Mammal

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From Wikipedia, the free encyclopedia

For other uses, see Mammal (disambiguation).
Several terms redirect here. For other uses, see Mammalian (film) and Mammalia (journal).

Mammals

Temporal range: Late Triassic –

Recent; 225 or 167–0

Ma See discussion of dates in text

PreꞒ

Pg

N
 Scientific classification

Kingdom: Animalia

Phylum: Chordata

Clade: Amniota

Clade: Synapsida

Clade: Mammaliaformes

Class: Mammalia
Linnaeus, 1758

Living subgroups

 Monotremata
 Theria
o Marsup
 ialia
o Placent
alia

A mammal (from Latin mamma 'breast')[1] is a vertebrate animal of the class Mammalia (/mə

ˈmeɪli.ə/). Mammals are characterized by the presence of milk-producing mammary glands for
feeding their young, a neocortex region of the brain, fur or hair, and three middle ear bones. These
characteristics distinguish them from reptiles and birds, which they diverged from in
the Carboniferous Period over 300 million years ago. Around 6,400 extant species of mammals have
been described and divided into 29 orders.
The largest orders of mammals, by number of species, are the rodents, bats,
and Eulipotyphla (including hedgehogs, moles and shrews). The next three are
the Primates (including humans, monkeys and lemurs), the even-toed
ungulates (including pigs, camels and whales), and the Carnivora (including cats, dogs and seals).
Mammals are the only living members of Synapsida; this clade, together with Sauropsida (reptiles
and birds), constitutes the larger Amniota clade. The early synapsids were sphenacodonts, a group
that included the famous Dimetrodon. The synapsids split into several diverse groups of non-
mammalian synapsids—traditionally and incorrectly referred to as mammal-like reptiles or by the
term pelycosaurs, and now known as stem mammals or protomammals—before giving rise
to therapsids during the beginning of the Middle Permian period. Mammals originated
from cynodonts, an advanced group of therapsids, during the Late Triassic-Early Jurassic. The
modern mammalian orders arose in the Paleogene and Neogene periods of the Cenozoic era, after
the extinction of non-avian dinosaurs, and have been the dominant[vague] terrestrial animal group from
66 million years ago to the present.
The basic mammalian body type is quadruped, and most mammals use their four extremities
for terrestrial locomotion; but in some, the extremities are adapted for life at sea, in the air, in
trees, underground, or on two legs. Mammals range in size from the 30–40 mm (1.2–
1.6 in) bumblebee bat to the 30 m (98 ft) 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 are egg-
laying mammals. The most species-rich group of mammals, the cohort called placentals, have
a placenta, which enables the feeding of the fetus during gestation.
Most mammals are intelligent, with some possessing large brains, self-awareness, and tool use.
Mammals can communicate and vocalize in several ways, including the production
of ultrasound, scent-marking, alarm signals, singing, and echolocation. Mammals can organize
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 of many types of mammals by humans played a major role in the Neolithic Revolution,
and resulted in farming 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 civilizations.
Domesticated mammals provided, and continue to provide, power for transport and agriculture, as
well as food (meat and dairy products), fur, and leather. Mammals are also hunted and raced for
sport, and are used as model organisms in science. Mammals have been depicted
in art since Paleolithic times, and appear in literature, film, mythology, and religion. Decline in
numbers and extinction of many mammals is primarily driven by human poaching and habitat
destruction, primarily deforestation.
Classification
Main article: Mammal classification
See also: List of placental mammals, List of monotremes and marsupials, List of mammal genera,
and List of mammal species

Over 70% of mammal species come from the orders Rodentia, rodents (blue); Chiroptera, bats (red); and Soricomorpha, shrews
(yellow).

