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Mammal - W
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Mammal
A mammal (from Latin mamma 'breast')[1] is a vertebrate
Mammals
animal of the class Mammalia (/məˈmeɪli.ə/). Mammals are
characterized by the presence of milk-producing mammary Temporal range:
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.
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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
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lab work progressively outdated Simpson's classification, it remains the closest thing to an official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
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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
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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]
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Mammalia
Monotremata
Theria
Marsupialia
Placentalia Atlantogenata
Afrotheria
Xenarthra
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Boreoeutheria Euarchontoglires
Euarchonta
Glires
Laurasiatheria
Eulipotyphla
Scrotifera
Chiroptera
Euungulata
Artiodactyla
Perissodactyla
Ferae
Pholidota
Carnivora
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 cladogram takes Mammalia to
be the crown group.[21]
Mammaliaformes
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Morganucodontidae
Docodonta
Haldanodon
Mammalia
Australosphenida (incl. Monotremata)
Fruitafossor
Haramiyavia
Multituberculata
Tinodon
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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]
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]
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
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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 Restoration of Juramaia sinensis, the
therefore pregnancy was short, as in modern marsupials. This suggests oldest known Eutherian (160
that the placenta was a later development.[42] M.Y.A.)[39]
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]
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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]
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
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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
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 Sexual dimorphism in aurochs, the
indicated by mating systems, and the degree of male-biased size extinct wild ancestor of cattle
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
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
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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
Raccoon lungs being inflated is known as a bellows lung due to its resemblance to blacksmith
manually 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]
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
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]
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Sound production
As in all other tetrapods, mammals have a larynx that can
quickly open and close to produce sounds, and a
supralaryngeal vocal tract 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 A diagram of ultrasonic signals emitted by a bat,
through the mouth, and to form both oral and nasal sounds; and the echo from a nearby object
nasal sounds, such as a dog whine, are generally soft sounds,
and oral sounds, such as a dog bark, are generally loud.[101]
Some mammals have a large larynx and thus a low-pitched voice, namely
the hammer-headed bat (Hypsignathus monstrosus) where the larynx can
0:17
Beluga whale echolocation sounds 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
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
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Thermoregulation
Hair length is not a factor in thermoregulation: for example, some tropical Porcupines use their spines for
mammals such as sloths have the same length of fur length as some arctic defense.
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
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 A leopard's disruptively colored coat
monkeys, and opossums such as the Mexican mouse opossums and provides camouflage for this ambush
Derby's woolly opossums, have blue skin due to light diffraction in predator.
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]
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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
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 Goat kids stay with their mother until
functioning.[120] The testes of most mammals descend into the scrotum they are weaned.
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
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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]
Species lifespan
Among mammals, species maximum lifespan varies significantly (for example the shrew has a lifespan of two
years, whereas the oldest bowhead whale is recorded to be 211 years).[135] 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,[136] it was found that
DNA excision repair capability increased systematically with species lifespan among seven mammalian
species. Species lifespan was observed to be robustly correlated with the capacity to recognize DNA double-
strand breaks as well as the level of the DNA repair protein Ku80.[135] 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.[137] 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.[138] Three additional studies of a variety of
mammalian species also reported a correlation between species lifespan and DNA repair
capability.[139][140][141]
Locomotion
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Terrestrial
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 and running, but
they employ many other gaits occasionally, including a four-legged crawl in tight spaces.[147] Mammals show
a vast range of gaits, the order that they place and lift their appendages in locomotion. Gaits can be grouped
