Mammals () are a group of
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
animals constituting the
class
Class or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used differentl ...
Mammalia (), characterized by the presence of
mammary gland
A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in primat ...
s which in
females
Female (symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction.
A female has larger gametes than a male. Females a ...
produce
milk
Milk is a white liquid food produced by the mammary glands of mammals. It is the primary source of nutrition for young mammals (including breastfed human infants) before they are able to digestion, digest solid food. Immune factors and immune ...
for feeding (nursing) their young, a
neocortex
The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
(a region of the brain),
fur
Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
or
hair
Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals.
The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
, and three
middle ear bones. These characteristics distinguish them from
reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s (including
bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s) from which they
diverged in the
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
, over 300 million years ago. Around 6,400
extant
Extant is the opposite of the word extinct. It may refer to:
* Extant hereditary titles
* Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English
* Extant taxon, a taxon which is not extinct, ...
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of mammals have been described divided into 29
orders
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
. The largest
orders
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
, in terms of number of species, are the
rodents
Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are nat ...
,
bats
Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
, and
Eulipotyphla
Eulipotyphla (, which means "truly fat and blind") is an order of mammals suggested by molecular methods of phylogenetic reconstruction, which includes the laurasiatherian members of the now-invalid polyphyletic order Lipotyphla, but not the afr ...
(
hedgehog
A hedgehog is a spiny mammal of the subfamily Erinaceinae, in the eulipotyphlan family Erinaceidae. There are seventeen species of hedgehog in five genera found throughout parts of Europe, Asia, and Africa, and in New Zealand by introducti ...
s,
mole
Mole (or Molé) may refer to:
Animals
* Mole (animal) or "true mole", mammals in the family Talpidae, found in Eurasia and North America
* Golden moles, southern African mammals in the family Chrysochloridae, similar to but unrelated to Talpida ...
s,
shrew
Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
s, and others). The next three are the
Primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s (including
human
Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
s,
ape
Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its siste ...
s,
monkey
Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
s, and others), the
Artiodactyla
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
(
cetacean
Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
s and
even-toed ungulate
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
s), and the
Carnivora
Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...
(
cat
The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s,
dog
The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
s,
seals
Seals may refer to:
* Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly:
** Earless seal, or "true seal"
** Fur seal
* Seal (emblem), a device to impress an emblem, used as a means of a ...
, and others).
In terms of
cladistics
Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
, which reflects evolutionary history, mammals are the only living members of the
Synapsid
Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
a (synapsids); this
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, together with
Sauropsida
Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syna ...
(reptiles and birds), constitutes the larger
Amniota
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distingu ...
clade. The early synapsids were
sphenacodonts, a group that included the famous ''
Dimetrodon
''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
''. The synapsids split into several diverse groups of non-mammalian synapsids — traditionally and incorrectly referred to as mammal-like reptiles or by the term pelycosaurs, and now known as stem mammals or protomammals — before giving rise to
therapsids
Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
during the beginning of the
Middle Permian
The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0. ...
period. Mammals originated from
cynodont
The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variety ...
s, an advanced group of therapsids, during the Late
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
-Early
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
. The modern mammalian orders arose in the
Paleogene
The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...
and
Neogene
The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
periods of the
Cenozoic
The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
era, after the
extinction of non-avian dinosaurs, and have been the dominant terrestrial animal group from 66 million years ago to the present.
The basic body type is
quadruped
Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
, and most mammals use their four extremities for
terrestrial locomotion
Terrestrial locomotion has evolved as animals adapted from aquatic to terrestrial environments. Locomotion on land raises different problems than that in water, with reduced friction being replaced by the increased effects of gravity.
As viewed ...
; but in some, the extremities are adapted for
life at sea
Life is a quality that distinguishes matter that has biological processes, such as signaling and self-sustaining processes, from that which does not, and is defined by the capacity for growth, reaction to stimuli, metabolism, energy transf ...
, in
the air,
in trees,
underground
Underground most commonly refers to:
* Subterranea (geography), the regions beneath the surface of the Earth
Underground may also refer to:
Places
* The Underground (Boston), a music club in the Allston neighborhood of Boston
* The Underground (S ...
, or
on two legs. Mammals range in size from the
bumblebee bat
Kitti's hog-nosed bat (''Craseonycteris thonglongyai''), also known as the bumblebee bat, is a near-threatened species of bat and the only extant member of the family Craseonycteridae. It occurs in western Thailand and southeast Myanmar, where it ...
to the
blue whale
The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can ...
—possibly the largest animal to have ever lived. Maximum lifespan varies from two years for the shrew to 211 years for the
bowhead whale
The bowhead whale (''Balaena mysticetus'') is a species of baleen whale belonging to the family Balaenidae and the only living representative of the genus ''Balaena''. They are the only baleen whale endemic to the Arctic and subarctic waters, ...
. All modern mammals give birth to live young, except the five species of
monotreme
Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals (Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brain ...
s, which are egg-laying mammals. The most species-rich group of mammals, the
cohort
Cohort or cohortes may refer to:
* Cohort (educational group), a group of students working together through the same academic curriculum
* Cohort (floating point), a set of different encodings of the same numerical value
* Cohort (military unit), ...
called
placentals
Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
, have a
placenta
The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate mater ...
, which enables the feeding of the fetus during
gestation
Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
.
Most mammals are
intelligent, with some possessing large brains,
self-awareness
In philosophy of self, self-awareness is the experience of one's own personality or individuality. It is not to be confused with consciousness in the sense of qualia. While consciousness is being aware of one's environment and body and lifesty ...
, and
tool use
Tool use by animals is a phenomenon in which an animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, defence, communication, recreation or construction. Originally thought to be a skill possessed o ...
. Mammals can communicate and vocalize in several ways, including the production of
ultrasound
Ultrasound is sound waves with frequency, frequencies higher than the upper audible limit of human hearing range, hearing. Ultrasound is not different from "normal" (audible) sound in its physical properties, except that humans cannot hea ...
,
scent-marking
In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species) using agonistic behaviors or (less commonly) real physical aggression. A ...
,
alarm signal
In animal communication, an alarm signal is an antipredator adaptation in the form of signals emitted by social animals in response to danger. Many primates and birds have elaborate alarm calls for warning conspecifics of approaching predators. ...
s,
singing
Singing is the act of creating musical sounds with the voice. A person who sings is called a singer, artist or vocalist (in jazz and/or popular music). Singers perform music (arias, recitatives, songs, etc.) that can be sung with or without ...
, and
echolocation. Mammals can organize themselves into
fission-fusion societies,
harems
Harem (Persian: حرمسرا ''haramsarā'', ar, حَرِيمٌ ''ḥarīm'', "a sacred inviolable place; harem; female members of the family") refers to domestic spaces that are reserved for the women of the house in a Muslim family. A hare ...
, and
hierarchies
A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
—but can also be solitary and
territorial
A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal.
In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
. Most mammals are
polygynous
Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women.
Incidence
Polygyny is more widespread in Africa than in any ...
, but some can be
monogamous
Monogamy ( ) is a form of Dyad (sociology), dyadic Intimate relationship, relationship in which an individual has only one Significant other, partner during their lifetime. Alternately, only one partner at any one time (Monogamy#Serial monogamy, ...
or
polyandrous
Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...
.
Domestication
Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. ...
of many types of mammals by humans played a major role in the
Neolithic Revolution
The Neolithic Revolution, or the (First) Agricultural Revolution, was the wide-scale transition of many human cultures during the Neolithic period from a lifestyle of hunting and gathering to one of agriculture and settlement, making an incre ...
, and resulted in
farming
Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
replacing
hunting and gathering
A traditional hunter-gatherer or forager is a human living an ancestrally derived lifestyle in which most or all food is obtained by foraging, that is, by gathering food from local sources, especially edible wild plants but also insects, fungi, ...
as the primary source of food for humans. This led to a major restructuring of human societies from nomadic to sedentary, with more co-operation among larger and larger groups, and ultimately the development of the first
civilization
A civilization (or civilisation) is any complex society characterized by the development of a state, social stratification, urbanization, and symbolic systems of communication beyond natural spoken language (namely, a writing system).
Ci ...
s. Domesticated mammals provided, and continue to provide, power for transport and agriculture, as well as food (
meat
Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
and
dairy product
Dairy products or milk products, also known as lacticinia, are food products made from (or containing) milk. The most common dairy animals are cow, water buffalo, nanny goat, and ewe. Dairy products include common grocery store food items in th ...
s),
fur
Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
, and
leather
Leather is a strong, flexible and durable material obtained from the tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffalo, pigs and hogs, ...
. Mammals are also
hunted and raced for sport, and are used as
model organism
A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...
s in science. Mammals have been depicted in
art
Art is a diverse range of human activity, and resulting product, that involves creative or imaginative talent expressive of technical proficiency, beauty, emotional power, or conceptual ideas.
There is no generally agreed definition of wha ...
since
Paleolithic
The Paleolithic or Palaeolithic (), also called the Old Stone Age (from Greek: παλαιός ''palaios'', "old" and λίθος ''lithos'', "stone"), is a period in human prehistory that is distinguished by the original development of stone too ...
times, and appear in literature, film, mythology, and religion. Decline in numbers and
extinction
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
of many mammals is primarily driven by human
poaching
Poaching has been defined as the illegal hunting or capturing of wild animals, usually associated with land use rights.
Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set a ...
and
habitat destruction
Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, primarily
deforestation
Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
.
Classification
![Mammal species pie chart](https://upload.wikimedia.org/wikipedia/commons/0/00/Mammal_species_pie_chart.svg)
Mammal classification has been through several revisions since
Carl Linnaeus
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
initially defined the class, and at present, no classification system is universally accepted. McKenna & Bell (1997) and Wilson & Reeder (2005) provide useful recent compendiums.
Simpson
Simpson most often refers to:
* Simpson (name), a British surname
*''The Simpsons'', an animated American sitcom
**The Simpson family, central characters of the series ''The Simpsons''
Simpson may also refer to:
Organizations Schools
*Simpso ...
(1945)
provides
systematics
Biological systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: cladograms, phylogenetic tre ...
of mammal origins and relationships that had been taught universally until the end of the 20th century.
However, since 1945, a large amount of new and more detailed information has gradually been found: The
paleontological record has been recalibrated, and the intervening years have seen much debate and progress concerning the theoretical underpinnings of systematization itself, partly through the new concept of
cladistics
Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
. Though fieldwork and lab work progressively outdated Simpson's classification, it remains the closest thing to an official classification of mammals, despite its known issues.
Most mammals, including the six most species-rich
orders
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
, belong to the placental group. The three largest orders in numbers of species are
Rodent
Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
ia:
mice
A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
,
rat
Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' ( pack rats), ''Bandicota'' (bandicoot ...
s,
porcupine
Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethizont ...
s,
beaver
Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers ar ...
s,
capybara
The capybaraAlso called capivara (in Brazil), capiguara (in Bolivia), chigüire, chigüiro, or fercho (in Colombia and Venezuela), carpincho (in Argentina, Paraguay and Uruguay) and ronsoco (in Peru). or greater capybara (''Hydrochoerus hydro ...
s, and other gnawing mammals;
Chiroptera
Bats are mammals of the Order (biology), order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in fli ...
: bats; and
Soricomorpha
Soricomorpha (from Greek "shrew-form") is a formerly used taxon within the class of mammals. In the past it formed a significant group within the former order Insectivora. However, Insectivora was shown to be polyphyletic and various new orders w ...
:
shrew
Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
s,
moles Moles can refer to:
*Moles de Xert, a mountain range in the Baix Maestrat comarca, Valencian Community, Spain
*The Moles (Australian band)
*The Moles, alter ego of Scottish band Simon Dupree and the Big Sound
People
*Abraham Moles, French engineer ...
, and
solenodon
Solenodons (from el, τέλειος , 'channel' or 'pipe' and el, ὀδούς , 'tooth') are venomous, nocturnal, burrowing, insectivorous mammals belonging to the family Solenodontidae . The two living solenodon species are the Cuban solen ...
s. The next three biggest orders, depending on the
biological classification
In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
scheme used, are the
Primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s:
ape
Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its siste ...
s,
monkey
Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
s, and
lemur
Lemurs ( ) (from Latin ''lemures'' – ghosts or spirits) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea (), divided into 8 Family (biology), families and consisting of 15 genera and around 100 exist ...
s; the
Cetartiodactyla
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
:
whale
Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s and
even-toed ungulates
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
; and the
Carnivora
Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...
which includes
cat
The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s,
dog
The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
s,
weasel
Weasels are mammals of the genus ''Mustela'' of the family Mustelidae. The genus ''Mustela'' includes the least weasels, polecats, stoats, ferrets and European mink. Members of this genus are small, active predators, with long and slender bo ...
s,
bear
Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the Nor ...
s,
seals
Seals may refer to:
* Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly:
** Earless seal, or "true seal"
** Fur seal
* Seal (emblem), a device to impress an emblem, used as a means of a ...
, and allies.
According to ''
Mammal Species of the World
''Mammal Species of the World: A Taxonomic and Geographic Reference'' is a standard reference work in mammalogy giving descriptions and bibliographic data for the known species of mammals. It is now in its third edition, published in late 2005, ...
'', 5,416 species were identified in 2006. These were grouped into 1,229
genera
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
, 153
families
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
and 29 orders.
In 2008, the
International Union for Conservation of Nature
The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN) completed a five-year Global Mammal Assessment for its
IUCN Red List
The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
, which counted 5,488 species. According to research published in the ''
Journal of Mammalogy
The ''Journal of Mammalogy'' is a bimonthly peer-reviewed scientific journal published by Oxford University Press on behalf of the American Society of Mammalogists. Both the society and the journal were established in 1919. The journal covers rese ...
'' in 2018, the number of recognized mammal species is 6,495, including 96 recently extinct.
Definitions
The word "
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), 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 ...
" is modern, from the scientific name ''Mammalia'' coined by Carl Linnaeus in 1758, derived from the
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''
mamma
Mama(s) or Mamma or Momma may refer to:
Roles
*Mother, a female parent
*Mama-san, in Japan and East Asia, a woman in a position of authority
*Mamas, a name for female associates of the Hells Angels
Places
*Mama, Russia, an urban-type settlement ...
'' ("teat, pap"). In an influential 1988 paper, Timothy Rowe defined Mammalia
phylogenetically
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
as the
crown group
In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
of mammals, the
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
consisting of the
most recent common ancestor
In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of living
monotreme
Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals (Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brain ...
s (
echidna
Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the ...
s and
platypus
The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal Endemic (ecology), endemic to Eastern states of Australia, eastern Australia, including Tasmania. The platypu ...
es) and
Theria
Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes.
Ch ...
n mammals (
marsupial
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...
s and
placental
Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupial
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsup ...
s) and all descendants of that ancestor. Since this ancestor lived in the
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
period, Rowe's definition excludes all animals from the earlier
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, despite the fact that Triassic fossils in the
Haramiyida
Haramiyida ("thief" from Arabic الحرامية (al ḥarāmiyah), "thief, bandit") is a possibly polyphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. Their teeth, which are by far the most common remains ...
have been referred to the Mammalia since the mid-19th century. If Mammalia is considered as the crown group, its origin can be roughly dated as the first known appearance of animals more closely related to some extant mammals than to others. ''
Ambondro'' is more closely related to monotremes than to therian mammals while ''
Amphilestes
''Amphilestes'' is a genus of extinct eutriconodont mammal from the Middle Jurassic of the United Kingdom. It was one of the first Mesozoic mammals discovered and described.
Discovery
The first specimen of ''Amphilestes'' was discovered along ...
'' and ''
Amphitherium
''Amphitherium'' is an extinct genus of stem cladotherian mammal that lived during the Middle Jurassic of England. It was one of the first Mesozoic mammals ever described. A recent phylogenetic study found it to be the sister taxon of ''Palaeoxo ...
'' are more closely related to the therians; as fossils of all three genera are dated about in the
Middle Jurassic
The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
, this is a reasonable estimate for the appearance of the crown group.