  Rodentia   Afrosoricida
  Chiroptera   Erinaceomorpha
  Soricomorpha   Cingulata
  Primates   Peramelemorphia
  Carnivora   Scandentia
  Artiodactyla   Perissodactyla
  Diprotodontia   Macroscelidea
  Lagomorpha   Pilosa
  Didelphimorphia   Monotremata
  Cetacea   Sirenia
  Dasyuromorphia   Proboscidea

Mammal classification has been through several revisions since Carl Linnaeus initially defined the
class, and at present, no classification system is universally accepted. McKenna & Bell (1997) and
Wilson & Reeder (2005) provide useful recent compendiums. [2] Simpson (1945)
[3]
 provides systematics of mammal origins and relationships that had been taught universally until the
end of the 20th century. However, since 1945, a large amount of new and more detailed information
has gradually been found: The paleontological record has been recalibrated, and the intervening
years have seen much debate and progress concerning the theoretical underpinnings of
systematization itself, partly through the new concept of cladistics. 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.[4]
Most mammals, including the six most species-rich orders, belong to the placental group. The three
largest orders in numbers of species are Rodentia: mice, rats, porcupines, beavers, capybaras, and
other gnawing mammals; Chiroptera: bats; and Soricomorpha: shrews, moles, and solenodons. The
next three biggest orders, depending on the biological classification scheme used, are
the Primates: apes, monkeys, and lemurs; the Cetartiodactyla: whales and even-toed ungulates; and
the Carnivora which includes cats, dogs, weasels, bears, seals, and allies.[5] According to Mammal
Species of the World, 5,416 species were identified in 2006. These were grouped into 1,229 genera,
153 families and 29 orders.[5] In 2008, the International Union for Conservation of Nature (IUCN)
completed a five-year Global Mammal Assessment for its IUCN Red List, which counted
5,488 species.[6] According to research published in the Journal of Mammalogy in 2018, the number of
recognized mammal species is 6,495, including 96 recently extinct.[7]
Definitions
The word "mammal" is modern, from the scientific name Mammalia coined by Carl Linnaeus in 1758,
derived from the Latin mamma ("teat, pap"). In an influential 1988 paper, Timothy Rowe defined
Mammalia phylogenetically as the crown group of mammals, the clade consisting of the most recent
common ancestor of living monotremes (echidnas and platypuses) and Therian mammals
(marsupials and placentals) and all descendants of that ancestor. [8] Since this ancestor lived in
the Jurassic period, Rowe's definition excludes all animals from the earlier Triassic, despite the fact
that Triassic fossils in the Haramiyida have been referred to the Mammalia since the mid-19th
century.[9] 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 167 million years ago in the Middle Jurassic, this is a reasonable estimate for
the appearance of the crown group.[10]
T.S. Kemp has provided a more traditional definition: "Synapsids that possess a dentary–
squamosal 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.[11] The earliest known synapsid satisfying
Kemp's definitions is Tikitherium, dated 225 Ma, so the appearance of mammals in this broader
sense can be given this Late Triassic date.[12][13]
McKenna/Bell classification
See also: Mammal classification §  McKenna/Bell classification
In 1997, the mammals were comprehensively revised by Malcolm C. McKenna and Susan K. Bell,
which has resulted in the McKenna/Bell classification. The authors worked together
as paleontologists at the American Museum of Natural History. McKenna inherited the project from
Simpson and, with Bell, constructed a completely updated hierarchical system, covering living and
extinct taxa, that reflects the historical genealogy of Mammalia. [4] Their 1997 book, Classification of
Mammals above the Species Level,[14] is a comprehensive work on the systematics, relationships and
occurrences of all mammal taxa, living and extinct, down through the rank of genus, though molecular
genetic data challenge several of the groupings.
In the following list, extinct groups are labelled with a dagger (†).
Class Mammalia