into categories according to their patterns of support sequence. For quadrupeds, there are three main
categories: walking gaits, running gaits and leaping gaits.[148] 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.[147]
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).[149] Many arboreal
species, such as tree porcupines, silky anteaters, spider monkeys, and possums, 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.[149] Size relating to weight affects gliding animals such as the sugar
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glider.[150] Some species of primate, bat and all species of sloth 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.[149]
Aerial
The wings of bats are much thinner and consist of more bones than those of birds, allowing bats to maneuver
more accurately and fly with more lift and less drag.[152][153] By folding the wings inwards towards their body
on the upstroke, they use 35% less energy during flight than birds.[154] 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.[155]
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.[156]
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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 gophers, 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.[158]
Many fossorial mammals such as shrews, hedgehogs, and moles were classified under the now obsolete order
Insectivora.[159]
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.[160][161] The flukes of
sirenians are raised up and down in long strokes to move the animal
A pod of short-beaked common
forward, and can be twisted to turn. The forelimbs are paddle-like flippers dolphins swimming
which aid in turning and slowing.[162]
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.[163][164] 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.[165] They rely on their fore-flippers for locomotion
in a wing-like manner similar to penguins and sea turtles.[166] Fore-flipper movement is not continuous, and
the animal glides between each stroke.[164] 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;[163] the hind-flippers serve as stabilizers.[165] Other semi-aquatic mammals include beavers,
hippopotamuses, otters and platypuses.[167] Hippos are very large semi-aquatic mammals, and their barrel-
shaped bodies have graviportal skeletal structures,[168] adapted to carrying their enormous weight, and their
specific gravity allows them to sink and move along the bottom of a river.[169]
Behavior
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Many mammals communicate by vocalizing. Vocal communication serves many purposes, including in
mating rituals, as warning calls,[171] to indicate food sources, and for social purposes. Males often call during
mating rituals to ward off other males and to attract females, as in the roaring of lions and red deer.[172] The
songs of the humpback whale may be signals to females;[173] they have different
dialects in different regions of the ocean.[174] Social vocalizations include the
territorial calls of gibbons, and the use of frequency in greater spear-nosed bats
to distinguish between groups.[175] 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.[170] Prairie dogs similarly have complex calls that
signal the type, size, and speed of an approaching predator.[176] Elephants
communicate socially with a variety of sounds including snorting, screaming, Vervet monkeys use at least
trumpeting, roaring and rumbling. Some of the rumbling calls are infrasonic, four distinct alarm calls for
below the hearing range of humans, and can be heard by other elephants up to 6 different predators.[170]
[177]
miles (9.7 km) away at still times near sunrise and sunset.
Orca calling including occasional condition and their ability to escape,[178][179] or when white-tailed deer
echolocation clicks and other prey mammals flag with conspicuous tail markings when
alarmed, informing the predator that it has been detected.[180] 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.[181][182][183] Microbats and
toothed whales including oceanic dolphins vocalize both socially and in echolocation.[184][185][186]
Feeding
To maintain a high constant body temperature is energy expensive—
mammals therefore need a nutritious and plentiful diet. While the earliest
mammals were probably predators, different species have since adapted
to meet their dietary requirements in a variety of ways. Some eat other
animals—this is a carnivorous diet (and includes insectivorous diets).
Other mammals, called herbivores, eat plants, which contain complex
carbohydrates such as cellulose. An herbivorous diet includes subtypes
A short-beaked echidna foraging for
such as granivory (seed eating), folivory (leaf eating), frugivory (fruit insects
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 or in a large cecum.[91] Some mammals are coprophagous,
consuming feces to absorb the nutrients not digested when the food was first ingested.[87]: 131–137 An
omnivore eats both prey and plants. Carnivorous mammals have a simple digestive tract because the proteins,
lipids and minerals found in meat require little in the way of specialized digestion. Exceptions to this include
baleen whales who also house gut flora in a multi-chambered stomach, like terrestrial herbivores.[187]
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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 18 ounces (510 g; 1.1 lb) 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).[188]
Furthermore, mammals that weigh more than 18 ounces (510 g; 1.1 lb) 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 (ants or termites).[189]
Some mammals are omnivores and display varying degrees of carnivory and
herbivory, generally leaning in favor of one more than the other. Since plants
and meat are digested differently, there is a preference for one over the
other, as in bears where some species may be mostly carnivorous and others
mostly herbivorous.[191] They are grouped into three categories:
mesocarnivory (50–70% meat), hypercarnivory (70% and greater of meat),
and hypocarnivory (50% or less of meat). The dentition of hypocarnivores
consists of dull, triangular carnassial teeth meant for grinding food.