T.S. Kemp has provided a more traditional definition: "
Synapsid
Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s that possess a
dentary
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
–
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone.
In most tetrapods, the squamosal and quadratojugal The quadratojugal is a skull bone present in many vertebrates, including ...
jaw articulation and
occlusion
Occlusion may refer to:
Health and fitness
* Occlusion (dentistry), the manner in which the upper and lower teeth come together when the mouth is closed
* Occlusion miliaria, a skin condition
* Occlusive dressing, an air- and water-tight trauma ...
between upper and lower molars with a transverse component to the movement" or, equivalently in Kemp's view, the clade originating with the last common ancestor of ''
Sinoconodon
''Sinoconodon'' is an extinct genus of mammaliamorphs that appears in the fossil record of the Lufeng Formation of China in the Sinemurian stage of the Early Jurassic period, about 193 million years ago. While sharing many plesiomorphic traits wi ...
'' and living mammals. The earliest known synapsid satisfying Kemp's definitions is ''
Tikitherium
''Tikitherium'' is an extinct genus of mammaliaforms from the Late Triassic. It is thought to be an insectivore and a close relative to Docodonta. ''Tikitherium'' refers to Tiki, the village located near the Tiki Formation where the specimen wa ...
'', dated , so the appearance of mammals in this broader sense can be given this
Late Triassic
The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
date.
McKenna/Bell classification
In 1997, the mammals were comprehensively revised by
Malcolm C. McKenna and Susan K. Bell, which has resulted in the McKenna/Bell classification. The authors worked together as
paleontologist
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
s at the
American Museum of Natural History
The American Museum of Natural History (abbreviated as AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. In Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 26 inter ...
. McKenna inherited the project from Simpson and, with Bell, constructed a completely updated hierarchical system, covering living and extinct taxa, that reflects the historical genealogy of Mammalia.
[ Their 1997 book, ''Classification of Mammals above the Species Level'',] is a comprehensive work on the systematics, relationships and occurrences of all mammal taxa, living and extinct, down through the rank of genus, though molecular genetic data challenge several of the groupings.
In the following list, extinct
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
groups are labelled with a dagger
A dagger is a fighting knife with a very sharp point and usually two sharp edges, typically designed or capable of being used as a thrusting or stabbing weapon.State v. Martin, 633 S.W.2d 80 (Mo. 1982): This is the dictionary or popular-use de ...
(†).
Class Mammalia
* Subclass Prototheria
Prototheria (; from Greek πρώτος, ''prōtos'', first, + θήρ, ''thēr'', wild animal) is a subclass to which the orders Monotremata, Morganucodonta, Docodonta, Triconodonta and Multituberculata have been assigned, although the validity ...
: monotremes: echidna
Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the ...
s and the platypus
The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal Endemic (ecology), endemic to Eastern states of Australia, eastern Australia, including Tasmania. The platypu ...
* Subclass Theriiformes
Theriiformes is a clade of mammals. The term was coined by Timothy B. Rowe in his doctoral dissertation, and is defined as the clade formed by the most recent common ancestor of multituberculates and theria
Theria (; Greek: , wild beast) i ...
: live-bearing mammals and their prehistoric relatives
** Infraclass †Allotheria
Allotheria (meaning "other beasts", from the Greek , '–other and , '–wild animal) is an extinct branch of successful Mesozoic mammals. The most important characteristic was the presence of lower molariform teeth equipped with two longitudin ...
: multituberculates
** Infraclass †Eutriconodonta
Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack ''et al.'' in 1973 as a replacement name for the ...
: eutriconodonts
** Infraclass Holotheria
Mammalia is a class of animal within the phylum Chordata. Mammal classification has been through several iterations since Carl Linnaeus initially defined the class. No classification system is universally accepted; McKenna & Bell (1997) and Wilson ...
: modern live-bearing mammals and their prehistoric relatives
*** Superlegion †Kuehneotheria
Kuehneotheriidae is an extinct family of mammaliaforms traditionally placed within 'Symmetrodonta', though now generally considered more basal than true symmetrodonts. All members of Kuehneotheriidae which have been found so far are represented o ...
*** Supercohort Theria
Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes.
Ch ...
: live-bearing mammals
**** Cohort Marsupialia
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a pou ...
: marsupials
***** Magnorder Australidelphia
Australidelphia is the superorder that contains roughly three-quarters of all marsupials, including all those native to Australasia and a single species — the monito del monte — from South America. All other American marsupials are members of ...
: Australian marsupials and the monito del monte
The monito del monte or colocolo opossum, ''Dromiciops gliroides'', also called ''chumaihuén'' in Mapudungun, is a diminutive marsupial native only to southwestern South America (Argentina and Chile). It is the only extant species in the ancient ...
***** Magnorder Ameridelphia
Ameridelphia is traditionally a superorder that includes all marsupials living in the Americas except for the Monito del monte (''Dromiciops''). It is now regarded as a paraphyletic group.
Orders
The orders within this group are listed below:
...
: New World marsupials. Now considered paraphyletic, with shrew opossum
The family (biology), family Caenolestidae contains the seven surviving species of shrew opossum: small, shrew-like marsupials that are confined to the Andes mountains of South America. The order is thought to have diverged from the ancestral mar ...
s being closer to australidelphians.
**** Cohort Placentalia
Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguished ...
: placentals
***** Magnorder Xenarthra
Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Ex ...
: xenarthrans
***** Magnorder Epitheria
Epitherians comprise all the placental mammals except the Xenarthra. They are primarily characterized by having a stirrup-shaped stapes in the middle ear, which allows for passage of a blood vessel. This is in contrast to the column-shaped stapes ...
: epitheres
****** Superorder †Leptictida
Leptictida (''leptos iktis'' "small/slender weasel") is a possibly paraphyletic extinct order of eutherian mammals. Their classification is contentious: according to cladistic studies, they may be (distantly) related to Euarchontoglires (rodents ...
****** Superorder Preptotheria
Preptotheria is a superorder of placental mammals proposed by McKenna & Bell in their classification of mammals.
Classification
The Linnean taxonomy of Preptotheria according to the scheme of McKenna & Bell (1997):
Cohort Placentalia
Magnorde ...
******* Grandorder Anagalida
Anagaloidea is an extinct order of placental mammals that first appeared during the Paleocene epoch.
Taxonomy
According to the traditional (morphological) view, Anagaloidea is part of the superorder Anagalida, along with the elephant shrews, rod ...
: lagomorphs
The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae ( hares and rabbits) and the Ochotonidae (pikas). The name of the order is derived from the Ancient Greek ''lagos'' (λα ...
, rodent
Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
s and elephant shrew
Elephant shrews, also called jumping shrews or sengis, are small insectivorous mammals native to Africa, belonging to the family Macroscelididae, in the order Macroscelidea. Their traditional common English name "elephant shrew" comes from a pe ...
s
******* Grandorder Ferae
Ferae ( , , "wild beasts") is a mirorder of placental mammalsMalcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'' in Columbia University Press, New York (1997), 631 Seiten. that groups together clades Pan-Ca ...
: carnivora
Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...
ns, pangolin
Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smut ...
s, †creodont
Creodonta ("meat teeth") is a former order of extinct carnivorous placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ance ...
s and relatives
******* Grandorder Lipotyphla
Lipotyphla is a formerly used order of mammals, including the members of the order Eulipotyphla (i.e. the solenodons, family Solenodontidae; hedgehogs and gymnures, family Erinaceidae; desmans, moles, and shrew-like moles, family Talpidae; and tru ...
: insectivora
The order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order Eulipotyphla, w ...
ns
******* Grandorder Archonta
The Archonta are a now-abandoned group of mammals, considered a superorder in some classifications, which consists of these orders:
*Primates
* Plesiadapiformes (extinct primate-like archontans)
* Scandentia (treeshrews)
* Dermoptera (colugos)
W ...
: bat
Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s, primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s, colugo
Colugos () are arboreal gliding mammals that are native to Southeast Asia. Their closest evolutionary relatives are primates. There are just two living species of colugos: the Sunda flying lemur (''Galeopterus variegatus'') and the Philippine fl ...
s and treeshrew
The treeshrews (or tree shrews or banxrings) are small mammals native to the tropical forests of South and Southeast Asia. They make up the entire order Scandentia, which split into two families: the Tupaiidae (19 species, "ordinary" treeshrews) ...
s (now considered paraphyletic, with bats being closer to other groups)
******* Grandorder Ungulata
Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
: ungulates
******** Order Tubulidentata
Orycteropodidae is a family of afrotherian mammals. Although there are many fossil species, the only species surviving today is the aardvark, ''Orycteropus afer''. Orycteropodidae is recognized as the only family within the order Tubulidentata ...
''incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'': aardvark
The aardvark ( ; ''Orycteropus afer'') is a medium-sized, burrowing, nocturnal mammal native to Africa. It is the only living species of the order Tubulidentata, although other prehistoric species and genera of Tubulidentata are known. Unlike ...
******** Mirorder Eparctocyona: †condylarth
Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wast ...
s, whale
Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s and artiodactyls
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
(even-toed ungulates)
******** Mirorder †Meridiungulata
South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals of controversial affinities that were indigenous to South America prior to the Great American Biotic Interchange. They comprise five major groups co ...
: South American ungulates
******** Mirorder Altungulata
Altungulata or Pantomesaxonia (''sensu'' and later authors) is an invalid clade (mirorder) of ungulate mammals comprising the perissodactyls, hyracoids, and tethytheres (sirenians, proboscideans, and related extinct taxa).
The name "Pantomesaxo ...
: perissodactyls
Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...
(odd-toed ungulates), elephant
Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae an ...
s, manatee
Manatees (family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing three of the four living species ...
s and hyrax
Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simil ...
es
Molecular classification of placentals
As of the early 21st century, molecular studies based on DNA analysis have suggested new relationships among mammal families. Most of these findings have been independently validated by retrotransposon
Retrotransposons (also called Class I transposable elements or transposons via RNA intermediates) are a type of genetic component that copy and paste themselves into different genomic locations (transposon) by converting RNA back into DNA through ...
presence/absence data. Classification systems based on molecular studies reveal three major groups or lineages of placental mammals—Afrotheria
Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a clade of mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephant shrews (also know ...
, Xenarthra
Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Ex ...
and Boreoeutheria
Boreoeutheria (, "northern true beasts") is a magnorder of placental mammals that groups together superorders Euarchontoglires and Laurasiatheria. With a few exceptionsExceptional clades whose males lack the usual boreoeutherian scrotum are moles ...
—which diverged in the Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
. The relationships between these three lineages is contentious, and all three possible hypotheses have been proposed with respect to which group is basal. These hypotheses are Atlantogenata
Atlantogenata is a proposed clade of mammals containing the cohorts or superorders Xenarthra and Afrotheria. These groups originated and radiated in the South American and African continents, respectively, presumably in the Cretaceous. Together ...
(basal Boreoeutheria), Epitheria
Epitherians comprise all the placental mammals except the Xenarthra. They are primarily characterized by having a stirrup-shaped stapes in the middle ear, which allows for passage of a blood vessel. This is in contrast to the column-shaped stapes ...
(basal Xenarthra) and Exafroplacentalia
Exafroplacentalia or Notolegia is a clade of placental mammals proposed in 2001 on the basis of molecular research.
Exafroplacentalia places Xenarthra as a sister group to the Boreoeutheria (comprising Laurasiatheria and Euarchontoglires), thus ...
(basal Afrotheria). Boreoeutheria in turn contains two major lineages—Euarchontoglires
Euarchontoglires (synonymous with Supraprimates) is a clade and a superorder of mammals, the living members of which belong to one of the five following groups: rodents, lagomorphs, treeshrews, colugos, and primates.
Evolutionary affinities wit ...
and Laurasiatheria
Laurasiatheria ("laurasian beasts") is a superorder of placental mammals that groups together true insectivores ( eulipotyphlans), bats ( chiropterans), carnivorans, pangolins ( pholidotes), even-toed ungulates (artiodactyls), odd-toed ungulates ...
.
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
Nuclear may refer to:
Physics
Relating to the nucleus of the atom:
* Nuclear engineering
*Nuclear physics
*Nuclear power
*Nuclear reactor
*Nuclear weapon
*Nuclear medicine
*Radiation therapy
*Nuclear warfare
Mathematics
*Nuclear space
*Nuclear ...
or mitochondrial
A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...
) and varying interpretations of paleogeographic
Palaeogeography (or paleogeography) is the study of historical geography, generally physical landscapes. Palaeogeography can also include the study of human or cultural environments. When the focus is specifically on landforms, the term paleo ...
data.[
The ]cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
above is based on Tarver ''et al.''. (2016)
Evolution
Origins
Synapsida, a clade that contains mammals and their extinct relatives, originated during the Pennsylvanian subperiod (~323 million to ~300 million years ago), when they split from the reptile lineage. Crown group mammals evolved from earlier mammaliaforms
Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recen ...
during the Early Jurassic
The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
. The cladogram takes Mammalia to be the crown group.
Evolution from older amniotes
The first fully terrestrial vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s were amniote
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s. Like their amphibious early tetrapod
Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
predecessors, they had lungs and limbs. Amniotic eggs, however, have internal membranes that allow the developing embryo
An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
to breathe but keep water in. Hence, amniotes can lay eggs on dry land, while amphibian
Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s generally need to lay their eggs in water.
The first amniotes apparently arose in the Pennsylvanian subperiod of the Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
. They descended from earlier reptiliomorph
Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians ( lissamphibians). It was de ...
amphibious tetrapods, which lived on land that was already inhabited by insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and other invertebrates as well as fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s, moss
Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es and other plants. Within a few million years, two important amniote lineages became distinct: the synapsid
Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s, which would later include the common ancestor of the mammals; and the sauropsid
Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syna ...
s, which now include turtle
Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
s, lizard
Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s, snake
Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
s, crocodilian
Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
s and dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s (including bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s). Synapsids have a single hole (temporal fenestra
An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets, ...
) low on each side of the skull. Primitive synapsids included the largest and fiercest animals of the early Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
such as Dimetrodon
''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
. Nonmammalian synapsids were traditionally – and incorrectly – called "mammal-like reptiles" or pelycosaurs
Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an order, but this is no ...
; we now know they were neither reptiles nor part of reptile lineage.[
]Therapsid
Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s, a group of synapsids, evolved in the Middle Permian
The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0. ...
, about 265 million years ago, and became the dominant land vertebrates. They differ from basal eupelycosaurs in several features of the skull and jaws, including: larger skulls and incisor
Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s which are equal in size in therapsids, but not for eupelycosaurs.[ The therapsid lineage leading to mammals went through a series of stages, beginning with animals that were very similar to their early synapsid ancestors and ending with ]probainognathia
Probainognathia is one of the two major subgroups of the clade Eucynodontia, the other being Cynognathia. The earliest forms were carnivorous and insectivorous, though some groups eventually also evolved herbivorous diets. The earliest and most ...
n cynodont
The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variety ...
s, some of which could easily be mistaken for mammals. Those stages were characterized by:
* The gradual development of a bony secondary palate
The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity.
A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
.
* Abrupt acquisition of endothermy
An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inste ...
among Mammaliamorpha
Mammaliamorpha is a clade of cynodonts. It contains the clades Tritylodontidae and Mammaliaformes, as well as a few genera that do not belong to either of these groups. The family Tritheledontidae has also been placed in Mammaliamorpha by some ph ...
, thus prior to the origin of mammals by 30-50 millions of years '.
* Progression towards an erect limb posture, which would increase the animals' stamina by avoiding Carrier's constraint
Carrier's constraint is the observation that air-breathing vertebrates which have two lungs and flex their bodies sideways during locomotion find it very difficult to move and breathe at the same time, because the sideways flexing expands one lung ...
. But this process was slow and erratic: for example, all herbivorous nonmammaliaform therapsids retained sprawling limbs (some late forms may have had semierect hind limbs); Permian carnivorous therapsids had sprawling forelimbs, and some late Permian ones also had semisprawling hindlimbs. In fact, modern monotremes still have semisprawling limbs.
