Mammal

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This article is about the animal class. For other uses, see Mammal (disambiguation).
"Mammalian" redirects here. For the 2010 documentary film, see Mammalian (film).
"Mammalia" redirects here. For the journal, see Mammalia (journal).

Mammals

Temporal range: Late Triassic–Recent; 225 or 167–0

Ma See discussion of dates in text

PreЄ

Pg

N
 Scientific classification

Kingdom: Animalia

Phylum: Chordata

Clade: Amniota

Clade: Synapsida

Clade: Mammaliaformes

Class: Mammalia
Linnaeus, 1758
 Living subgroups

 Subclass Yinotheria
 Infraclass Australosphenida
 Order Monotremata
 Subclass Theriiformes
 Infraclass Holotheria
 Superlegion Trechnotheria
 Legion Cladotheria
 Supercohort Theria
 Cohort Marsupialia
 Cohort Placentalia

Mammals (from Latin mamma "breast") are vertebrate animals constituting

the class Mammalia (/məˈmeɪliə/), and characterized by the presence of mammary glands which
in females produce milk for feeding (nursing) their young, a neocortex(a region of the
brain), fur or hair, and three middle ear bones. These characteristics distinguish them
from reptiles and birds, from which they diverged in the late Triassic, 201–227 million years ago.
There are around 5,450 species of mammals. The largest orders are
the rodents, bats and Soricomorpha (shrews and others). The next three are
the Primates (humans, apes, monkeys, and others), the Cetartiodactyla (whales and even-toed
ungulates), and the Carnivora (cats, dogs, seals, and others).
In cladistics, which reflect evolution, mammals are classified as endothermic amniotes. They are the
only living Synapsida, which together with the Sauropsida (reptiles and birds) form
the Amniota clade. The early synapsid mammalian ancestors were sphenacodont pelycosaurs, a
group that produced the non-mammalian Dimetrodon. At the end of the Carboniferous period around
300 million years ago, this group diverged from the sauropsid line that led to today's reptiles and
birds. The line following the stem group Sphenacodontia split off several diverse groups of non-
mammalian synapsids—sometimes referred to as mammal-like reptiles—before giving rise to the
proto-mammals (Therapsida) in the early Mesozoic era. 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 among the dominant terrestrial animal groups from 66 million years ago to
the present.
The basic 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-
meter (98 ft) blue whale—the largest animal on the planet. 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 different 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 farmingreplacing 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 Palaeolithic 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.

Contents

 1Classification
o 1.1Definitions
o 1.2McKenna/Bell classification
o 1.3Molecular classification of placentals
 2Evolution
o 2.1Origins
o 2.2Evolution from amniotes
o 2.3First mammals
o 2.4Earliest appearances of features
o 2.5Rise of the mammals
 3Anatomy
o 3.1Distinguishing features
o 3.2Biological systems
o 3.3Sound production
o 3.4Fur
 3.4.1Function
 3.4.2Types
 3.4.3Thermoregulation
 3.4.4Coloration
o 3.5Reproductive system
o 3.6Endothermy
o 3.7Species lifespan
o 3.8Locomotion
 3.8.1Terrestrial
 3.8.2Arboreal
 3.8.3Aerial
 3.8.4Fossorial and subterranean
 3.8.5Aquatic
 4Behavior
o 4.1Communication and vocalization
o 4.2Feeding
o 4.3Intelligence
o 4.4Social structure
 5Humans and other mammals
 o 5.1In human culture
o 5.2Uses and importance
o 5.3Hybrids
o 5.4Threats
 6Notes
 7See also
 8References
 9Further reading
 10External links

Classification[edit]
Main article: Mammal classification
See also: List of placental mammals, List of monotremes and marsupials, List of mammal genera,
and List of mammal species

The orders Rodentia (blue), Chiroptera (red) and Soricomorpha (yellow) together make up over 70% of
mammal species.