Hypercarnivores, however, have conical teeth and sharp carnassials meant
for slashing, and in some cases strong jaws for bone-crushing, as in the case
of hyenas, allowing them to consume bones; some extinct groups, notably
the Machairodontinae, had saber-shaped canines.[190]
Many mammals, in the absence of sufficient food requirements in an environment, suppress their metabolism
and conserve energy in a process known as hibernation.[195] 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.[196] 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 −2.9 °C (26.8 °F),
however the head and neck always stay above 0 °C (32 °F).[197] A few mammals in hot environments aestivate
in times of drought or extreme heat, for example the fat-tailed dwarf lemur (Cheirogaleus medius).[198]
Intelligence
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In intelligent mammals, such as primates, the cerebrum is larger relative to the rest of the brain. Intelligence
itself is not easy to define, but indications of intelligence include the ability to learn, matched with behavioral
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 colonize a fresh habitat. In some mammals, food gathering
appears to be related to intelligence: a deer feeding on plants has a brain smaller than a cat, which must think
to outwit its prey.[189]
Social structure
Eusociality is the highest level of social organization. These societies have an overlap of adult generations, the
division of reproductive labor and cooperative caring of young. Usually insects, such as bees, ants and
termites, have eusocial behavior, but it is demonstrated in two rodent species: the naked mole-rat[213] and the
Damaraland mole-rat.[214]
Presociality is when animals exhibit more than just sexual interactions with members of the same species, but
fall short of qualifying as eusocial. That is, presocial animals can display communal living, cooperative care of
young, or primitive division of reproductive labor, but they do not display all of the three essential traits of
eusocial animals. Humans and some species of Callitrichidae (marmosets and tamarins) are unique among
primates in their degree of cooperative care of young.[215] Harry Harlow set up an experiment with rhesus
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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.[216]
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.[222] A solitary animal, while foraging, can also be
less conspicuous to predators or prey.[223]
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.[228][229] 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.[230] Lek mating occurs when males congregate around females
and try to attract them with various courtship displays and vocalizations, as in harbor seals.[231]
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.
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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 (M. fuscata) do.[232]
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.[233][234][235] Mammals such as mammoths, horses and deer are
among the earliest subjects of art, being found in Upper Paleolithic cave
paintings such as at Lascaux.[236] Major artists such as Albrecht Dürer,
George Stubbs and Edwin Landseer are known for their portraits of
mammals.[237] Many species of mammals have been hunted for sport and Upper Paleolithic cave painting of a
for food; deer and wild boar are especially popular as game variety of large mammals, Lascaux,
[238][239][240] c. 17,300 years old.
animals. Mammals such as horses and dogs are widely raced
for sport, often combined with betting on the outcome.[241][242] There is a
tension between the role of animals as companions to humans, and their existence as individuals with rights
of their own.[243] Mammals further play a wide variety of roles in literature,[244][245][246] film,[247]
mythology, and religion.[248][249][250]
Domestic mammals form a large part of the livestock raised for meat across the
world. They include (2009) around 1.4 billion cattle, 1 billion sheep, 1 billion
Cattle have been kept for
domestic pigs,[254][255] and (1985) over 700 million rabbits.[256] Working
milk for thousands of years.
domestic animals including cattle and horses have been used for work and
transport from the origins of agriculture, their numbers declining with the arrival
of mechanised transport and agricultural machinery. 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,
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again mainly in rural areas. In mountainous regions unsuitable for wheeled vehicles, pack animals continue to
transport goods.[257] Mammal skins provide leather for shoes, clothing and upholstery. Wool from mammals
including sheep, goats and alpacas has been used for centuries for clothing.[258][259]
Hybrids
Hybrids are offspring resulting from the
breeding of two genetically distinct individuals,
which usually will result in a high degree of
heterozygosity, though hybrid and heterozygous
are not synonymous. The deliberate or
accidental hybridizing of two or more species of
closely related animals through captive
breeding is a human activity which has been in A true quagga, 1870 (left) vs. a bred-back quagga, 2014 (right)
existence for millennia and has grown for
economic purposes.[269] Hybrids between
different subspecies within a species (such as between the Bengal tiger and Siberian tiger) are known as intra-
specific hybrids. Hybrids between different species within the same genus (such as between lions and tigers)
are known as interspecific hybrids or crosses. Hybrids between different genera (such as between sheep and