* The dentary
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
gradually became the main bone of the lower jaw which, by the Triassic, progressed towards the fully mammalian jaw (the lower consisting only of the dentary) and middle ear (which is constructed by the bones that were previously used to construct the jaws of reptiles).
First mammals
The Permian–Triassic extinction event
The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
about 252 million years ago, which was a prolonged event due to the accumulation of several extinction pulses, ended the dominance of carnivorous therapsids. In the early Triassic, most medium to large land carnivore niches were taken over by archosaur
Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
s which, over an extended period (35 million years), came to include the crocodylomorphs
Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction.
During Mesozoic and early Cenozoic times, cr ...
, the pterosaur
Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
s and the dinosaurs; however, large cynodonts like ''Trucidocynodon
''Trucidocynodon'' is an extinct genus of ecteniniid cynodonts from Upper Triassic of Brazil. It contains a single species, ''Trucidocynodon riograndensis''. Fossils of ''Trucidocynodon'' were discovered in Santa Maria Formation outcrops in Pal ...
'' and traversodontids
Traversodontidae is an extinct family of herbivorous cynodonts. Traversodonts were primarily Gondwanan, with many species known from Africa and South America. Recently, traversodonts have also been found from Europe and eastern North America. ...
still occupied large sized carnivorous and herbivorous niches respectively. By the Jurassic, the dinosaurs had come to dominate the large terrestrial herbivore niches as well.
The first mammals (in Kemp's sense) appeared in the Late Triassic epoch (about 225 million years ago), 40 million years after the first therapsids. They expanded out of their nocturnal insectivore
A robber fly eating a hoverfly
An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects.
The first vertebrate insectivores wer ...
niche from the mid-Jurassic onwards; The Jurassic ''Castorocauda
''Castorocauda'' is an extinct, semi-aquatic, superficially otter-like genus of docodont mammaliaforms with one species, ''C. lutrasimilis''. It is part of the Yanliao Biota, found in the Daohugou Beds of Inner Mongolia, China dating to the M ...
'', for example, was a close relative of true mammals that had adaptations for swimming, digging and catching fish. Most, if not all, are thought to have remained nocturnal (the nocturnal bottleneck
The nocturnal bottleneck hypothesis is a hypothesis to explain several mammalian traits. In 1942, Gordon Lynn Walls described this concept which states that placental mammals were mainly or even exclusively nocturnal through most of their evolu ...
), accounting for much of the typical mammalian traits. The majority of the mammal species that existed in the Mesozoic Era
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising ...
were multituberculates, eutriconodonts and spalacotheriid
Spalacotheriidae is a family of extinct mammals belonging to the paraphyletic group ' Symmetrodonta'. They lasted from the Early Cretaceous to the Campanian in North America, Europe, Asia and North Africa.
Spalacotheriids are characterised by ha ...
s. The earliest known metatherian
Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well ...
is ''Sinodelphys
''Sinodelphys'' is an extinct eutherian from the Early Cretaceous, estimated to be 125 million years old. It was discovered and described in 2003 in rocks of the Yixian Formation in Liaoning Province, China, by a team of scientists including Zhe ...
'', found in 125 million-year-old Early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
shale
Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
in China's northeastern Liaoning Province
Liaoning () is a coastal provinces of China, province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and i ...
. The fossil is nearly complete and includes tufts of fur and imprints of soft tissues.
The oldest known fossil among the Eutheria
Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials.
Eutherians are distinguished from noneutherians by various phenotypic tra ...
("true beasts") is the small shrewlike ''Juramaia sinensis
''Juramaia'' is an extinct genus of very basal eutherian mammal known from the Late Jurassic ( Oxfordian stage) deposits of western Liaoning, China. It is a small shrew-like mammal with a body length of approximately 70–100 mm, making it ...
'', or "Jurassic mother from China", dated to 160 million years ago in the late Jurassic. A later eutherian relative, ''Eomaia
''Eomaia'' ("dawn mother") is a genus of extinct fossil mammals containing the single species ''Eomaia scansoria'', discovered in rocks that were found in the Yixian Formation, Liaoning Province, China, and dated to the Barremian Age of the Lower ...
'', dated to 125 million years ago in the early Cretaceous, possessed some features in common with the marsupials but not with the placentals, evidence that these features were present in the last common ancestor of the two groups but were later lost in the placental lineage. In particular, the epipubic bone
Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them).
They first occ ...
s extend forwards from the pelvis. These are not found in any modern placental, but they are found in marsupials, monotremes, other nontherian mammals and ''Ukhaatherium
''Ukhaatherium'' is a now extinct species of mammal that lived during the upper Cretaceous about 84 to 72 million years ago in today's East Asia. It is known above all from the fossil locality Ukhaa Tolgod, Mongolia. An adult ''Ukhaatherium'' h ...
'', an early Cretaceous animal in the eutherian order Asioryctitheria
Asioryctitheria ("Asian digging beasts") is an extinct order of early eutherians.
Skull structure
With the exception of '' Prokennalestes'', these advanced forms lacked a Meckelian groove. Furthermore, they were equipped with double-rooted can ...
. This also applies to the multituberculates. They are apparently an ancestral feature, which subsequently disappeared in the placental lineage. These epipubic bones seem to function by stiffening the muscles during locomotion, reducing the amount of space being presented, which placentals require to contain their fetus
A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal deve ...
during gestation periods. A narrow pelvic outlet indicates that the young were very small at birth and therefore pregnancy
Pregnancy is the time during which one or more offspring develops ( gestates) inside a woman's uterus (womb). A multiple pregnancy involves more than one offspring, such as with twins.
Pregnancy usually occurs by sexual intercourse, but ca ...
was short, as in modern marsupials. This suggests that the placenta was a later development.
One of the earliest known monotremes was ''Teinolophos
''Teinolophos'' is a prehistoric species of monotreme, or egg-laying mammal, from the Teinolophidae. It is known from four specimens, each consisting of a partial lower jawbone collected from the Wonthaggi Formation at Flat Rocks, Victoria (Aus ...
'', which lived about 120 million years ago in Australia. Monotremes have some features which may be inherited from the original amniotes such as the same orifice to urinate, defecate and reproduce (cloaca
In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
)—as lizards and birds also do— and they lay eggs
Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
which are leathery and uncalcified.
Earliest appearances of features
''Hadrocodium
''Hadrocodium wui'' is an extinct mammaliaform that lived during the Sinemurian stage of the Early Jurassic approximately in the Lufeng Formation of the Lufeng Basin in what is now the Yunnan province in south-western China
(, paleocoordinates ...
'', whose fossils date from approximately 195 million years ago, in the early Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
, provides the first clear evidence of a jaw joint formed solely by the squamosal and dentary bones; there is no space in the jaw for the articular, a bone involved in the jaws of all early synapsids.
The earliest clear evidence of hair or fur is in fossils of ''Castorocauda
''Castorocauda'' is an extinct, semi-aquatic, superficially otter-like genus of docodont mammaliaforms with one species, ''C. lutrasimilis''. It is part of the Yanliao Biota, found in the Daohugou Beds of Inner Mongolia, China dating to the M ...
'' and ''Megaconus
''Megaconus'' is an extinct genus of allotherian mammal from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. The type and only species, ''Megaconus mammaliaformis'' was first described in the journal ''Nature'' in 2013. ''Meg ...
'', from 164 million years ago in the mid-Jurassic. In the 1950s, it was suggested that the foramina (passages) in the maxilla
The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
e and premaxilla
The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
e (bones in the front of the upper jaw) of cynodonts were channels which supplied blood vessels and nerves to vibrissae (whiskers
Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to touch, sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are c ...
) and so were evidence of hair or fur; it was soon pointed out, however, that foramina do not necessarily show that an animal had vibrissae, as the modern lizard ''Tupinambis
''Tupinambis'' is a lizard genus which belongs to the family Teiidae and contains eight described species. These large lizards are commonly referred to as tegus (''teiús'' in Portuguese). '' T. merianae'' (Argentine black and white tegu), '' T. ...
'' has foramina that are almost identical to those found in the nonmammalian cynodont ''Thrinaxodon
''Thrinaxodon'' is an extinct genus of cynodonts, most commonly regarded by its species ''T. liorhinus'' which lived in what are now South Africa and Antarctica during the Early Triassic. ''Thrinaxodon'' lived just after the Permian–Triassic m ...
''. Popular sources, nevertheless, continue to attribute whiskers to ''Thrinaxodon''. Studies on Permian coprolites
A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is de ...
suggest that non-mammalian synapsids
Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the Sauropsida, sauropsids, the group that inc ...
of the epoch already had fur, setting the evolution of hairs possibly as far back as dicynodont
Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typicall ...
s.
When endothermy
An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inste ...
first appeared in the evolution of mammals is uncertain, though it is generally agreed to have first evolved in non-mammalian therapsids
Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
.[ Modern monotremes have lower body temperatures and more variable metabolic rates than marsupials and placentals,] but there is evidence that some of their ancestors, perhaps including ancestors of the therians, may have had body temperatures like those of modern therians. Likewise, some modern therians like afrotheres and xenarthrans have secondarily developed lower body temperatures.
The evolution of erect limbs in mammals is incomplete—living and fossil monotremes have sprawling limbs. The parasagittal (nonsprawling) limb posture appeared sometime in the late Jurassic or early Cretaceous; it is found in the eutherian ''Eomaia'' and the metatherian ''Sinodelphys'', both dated to 125 million years ago. Epipubic
Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them).
They first occ ...
bones, a feature that strongly influenced the reproduction of most mammal clades, are first found in Tritylodontidae
Tritylodontidae ("three-knob teeth", named after the shape of their cheek teeth) is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, bearing several mammalian traits like erect limbs, endothermy and detail ...
, suggesting that it is a synapomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
between them and mammaliformes
Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent comm ...
. They are omnipresent in non-placental mammaliformes, though ''Megazostrodon
''Megazostrodon'' is an extinct genus of basal mammaliaforms belonging to the order Morganucodonta. It is approximately 200 million years old. '' and ''Erythrotherium
''Erythrotherium'' (meaning "red beast") is an extinct genus of basal mammaliaforms from the Late Triassic to Lower Jurassic. It is related to ''Morganucodon''. Only one species is recorded, ''Erythrotherium parringtoni'', from Red Beds, Stormber ...
'' appear to have lacked them.
It has been suggested that the original function of lactation
Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proces ...
(milk
Milk is a white liquid food produced by the mammary glands of mammals. It is the primary source of nutrition for young mammals (including breastfed human infants) before they are able to digestion, digest solid food. Immune factors and immune ...
production) was to keep eggs moist. Much of the argument is based on monotremes, the egg-laying mammals. In human females, mammary glands become fully developed during puberty, regardless of pregnancy.
Rise of the mammals
Therian mammals took over the medium- to large-sized ecological niches in the Cenozoic
The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
, after the Cretaceous–Paleogene extinction event
The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
approximately 66 million years ago emptied ecological space once filled by non-avian dinosaurs and other groups of reptiles, as well as various other mammal groups, and underwent an exponential increase in body size (megafauna
In terrestrial zoology, the megafauna (from Greek μέγας ''megas'' "large" and New Latin ''fauna'' "animal life") comprises the large or giant animals of an area, habitat, or geological period, extinct and/or extant. The most common threshold ...
). Then mammals diversified very quickly; both birds and mammals show an exponential rise in diversity. For example, the earliest known bat dates from about 50 million years ago, only 16 million years after the extinction of the non-avian dinosaurs.
Molecular phylogenetic studies initially suggested that most placental orders diverged about 100 to 85 million years ago and that modern families appeared in the period from the late Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
through the Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
. However, no placental fossils have been found from before the end of the Cretaceous. The earliest undisputed fossils of placentals comes from the early Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
, after the extinction of the non-avian dinosaurs. In particular, scientists have identified an early Paleocene animal named ''Protungulatum donnae
''Protungulatum'' ('first ungulate') is a extinct genus of pan-euungulate mammals within extinct family Protungulatidae, and also one of the earliest known placental mammals in the fossil record, that lived in North America from the Late Creta ...
'' as one of the first placental mammals. however it has been reclassified as a non-placental eutherian. Recalibrations of genetic and morphological diversity rates have suggested a Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
origin for placentals, and a Paleocene origin for most modern clades.
The earliest known ancestor of primates is ''Archicebus achilles
''Archicebus'' is a genus of fossil primates that lived in the early Eocene forests (~55.8–54.8 million years ago) of what is now Jingzhou in the Hubei Province in central China, discovered in 2003. The only known species, ''A. achil ...
'' from around 55 million years ago. This tiny primate weighed 20–30 grams (0.7–1.1 ounce) and could fit within a human palm.
Anatomy
Distinguishing features
Living mammal species can be identified by the presence of sweat gland
Sweat glands, also known as sudoriferous or sudoriparous glands, , are small tubular structures of the skin that produce sweat. Sweat glands are a type of exocrine gland, which are glands that produce and secrete substances onto an epithelial sur ...
s, including those that are specialized to produce milk to nourish their young. In classifying fossils, however, other features must be used, since soft tissue glands and many other features are not visible in fossils.
Many traits shared by all living mammals appeared among the earliest members of the group:
* Jaw joint – The dentary
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
(the lower jaw bone, which carries the teeth) and the squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone.
In most tetrapods, the squamosal and quadratojugal The quadratojugal is a skull bone present in many vertebrates, including ...
(a small cranial
Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
bone) meet to form the joint. In most gnathostomes
Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...
, including early therapsids
Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
, the joint consists of the articular
The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals.
Anatomy
In most vertebrates, the articular bone is connected to two oth ...
(a small bone at the back of the lower jaw) and quadrate
Quadrate may refer to:
* Quadrate bone
* Quadrate (heraldry)
* Quadrate lobe of liver
* Quadrate tubercle
The quadrate tubercle is a small tubercle found upon the upper part of the femur. It serves as a point of insertion of the quadratus femori ...
(a small bone at the back of the upper jaw).[
* ]Middle ear
The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear).
The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
– In crown-group mammals, sound is carried from the eardrum
In the anatomy of humans and various other tetrapods, the eardrum, also called the tympanic membrane or myringa, is a thin, cone-shaped membrane that separates the external ear
The outer ear, external ear, or auris externa is the extern ...
by a chain of three bones, the malleus
The malleus, or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibrations fro ...
, the incus
The ''incus'' (plural incudes) or anvil is a bone in the middle ear. The anvil-shaped small bone is one of three ossicles in the middle ear. The ''incus'' receives vibrations from the ''malleus'', to which it is connected laterally, and transmit ...
and the stapes
The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foot ...
. Ancestrally, the malleus and the incus are derived from the articular and the quadrate bones that constituted the jaw joint of early therapsids.
* Tooth replacement – Teeth can be replaced once (diphyodonty A diphyodont is any animal with two ss of teeth, initially the ''deciduous'' set and consecutively the ''permanent'' set. Most mammals are diphyodonts—as to chew their food they need a strong, durable and complete set of teeth.
Diphyodonts contra ...
) or (as in toothed whales and murid
In Sufism, a ''murīd'' (Arabic مُرِيد 'one who seeks') is a novice committed to spiritual enlightenment by ''sulūk'' (traversing a path) under a spiritual guide, who may take the title murshid, '' pir'' or ''shaykh''. A '' sālik'' or Su ...
rodents) not at all (monophyodont
A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, teari ...
y). Elephants, manatees, and kangaroos continually grow new teeth throughout their life (polyphyodont
A polyphyodont is any animal whose teeth are continually replaced. In contrast, diphyodonts are characterized by having only two successive sets of teeth.
Polyphyodonts include most toothed fishes, many reptiles such as crocodiles and geckos, and ...
y).
* Prismatic enamel – The enamel coating on the surface of a tooth consists of prisms, solid, rod-like structures extending from the dentin
Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
to the tooth's surface.
* Occipital condyle
The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra.
The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
s – Two knobs at the base of the skull fit into the topmost neck vertebra
In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
; most other tetrapod
Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s, in contrast, have only one such knob.