 Subclass Prototheria: monotremes: echidnas and the platypus

 Subclass Theriiformes: live-bearing mammals and their prehistoric relatives
o Infraclass †Allotheria: multituberculates
o Infraclass †Eutriconodonta: eutriconodonts
o Infraclass Holotheria: modern live-bearing mammals and their prehistoric
relatives
 Superlegion †Kuehneotheria
 Supercohort Theria: live-bearing mammals
 Cohort Marsupialia: marsupials
  Magnorder Australidelphia: Australian marsupials
and the monito del monte
 Magnorder Ameridelphia: New World marsupials.
Now considered paraphyletic, with shrew
opossums being closer to australidelphians.[15]
 Cohort Placentalia: placentals
 Magnorder Xenarthra: xenarthrans
 Magnorder Epitheria: epitheres
 Superorder †Leptictida
 Superorder Preptotheria
 Grandorder Anagalida: lago
morphs, rodents and elepha
nt shrews
 Grandorder Ferae: carnivora
ns, pangolins,
†creodonts and relatives
 Grandorder Lipotyphla: inse
ctivorans
 Grandorder Archonta: bats, 
primates, colugos and trees
hrews (now considered
paraphyletic, with bats being
closer to other groups)
 Grandorder Ungulata:
ungulates
 Order Tubulidenta
ta incertae
sedis: aardvark
 Mirorder Eparctoc
yona:
†condylarths, wha
les and artiodactyl
s (even-toed
ungulates)
 Mirorder
†Meridiungulata:
South American
ungulates
 Mirorder Altungula
ta: perissodactyls 
(odd-toed
ungulates), eleph
ants, manatees a
nd hyraxes
Molecular classification of placentals
Genus-level molecular phylogeny of 116 extant mammals inferred from the gene tree information of 14,509 coding DNA
sequences.[16] The major clades are colored: Marsupials (magenta), Xenarthrans (orange), afrotherians (red), laurasiatherians
(green), and euarchontoglires (blue).

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.[17] Classification systems based on molecular studies
reveal three major groups or lineages of placental mammals—
Afrotheria, Xenarthra and Boreoeutheria—which diverged in the Cretaceous. 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).[18] 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)[19] and
varying interpretations of paleogeographic data.[18]
Mammali
a Monotremata 
Theria

Marsupialia 
Placentalia Atlantogenata

Afrotheria   

Xenarthra   
Boreoeutheria Euarchontoglires

Euarchonta   

Glires   
Laurasiatheria
Eulipotyphla 
The cladogram above is based on Tarver et al.. (2016)[20]

Evolution
Main article: Evolution of mammals
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
cladogram takes Mammalia to be the crown group.[21]
Mammaliaformes
Morganucodontidae 

Docodonta 

Haldanodon
Mammalia
Australosphenida (incl. Monotremata) 

Fruitafossor

Haramiyavia

Multituberculata 

Tinodon

Eutriconodonta (incl. Gobiconodonta) 

Trechnotheria (incl. Theria) 

Evolution from older amniotes

The original synapsid skull structure contains one temporal opening behind the orbitals, in a fairly low position on the skull (lower
right in this image). This opening might have assisted in containing the jaw muscles of these organisms which could have
increased their biting strength.

The first fully terrestrial vertebrates were amniotes. Like their amphibious

early tetrapod predecessors, they had lungs and limbs. Amniotic eggs, however, have internal
membranes that allow the developing embryo to breathe but keep water in. Hence, amniotes can lay
eggs on dry land, while amphibians generally need to lay their eggs in water.
The first amniotes apparently arose in the Pennsylvanian subperiod of the Carboniferous. They
descended from earlier reptiliomorph amphibious tetrapods,[22] which lived on land that was already
inhabited by insects and other invertebrates as well as ferns, mosses 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 turtles, lizards, snakes, crocodilians and dinosaurs (including birds).[23] 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 such as Dimetrodon.[24] Nonmammalian synapsids were
traditionally—and incorrectly—called "mammal-like reptiles" or pelycosaurs; we now know they were
neither reptiles nor part of reptile lineage. [25][26]
Therapsids, a group of synapsids, evolved in the Middle Permian, about 265 million years ago, and
became the dominant land vertebrates. [25] 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.[25] The therapsid lineage leading to mammals went through a series of stages,
beginning with animals that were very similar to their early synapsid ancestors and ending
with probainognathian cynodonts, some of which could easily be mistaken for mammals. Those
stages were characterized by:[27]