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goats) are known as intergeneric hybrids.[270] 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 recognized as species, such as the red wolf (though this is
controversial).[271]
Artificial selection, 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 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. As a result, bred-back breeds are at best vague look-alikes of extinct wildtypes, as
Heck cattle are of the aurochs.[272]
Purebred wild species evolved to a specific ecology can be threatened with extinction[273] through the process
of genetic pollution, the uncontrolled hybridization, introgression genetic swamping which leads to
homogenization or out-competition from the heterosic hybrid species.[274] 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.[275] 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 is a normal
evolutionary process, nevertheless, hybridization threatens the existence of rare species.[276][277]
Threats
The loss of species from ecological communities, defaunation,
is primarily driven by human activity.[278] This has resulted
in empty forests, ecological communities depleted of large
vertebrates.[279][280] 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.[281][282] The 2019
Global Assessment Report on Biodiversity and Ecosystem
Services by IPBES states that the total biomass of wild
mammals has declined by 82 percent since the beginning of
human civilization.[283][284] Wild animals make up just 4%
of mammalian biomass on earth, while humans and their
domesticated animals make up 96%.[260]
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world.[292][293] Scientists claim that the growing demand for meat is contributing to biodiversity loss as this is
a significant driver of deforestation and habitat destruction; species-rich habitats, such as significant portions
of the Amazon rainforest, are being converted to agricultural land for meat production.[294][295][296] Another
influence is over-hunting and poaching, which can reduce the overall population of game animals,[297]
especially those located near villages,[298] as in the case of peccaries.[299] The effects of poaching can
especially be seen in the ivory trade with African elephants.[300] Marine mammals are at risk from
entanglement from fishing gear, notably cetaceans, with discard mortalities ranging from 65,000 to 86,000
individuals annually.[301]
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.[302] Another notable conservation organization is the IUCN,
which has a membership of over 1,200 governmental and non-governmental organizations.[303]
Recent extinctions can be directly attributed to human influences.[304][278] The IUCN characterizes 'recent'
extinction as those that have occurred past the cut-off point of 1500,[305] and around 80 mammal species
have gone extinct since that time and 2015.[306] Some species, such as the Père David's deer[307] 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.[308]: 318 Other
populations are only locally extinct (extirpated), still existing elsewhere, but reduced in
distribution,[308]: 75–77 as with the extinction of gray whales in the Atlantic.[309]
Notes
a. Decreased latency to approach the mirror, repetitious head circling and close viewing of the marked areas
were considered signs of self-recognition since they do not have arms and cannot touch the marked
areas.[211]
b. Diamond discussed this matter further in his 1997 book Guns, Germs, and Steel.[251]
See also
List of mammal genera – living mammals
List of mammalogists
List of monotremes and marsupials
List of placental mammals
List of prehistoric mammals
List of threatened mammals of the United States
Lists of mammals by population size
Lists of mammals by region
Mammals described in the 2000s
Mammals in culture
Small mammal
External resources
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Further reading
Brown WM (2001). "Natural selection of mammalian brain components". Trends in Ecology and Evolution.
16 (9): 471–473. doi:10.1016/S0169-5347(01)02246-7 (https://doi.org/10.1016%2FS0169-5347%2801%2
902246-7).
McKenna MC, Bell SK (1997). Classification of Mammals Above the Species Level (https://books.google.c
om/books?id=id=zS7FZkzIw-cC). New York: Columbia University Press. ISBN 978-0-231-11013-6.
OCLC 37345734 (https://www.worldcat.org/oclc/37345734).
Nowak RM (1999). Walker's mammals of the world (https://books.google.com/books?id=T37sFCl43E8C)
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External links
Biodiversitymapping.org – All mammal orders in the world with distribution maps (http://www.biodiversitym
apping.org/mammals.htm) Archived (https://web.archive.org/web/20160926130232/http://www.biodiversity
mapping.org/mammals.htm) 2016-09-26 at the Wayback Machine
Paleocene Mammals (http://paleocene-mammals.de/), a site covering the rise of the mammals,
paleocene-mammals.de
Evolution of Mammals (http://www.enchantedlearning.com/subjects/mammals/Evolution.shtml), a brief
introduction to early mammals, enchantedlearning.com
European Mammal Atlas EMMA (http://www.european-mammals.org/php/mapmaker.php) from Societas
Europaea Mammalogica, European-mammals.org
Marine Mammals of the World (https://swfsc.noaa.gov/uploadedFiles/Divisions/PRD/Publications/Jefferso
netal93(14).pdf) – An overview of all marine mammals, including descriptions, both fully aquatic and semi-
aquatic, noaa.gov
Mammalogy.org (http://www.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
https://en.wikipedia.org/wiki/Mammal Page 53 of 53