming or other agriculture. There are list of agricultural machinery, many types of such equipment, from hand tools and power tools to tractor ...
. 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 a working animal used to transport goods or materials by carrying them, usually on its back. Domestic animals of many species are used in this way, among them alpacas, Bact ...
s continue to transport goods. Mammal skins provide
leather Leather is a strong, flexible and durable material obtained from the tanning (leather), tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffal ...
for
shoe A shoe is an item of footwear intended to protect and comfort the human foot. Though the human foot can adapt to varied terrains and climate conditions, it is vulnerable, and shoes provide protection. Form was originally tied to function, but ...
s,
clothing Clothing (also known as clothes, garments, dress, apparel, or attire) is any item worn on a human human body, body. Typically, clothing is made of fabrics or textiles, but over time it has included garments made from animal skin and other thin s ...
and upholstery.
Wool Wool is the textile fiber 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 some properties similar to animal w ...
from mammals including sheep, goats and
alpaca The alpaca (''Lama pacos'') is a species of South American camelid mammal. Traditionally, alpacas were kept in herds that grazed on the level heights of the Andes of Southern Peru, Western Bolivia, Ecuador, and Northern Chile. More recentl ...
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 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 The word 'safety' entered the English language in the 1 ...
. 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 Causality, cause-and-effect by demonstrating what outcome o ...
each year. A knockout mouse is a genetically modified mouse with an inactivated
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
, replaced or disrupted with an artificial piece of DNA. They enable the study of sequenced 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 hybridising 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 and the nominate tiger subspecies. It ranks among the largest wild cats alive today. It is estimated to have been present in the Indian subcontinent since the Late ...
and
Siberian tiger The Siberian tiger or Amur tiger is a population of the tiger subspecies ''Panthera tigris tigris'' native to Northeast China, the Russian Far East, and possibly North Korea. It once ranged throughout the Korea, Korean Peninsula, but currently ...
) 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 zones, 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 recognised as species, such as the red wolf (though this is controversial).
Artificial selection Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ...
, the deliberate selective breeding of domestic animals, is being used to breed back recently extinct animals in an attempt to achieve an animal breed with a
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
that resembles that extinct wildtype 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 an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
. As a result, bred-back breeds are at best vague look-alikes of extinct wildtypes, as Heck cattle are of the aurochs. Purebred wild species evolved to a specific ecology can be threatened with extinction through the process of genetic pollution, the uncontrolled hybridisation, introgression genetic swamping which leads to homogenisation 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 can swamp the rarer gene pool and create hybrids, depleting the purebred gene pool. For example, the endangered wild water buffalo is most threatened with extinction by genetic pollution from the domestic water buffalo. Such extinctions are not always apparent from a morphological standpoint. Some degree of
gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
is a normal evolutionary process, nevertheless, hybridisation 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 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, into extinction. The 2019 '' Global Assessment Report on Biodiversity and Ecosystem Services'' by IPBES states that the total
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
of wild mammals has declined by 82 per cent since the beginning of human civilisation. Wild animals make up just 4% of mammalian
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
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 ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
, giraffes, and species of
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s and pangolins. According to the WWF's 2020 '' Living Planet Report'', vertebrate
wildlife Wildlife refers to domestication, undomesticated animals and uncultivated plant species which can exist in their natural habitat, but has come to include all organisms that grow or live wilderness, wild in an area without being species, introdu ...
populations have declined by 68% since 1970 as a result of human activities, particularly
overconsumption Overconsumption describes a situation where consumers overuse their available goods and services to where they can't, or don't want to, replenish or reuse them. In microeconomics, this is the point where the marginal cost of a consumer is greater ...
,
population growth Population growth is the increase in the number of people in a population or dispersed group. The World population, global population has grown from 1 billion in 1800 to 8.2 billion in 2025. Actual global human population growth amounts to aroun ...
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 arable farming, crop plants and of Animal husbandry, animals, with higher levels ...
, which is evidence that humans have triggered a sixth mass extinction event. Hunting alone threatens hundreds of mammalian species around the world. Scientists claim that the growing demand for
meat Meat is animal Tissue (biology), tissue, often muscle, that is eaten as food. Humans have hunted and farmed other animals for meat since prehistory. The Neolithic Revolution allowed the domestication of vertebrates, including chickens, sheep, ...
is contributing to biodiversity loss as this is a significant driver of
deforestation Deforestation or forest clearance is the removal and destruction 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. Ab ...
and habitat destruction; species-rich habitats, such as significant portions of the
Amazon rainforest The Amazon rainforest, also called the Amazon jungle or Amazonia, is a Tropical and subtropical moist broadleaf forests, moist broadleaf tropical rainforest in the Amazon biome that covers most of the Amazon basin of South America. This basin ...
, are being converted to agricultural land for meat production. Another influence is over-hunting and
poaching Poaching is 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 against the huntin ...
, which can reduce the overall population of game animals, especially those located near villages, as in the case of peccaries. 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 Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
, 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, otherwise known as the Rio Accord, which includes 189 signatory countries that are focused on identifying endangered species and habitats. Another notable conservation organisation is the IUCN, which has a membership of over 1,200 governmental and non-governmental organisations. Recent extinctions can be directly attributed to human influences. The IUCN characterises '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 ecologically extinct, surviving in such low numbers that they essentially have no impact on the ecosystem. Other populations are only locally extinct (extirpated), still existing elsewhere, but reduced in distribution, as with the extinction of grey whales in the Atlantic.


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 endangered mammals * Lists of mammals by population size * Lists of mammals by region * Mammals described in the 2000s * Mammals in culture * Small mammal


Notes


References

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Further reading

* * * * * * * * *


External links


ASM Mammal Diversity Database


* [https://paleocene-mammals.de/ 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