 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 [28].
 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. [29] In
the early Triassic, most medium to large land carnivore niches were taken over
by archosaurs[30] which, over an extended period (35 million years), came to include
the crocodylomorphs,[31] the pterosaurs and the dinosaurs;[32] 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.[33]
The first mammals (in Kemp's sense) appeared in the Late Triassic epoch (about 225 million years
ago), 40 million years after the first therapsids. They expanded out of their nocturnal insectivore niche
from the mid-Jurassic onwards;[34] The Jurassic Castorocauda, for example, was a close relative of
true mammals that had adaptations for swimming, digging and catching fish. [35] Most, if not all, are
thought to have remained nocturnal (the nocturnal bottleneck), accounting for much of the typical
mammalian traits.[36] The majority of the mammal species that existed in the Mesozoic Era were
multituberculates, eutriconodonts and spalacotheriids.[37] The earliest
known metatherian is Sinodelphys, found in 125 million-year-old Early Cretaceous shale in China's
northeastern Liaoning Province. The fossil is nearly complete and includes tufts of fur and imprints of
soft tissues.[38]

Restoration of Juramaia sinensis, the oldest known Eutherian (160 M.Y.A.)[39]

The oldest known fossil among the Eutheria ("true beasts") is the small shrewlike Juramaia sinensis,
or "Jurassic mother from China", dated to 160 million years ago in the late Jurassic.[39] 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.[40] 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.[41] 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 during gestation periods. A narrow pelvic outlet indicates that the young were
very small at birth and therefore pregnancy was short, as in modern marsupials. This suggests that
the placenta was a later development. [42]
One of the earliest known monotremes was Teinolophos, which lived about 120 million years ago in
Australia.[43] Monotremes have some features which may be inherited from the original amniotes such
as the same orifice to urinate, defecate and reproduce (cloaca)—as lizards and birds also do—[44] and
they lay eggs which are leathery and uncalcified.[45]
Earliest appearances of features
Hadrocodium, whose fossils date from approximately 195 million years ago, in the early Jurassic,
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. [46]
Fossil of Thrinaxodon at the National Museum of Natural History

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;[47][48] 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.[26][49] Popular sources, nevertheless, continue to attribute
whiskers to Thrinaxodon.[50] Studies on Permian coprolites suggest that non-mammalian synapsids of
the epoch already had fur, setting the evolution of hairs possibly as far back as dicynodonts.[51]
When endothermy first appeared in the evolution of mammals is uncertain, though it is generally
agreed to have first evolved in non-mammalian therapsids.[51][52] Modern monotremes have lower body
temperatures and more variable metabolic rates than marsupials and placentals, [53] 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. [54] Likewise, some modern therians like afrotheres
and xenarthrans have secondarily developed lower body temperatures. [55]
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.[56] Epipubic bones, a feature that strongly influenced the reproduction of most
mammal clades, are first found in Tritylodontidae, suggesting that it is a synapomorphy between them
and mammaliformes. They are omnipresent in non-placental mammaliformes,
though Megazostrodon and Erythrotherium appear to have lacked them.[57]
It has been suggested that the original function of lactation (milk production) was to keep eggs moist.
Much of the argument is based on monotremes, the egg-laying mammals. [58][59] In human females,
mammary glands become fully developed during puberty, regardless of pregnancy. [60]
Rise of the mammals
Therian mammals took over the medium- to large-sized ecological niches in the Cenozoic, after
the Cretaceous–Paleogene extinction event 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,[61] and underwent an exponential increase in body size (megafauna).[62] Then
mammals diversified very quickly; both birds and mammals show an exponential rise in diversity.
[61]
 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.[63]
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 through
the Miocene.[64] However, no placental fossils have been found from before the end of the Cretaceous.
[65]
 The earliest undisputed fossils of placentals come from the early Paleocene, after the extinction of
the non-avian dinosaurs.[65] (Scientists identified an early Paleocene animal named Protungulatum
donnae as one of the first placental mammals,[66] but it has since been reclassified as a non-placental
eutherian.)[67] Recalibrations of genetic and morphological diversity rates have suggested a Late
Cretaceous origin for placentals, and a Paleocene origin for most modern clades. [68]
The earliest known ancestor of primates is Archicebus achilles[69] from around 55 million years ago.
[69]
 This tiny primate weighed 20–30 grams (0.7–1.1 ounce) and could fit within a human palm. [69]