For the most part, these characteristics were not present in the Triassic ancestors of the mammals. Nearly all mammaliaforms possess an epipubic bone, the exception being modern placentals.[
]
Sexual dimorphism
On average, male mammals are larger than females, with males being at least 10% larger than females in over 45% of investigated species. Most mammalian orders also exhibit male-biased sexual dimorphism
Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, although some orders do not show any bias or are significantly female-biased (Lagomorpha
The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (hares and rabbits) and the Ochotonidae (pikas). The name of the order is derived from the Ancient Greek ''lagos'' (λαγ ...
). Sexual size dimorphism increases with body size across mammals ( Rensch's rule), suggesting that there are parallel selection pressures on both male and female size. Male-biased dimorphism relates to sexual selection on males through male–male competition for females, as there is a positive correlation between the degree of sexual selection, as indicated by mating systems, and the degree of male-biased size dimorphism. The degree of sexual selection is also positively correlated with male and female size across mammals. Further, parallel selection pressure on female mass is identified in that age at weaning is significantly higher in more polygynous
Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women.
Incidence
Polygyny is more widespread in Africa than in any ...
species, even when correcting for body mass. Also, the reproductive rate is lower for larger females, indicating that fecundity selection selects for smaller females in mammals. Although these patterns hold across mammals as a whole, there is considerable variation across orders.
Biological systems
The majority of mammals have seven cervical vertebrae
In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
(bones in the neck). The exceptions are the manatee
Manatees (family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing three of the four living species ...
and the two-toed sloth
''Choloepus'' is a genus of xenarthran mammals of Central and South America within the monotypic family Choloepodidae, consisting of two-toed sloths, sometimes also called two-fingered sloths. The two species of ''Choloepus'' (which means "lame ...
, which have six, and the three-toed sloth
The three-toed or three-fingered sloths are arboreal neotropical mammals . They are the only members of the genus ''Bradypus'' and the family Bradypodidae. The four living species of three-toed sloths are the brown-throated sloth, the maned sloth ...
which has nine. All mammalian brains possess a neocortex
The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
, a brain region unique to mammals. Placental brains have a corpus callosum
The corpus callosum (Latin for "tough body"), also callosal commissure, is a wide, thick nerve tract, consisting of a flat bundle of commissural fibers, beneath the cerebral cortex in the brain. The corpus callosum is only found in placental mam ...
, unlike monotremes and marsupials.
The lung
The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...
s of mammals are spongy and honeycombed. Breathing is mainly achieved with the diaphragm
Diaphragm may refer to:
Anatomy
* Thoracic diaphragm, a thin sheet of muscle between the thorax and the abdomen
* Pelvic diaphragm or pelvic floor, a pelvic structure
* Urogenital diaphragm or triangular ligament, a pelvic structure
Other
* Diap ...
, which divides the thorax from the abdominal cavity, forming a dome convex to the thorax. Contraction of the diaphragm flattens the dome, increasing the volume of the lung cavity. Air enters through the oral and nasal cavities, and travels through the larynx, trachea and bronchi
A bronchus is a passage or airway in the lower respiratory tract that conducts air into the lungs. The first or primary bronchi pronounced (BRAN-KAI) to branch from the trachea at the carina are the right main bronchus and the left main bronchus. ...
, and expands the alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit.
Uses in anatomy and zoology
* Pulmonary alveolus, an air sac in the lungs
** Alveolar cell or pneumocyte
** Alveolar duct
** Alveolar macrophage
* ...
. Relaxing the diaphragm has the opposite effect, decreasing the volume of the lung cavity, causing air to be pushed out of the lungs. During exercise, the abdominal wall contracts
A contract is a legally enforceable agreement between two or more parties that creates, defines, and governs mutual rights and obligations between them. A contract typically involves the transfer of goods, services, money, or a promise to tran ...
, increasing pressure on the diaphragm, which forces air out quicker and more forcefully. The rib cage
The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels.
The sternum, together known as the thoracic cage, is a semi- ...
is able to expand and contract the chest cavity through the action of other respiratory muscles. Consequently, air is sucked into or expelled out of the lungs, always moving down its pressure gradient.[ This type of lung is known as a bellows lung due to its resemblance to blacksmith ]bellows
A bellows or pair of bellows is a device constructed to furnish a strong blast of air. The simplest type consists of a flexible bag comprising a pair of rigid boards with handles joined by flexible leather sides enclosing an approximately airtigh ...
.
The mammalian heart
The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
has four chambers, two upper atria, the receiving chambers, and two lower ventricles, the discharging chambers. The heart has four valves, which separate its chambers and ensures blood flows in the correct direction through the heart (preventing backflow). After gas exchange
Gas exchange is the physical process by which gases move passively by Diffusion#Diffusion vs. bulk flow, diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liqui ...
in the pulmonary capillaries (blood vessels in the lungs), oxygen-rich blood returns to the left atrium via one of the four pulmonary vein
The pulmonary veins are the veins that transfer oxygenated blood from the lungs to the heart. The largest pulmonary veins are the four ''main pulmonary veins'', two from each lung that drain into the left atrium of the heart. The pulmonary vein ...
s. Blood flows nearly continuously back into the atrium, which acts as the receiving chamber, and from here through an opening into the left ventricle. Most blood flows passively into the heart while both the atria and ventricles are relaxed, but toward the end of the ventricular relaxation period, the left atrium will contract, pumping blood into the ventricle. The heart also requires nutrients and oxygen found in blood like other muscles, and is supplied via coronary arteries
The coronary arteries are the arterial blood vessels of coronary circulation, which transport oxygenated blood to the heart muscle. The heart requires a continuous supply of oxygen to function and survive, much like any other tissue or organ of ...
.
The integumentary system
The integumentary system is the set of organs forming the outermost layer of an animal's body. It comprises the skin and its appendages, which act as a physical barrier between the external environment and the internal environment that it serves ...
(skin) is made up of three layers: the outermost epidermis
The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
, the dermis
The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
and the hypodermis
The subcutaneous tissue (), also called the hypodermis, hypoderm (), subcutis, superficial fascia, is the lowermost layer of the integumentary system in vertebrates. The types of cells found in the layer are fibroblasts, adipose cells, and macro ...
. The epidermis is typically 10 to 30 cells thick; its main function is to provide a waterproof layer. Its outermost cells are constantly lost; its bottommost cells are constantly dividing and pushing upward. The middle layer, the dermis, is 15 to 40 times thicker than the epidermis. The dermis is made up of many components, such as bony structures and blood vessels. The hypodermis is made up of adipose tissue
Adipose tissue, body fat, or simply fat is a loose connective tissue composed mostly of adipocytes. In addition to adipocytes, adipose tissue contains the stromal vascular fraction (SVF) of cells including preadipocytes, fibroblasts, vascular e ...
, which stores lipids and provides cushioning and insulation. The thickness of this layer varies widely from species to species;[ ]marine mammal
Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their reli ...
s require a thick hypodermis (blubber
Blubber is a thick layer of vascularized adipose tissue under the skin of all cetaceans, pinnipeds, penguins, and sirenians.
Description
Lipid-rich, collagen fiber-laced blubber comprises the hypodermis and covers the whole body, except for pa ...
) for insulation, and right whale
Right whales are three species of large baleen whales of the genus ''Eubalaena'': the North Atlantic right whale (''E. glacialis''), the North Pacific right whale (''E. japonica'') and the Southern right whale (''E. australis''). They are clas ...
s have the thickest blubber at . Although other animals have features such as whiskers, feathers
Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier e ...
, setae
In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms.
Animal setae
Protostomes
Annelid setae are stiff bristles present on the body. Th ...
, or cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
that superficially resemble it, no animals other than mammals have hair
Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals.
The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
. It is a definitive characteristic of the class, though some mammals have very little.
Herbivores have developed a diverse range of physical structures to facilitate the consumption of plant material. To break up intact plant tissues, mammals have developed teeth
A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
structures that reflect their feeding preferences. For instance, frugivore
A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance an ...
s (animals that feed primarily on fruit) and herbivores that feed on soft foliage have low-crowned teeth specialized for grinding foliage and seed
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s. Grazing
In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and other ...
animals that tend to eat hard, silica
Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
-rich grasses, have high-crowned teeth, which are capable of grinding tough plant tissues and do not wear down as quickly as low-crowned teeth. Most carnivorous mammals have carnassial
Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
iforme teeth (of varying length depending on diet), long canines and similar tooth replacement patterns.
The stomach of even-toed ungulate
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
s (Artiodactyla) is divided into four sections: the rumen
The rumen, also known as a paunch, is the largest stomach compartment in ruminants and the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. The rumen's microbial favoring environment allow ...
, the reticulum
Reticulum is a small, faint constellation in the southern sky. Its name is Latin for a small net, or reticle—a net of crosshairs at the focus of a telescope eyepiece that is used to measure star positions. The constellation is best viewed bet ...
, the omasum
The omasum, also known as the bible, the fardel, the manyplies and the psalterium, is the third compartment of the stomach in ruminants. The omasum comes after the rumen and reticulum and before the abomasum. Different ruminants have different oma ...
and the abomasum
The abomasum, also known as the maw,[The Cham ...](_blank)
(only ruminant
Ruminants (suborder Ruminantia) are ungulate, hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principally ...
s have a rumen). After the plant material is consumed, it is mixed with saliva in the rumen and reticulum and separates into solid and liquid material. The solids lump together to form a bolus
Bolus may refer to:
Geography
* Bolus, Iran, a village in Ardabil Province, Iran
* Bolus, or Baulus, an Anatolian village on the site of ancient Berissa
Medicine
* Bolus (digestion), a ball-shaped mass moving through the digestive tract
* Bolus ...
(or cud
Cud is a portion of food that returns from a ruminant's stomach to the mouth to be chewed for the second time. More precisely, it is a bolus of semi-degraded food regurgitated from the reticulorumen of a ruminant. Cud is produced during the phy ...
), and is regurgitated. When the bolus enters the mouth, the fluid is squeezed out with the tongue and swallowed again. Ingested food passes to the rumen and reticulum where cellulolytic microbe
A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s (bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, protozoa
Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
and fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
) produce cellulase
Cellulase (EC 3.2.1.4; systematic name 4-β-D-glucan 4-glucanohydrolase) is any of several enzymes produced chiefly by fungi, bacteria, and protozoans that catalyze cellulolysis, the decomposition of cellulose and of some related polysaccharide ...
, which is needed to break down the cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
in plants. Perissodactyls
Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...
, in contrast to the ruminants, store digested food that has left the stomach in an enlarged cecum
The cecum or caecum is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix (anatomy), appendix, to which i ...
, where it is fermented by bacteria. Carnivora have a simple stomach adapted to digest primarily meat, as compared to the elaborate digestive systems of herbivorous animals, which are necessary to break down tough, complex plant fibers. The caecum
The cecum or caecum is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix, to which it is joined). The w ...
is either absent or short and simple, and the large intestine is not sacculated or much wider than the small intestine.
The mammalian excretory system
The excretory system is a passive biological system that removes excess, unnecessary materials from the body fluids of an organism, so as to help maintain internal chemical homeostasis and prevent damage to the body. The dual function of excreto ...
involves many components. Like most other land animals, mammals are ureotelic
Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds, w ...
, and convert ammonia
Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
into urea
Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid.
Urea serves an important r ...
, which is done by the liver
The liver is a major Organ (anatomy), organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of proteins and biochemicals necessary for ...
as part of the urea cycle
The urea cycle (also known as the ornithine cycle) is a cycle of biochemical reactions that produces urea (NH2)2CO from ammonia (NH3). Animals that use this cycle, mainly amphibians and mammals, are called ureotelic.
The urea cycle converts highl ...
. Bilirubin
Bilirubin (BR) (Latin for "red bile") is a red-orange compound that occurs in the normal catabolic pathway that breaks down heme in vertebrates. This catabolism is a necessary process in the body's clearance of waste products that arise from the ...
, a waste product derived from blood cell
A blood cell, also called a hematopoietic cell, hemocyte, or hematocyte, is a cell produced through hematopoiesis and found mainly in the blood. Major types of blood cells include red blood cells (erythrocytes), white blood cells (leukocytes), ...
s, is passed through bile
Bile (from Latin ''bilis''), or gall, is a dark-green-to-yellowish-brown fluid produced by the liver of most vertebrates that aids the digestion of lipids in the small intestine. In humans, bile is produced continuously by the liver (liver bile ...
and urine
Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excretion, excreted from the body through the urethra.
Cel ...
with the help of enzymes excreted by the liver. The passing of bilirubin via bile through the intestinal tract
The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and ...
gives mammalian feces
Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
a distinctive brown coloration. Distinctive features of the mammalian kidney include the presence of the renal pelvis
The renal pelvis or pelvis of the kidney is the funnel-like dilated part of the ureter in the kidney. It is formed by the covnvergence of the major calyces, acting as a funnel for urine flowing from the major calyces to the ureter. It has a mucous ...
and renal pyramid
The renal medulla is the innermost part of the kidney. The renal medulla is split up into a number of sections, known as the renal pyramids. Blood enters into the kidney via the renal artery, which then splits up to form the segmental arteries whi ...
s, and of a clearly distinguishable cortex
Cortex or cortical may refer to:
Biology
* Cortex (anatomy), the outermost layer of an organ
** Cerebral cortex, the outer layer of the vertebrate cerebrum, part of which is the ''forebrain''
*** Motor cortex, the regions of the cerebral cortex i ...
and medulla
Medulla or Medullary may refer to:
Science
* Medulla oblongata, a part of the brain stem
* Renal medulla, a part of the kidney
* Adrenal medulla, a part of the adrenal gland
* Medulla of ovary, a stroma in the center of the ovary
* Medulla of th ...
, which is due to the presence of elongated loops of Henle
In the kidney, the loop of Henle () (or Henle's loop, Henle loop, nephron loop or its Latin counterpart ''ansa nephroni'') is the portion of a nephron that leads from the proximal convoluted tubule to the distal convoluted tubule. Named after its ...
. Only the mammalian kidney has a bean shape, although there are some exceptions, such as the multilobed reniculate kidney The reniculate kidney is a multilobed kidney found in marine and aquatic mammals such as pinnipeds (seals, sea lions and walruses) and cetaceans (dolphins and whales) but absent in terrestrial mammals except bears. Kidneys of this morphology
Morph ...
s of pinnipeds, cetacea
Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
ns and bears.[ Most adult placental mammals have no remaining trace of the ]cloaca
In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
. In the embryo, the embryonic cloaca
The cloaca is a structure in the development of the urinary and reproductive organs.
The hind-gut is at first prolonged backward into the body-stalk as the tube of the allantois; but, with the growth and flexure of the tail-end of the embryo, ...
divides into a posterior region that becomes part of the anus, and an anterior region that has different fates depending on the sex of the individual: in females, it develops into the vestibule that receives the urethra
The urethra (from Greek οὐρήθρα – ''ourḗthrā'') is a tube that connects the urinary bladder to the urinary meatus for the removal of urine from the body of both females and males. In human females and other primates, the urethra con ...
and vagina
In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
, while in males it forms the entirety of the penile urethra.[ However, the ]tenrec
A tenrec is any species of mammal within the afrotherian family Tenrecidae endemic to Madagascar. Tenrecs are wildly diverse; as a result of convergent evolution some resemble hedgehogs, shrews, opossums, rats, and mice. They occupy aquatic, a ...
s, golden mole
Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae and as such they are taxonomically distinct from the true moles, family Talpidae, and other mole-like families, all o ...
s, and some shrew
Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
s retain a cloaca as adults. In marsupials, the genital tract is separate from the anus, but a trace of the original cloaca does remain externally. Monotremes, which translates from Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
into "single hole", have a true cloaca.
Sound production
As in all other tetrapods, mammals have a larynx
The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
that can quickly open and close to produce sounds, and a supralaryngeal vocal tract
The vocal tract is the cavity in human bodies and in animals where the sound produced at the sound source (larynx in mammals; syrinx (biology), syrinx in birds) is filtered.