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

Mammal classification has been through several iterations since Carl Linnaeus initially defined the
class. No classification system is universally accepted; McKenna & Bell (1997) and Wilson & Reader
(2005) provide useful recent compendiums.[1]George Gaylord Simpson's "Principles of Classification
and a Classification of Mammals" (AMNH Bulletin v. 85, 1945) provides systematics of mammal
origins and relationships that were universally taught until the end of the 20th century. Since
Simpson's classification, 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 field work gradually made Simpson's
classification outdated, it remains the closest thing to an official classification of mammals.[2]
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 including the apes, monkeys and lemurs;
the Cetartiodactyla including whales and even-toed ungulates; and the Carnivora which
includes cats, dogs, weasels, bears, seals and allies.[3] 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.[3] 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.[4] According to a
research published in the Journal of Mammalogy in 2018, the number of recognized mammal
species is 6,495 species included 96 recently extinct.[5]
Definitions [edit]
The word "mammal" is modern, from the scientific name Mammalia coined by Carl Linnaeus in 1758,
derived from the Latinmamma ("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.[6] Since this ancestor lived in
the Jurassicperiod, Rowe's definition excludes all animals from the earlier Triassic, despite the fact
that Triassic fossils in the Haramiyidahave been referred to the Mammalia since the mid-19th
century.[7] 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.[8]
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.[9] 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.[10][11]
McKenna/Bell classification[edit]
In 1997, the mammals were comprehensively revised by Malcolm C. McKenna and Susan K. Bell,
which has resulted in the McKenna/Bell classification. Their 1997 book, Classification of Mammals
above the Species Level,[12] 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 higher level groupings. The authors worked together
as paleontologists at the American Museum of Natural History, New York. 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.[2]
Extinct groups are represented by a dagger (†).
Class Mammalia
 Subclass Prototheria: monotremes: echidnas and the platypus
 Subclass Theriiformes: live-bearing mammals and their prehistoric relatives
 Infraclass †Allotheria: multituberculates
 Infraclass †Eutriconodonta: eutriconodonts
 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.[13]
 Cohort Placentalia: placentals
 Magnorder Xenarthra: xenarthrans
 Magnorder Epitheria: epitheres
 Superorder †Leptictida
 Superorder Preptotheria
 Grandorder Anagalida: lagomorphs, rodents and elephant shrews
 Grandorder Ferae: carnivorans, pangolins, †creodonts and relatives
 Grandorder Lipotyphla: insectivorans
 Grandorder Archonta: bats, primates, colugos and treeshrews
 Grandorder Ungulata: ungulates
 Order Tubulidentata incertae sedis: aardvark
 Mirorder Eparctocyona:
†condylarths, whales and artiodactyls (even-toed ungulates)
 Mirorder †Meridiungulata: South American ungulates
 Mirorder Altungulata: perissodactyls (odd-toed
ungulates), elephants, manatees and hyraxes
Molecular classification of placentals[edit]
Molecular studies based on DNA analysis have suggested new relationships among mammal
families over the last few years. Most of these findings have been independently validated
by retrotransposon presence/absence data.[14] 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 different hypotheses have been
proposed with respect to which group is basal. These hypotheses are Atlantogenata (basal
Boreoeutheria), Epitheria (basal Xenarthra) and Exafroplacentalia (basal
Afrotheria).[15] 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)[16] and
varying interpretations of paleogeographic data.[15]
Mamm  
alia Monotremata
 
  Ther  
ia Marsupialia
 
  Placentalia
   Atlantogenata  
   
Afrotheria
 
  Xenarthra

Boreoeutheria Euarchontoglires  
     

Euarchonta
 
 
Glires

Laurasiatheria  
    Eulipotyphla
Scrotifera  
    Chiroptera
  Euungulata  
     
Cetartiodactyla
 
 
Perissodactyla

Ferae  
    Pholidota
 
 
Carnivora

The cladogram above is based on Tarver et al. (2016)[17]

Group I: Superorder Afrotheria[18]