Anatomy
Distinguishing features
Living mammal species can be identified by the presence of sweat glands, including those that are
specialized to produce milk to nourish their young.[70] In classifying fossils, however, other features
must be used, since soft tissue glands and many other features are not visible in fossils. [71]
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, 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).[46]
 Middle ear – In crown-group mammals, sound is carried from the eardrum by a chain of
three bones, the malleus, the incus 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.[72]
 Tooth replacement – Teeth can be replaced once (diphyodonty) or (as in toothed whales
and murid rodents) not at all (monophyodonty).[73] Elephants, manatees, and kangaroos
continually grow new teeth throughout their life (polyphyodonty).[74]
 Prismatic enamel – The enamel coating on the surface of a tooth consists of prisms, solid,
rod-like structures extending from the dentin to the tooth's surface.[75]
 Occipital condyles – Two knobs at the base of the skull fit into the topmost neck vertebra;
most other tetrapods, in contrast, have only one such knob.[76]
For the most part, these characteristics were not present in the Triassic ancestors of the mammals.
[77]
 Nearly all mammaliaforms possess an epipubic bone, the exception being modern placentals. [78]
Sexual dimorphism

Sexual dimorphism in aurochs, the extinct wild ancestor of cattle

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, 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. [79]
Biological systems
Main article: Biological system
Raccoon lungs being inflated manually

The majority of mammals have seven cervical vertebrae (bones in the neck). The exceptions are
the manatee and the two-toed sloth, which have six, and the three-toed sloth which has nine.[80] All
mammalian brains possess a neocortex, a brain region unique to mammals.[81] Placental brains have
a corpus callosum, unlike monotremes and marsupials.[82]
The lungs of mammals are spongy and honeycombed. Breathing is mainly achieved with
the diaphragm, which divides the thorax from the abdominal cavity, forming a dome convex to the
thorax. Contraction of the diaphragm flattens the dome, increasing the volume of the lung cavity. Air
enters through the oral and nasal cavities, and travels through the larynx, trachea and bronchi, and
expands the alveoli. Relaxing the diaphragm has the opposite effect, decreasing the volume of the
lung cavity, causing air to be pushed out of the lungs. During exercise, the abdominal wall contracts,
increasing pressure on the diaphragm, which forces air out quicker and more forcefully. The rib
cage 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.[83][84] This type of lung is known as a bellows lung due to its resemblance to
blacksmith bellows.[84]
The mammalian heart has four chambers, two upper atria, the receiving chambers, and two
lower ventricles, the discharging chambers.[85] 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.[86]
 Didactic models of a mammalian heart

Mammal skin: (1) hair, (2) epidermis, (3) sebaceous gland, (4) Arrector pili muscle, (5) dermis, (6) hair follicle, (7) sweat gland.
Not labeled, the bottom layer: hypodermis, showing round adipocytes

The integumentary system (skin) is made up of three layers: the outermost epidermis, the dermis 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, which stores lipids and provides cushioning and insulation. The
thickness of this layer varies widely from species to species; [87]: 97  marine mammals require a thick
hypodermis (blubber) for insulation, and right whales have the thickest blubber at 20 inches (51 cm).
[88]
 Although other animals have features such as whiskers, feathers, setae, or cilia that superficially
resemble it, no animals other than mammals have hair. It is a definitive characteristic of the class,
though some mammals have very little.[87]: 61 
The carnassials (teeth in the very back of the mouth) of the insectivorous aardwolf (left) vs. that of a gray wolf (right) which
consumes large vertebrates

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 specialized for grinding foliage
and seeds. Grazing animals that tend to eat hard, silica-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.
[89]
 Most carnivorous mammals have carnassialiforme teeth (of varying length depending on diet), long
canines and similar tooth replacement patterns.[90]
The stomach of even-toed ungulates (Artiodactyla) is divided into four sections: the rumen,
the reticulum, the omasum and the abomasum (only ruminants 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, protozoa and fungi)
produce cellulase, which is needed to break down the cellulose in plants.[91] Perissodactyls, in contrast
to the ruminants, store digested food that has left the stomach in an enlarged cecum, where it is
fermented by bacteria.[92] Carnivora have a simple stomach adapted to digest primarily meat, as
compared to the elaborate digestive systems of herbivorous animals, which are necessary to break
down tough, complex plant fibers. The caecum is either absent or short and simple, and the large
intestine is not sacculated or much wider than the small intestine.[93]