In birds it consists of the Vertebrate trachea, trachea, the Syrinx (bio ...
which filters this sound. The lungs and surrounding musculature provide the air stream and pressure required to phonate
The term phonation has slightly different meanings depending on the subfield of phonetics. Among some phoneticians, ''phonation'' is the process by which the vocal folds produce certain sounds through quasi-periodic vibration. This is the defini ...
. The larynx controls the pitch and volume
Volume is a measure of occupied three-dimensional space. It is often quantified numerically using SI derived units (such as the cubic metre and litre) or by various imperial or US customary units (such as the gallon, quart, cubic inch). The de ...
of sound, but the strength the lungs exert to exhale
Exhalation (or expiration) is the flow of the breath out of an organism. In animals, it is the movement of air from the lungs out of the airways, to the external environment during breathing.
This happens due to elastic properties of the lungs, ...
also contributes to volume. More primitive mammals, such as the echidna, can only hiss, as sound is achieved solely through exhaling through a partially closed larynx. Other mammals phonate using vocal fold
The human voice consists of sound made by a human being
Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and exceptional cognitive skills due to a large and compl ...
s. The movement or tenseness of the vocal folds can result in many sounds such as purr
A purr is a tonal fluttering sound made by some species of felids and two species of genets. It varies in loudness and tone among species and in the same animal. Felids are a family of mammals that belong to the order Carnivora and are infor ...
ing and screaming
A scream is a loud speech production, vocalization in which air is passed through the vocal cords with greater force than is used in regular or close-distance vocalisation. This can be performed by any creature possessing lungs, including human ...
. 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
Nasal is an adjective referring to the nose, part of human or animal anatomy. It may also be shorthand for the following uses in combination:
* With reference to the human nose:
** Nasal administration, a method of pharmaceutical drug delivery
** ...
sounds; nasal sounds, such as a dog whine, are generally soft sounds, and oral sounds, such as a dog bark, are generally loud.[
Some mammals have a large larynx and thus a low-pitched voice, namely the ]hammer-headed bat
The hammer-headed bat ('), also known as hammer-headed fruit bat and big-lipped bat, is a megabat widely distributed in West and Central Africa. It is the only member of the genus ''Hypsignathus'', which is part of the tribe Epomophorini along w ...
(''Hypsignathus monstrosus'') where the larynx can take up the entirety of the thoracic cavity
The thoracic cavity (or chest cavity) is the chamber of the body of vertebrates that is protected by the thoracic wall (rib cage and associated skin, muscle, and fascia). The central compartment of the thoracic cavity is the mediastinum. There ...
while pushing the lungs, heart, and trachea into the abdomen
The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
. Large vocal pads can also lower the pitch, as in the low-pitched roars of big cat
The term "big cat" is typically used to refer to any of the five living members of the genus '' Panthera'', namely the tiger, lion, jaguar, leopard, and snow leopard.
Despite enormous differences in size, various cat species are quite similar ...
s. The production of infrasound
Infrasound, sometimes referred to as low status sound, describes sound waves with a frequency below the lower limit of human audibility (generally 20 Hz). Hearing becomes gradually less sensitive as frequency decreases, so for humans to perce ...
is possible in some mammals such as the African elephant (''Loxodonta'' spp.) and baleen whale
Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in their ...
s. Small mammals with small larynxes have the ability to produce ultrasound
Ultrasound is sound waves with frequency, frequencies higher than the upper audible limit of human hearing range, hearing. Ultrasound is not different from "normal" (audible) sound in its physical properties, except that humans cannot hea ...
, which can be detected by modifications to the middle ear
The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear).
The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
and cochlea
The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory org ...
. Ultrasound is inaudible to birds and reptiles, which might have been important during the Mesozoic, when birds and reptiles were the dominant predators. This private channel is used by some rodents in, for example, mother-to-pup communication, and by bats when echolocating. Toothed whales also use echolocation, but, as opposed to the vocal membrane that extends upward from the vocal folds, they have a melon
A melon is any of various plants of the family Cucurbitaceae with sweet, edible, and fleshy fruit. The word "melon" can refer to either the plant or specifically to the fruit. Botanically, a melon is a kind of berry, specifically a "pepo". The ...
to manipulate sounds. Some mammals, namely the primates, have air sacs attached to the larynx, which may function to lower the resonances or increase the volume of sound.
The vocal production system is controlled by the cranial nerve nuclei
A cranial nerve nucleus is a collection of neurons (gray matter) in the brain stem that is associated with one or more of the cranial nerves. Axons carrying information to and from the cranial nerves form a synapse first at these nuclei. Lesions ...
in the brain, and supplied by the recurrent laryngeal nerve
The recurrent laryngeal nerve (RLN) is a branch of the vagus nerve (cranial nerve X) that supplies all the intrinsic muscles of the larynx, with the exception of the cricothyroid muscles. There are two recurrent laryngeal nerves, right and le ...
and the superior laryngeal nerve
The superior laryngeal nerve is a branch of the vagus nerve. It arises from the middle of the inferior ganglion of vagus nerve and in its course receives a branch from the superior cervical ganglion of the sympathetic nervous system.
The superi ...
, branches of the vagus nerve
The vagus nerve, also known as the tenth cranial nerve, cranial nerve X, or simply CN X, is a cranial nerve that interfaces with the parasympathetic control of the heart, lungs, and digestive tract. It comprises two nerves—the left and right ...
. The vocal tract is supplied by the hypoglossal nerve
The hypoglossal nerve, also known as the twelfth cranial nerve, cranial nerve XII, or simply CN XII, is a cranial nerve that innervates all the extrinsic and intrinsic muscles of the tongue except for the palatoglossus, which is innervated by ...
and facial nerve
The facial nerve, also known as the seventh cranial nerve, cranial nerve VII, or simply CN VII, is a cranial nerve that emerges from the pons of the brainstem, controls the muscles of facial expression, and functions in the conveyance of tas ...
s. Electrical stimulation of the periaqueductal gray
The periaqueductal gray (PAG, also known as the central gray) is a brain region that plays a critical role in autonomic function, motivated behavior and behavioural responses to threatening stimuli. PAG is also the primary control center for d ...
(PEG) region of the mammalian midbrain
The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', " ...
elicit vocalizations. The ability to learn new vocalizations is only exemplified in humans, seals, cetaceans, elephants and possibly bats; in humans, this is the result of a direct connection between the motor cortex
The motor cortex is the region of the cerebral cortex believed to be involved in the planning, control, and execution of voluntary movements.
The motor cortex is an area of the frontal lobe located in the posterior precentral gyrus immediately a ...
, which controls movement, and the motor neuron
A motor neuron (or motoneuron or efferent neuron) is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly or indirectl ...
s in the spinal cord.[
]
Fur
The primary function of the fur of mammals is thermoregulation. Others include protection, sensory purposes, waterproofing, and camouflage. Different types of fur serve different purposes:[
* Definitive – which may be ]shed
A shed is typically a simple, single-story roofed structure that is used for hobbies, or as a workshop in a back garden or on an allotment. Sheds vary considerably in their size and complexity of construction, from simple open-sided ones de ...
after reaching a certain length
* Vibrissae – sensory hairs, most commonly whisker
Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are coarser ...
s
* Pelage – guard hairs, under-fur, and awn hair
Awn hairs are the intermediate hairs in a mammal's coat. They are shorter than the guard hairs and longer than the down hairs. They help with insulation and protect the down hairs underneath. Most of the visible coat is made of this kind of hair ...
* Spines – stiff guard hair used for defense (such as in porcupine
Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethizont ...
s)
* Bristle
A bristle is a stiff hair or feather (natural or artificial), either on an animal, such as a pig, a plant, or on a tool such as a brush or broom.
Synthetic types
Synthetic materials such as nylon are also used to make bristles in items such as br ...
s – long hairs usually used in visual signals. (such as a lion's mane)
* Velli – often called "down fur" which insulates newborn mammals
* Wool
Wool is the textile fibre obtained from sheep and other mammals, especially goats, rabbits, and camelids. The term may also refer to inorganic materials, such as mineral wool and glass wool, that have properties similar to animal wool.
As ...
– long, soft and often curly
Thermoregulation
Hair length is not a factor in thermoregulation: for example, some tropical mammals such as sloths have the same length of fur length as some arctic mammals but with less insulation; and, conversely, other tropical mammals with short hair have the same insulating value as arctic mammals. The denseness of fur can increase an animal's insulation value, and arctic mammals especially have dense fur; for example, the musk ox
Musk (Persian: مشک, ''Mushk'') is a class of aromatic substances commonly used as base notes in perfumery. They include glandular secretions from animals such as the musk deer, numerous plants emitting similar fragrances, and artificial sub ...
has guard hairs measuring as well as a dense underfur, which forms an airtight coat, allowing them to survive in temperatures of .[ Some desert mammals, such as camels, use dense fur to prevent solar heat from reaching their skin, allowing the animal to stay cool; a camel's fur may reach in the summer, but the skin stays at .][ ]Aquatic mammal
Aquatic and semiaquatic mammals are a diverse group of mammals that dwell partly or entirely in bodies of water. They include the various marine mammals who dwell in oceans, as well as various freshwater species, such as the European otter. Th ...
s, conversely, trap air in their fur to conserve heat by keeping the skin dry.[
]
Coloration
Mammalian coats are colored for a variety of reasons, the major selective pressures including camouflage
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
, sexual selection
Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
, communication, and thermoregulation. Coloration in both the hair and skin of mammals is mainly determined by the type and amount of melanin
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
; eumelanin
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amin ...
s for brown and black colors and pheomelanin
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
for a range of yellowish to reddish colors, giving mammals an earth tone
Earth tone is a color scheme with multiple meanings. In its narrowest sense, it refers to "any color containing some brown" – the color of ground or soil (earth). It can also refer to "natural colors" (colors found in nature) such as brown soil, ...
. Some mammals have more vibrant colors; certain monkeys such mandrill
The mandrill (''Mandrillus sphinx'') is a large Old World monkey native to west central Africa. It is one of the most colorful mammals in the world, with red and blue skin on its face and posterior. The species is sexually dimorphic, as males ...
s and vervet monkey
The vervet monkey (''Chlorocebus pygerythrus''), or simply vervet, is an Old World monkey of the family Cercopithecidae native to Africa. The term "vervet" is also used to refer to all the members of the genus ''Chlorocebus''. The five distinct ...
s, and opossums such as the Mexican mouse opossum
The Mexican mouse opossum (''Marmosa mexicana'') is a species of North American opossum in the family Didelphidae.
Description
''Marmosa mexicana'' is a small to moderate-sized reddish-brown marsupial, varying from bright to dull coloration. Ha ...
s and Derby's woolly opossum
Derby's woolly opossum (''Caluromys derbianus''), or the Central American woolly opossum, is an opossum found in deciduous and moist evergreen forests of Central America, from southern Mexico to western Ecuador and Colombia. It was first describ ...
s, have blue skin due to light diffraction in collagen
Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
fibers. Many sloths appear green because their fur hosts green algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
; this may be a symbiotic
Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
relation that affords camouflage
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
to the sloths.
Camouflage is a powerful influence in a large number of mammals, as it helps to conceal individuals from predators or prey. In arctic and subarctic mammals such as the arctic fox
The Arctic fox (''Vulpes lagopus''), also known as the white fox, polar fox, or snow fox, is a small fox native to the Arctic regions of the Northern Hemisphere and common throughout the Arctic tundra biome. It is well adapted to living in co ...
(''Alopex lagopus''), collared lemming
''Dicrostonyx'' is a genus of rodent in the family Cricetidae. It contains the collared lemmings. They are the only North American rodents that turn completely white in winter.
It contains the following species:
* Northern collared lemming (''Dic ...
(''Dicrostonyx groenlandicus''), stoat
The stoat (''Mustela erminea''), also known as the Eurasian ermine, Beringian ermine and ermine, is a mustelid native to Eurasia and the northern portions of North America. Because of its wide circumpolar distribution, it is listed as Least Conc ...
(''Mustela erminea''), and snowshoe hare
The snowshoe hare (''Lepus americanus''), also called the varying hare or snowshoe rabbit, is a species of hare found in North America. It has the name "snowshoe" because of the large size of its hind feet. The animal's feet prevent it from sink ...
(''Lepus americanus''), seasonal color change between brown in summer and white in winter is driven largely by camouflage. Some arboreal mammals, notably primates and marsupials, have shades of violet, green, or blue skin on parts of their bodies, indicating some distinct advantage in their largely arboreal
Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
habitat due to convergent evolution
Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.
Aposematism
Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
, 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
Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or ginge ...
and the powerful and aggressive honey badger
The honey badger (''Mellivora capensis''), also known as the ratel ( or ), is a mammal widely distributed in Africa, Southwest Asia, and the Indian subcontinent. Because of its wide range and occurrence in a variety of habitats, it is listed ...
. Coat color is sometimes sexually dimorphic
Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, as in many primate species. Differences in female and male coat color may indicate nutrition and hormone levels, important in mate selection. Coat color may influence the ability to retain heat, depending on how much light is reflected. Mammals with a darker colored coat can absorb more heat from solar radiation, and stay warmer, and some smaller mammals, such as vole
Voles are small rodents that are relatives of lemmings and hamsters, but with a stouter body; a longer, hairy tail; a slightly rounder head; smaller eyes and ears; and differently formed molars (high-crowned with angular cusps instead of low-c ...
s, have darker fur in the winter. The white, pigmentless fur of arctic mammals, such as the polar bear, may reflect more solar radiation directly onto the skin.[ The dazzling black-and-white striping of ]zebra
Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: the Grévy's zebra (''Equus grevyi''), plains zebra (''E. quagga''), and the mountain zebra (''E. zeb ...
s appear to provide some protection from biting flies.
Reproductive system
Mammals are solely gonochoric
In biology, gonochorism is a sexual system where there are only two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric.
Gonochorism c ...
(an animal is born with either male or female genitalia, as opposed to hermaphrodite
In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes.
Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
s where there is no such schism). In male placentals, the penis
A penis (plural ''penises'' or ''penes'' () is the primary sexual organ that male animals use to inseminate females (or hermaphrodites) during copulation. Such organs occur in many animals, both vertebrate and invertebrate, but males do n ...
is used both for urination and copulation. Depending on the species, an erection
An erection (clinically: penile erection or penile tumescence) is a physiological phenomenon in which the penis becomes firm, engorged, and enlarged. Penile erection is the result of a complex interaction of psychological, neural, vascular, ...
may be fueled by blood flow into vascular, spongy tissue or by muscular action. A penis may be contained in a prepuce
Prepuce , or as an adjective, preputial , refers to two homologous structures of male and female genitals:
*Clitoral hood, skin surrounding and protecting the head of the clitoris
*Foreskin, skin surrounding and protecting the head of the penis in ...
when not erect, and some placentals also have a penis bone (baculum
The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a bone found in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as the ...
). Marsupials typically have forked penises, while the echidna
Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the ...
penis generally has four heads with only two functioning. The testes
A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoster ...
of most mammals descend into the scrotum
The scrotum or scrotal sac is an anatomical male reproductive structure located at the base of the penis that consists of a suspended dual-chambered sac of skin and smooth muscle. It is present in most terrestrial male mammals. The scrotum cont ...
which is typically posterior to the penis but is often anterior in marsupials. Female mammals generally have a clitoris
The clitoris ( or ) is a female sex organ present in mammals, ostriches and a limited number of other animals. In humans, the visible portion – the glans – is at the front junction of the labia minora (inner lips), above the ope ...
, labia majora
The labia majora (singular: ''labium majus'') are two prominent longitudinal cutaneous folds that extend downward and backward from the mons pubis to the perineum. Together with the labia minora they form the labia of the vulva.
The labia majo ...
and labia minora
The labia minora (Latin for 'smaller lips', singular: ''labium minus'', 'smaller lip'), also known as the inner labia, inner lips, vaginal lips or nymphae are two flaps of skin on either side of the human vaginal opening in the vulva, situated be ...
on the outside, while the internal system contains paired oviduct
The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, o ...
s, 1–2 uteri
The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uteru ...