 Clade Afroinsectiphilia
 Order Macroscelidea: elephant shrews (Africa)
 Order Afrosoricida: tenrecs and golden moles (Africa)
 Order Tubulidentata: aardvark (Africa south of the Sahara)
 Clade Paenungulata
 Order Hyracoidea: hyraxes or dassies (Africa, Arabia)
 Order Proboscidea: elephants (Africa, Southeast Asia)
 Order Sirenia: dugong and manatees (cosmopolitan tropical)
Group II: Superorder Xenarthra[18]

 Order Pilosa: sloths and anteaters (neotropical)

 Order Cingulata: armadillos and extinct relatives (Americas)
Group III: Magnaorder Boreoeutheria[18]

 Superorder: Euarchontoglires (Supraprimates)

 Grandorder Euarchonta
 Order Scandentia: treeshrews (Southeast Asia).
 Order Dermoptera: flying lemurs or colugos (Southeast Asia)
 Order Primates: lemurs, bushbabies, monkeys, apes, humans (cosmopolitan)
 Grandorder Glires
 Order Lagomorpha: pikas, rabbits, hares (Eurasia, Africa, Americas)
 Order Rodentia: rodents (cosmopolitan)
 Superorder: Laurasiatheria
 Order Eulipotyphla: shrews, hedgehogs, moles, solenodons
 Clade Scrotifera
 Order Chiroptera: bats (cosmopolitan)
 Clade Fereuungulata
 Clade Ferae
 Order Pholidota: pangolins or scaly anteaters (Africa, South Asia)
 Order Carnivora: carnivores (cosmopolitan), including cats and dogs
 Clade Euungulata
 Order Cetartiodactyla: cetaceans (whales, dolphins and porpoises) and even-
toed ungulates, including pigs, cattle, deer and giraffes
 Order Perissodactyla: odd-toed ungulates,
including horses, donkeys, zebras, tapirs and rhinoceroses

Evolution[edit]
Main article: Evolution of mammals
Origins[edit]
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 reptilian
and avian lineages. Crown group mammals evolved from earlier mammaliaforms during the Early
Jurassic. The cladogram takes Mammalia to be the crown group.[19]
Mammaliaformes  
    Morganucodontidae
   
Docodonta
   
   
Haldanodon
   
Mammalia  
   
Australosphenida (incl. Monotremata)
    
Fruitafossor
   
     
      Haramiyavia
 
  Multituberculata

 
Tinodon
 
 
 
Eutriconodonta (incl. Gobiconodonta)
 
  Trechnotheria (incl. Theria)

Evolution from amniotes[edit]

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 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 reptiliomorphamphibious tetrapods,[20] 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, dinosaurs and birds.[21]Synapsids have a single hole
(temporal fenestra) low on each side of the skull. One synapsid group, the pelycosaurs, included the
largest and fiercest animals of the early Permian.[22] Nonmammalian synapsids are sometimes called
"mammal-like reptiles".[23][24]
Therapsids, a group of synapsids, descended from pelycosaurs in the Middle Permian, about 265
million years ago, and became the dominant land vertebrates.[23] 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.[23] The therapsid lineage
leading to mammals went through a series of stages, beginning with animals that were very similar
to their pelycosaur ancestors and ending with probainognathian cynodonts, some of which could
easily be mistaken for mammals. Those stages were characterized by:[25]
 The gradual development of a bony secondary palate.
 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[edit]
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.[26] In the early Triassic, most medium to large land carnivore niches were taken over
by archosaurs[27] which, over an extended period (35 million years), came to include
the crocodylomorphs,[28] the pterosaurs and the dinosaurs;[29] 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.[30]
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;[31] The Jurassic Castorocauda, for
example, was a close relative of true mammals that had adaptations for swimming, digging and
catching fish.[32] Most, if not all, are thought to have remained nocturnal (the Nocturnal bottleneck),
accounting for much of the typical mammalian traits.[33] The majority of the mammal species that
existed in the Mesozoic Era were multituberculates, eutriconodonts and spalacotheriids.[34] 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.[35]