Bovine kidney

The mammalian excretory system involves many components. Like most other land animals,
mammals are ureotelic, and convert ammonia into urea, which is done by the liver as part of the urea
cycle.[94] Bilirubin, a waste product derived from blood cells, is passed through bile and urine with the
help of enzymes excreted by the liver.[95] The passing of bilirubin via bile through the intestinal
tract gives mammalian feces a distinctive brown coloration.[96] Distinctive features of the mammalian
kidney include the presence of the renal pelvis and renal pyramids, and of a clearly
distinguishable cortex 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.[97][98] Most adult placental mammals
have no remaining trace of the cloaca. In the embryo, the embryonic cloaca divides into a posterior
region that becomes part of the anus, and an anterior region that has different fates depending on the
sex of the individual: in females, it develops into the vestibule that receives the urethra and vagina,
while in males it forms the entirety of the penile urethra.[98] However, the tenrecs, golden moles, and
some shrews retain a cloaca as adults.[99] In marsupials, the genital tract is separate from the anus,
but a trace of the original cloaca does remain externally. [98] Monotremes, which translates
from Greek into "single hole", have a true cloaca.[100]
Sound production

A diagram of ultrasonic signals emitted by a bat, and the echo from a nearby object

As in all other tetrapods, mammals have a larynx that can quickly open and close to produce sounds,
and a supralaryngeal vocal tract 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 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 purring and screaming. Mammals can change the
position of the larynx, allowing them to breathe through the nose while swallowing through the mouth,
and to form both oral and nasal sounds; nasal sounds, such as a dog whine, are generally soft
sounds, and oral sounds, such as a dog bark, are generally loud. [101]
0:17

Beluga whale echolocation sounds

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.[102] Large vocal pads can also lower the pitch,
as in the low-pitched roars of big cats.[103] The production of infrasound is possible in some mammals
such as the African elephant (Loxodonta spp.) and baleen whales.[104][105] Small mammals with small
larynxes have the ability to produce ultrasound, which can be detected by modifications to the middle
ear and cochlea. 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.[101]
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
vocal tract is supplied by the hypoglossal nerve and facial nerves. Electrical stimulation of
the periaqueductal gray (PEG) region of the mammalian midbrain elicit vocalizations. The ability to
learn new vocalizations is only exemplified in humans, seals, cetaceans, elephants and possibly bats;
in humans, this is the result of a direct connection between the motor cortex, which controls
movement, and the motor neurons in the spinal cord.[101]
Fur
Main article: Fur

Porcupines use their spines for defense.

The primary function of the fur of mammals is thermoregulation. Others include protection, sensory
purposes, waterproofing, and camouflage.[106] Different types of fur serve different purposes: [87]: 99 

 Definitive – which may be shed after reaching a certain length

 Vibrissae – sensory hairs, most commonly whiskers
 Pelage – guard hairs, under-fur, and awn hair
 Spines – stiff guard hair used for defense (such as in porcupines)
 Bristles – long hairs usually used in visual signals. (such as a lion's mane)
 Velli – often called "down fur" which insulates newborn mammals
 Wool – long, soft and often curly
Thermoregulation
Hair length is not a factor in thermoregulation: for example, some tropical mammals such as sloths
have the same length of fur length as some arctic mammals but with less insulation; and, conversely,
other tropical mammals with short hair have the same insulating value as arctic mammals. The
denseness of fur can increase an animal's insulation value, and arctic mammals especially have
dense fur; for example, the musk ox has guard hairs measuring 30 cm (12 in) as well as a dense
underfur, which forms an airtight coat, allowing them to survive in temperatures of −40 °C (−40 °F).[87]: 
162–163 
 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 70 °C (158 °F) in the summer, but the
skin stays at 40 °C (104 °F).[87]: 188  Aquatic mammals, conversely, trap air in their fur to conserve heat by
keeping the skin dry.[87]: 162–163 
A leopard's disruptively colored coat provides camouflage for this ambush predator.