, 1–2 cervices
The cervix or cervix uteri (Latin, 'neck of the uterus') is the lower part of the uterus (womb) in the human female reproductive system. The cervix is usually 2 to 3 cm long (~1 inch) and roughly cylindrical in shape, which changes during ...
and a vagina
In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
. Marsupials have two lateral vaginas and a medial vagina. The "vagina" of monotremes is better understood as a "urogenital sinus". The uterine systems of placental mammals can vary between a duplex, where there are two uteri and cervices which open into the vagina, a bipartite, where two uterine horn
The uterine horns (cornua of uterus) are the points in the upper uterus where the fallopian tubes exit to meet the ovaries. They are one of the points of attachment for the round ligament of uterus (the other being the mons pubis). They also provi ...
s have a single cervix that connects to the vagina, a bicornuate
A bicornuate uterus or bicornate uterus (from the Latin ''cornū'', meaning "horn"), is a type of mullerian anomaly in the human uterus, where there is a deep indentation at the fundus (top) of the uterus.
Pathophysiology
A bicornuate uterus ...
, which consists where two uterine horns that are connected distally but separate medially creating a Y-shape, and a simplex, which has a single uterus.
The ancestral condition for mammal reproduction is the birthing of relatively undeveloped, either through direct vivipary
In plants, vivipary occurs when seeds or embryos begin to develop before they detach from the parent. Plants such as some Iridaceae and Agavoideae grow cormlets in the axils of their inflorescences. These fall and in favourable circumstances the ...
or a short period as soft-shelled eggs. This is likely due to the fact that the torso could not expand due to the presence of epipubic bones
Epipubic bones are a pair of bones projecting forward from the pelvic bones of modern marsupials, monotremes and fossil mammals like multituberculates, and even basal eutherians (the ancestors of placental mammals, who lack them).
They first occu ...
. The oldest demonstration of this reproductive style is with ''Kayentatherium
''Kayentatherium'' is an extinct genus of tritylodontid cynodonts that lived during the Early Jurassic. It is one of two tritylodonts from the Kayenta Formation of northern Arizona, United States.
''Kayentatherium'' means "Kayenta Beast", and is ...
'', which produced undeveloped perinate
A perinate is a member of a viviparous species from approximately one month before birth to one month after it.
See also
* Fetus
* Neonate
* Prenatal and perinatal psychology
Prenatal psychology can be seen as a part of developmental psychology, ...
s, but at much higher litter sizes than any modern mammal, 38 specimens. Most modern mammals are viviparous
Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
, giving birth to live young. However, the five species of monotreme, the platypus and the four species of echidna, lay eggs. The monotremes have a sex determination system
A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. Most organisms that create their offspring using sexual reproduction have two sexes.
In some species there are hermaphr ...
different from that of most other mammals. In particular, the sex chromosome
A sex chromosome (also referred to as an allosome, heterotypical chromosome, gonosome, heterochromosome, or idiochromosome) is a chromosome that differs from an ordinary autosome in form, size, and behavior. The human sex chromosomes, a typical ...
s of a platypus are more like those of a chicken than those of a therian mammal.
Viviparous mammals are in the subclass Theria; those living today are in the marsupial and placental infraclasses. Marsupials have a short gestation
Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
period, typically shorter than its estrous cycle
The estrous cycle (, originally ) is the set of recurring physiological changes that are induced by reproductive hormones in most mammalian therian females. Estrous cycles start after sexual maturity in females and are interrupted by anestrous p ...
and generally giving birth to a number of undeveloped newborns that then undergo further development; in many species, this takes place within a pouch-like sac, the marsupium, located in the front of the mother's abdomen
The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
. This is the plesiomorphic
In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades.
Plesiomorphy, symplesiomorphy, apomorphy, and ...
condition among viviparous mammals; the presence of epipubic bones in all non-placental mammals prevents the expansion of the torso needed for full pregnancy. Even non-placental eutherians probably reproduced this way. The placentals give birth to relatively complete and developed young, usually after long gestation periods. They get their name from the placenta
The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate mater ...
, which connects the developing fetus to the uterine wall to allow nutrient uptake. In placental mammals, the epipubic is either completely lost or converted into the baculum; allowing the torso to be able to expand and thus birth developed offspring.
The mammary gland
A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in primat ...
s of mammals are specialized to produce milk, the primary source of nutrition for newborns. The monotremes branched early from other mammals and do not have the nipple
The nipple is a raised region of tissue on the surface of the breast from which, in females, milk leaves the breast through the lactiferous ducts to feed an infant. The milk can flow through the nipple passively or it can be ejected by smooth mu ...
s seen in most mammals, but they do have mammary glands. The young lick the milk from a mammary patch on the mother's belly. Compared to placental mammals, the milk of marsupials changes greatly in both production rate and in nutrient composition, due to the underdeveloped young. In addition, the mammary glands have more autonomy allowing them to supply separate milks to young at different development stages. Lactose
Lactose is a disaccharide sugar synthesized by galactose and glucose subunits and has the molecular formula C12H22O11. Lactose makes up around 2–8% of milk (by mass). The name comes from ' (gen. '), the Latin word for milk, plus the suffix '' - ...
is the main sugar in placental mammal milk while monotreme and marsupial milk is dominated by oligosaccharide
An oligosaccharide (/ˌɑlɪgoʊˈsækəˌɹaɪd/; from the Greek ὀλίγος ''olígos'', "a few", and σάκχαρ ''sácchar'', "sugar") is a saccharide polymer containing a small number (typically two to ten) of monosaccharides (simple sugar ...
s. Weaning
Weaning is the process of gradually introducing an infant human or another mammal to what will be its adult diet while withdrawing the supply of its mother's milk.
The process takes place only in mammals, as only mammals produce milk. The infan ...
is the process in which a mammal becomes less dependent on their mother's milk and more on solid food.
Endothermy
Nearly all mammals are endothermic
In thermochemistry, an endothermic process () is any thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, H.P., Butler, L. J. (2015).''Principle of Modern Chemistry'', Brooks Cole. p. ...
("warm-blooded"). Most mammals also have hair to help keep them warm. Like birds, mammals can forage or hunt in weather and climates too cold for ectotherm
An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...
ic ("cold-blooded") reptiles and insects. Endothermy requires plenty of food energy, so mammals eat more food per unit of body weight than most reptiles. Small insectivorous mammals eat prodigious amounts for their size. A rare exception, the naked mole-rat
The naked mole-rat (''Heterocephalus glaber''), also known as the sand puppy, is a burrowing rodent native to the Horn of Africa and parts of Kenya, notably in Somali regions. It is closely related to the blesmols and is the only species in th ...
produces little metabolic heat, so it is considered an operational poikilotherm
A poikilotherm () is an animal whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important stressors is temperature change, which can lead to alterations in membrane ...
. Birds are also endothermic, so endothermy is not unique to mammals.
Species lifespan
Among mammals, species maximum lifespan varies significantly (for example the shrew
Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
has a lifespan of two years, whereas the oldest bowhead whale
The bowhead whale (''Balaena mysticetus'') is a species of baleen whale belonging to the family Balaenidae and the only living representative of the genus ''Balaena''. They are the only baleen whale endemic to the Arctic and subarctic waters, ...
is recorded to be 211 years). Although the underlying basis for these lifespan differences is still uncertain, numerous studies indicate that the ability to repair DNA damage is an important determinant of mammalian lifespan. In a 1974 study by Hart and Setlow, it was found that DNA excision repair capability increased systematically with species lifespan among seven mammalian species. Species lifespan was observed to be robustly correlated with the capacity to recognize DNA double-strand breaks as well as the level of the DNA repair protein Ku80
Ku80 is a protein that, in humans, is encoded by the ''XRCC5'' gene. Together, Ku70 and Ku80 make up the Ku heterodimer, which binds to DNA double-strand break ends and is required for the non-homologous end joining (NHEJ) pathway of DNA repair ...
. In a study of the cells from sixteen mammalian species, genes employed in DNA repair were found to be up-regulated in the longer-lived species. The cellular level of the DNA repair enzyme poly ADP ribose polymerase
Poly (ADP-ribose) polymerase (PARP) is a family of proteins involved in a number of cellular processes such as DNA repair, genomic stability, and programmed cell death.
Members of PARP family
The PARP family comprises 17 members (10 putative) ...
was found to correlate with species lifespan in a study of 13 mammalian species. Three additional studies of a variety of mammalian species also reported a correlation between species lifespan and DNA repair capability.
Locomotion
Terrestrial
Most vertebrates—the amphibians, the reptiles and some mammals such as humans and bears—are plantigrade
151px, Portion of a human skeleton, showing plantigrade habit
In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. T ...
, walking on the whole of the underside of the foot. Many mammals, such as cats and dogs, are digitigrade
In terrestrial vertebrates, digitigrade () locomotion is walking or running on the toes (from the Latin ''digitus'', 'finger', and ''gradior'', 'walk'). A digitigrade animal is one that stands or walks with its toes (metatarsals) touching the groun ...
, walking on their toes, the greater stride length allowing more speed. Digitigrade mammals are also often adept at quiet movement. Some animals such as horse
The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million y ...
s are unguligrade
Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
, walking on the tips of their toes. This even further increases their stride length and thus their speed. A few mammals, namely the great apes, are also known to walk on their knuckles, at least for their front legs. Giant anteater
The giant anteater (''Myrmecophaga tridactyla'') is an insectivorous mammal native to Central and South America. It is one of four living species of anteaters, of which it is the largest member. The only extant member of the genus ''Myrmecopha ...
s and platypuses are also knuckle-walkers. Some mammals are bipeds
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
, using only two limbs for locomotion, which can be seen in, for example, humans and the great apes. Bipedal species have a larger field of vision than quadrupeds, conserve more energy and have the ability to manipulate objects with their hands, which aids in foraging. Instead of walking, some bipeds hop, such as kangaroos and kangaroo rat
Kangaroo rats, small mostly nocturnal rodents of genus ''Dipodomys'', are native to arid areas of western North America. The common name derives from their bipedal form. They hop in a manner similar to the much larger kangaroo, but developed thi ...
s.
Animals will use different gaits for different speeds, terrain and situations. For example, horses show four natural gaits, the slowest horse gait
Horses can use various gaits (patterns of leg movement) during locomotion across solid ground, either naturally or as a result of specialized training by humans.Ensminger, M. E. ''Horses and Horsemanship'' 6th edition USA: Interstate Publisher ...
is the walk
Walking (also known as ambulation) is one of the main gaits of terrestrial locomotion among legged animals. Walking is typically slower than running and other gaits. Walking is defined by an 'inverted pendulum' gait in which the body vaults ov ...
, then there are three faster gaits which, from slowest to fastest, are the trot
The trot is a ten-beat diagonal horse gait where the diagonal pairs of legs move forward at the same time with a moment of suspension between each beat. It has a wide variation in possible speeds, but averages about . A very slow trot is someti ...
, the canter
The canter and gallop are variations on the fastest gait that can be performed by a horse or other equine. The canter is a controlled three-beat gait, while the gallop is a faster, four-beat variation of the same gait. It is a natural gait pos ...
and the gallop
The canter and gallop are variations on the fastest gait that can be performed by a horse or other equine. The canter is a controlled three-beat gait, while the gallop is a faster, four-beat variation of the same gait. It is a natural gait pos ...
. Animals may also have unusual gaits that are used occasionally, such as for moving sideways or backwards. For example, the main human gaits are bipedal walking
Walking (also known as ambulation) is one of the main gaits of terrestrial locomotion among legged animals. Walking is typically slower than running and other gaits. Walking is defined by an 'inverted pendulum' gait in which the body vaults o ...
and running
Running is a method of terrestrial locomotion allowing humans and other animals to move rapidly on foot. Running is a type of gait characterized by an aerial phase in which all feet are above the ground (though there are exceptions). This is ...
, but they employ many other gaits occasionally, including a four-legged crawl
Crawl, The Crawl, or crawling may refer to:
Biology
* Crawling (human), any of several types of human quadrupedal gait
* Limbless locomotion, the movement of limbless animals over the ground
* Undulatory locomotion, a type of motion characterize ...
in tight spaces. Mammals show a vast range of gait
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency. Di ...
s, the order that they place and lift their appendages in locomotion. Gaits can be grouped into categories according to their patterns of support sequence. For quadrupeds, there are three main categories: walking gaits, running gaits and leaping gaits
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency. Dif ...
. Walking is the most common gait, where some feet are on the ground at any given time, and found in almost all legged animals. Running is considered to occur when at some points in the stride all feet are off the ground in a moment of suspension.[
]
Arboreal
Arboreal animals frequently have elongated limbs that help them cross gaps, reach fruit or other resources, test the firmness of support ahead and, in some cases, to brachiate
Brachiation (from "brachium", Latin for "arm"), or arm swinging, is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb. ...
(swing between trees). Many arboreal species, such as tree porcupines, silky anteater
The silky anteater, also known as the pygmy anteater, has traditionally been considered a single species of anteater, ''Cyclopes didactylus'', in the genus ''Cyclopes'', the only living genus in the family Cyclopedidae. Found in southern Mexico, ...
s, spider monkeys, and possums
Possum may refer to:
Animals
* Phalangeriformes, or possums, any of a number of arboreal marsupial species native to Australia, New Guinea, and Sulawesi
** Common brushtail possum (''Trichosurus vulpecula''), a common possum in Australian urban a ...
, use prehensile tail
A prehensile tail is the tail of an animal that has adapted to grasp or hold objects. Fully prehensile tails can be used to hold and manipulate objects, and in particular to aid arboreal creatures in finding and eating food in the trees. If the ta ...
s to grasp branches. In the spider monkey, the tip of the tail has either a bare patch or adhesive pad, which provides increased friction. Claws can be used to interact with rough substrates and reorient the direction of forces the animal applies. This is what allows squirrel
Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
s to climb tree trunks that are so large to be essentially flat from the perspective of such a small animal. However, claws can interfere with an animal's ability to grasp very small branches, as they may wrap too far around and prick the animal's own paw. Frictional gripping is used by primates, relying upon hairless fingertips. Squeezing the branch between the fingertips generates frictional force that holds the animal's hand to the branch. However, this type of grip depends upon the angle of the frictional force, thus upon the diameter of the branch, with larger branches resulting in reduced gripping ability. To control descent, especially down large diameter branches, some arboreal animals such as squirrels have evolved highly mobile ankle joints that permit rotating the foot into a 'reversed' posture. This allows the claws to hook into the rough surface of the bark, opposing the force of gravity. Small size provides many advantages to arboreal species: such as increasing the relative size of branches to the animal, lower center of mass, increased stability, lower mass (allowing movement on smaller branches) and the ability to move through more cluttered habitat.[ Size relating to weight affects gliding animals such as the ]sugar glider
The sugar glider (''Petaurus breviceps'') is a small, omnivorous, arboreal, and nocturnal gliding possum belonging to the marsupial infraclass. The common name refers to its predilection for sugary foods such as sap and nectar and its ability ...
. Some species of primate, bat and all species of sloth
Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
achieve passive stability by hanging beneath the branch. Both pitching and tipping become irrelevant, as the only method of failure would be losing their grip.[
]
Aerial
Bats are the only mammals that can truly fly. They fly through the air at a constant speed by moving their wings up and down (usually with some fore-aft movement as well). Because the animal is in motion, there is some airflow relative to its body which, combined with the velocity of the wings, generates a faster airflow moving over the wing. This generates a lift force vector pointing forwards and upwards, and a drag force vector pointing rearwards and upwards. The upwards components of these counteract gravity, keeping the body in the air, while the forward component provides thrust to counteract both the drag from the wing and from the body as a whole.