Restoration of Juramaia sinensis, the oldest known Eutherian (160 M.Y.A.)[36]

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.[36] 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.[37] 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.[38] 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 fetusduring 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.[39]
One of the earliest known monotremes was Teinolophos, which lived about 120 million years ago in
Australia.[40] 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—
[41]
and they lay eggs which are leathery and uncalcified.[42]
Earliest appearances of features[edit]
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.[43]

Foramina in the upper jaw are not indicative of whiskers, as in the red tegu (Tupinambis rufescens).

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;[44][45] 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.[24][46] Popular sources, nevertheless, continue to attribute
whiskers to Thrinaxodon.[47] 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.[48]
When endothermy first appeared in the evolution of mammals is uncertain, though it is generally
agreed to have first evolved in non-mammalian therapsids.[48][49] Modern monotremes have lower
body temperatures and more variable metabolic rates than marsupials and placentals,[50] 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.[51] Likewise, some modern therians like afrotheres
and xenarthrans have secondarily developed lower body temperatures.[52]
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.[53] 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.[54]
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.[55][56]
Rise of the mammals[edit]
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,[57] and underwent an exponential increase in body size (megafauna).[58] Then
mammals diversified very quickly; both birds and mammals show an exponential rise in
diversity.[57] For example, the earliest known bat dates from about 50 million years ago, only 16
million years after the extinction of the dinosaurs.[59]
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.[60] However, no placental fossils have been found from before the end of the
Cretaceous.[61] The earliest undisputed fossils of placentals comes from the early Paleocene, after
the extinction of the dinosaurs.[61] In particular, scientists have identified an early Paleocene animal
named Protungulatum donnae as one of the first placental mammals.[62] however it has been
reclassified as a non-placental eutherian.[63] Recalibrations of genetic and morphological diversity
rates have suggested a Late Cretaceous origin for placentals, and a Paleocene origin for most
modern clades.[64]
The earliest known ancestor of primates is Archicebus achilles[65] from around 55 million years
ago.[65] This tiny primate weighed 20–30 grams (0.7–1.1 ounce) and could fit within a human palm.[65]

Anatomy[edit]
Distinguishing features[edit]
Living mammal species can be identified by the presence of sweat glands, including those that are
specialized to produce milk to nourish their young.[66] In classifying fossils, however, other features
must be used, since soft tissue glands and many other features are not visible in fossils.[67]
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).[43]
 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.[68]
 Tooth replacement – Teeth are replaced once or (as in toothed whales and murid rodents) not
at all, rather than being replaced continually throughout life.[69]
 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.[70]
 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.[71]
For the most part, these characteristics were not present in the Triassic ancestors of the
mammals.[72] Nearly all mammaliaforms possess an epipubic bone, the exception being modern
placentals.[73]
Biological systems[edit]
Main article: Biological system

Raccoon lungs being inflated manually

The majority of mammals have seven cervical vertebrae (bones in the neck), including bats, giraffes,
whales and humans. The exceptions are the manatee and the two-toed sloth, which have just six,
and the three-toed sloth which has nine cervical vertebrae.[74] All mammalian brains possess
a neocortex, a brain region unique to mammals.[75] Placental mammals have a corpus callosum,
unlike monotremes and marsupials.[76]
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.[77][78] This type of lung is known as a bellows lung due to its resemblance to
blacksmith bellows.[78]
The mammalian heart has four chambers, two upper atria, the receiving chambers, and two
lower ventricles, the discharging chambers.[79] 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.[80]
 Didactic models of a mammalian heart
The integumentary system 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;[81]:97 marine
mammals require a thick hypodermis (blubber) for insulation, and right whales have the thickest
blubber at 20 inches (51 cm).[82] 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, careful
examination reveals the characteristic, often in obscure parts of their bodies.[81]:61