Coloration
Mammalian coats are colored for a variety of reasons, the major selective pressures
including camouflage, sexual selection, 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 colors and pheomelanin for a range of yellowish to reddish colors, giving mammals
an earth tone.[107][108] Some mammals have more vibrant colors; certain monkeys
such mandrills 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 fibers.
[109]
 Many sloths appear green because their fur hosts green algae; this may be a symbiotic relation
that affords camouflage to the sloths.[110]
Camouflage is a powerful influence in a large number of mammals, as it helps to conceal individuals
from predators or prey.[111] 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.[112] 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.[109]
Aposematism, warning off possible predators, is the most likely explanation of the black-and-white
pelage of many mammals which are able to defend themselves, such as in the foul-
smelling skunk and the powerful and aggressive honey badger.[113] Coat color is sometimes sexually
dimorphic, as in many primate species.[114] Differences in female and male coat color may indicate
nutrition and hormone levels, important in mate selection. [115] Coat color may influence the ability to
retain heat, depending on how much light is reflected. Mammals with a darker colored coat can
absorb more heat from solar radiation, and stay warmer, and some smaller mammals, such as voles,
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. [87]: 166–167 [106] The dazzling black-and-white striping
of zebras appear to provide some protection from biting flies. [116]
Reproductive system
Main article: Mammalian reproduction

Goat kids stay with their mother until they are weaned.

Mammals are solely gonochoric (an animal is born with either male or female genitalia, as opposed
to hermaphrodites where there is no such schism).[117] In male placentals, the penis is used both for
urination and copulation. Depending on the species, an erection may be fueled by blood flow into
vascular, spongy tissue or by muscular action. A penis may be contained in a prepuce when not
erect, and some placentals also have a penis bone (baculum).[118] Marsupials typically have forked
penises,[119] while the echidna penis generally has four heads with only two functioning. [120] The testes 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 clitoris, labia majora and labia minora on the
outside, while the internal system contains paired oviducts, 1–2 uteri, 1–2 cervices and a vagina.
Marsupials have two lateral vaginas and a medial vagina. The "vagina" of monotremes is better
understood as a "urogenital sinus". The uterine systems of placental mammals can vary between a
duplex, where there are two uteri and cervices which open into the vagina, a bipartite, where
two uterine horns have a single cervix that connects to the vagina, a bicornuate, which consists
where two uterine horns that are connected distally but separate medially creating a Y-shape, and a
simplex, which has a single uterus.[121][122][87]: 220–221, 247 

Matschie's tree-kangaroo with young in pouch

The ancestral condition for mammal reproduction is the birthing of relatively undeveloped, 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.[123] 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 different from that of most
other mammals.[124] In particular, the sex chromosomes of a platypus are more like those of a chicken
than those of a therian mammal.[125]
Viviparous mammals are in the subclass Theria; those living today are in the marsupial and placental
infraclasses. Marsupials have a short gestation period, typically shorter than its estrous cycle 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. This is the plesiomorphic condition among viviparous mammals; the presence of
epipubic bones in all non-placental mammals prevents the expansion of the torso needed for full
pregnancy.[78] Even non-placental eutherians probably reproduced this way. [41] The placentals give birth
to relatively complete and developed young, usually after long gestation periods. [126] They get their
name from the placenta, which connects the developing fetus to the uterine wall to allow nutrient
uptake.[127] In placental mammals, the epipubic is either completely lost or converted into the baculum;
allowing the torso to be able to expand and thus birth developed offspring. [123]
The mammary glands of mammals are specialized to produce milk, the primary source of nutrition for
newborns. The monotremes branched early from other mammals and do not have the nipples seen in
most mammals, but they do have mammary glands. The young lick the milk from a mammary patch
on the mother's belly.[128] 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.[129] Lactose is the main sugar in placental mammal milk while monotreme and
marsupial milk is dominated by oligosaccharides.[130] Weaning is the process in which a mammal
becomes less dependent on their mother's milk and more on solid food. [131]
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. [132] 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.[133] Birds are also endothermic, so endothermy is
not unique to mammals.[134]