The wings of bats are much thinner and consist of more bones than those of birds, allowing bats to maneuver more accurately and fly with more lift and less drag. By folding the wings inwards towards their body on the upstroke, they use 35% less energy during flight than birds. The membranes are delicate, ripping easily; however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly. The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cell
Merkel cells, also known as Merkel-Ranvier cells or tactile epithelial cells, are oval-shaped mechanoreceptors essential for light touch sensation and found in the skin of vertebrates. They are abundant in highly sensitive skin like that of the f ...
s, also found on human fingertips. These sensitive areas are different in bats, as each bump has a tiny hair in the center, making it even more sensitive and allowing the bat to detect and collect information about the air flowing over its wings, and to fly more efficiently by changing the shape of its wings in response.
Fossorial and subterranean
A fossorial (from Latin ''fossor'', meaning "digger") is an animal adapted to digging which lives primarily, but not solely, underground. Some examples are badger
Badgers are short-legged omnivores in the family Mustelidae (which also includes the otters, wolverines, martens, minks, polecats, weasels, and ferrets). Badgers are a polyphyletic rather than a natural taxonomic grouping, being united b ...
s, and naked mole-rat
The naked mole-rat (''Heterocephalus glaber''), also known as the sand puppy, is a burrowing rodent native to the Horn of Africa and parts of Kenya, notably in Somali regions. It is closely related to the blesmols and is the only species in th ...
s. Many rodent
Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
species are also considered fossorial because they live in burrows for most but not all of the day. Species that live exclusively underground are subterranean, and those with limited adaptations to a fossorial lifestyle sub-fossorial. Some organisms are fossorial to aid in temperature regulation
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
while others use the underground habitat for protection from predator
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s or for food storage
Food storage is a way of decreasing the variability of the food supply in the face of natural, inevitable variability. p.507 It allows food to be eaten for some time (typically weeks to months) after harvest rather than solely immediately. I ...
.[Damiani, R, 2003, Earliest evidence of cynodont burrowing, The Royal Society Publishing, Volume 270, Issue 1525]
Fossorial mammals have a fusiform body, thickest at the shoulders and tapering off at the tail and nose. Unable to see in the dark burrows, most have degenerated eyes, but degeneration varies between species; pocket gopher
Pocket gophers, commonly referred to simply as gophers, are burrowing rodents of the family Geomyidae. The roughly 41 speciesSearch results for "Geomyidae" on thASM Mammal Diversity Database are all endemic to North and Central America. They are ...
s, for example, are only semi-fossorial and have very small yet functional eyes, in the fully fossorial marsupial mole
Marsupial moles, the Notoryctidae , are highly specialized marsupial mammals, known from two species found at the Australian interior.
* ''Notoryctes typhlops'' (southern marsupial mole, known as the ''itjaritjari'' by the Pitjantjatjara and Yan ...
the eyes are degenerated and useless, talpa moles have vestigial
Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
eyes and the cape golden mole
The Cape golden mole (''Chrysochloris asiatica'') is a small, insectivorous mammal of the family Chrysochloridae, the golden mole
Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family ...
has a layer of skin covering the eyes. External ears flaps are also very small or absent. Truly fossorial mammals have short, stout legs as strength is more important than speed to a burrowing mammal, but semi-fossorial mammals have cursorial
A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often u ...
legs. The front paws are broad and have strong claws to help in loosening dirt while excavating burrows, and the back paws have webbing, as well as claws, which aids in throwing loosened dirt backwards. Most have large incisors to prevent dirt from flying into their mouth.
Many fossorial mammals such as shrews, hedgehogs, and moles were classified under the now obsolete order Insectivora
The order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order Eulipotyphla, w ...
.
Aquatic
Fully aquatic mammals, the cetaceans and sirenia
The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct f ...
ns, have lost their legs and have a tail fin to propel themselves through the water. Flipper movement is continuous. Whales swim by moving their tail fin and lower body up and down, propelling themselves through vertical movement, while their flippers are mainly used for steering. Their skeletal anatomy allows them to be fast swimmers. Most species have a dorsal fin
A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through conv ...
to prevent themselves from turning upside-down in the water. The flukes of sirenians are raised up and down in long strokes to move the animal forward, and can be twisted to turn. The forelimbs are paddle-like flippers which aid in turning and slowing.
Semi-aquatic
In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below.
Semiaquatic animals
Semiaquatic animals include:
* Verte ...
mammals, like pinnipeds, have two pairs of flippers on the front and back, the fore-flippers and hind-flippers. The elbows and ankles are enclosed within the body. Pinnipeds have several adaptions for reducing drag. In addition to their streamlined bodies, they have smooth networks of muscle bundles in their skin that may increase laminar flow
In fluid dynamics, laminar flow is characterized by fluid particles following smooth paths in layers, with each layer moving smoothly past the adjacent layers with little or no mixing. At low velocities, the fluid tends to flow without lateral mi ...
and make it easier for them to slip through water. They also lack arrector pili
The arrector pili muscles, also known as hair erector muscles, are small muscles attached to hair follicles in mammals. Contraction of these muscles causes the hairs to stand on end, known colloquially as goose bumps (piloerection).
Structure
E ...
, so their fur can be streamlined as they swim.[ They rely on their fore-flippers for locomotion in a wing-like manner similar to ]penguin
Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
s and sea turtles
Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhead, ...
. Fore-flipper movement is not continuous, and the animal glides between each stroke. Compared to terrestrial carnivorans, the fore-limbs are reduced in length, which gives the locomotor muscles at the shoulder and elbow joints greater mechanical advantage;[ the hind-flippers serve as stabilizers.] Other semi-aquatic mammals include beavers, hippopotamus
The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
es, otter
Otters are carnivorous mammals in the subfamily Lutrinae. The 13 extant otter species are all semiaquatic, aquatic, or marine, with diets based on fish and invertebrates. Lutrinae is a branch of the Mustelidae family, which also includes wea ...
s and platypuses. Hippos are very large semi-aquatic mammals, and their barrel-shaped bodies have graviportal skeletal structures, adapted to carrying their enormous weight, and their specific gravity
Relative density, or specific gravity, is the ratio of the density (mass of a unit volume) of a substance to the density of a given reference material. Specific gravity for liquids is nearly always measured with respect to water (molecule), wa ...
allows them to sink and move along the bottom of a river.
Behavior
Communication and vocalization
Many mammals communicate by vocalizing. Vocal communication serves many purposes, including in mating rituals, as warning calls, to indicate food sources, and for social purposes. Males often call during mating rituals to ward off other males and to attract females, as in the roaring
A roar is a type of Animal communication#Auditory, animal vocalization that is deep and acoustic resonance, resonating. Many mammals have evolved to produce roars and other roar-like vocals for purposes such as long-distance communication and intim ...
of lion
The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'' native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphi ...
s and red deer
The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or hart, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of wes ...
. The songs
A song is a musical composition intended to be performed by the human voice. This is often done at distinct and fixed pitches (melodies) using patterns of sound and silence. Songs contain various forms, such as those including the repetition ...
of the humpback whale may be signals to females; they have different dialects in different regions of the ocean. Social vocalizations include the territorial
A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal.
In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
calls of gibbon
Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical rainforest from eastern Bangladesh to Northeast India ...
s, and the use of frequency in greater spear-nosed bat
The greater spear-nosed bat (''Phyllostomus hastatus'') is a bat species of the family Phyllostomidae from South and Central America. It is one of the larger bats of this region and is omnivorous.
Habitat
''Phyllostomus hastatus'' lives in tr ...
s to distinguish between groups. The vervet monkey
The vervet monkey (''Chlorocebus pygerythrus''), or simply vervet, is an Old World monkey of the family Cercopithecidae native to Africa. The term "vervet" is also used to refer to all the members of the genus ''Chlorocebus''. The five distinct ...
gives a distinct alarm call for each of at least four different predators, and the reactions of other monkeys vary according to the call. For example, if an alarm call signals a python, the monkeys climb into the trees, whereas the eagle alarm causes monkeys to seek a hiding place on the ground. Prairie dogs
Prairie dogs (genus ''Cynomys'') are herbivorous burrowing ground squirrels native to the grasslands of North America. Within the genus are five species: black-tailed, white-tailed, Gunnison's, Utah, and Mexican prairie dogs. In Mexico, p ...
similarly have complex calls that signal the type, size, and speed of an approaching predator. Elephants communicate socially with a variety of sounds including snorting, screaming, trumpeting, roaring and rumbling. Some of the rumbling calls are infrasonic
Infrasound, sometimes referred to as low status sound, describes sound waves with a frequency below the lower limit of human audibility (generally 20 Hz). Hearing becomes gradually less sensitive as frequency decreases, so for humans to perce ...
, below the hearing range of humans, and can be heard by other elephants up to away at still times near sunrise and sunset.
left, Orca calling including occasional echolocation clicks
Mammals signal by a variety of means. Many give visual anti-predator signals, as when deer and gazelle
A gazelle is one of many antelope species in the genus ''Gazella'' . This article also deals with the seven species included in two further genera, ''Eudorcas'' and ''Nanger'', which were formerly considered subgenera of ''Gazella''. A third f ...
stot, honestly indicating their fit condition and their ability to escape, or when white-tailed deer
The white-tailed deer (''Odocoileus virginianus''), also known as the whitetail or Virginia deer, is a medium-sized deer native to North America, Central America, and South America as far south as Peru and Bolivia. It has also been introduced t ...
and other prey mammals flag with conspicuous tail markings when alarmed, informing the predator that it has been detected. Many mammals make use of scent-marking
In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species) using agonistic behaviors or (less commonly) real physical aggression. A ...
, sometimes possibly to help defend territory, but probably with a range of functions both within and between species. Microbat
Microbats constitute the suborder Microchiroptera within the order Chiroptera (bats). Bats have long been differentiated into Megachiroptera (megabats) and Microchiroptera, based on their size, the use of echolocation by the Microchiroptera a ...
s and toothed whale
The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales possessing teeth, such as the beaked whales and sperm whales. Seventy-three species of t ...
s including oceanic dolphin
Oceanic dolphins or Delphinidae are a widely distributed family of dolphins that live in the sea. Close to forty extant species are recognised. They include several big species whose common names contain "whale" rather than "dolphin", such as the ...
s vocalize both socially and in echolocation.
Feeding
To maintain a high constant body temperature is energy expensive—mammals therefore need a nutritious and plentiful diet. While the earliest mammals were probably predators, different species have since adapted to meet their dietary requirements in a variety of ways. Some eat other animals—this is a carnivorous
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
diet (and includes insectivorous diets). Other mammals, called herbivore
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s, eat plants, which contain complex carbohydrate
In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or may ...
s such as cellulose. An herbivorous diet includes subtypes such as granivory
Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed predators) feed on the seeds of plants as a main or exclusive food source,Hulme, P.E. and Benkman, C.W. (2002) "Granivory", pp. 132 ...
(seed eating), folivory
In zoology, a folivore is a herbivore that specializes in eating leaves. Mature leaves contain a high proportion of hard-to-digest cellulose, less energy than other types of foods, and often toxic compounds.Jones, S., Martin, R., & Pilbeam, D. (1 ...
(leaf eating), frugivory
A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance an ...
(fruit eating), nectarivory (nectar eating), gummivory A gummivore is an omnivorous animal whose diet consists primarily of the gums and saps of trees (about 90%) and bugs for protein. Notable gummivores include arboreal, terrestrial primates like certain marmosets and lemurs. These animals that live o ...
(gum eating) and mycophagy
Fungivory or mycophagy is the process of organisms consuming fungi. Many different organisms have been recorded to gain their energy from consuming fungi, including birds, mammals, insects, plants, amoebas, gastropods, nematodes, bacteria and othe ...
(fungus eating). The digestive tract of an herbivore is host to bacteria that ferment these complex substances, and make them available for digestion, which are either housed in the multichambered stomach
The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
or in a large cecum. Some mammals are coprophagous
Coprophagia () or coprophagy () is the consumption of feces. The word is derived from the grc, κόπρος , "feces" and , "to eat". Coprophagy refers to many kinds of feces-eating, including eating feces of other species (heterospecifics), of ...
, consuming feces
Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
to absorb the nutrients not digested when the food was first ingested.[ An ]omnivore
An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
eats both prey and plants. Carnivorous mammals have a simple digestive tract
The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans a ...
because the protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s, lipid
Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
s and mineral
In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. ( ...
s found in meat require little in the way of specialized digestion. Exceptions to this include baleen whale
Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in their ...
s who also house gut flora
Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut mi ...
in a multi-chambered stomach, like terrestrial herbivores.
The size of an animal is also a factor in determining diet type (Allen's rule
Allen's rule is an ecogeographical rule formulated by Joel Asaph Allen in 1877, broadly stating that animals adapted to cold climates have thicker limbs and bodily appendages than animals adapted to warm climates. More specifically, it states that ...
). Since small mammals have a high ratio of heat-losing surface area to heat-generating volume, they tend to have high energy requirements and a high metabolic rate
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
. Mammals that weigh less than about are mostly insectivorous because they cannot tolerate the slow, complex digestive process of an herbivore. Larger animals, on the other hand, generate more heat and less of this heat is lost. They can therefore tolerate either a slower collection process (carnivores that feed on larger vertebrates) or a slower digestive process (herbivores). Furthermore, mammals that weigh more than usually cannot collect enough insects during their waking hours to sustain themselves. The only large insectivorous mammals are those that feed on huge colonies of insects (ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s or termite
Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
s).
Some mammals are omnivores and display varying degrees of carnivory and herbivory, generally leaning in favor of one more than the other. Since plants and meat are digested differently, there is a preference for one over the other, as in bears where some species may be mostly carnivorous and others mostly herbivorous. They are grouped into three categories: mesocarnivory (50–70% meat), hypercarnivory
A hypercarnivore is an animal which has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
(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 hyena
Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the clas ...
s, allowing them to consume bones; some extinct groups, notably the Machairodontinae
Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million until ...
, had saber-shaped canines
Canine may refer to:
Zoology and anatomy
* a dog-like Canid animal in the subfamily Caninae
** ''Canis'', a genus including dogs, wolves, coyotes, and jackals
** Dog, the domestic dog
* Canine tooth, in mammalian oral anatomy
People with the surn ...
.
Some physiological carnivores consume plant matter and some physiological herbivores consume meat. From a behavioral aspect, this would make them omnivores, but from the physiological standpoint, this may be due to zoopharmacognosy
Zoopharmacognosy is a behaviour in which non-human animals self-medicate by selecting and ingesting or topically applying plants, soils, insects, and psychoactive drugs to prevent or reduce the harmful effects of pathogens, toxins, and even ...
. Physiologically, animals must be able to obtain both energy and nutrients from plant and animal materials to be considered omnivorous. Thus, such animals are still able to be classified as carnivores and herbivores when they are just obtaining nutrients from materials originating from sources that do not seemingly complement their classification. For example, it is well documented that some ungulates such as giraffes, camels, and cattle, will gnaw on bones to consume particular minerals and nutrients. Also, cats, which are generally regarded as obligate carnivores, occasionally eat grass to regurgitate indigestible material (such as hairball
A hairball is a small collection of hair or fur formed in the stomach of animals, and uncommonly in humans, that is occasionally vomited up when it becomes too big. Hairballs are primarily a tight elongated cylinder of packed fur, but may includ ...
s), aid with hemoglobin production, and as a laxative.
Many mammals, in the absence of sufficient food requirements in an environment, suppress their metabolism and conserve energy in a process known as hibernation
Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
. In the period preceding hibernation, larger mammals, such as bears, become polyphagic
Polyphagia or hyperphagia is an abnormally strong, incessant sensation of hunger or desire to eat often leading to overeating. In contrast to an increase in appetite following exercise, polyphagia does not subside after eating and often leads to ...
to increase fat stores, whereas smaller mammals prefer to collect and stash food. The slowing of the metabolism is accompanied by a decreased heart and respiratory rate, as well as a drop in internal temperatures, which can be around ambient temperature in some cases. For example, the internal temperatures of hibernating arctic ground squirrel
The Arctic ground squirrel (''Urocitellus parryii'') (Inuktitut: ''ᓯᒃᓯᒃ, siksik'') is a species of ground squirrel native to the Arctic and Subarctic of North America and Asia. People in Alaska, particularly around the Aleutians, refer to ...
s can drop to , however the head and neck always stay above . A few mammals in hot environments aestivate
Aestivation ( la, aestas (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a ...
in times of drought or extreme heat, for example the fat-tailed dwarf lemur
The fat-tailed dwarf lemur (''Cheirogaleus medius''), also known as the lesser dwarf lemur, western fat-tailed dwarf lemur, or spiny forest dwarf lemur, is endemic to Madagascar.
Hibernation
Recent research has shown that ''C. medius'' hibernat ...
(''Cheirogaleus medius'').
Intelligence
In intelligent mammals, such as primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s, the cerebrum
The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. In ...
is larger relative to the rest of the brain. Intelligence
Intelligence has been defined in many ways: the capacity for abstraction, logic, understanding, self-awareness, learning, emotional knowledge, reasoning, planning, creativity, critical thinking, and problem-solving. More generally, it can b ...
itself is not easy to define, but indications of intelligence include the ability to learn, matched with behavioral flexibility. Rats
Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' (pack rats), ''Bandicota'' (bandicoot ...
, for example, are considered to be highly intelligent, as they can learn and perform new tasks, an ability that may be important when they first colonize a fresh habitat
In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
. In some mammals, food gathering appears to be related to intelligence: a deer feeding on plants has a brain smaller than a cat, which must think to outwit its prey.
Tool use by animals
Tool use by animals is a phenomenon in which an animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, defence, communication, recreation or construction. Originally thought to be a skill possessed o ...
may indicate different levels of learning
Learning is the process of acquiring new understanding, knowledge, behaviors, skills, value (personal and cultural), values, attitudes, and preferences. The ability to learn is possessed by humans, animals, and some machine learning, machines ...
and cognition
Cognition refers to "the mental action or process of acquiring knowledge and understanding through thought, experience, and the senses". It encompasses all aspects of intellectual functions and processes such as: perception, attention, thought, ...
. The sea otter
The sea otter (''Enhydra lutris'') is a marine mammal native to the coasts of the northern and eastern North Pacific Ocean. Adult sea otters typically weigh between , making them the heaviest members of the weasel family, but among the small ...
uses rocks as essential and regular parts of its foraging behaviour (smashing abalone
Abalone ( or ; via Spanish , from Rumsen ''aulón'') is a common name for any of a group of small to very large marine gastropod molluscs in the family (biology), family Haliotidae. Other common name
In biology, a common name of a taxon o ...
from rocks or breaking open shells), with some populations spending 21% of their time making tools. Other tool use, such as chimpanzee
The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s using twigs to "fish" for termites, may be developed by watching others use tools and may even be a true example of animal teaching. Tools may even be used in solving puzzles in which the animal appears to experience a "Eureka moment". Other mammals that do not use tools, such as dogs, can also experience a Eureka moment.
Brain size
The size of the brain is a frequent topic of study within the fields of anatomy, biological anthropology, animal science and evolution. Brain size is sometimes measured by weight and sometimes by volume (via MRI scans or by skull volume). Neur ...
was previously considered a major indicator of the intelligence of an animal. Since most of the brain is used for maintaining bodily functions, greater ratios of Brain-to-body mass ratio, brain to body mass may increase the amount of brain mass available for more complex cognitive tasks. Allometric analysis indicates that mammalian brain size scales at approximately the or exponent of the body mass. Comparison of a particular animal's brain size with the expected brain size based on such allometric analysis provides an encephalization quotient, encephalisation quotient that can be used as another indication of animal intelligence. Sperm whales have the largest brain mass of any animal on earth, averaging and in mature males.
Self-awareness appears to be a sign of abstract thinking. Self-awareness, although not well-defined, is believed to be a precursor to more advanced processes such as metacognition, metacognitive reasoning. The traditional method for measuring this is the mirror test, which determines if an animal possesses the ability of self-recognition. Mammals that have passed the mirror test include Asian elephants (some pass, some do not); chimpanzees; bonobos; orangutans; human
Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
s, from 18 months (mirror stage); bottlenose dolphins killer whales; and false killer whales.[
]
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 and the Damaraland mole-rat.
Presociality is when animals exhibit more than just sexual interactions with members of the same species, but fall short of qualifying as eusocial. That is, presocial animals can display communal living, cooperative care of young, or primitive division of reproductive 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. Harry Harlow set up an experiment with rhesus monkeys, presocial primates, in 1958; the results from this study showed that social encounters are necessary in order for the young monkeys to develop both mentally and sexually.
A fission-fusion society is a society that changes frequently in its size and composition, making up a permanent social group called the "parent group". Permanent social networks consist of all individual members of a community and often varies to track changes in their environment. In a fission–fusion society, the main parent group can fracture (fission) into smaller stable subgroups or individuals to adapt to Social environment, environmental or social circumstances. For example, a number of males may break off from the main group in order to hunt or forage for food during the day, but at night they may return to join (fusion) the primary group to share food and partake in other activities. Many mammals exhibit this, such as primates (for example orangutans and spider monkeys), elephants, spotted hyenas, lions, and dolphins.
Solitary animals defend a territory and avoid social interactions with the members of its species, except during breeding season. This is to avoid resource competition, as two individuals of the same species would occupy the same niche, and to prevent depletion of food. A solitary animal, while foraging, can also be less conspicuous to predators or prey.
In a dominance hierarchy, hierarchy, individuals are either dominant or submissive. A despotic hierarchy is where one individual is dominant while the others are submissive, as in wolves and lemurs, and a pecking order is a linear ranking of individuals where there is a top individual and a bottom individual. Pecking orders may also be ranked by sex, where the lowest individual of a sex has a higher ranking than the top individual of the other sex, as in hyenas. Dominant individuals, or alphas, have a high chance of reproductive success, especially in harems
Harem (Persian: حرمسرا ''haramsarā'', ar, حَرِيمٌ ''ḥarīm'', "a sacred inviolable place; harem; female members of the family") refers to domestic spaces that are reserved for the women of the house in a Muslim family. A hare ...
where one or a few males (resident males) have exclusive breeding rights to females in a group. Non-resident males can also be accepted in harems, but some species, such as the common vampire bat (''Desmodus rotundus''), may be more strict.
Some mammals are perfectly Monogamy in animals, monogamous, meaning that they pair bond, mate for life and take no other partners (even after the original mate's death), as with wolves, Eurasian beavers, and otters. There are three types of polygamy: either one or multiple dominant males have breeding rights (polygyny in animals, 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 lek mating, leks, are estimated to occur in up to 90% of mammals. Lek mating occurs when males congregate around females and try to attract them with various courtship displays and vocalizations, as in harbor seals.
All higher mammals (excluding monotremes) share two major adaptations for care of the young: live birth and lactation. These imply a group-wide choice of a degree of parental care. They may build nests and dig burrows to raise their young in, or feed and guard them often for a prolonged period of time. Many mammals are K-selected, and invest more time and energy into their young than do r-selected animals. When two animals mate, they both share an interest in the success of the offspring, though often to different extremes. Mammalian females exhibit some degree of maternal aggression, another example of parental care, which may be targeted against other females of the species or the young of other females; however, some mammals may "aunt" the infants of other females, and care for them. Mammalian males may play a role in child rearing, as with tenrecs, however this varies species to species, even within the same genus. For example, the males of the southern pig-tailed macaque (''Macaca nemestrina'') do not participate in child care, whereas the males of the Japanese macaque (''M. fuscata'') do.
Humans and other mammals
In human culture
Non-human mammals play a wide variety of roles in human culture. They are the most popular of pets, with tens of millions of dogs, cats and other animals including rabbits and mice kept by families around the world. Mammals such as mammoths, horses and deer are among the earliest subjects of art, being found in Upper Paleolithic cave paintings such as at Lascaux. Major artists such as Albrecht Dürer, George Stubbs and Edwin Landseer are known for their portraits of mammals. Many species of mammals have been hunted for sport and for food; deer and wild boar are especially popular as game (hunting), game animals. Mammals such as horse racing, horses and greyhound racing, dogs are widely raced for sport, often combined with gambling, betting on the outcome. There is a tension between the role of animals as companions to humans, and their existence as individuals with animal rights, rights of their own. Mammals further play a wide variety of roles in literature, film, mythology, and religion.
Uses and importance
The domestication of mammals was instrumental in the Neolithic Revolution, Neolithic development of agriculture and of civilization
A civilization (or civilisation) is any complex society characterized by the development of a state, social stratification, urbanization, and symbolic systems of communication beyond natural spoken language (namely, a writing system).
Ci ...
, causing farmers to replace hunter-gatherers around the world. This transition from hunting and gathering to pastoralism, herding flocks and agriculture, growing crops was a major step in human history. The new agricultural economies, based on domesticated mammals, caused "radical restructuring of human societies, worldwide alterations in biodiversity, and significant changes in the Earth's landforms and its atmosphere... momentous outcomes".
Domestication, Domestic mammals form a large part of the livestock raised for meat
Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
across the world. They include (2009) around 1.4 billion cattle, 1 billion sheep, 1 billion domestic pigs, and (1985) over 700 million rabbits. Working animal, Working 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, again mainly in rural areas. In mountainous regions unsuitable for wheeled vehicles, pack animals continue to transport goods. Mammal skins provide leather
Leather is a strong, flexible and durable material obtained from the tanning, or chemical treatment, of animal skins and hides to prevent decay. The most common leathers come from cattle, sheep, goats, equine animals, buffalo, pigs and hogs, ...
for shoes, clothing and upholstery. Wool
Wool is the textile fibre obtained from sheep and other mammals, especially goats, rabbits, and camelids. The term may also refer to inorganic materials, such as mineral wool and glass wool, that have properties similar to animal wool.
As ...
from mammals including sheep, goats and alpacas has been used for centuries for clothing.
Mammals serve a major role in science as animal model, experimental animals, both in fundamental biological research, such as in genetics, and in the development of new medicines, which must be tested exhaustively to demonstrate their Pharmacovigilance, safety. Millions of mammals, especially mice and rats, are used in animal testing, experiments each year. A knockout mouse is a genetically modified mouse with an inactivated gene, replaced or disrupted with an artificial piece of DNA. They enable the study of sequencing, sequenced genes whose functions are unknown. A small percentage of the mammals are non-human primates, used in research for their similarity to humans.
Despite the benefits domesticated mammals had for human development, humans have an increasingly detrimental effect on wild mammals across the world. It has been estimated that the mass of all ''wild'' mammals has declined to only 4% of all mammals, with 96% of mammals being humans and their livestock now (see figure). In fact, terrestrial wild mammals make up only 2% of all mammals.
Hybrids
Hybrids are offspring resulting from the breeding of two genetically distinct individuals, which usually will result in a high degree of heterozygosity, though hybrid and heterozygous are not synonymous. The deliberate or accidental hybridizing of two or more species of closely related animals through captive breeding is a human activity which has been in existence for millennia and has grown for economic purposes. Hybrids between different subspecies within a species (such as between the Bengal tiger 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 goats) are known as intergeneric hybrids. Natural hybrids will occur in hybrid zones, where two populations of species within the same genera or species living in the same or adjacent areas will interbreed with each other. Some hybrids have been recognized as species, such as the red wolf (though this is controversial).
Artificial selection, the deliberate selective breeding of domestic animals, is being used to breeding back, breed back Holocene extinction, 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
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
. As a result, bred-back breeds are at best vague look-alikes of extinct wildtypes, as Heck cattle are of the aurochs.
Purebred wild species evolved to a specific ecology can be threatened with extinction through the process of genetic pollution, the uncontrolled hybridization, introgression genetic swamping which leads to homogenization or Fitness (biology), out-competition from the heterosis, heterosic hybrid species. When new populations are imported or selectively bred by people, or when habitat modification brings previously isolated species into contact, extinction in some species, especially rare varieties, is possible. Interbreeding can swamp the rarer gene pool and create hybrids, depleting the purebred gene pool. For example, the endangered wild water buffalo is most threatened with extinction by genetic pollution from the Water buffalo, domestic water buffalo. Such extinctions are not always apparent from a morphology (biology), morphological standpoint. Some degree of gene flow is a normal evolutionary process, nevertheless, hybridization threatens the existence of rare species.
Threats
The loss of species from ecological communities, defaunation, is primarily driven by human activity.[ This has resulted in empty forests, ecological communities depleted of large vertebrates.] In the Quaternary extinction event, the mass die-off of megafaunal variety coincided with the appearance of humans, suggesting a human influence. One hypothesis is that humans hunted large mammals, such as the woolly mammoth, into extinction. The 2019 ''Global Assessment Report on Biodiversity and Ecosystem Services'' by IPBES states that the total Biomass (ecology), biomass of wild mammals has declined by 82 percent since the beginning of human civilization. Wild animals make up just 4% of mammalian Biomass (ecology), biomass on earth, while humans and their domesticated animals make up 96%.
Various species are predicted to List of critically endangered species, become extinct in the near future, among them the rhinoceros, giraffes, and species of primates and pangolins. According to the WWF's 2020 ''Living Planet Report'', vertebrate wildlife populations have declined by 68% since 1970 as a result of human activities, particularly overconsumption, population growth and intensive farming, which is evidence that humans have triggered a sixth mass extinction event. Hunting alone threatens hundreds of mammalian species around the world. Scientists claim that the growing demand for meat
Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
is contributing to biodiversity loss as this is a significant driver of deforestation
Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
and habitat destruction
Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
; species-rich habitats, such as significant portions of the Amazon rainforest, are being converted to agricultural land for meat production. Another influence is over-hunting and Species affected by poaching, poaching, which can reduce the overall population of game animals, especially those located near villages, as in the case of peccary, peccaries. The effects of poaching can especially be seen in the ivory trade with African elephants. Marine mammals are at risk from entanglement from fishing gear, notably Cetacean bycatch, cetaceans, with discard mortalities ranging from 65,000 to 86,000 individuals annually.
Attention is being given to endangered species globally, notably through the Convention on Biological Diversity, otherwise known as the Rio Accord, which includes 189 signatory countries that are focused on identifying endangered species and habitats. Another notable conservation organization is the IUCN, which has a membership of over 1,200 governmental and Non-governmental organization, non-governmental organizations.
List of recently extinct mammals, Recent extinctions can be directly attributed to human influences. The IUCN characterizes 'recent' extinction as those that have occurred past the cut-off point of 1500, and around 80 mammal species have gone extinct since that time and 2015. Some species, such as the Père David's deer are extinct in the wild, and survive solely in captive populations. Other species, such as the Florida panther, are Ecological extinction, ecologically extinct, surviving in such low numbers that they essentially have no impact on the ecosystem. Other populations are only Local extinction, locally extinct (extirpated), still existing elsewhere, but reduced in distribution,[ as with the extinction of gray whales in the Atlantic Ocean, Atlantic.]
Notes
See also
* List of mammal genera – living mammals
* List of mammalogists
* List of monotremes and marsupials
* List of placental mammals
* List of prehistoric mammals
* List of threatened mammals of the United States
* Lists of mammals by population, Lists of mammals by population size
* Lists of mammals by region
* Mammals described in the 2000s
* Mammals in culture
* Small mammal
References
Further reading
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External links
Biodiversitymapping.org – All mammal orders in the world with distribution maps
Paleocene Mammals
a site covering the rise of the mammals, paleocene-mammals.de
Evolution of Mammals
a brief introduction to early mammals, enchantedlearning.com
European Mammal Atlas EMMA
from Societas Europaea Mammalogica, European-mammals.org
Marine Mammals of the World
– An overview of all marine mammals, including descriptions, both fully aquatic and semi-aquatic, noaa.gov
Mammalogy.org
The American Society of Mammalogists was established in 1919 for the purpose of promoting the study of mammals, and this website includes a mammal image library
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Mammals,
Bathonian first appearances
Extant Middle Jurassic first appearances
Taxa named by Carl Linnaeus