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The
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of
mammals 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 o ...
has passed through many stages since the first appearance of their
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 ...
ancestors in the Pennsylvanian sub-period of the late
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 ''Carboniferou ...
period. By the mid-
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 per ...
, there were many synapsid species that looked like mammals. The lineage leading to today's mammals split up in the
Jurassic The Jurassic ( ) is a geologic period and 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 Jurassic constitutes the middle period of ...
; synapsids from this period include '' Dryolestes'', more closely related to extant
placental 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 ...
s and
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 ...
s than to
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 brai ...
s, as well as '' Ambondro'', more closely related to monotremes. Later on, 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 ...
n and
metatheria 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 w ...
n lineages separated; the metatherians are the animals more closely related to the marsupials, while the eutherians are those more closely related to the placentals. Since '' Juramaia'', the earliest known eutherian, lived 160 million years ago in the Jurassic, this divergence must have occurred in the same period. 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 ...
wiped out the non-avian dinosaurs (
birds 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 ...
being the only surviving dinosaurs) and several mammalian groups, placental and marsupial mammals diversified into many new forms and ecological niches throughout the
Paleogene The Paleogene ( ; also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period million years ago ( Mya) to the beginning o ...
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. ...
, by the end of which all modern
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 ...
had appeared. The synapsid lineage became distinct from 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 syn ...
lineage in the late Carboniferous period, between 320 and 315 million years ago. The only living synapsids are mammals, while the sauropsids gave rise to the
dinosaurs 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 the ...
, and today's reptiles and birds along with all the extinct amniotes more closely related to them than to mammals. Primitive synapsids were traditionally called
mammal-like reptiles 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 ...
or
pelycosaur 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 ...
s, but both are now seen as outdated and disfavored
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
terms, since they were not reptiles, nor part of reptile lineage. The modern term for these is stem mammals, and sometimes protomammals or paramammals. Throughout the
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 Paleo ...
period, the synapsids included the dominant
carnivore 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 s ...
s and several important
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 mouthpar ...
s. In the subsequent Triassic period, however, a previously obscure group of sauropsids, the
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-avia ...
s, became the dominant vertebrates. The mammaliaforms appeared during this period; their superior sense of smell, backed up by a large brain, facilitated entry into nocturnal niches with less exposure to archosaur predation. The nocturnal lifestyle may have contributed greatly to the development of mammalian traits such as
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 ...
and
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 fi ...
. Later in the
Mesozoic 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 of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
, after
theropod dinosaurs Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
replaced rauisuchians as the dominant carnivores, mammals spread into other
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s. For example, some became aquatic, some were gliders, and some even fed on juvenile dinosaurs. Most of the evidence consists of
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
. For many years, fossils of Mesozoic mammals and their immediate ancestors were very rare and fragmentary; but, since the mid-1990s, there have been many important new finds, especially in China. The relatively new techniques of
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
have also shed light on some aspects of mammalian evolution by estimating the timing of important divergence points for modern species. When used carefully, these techniques often, but not always, agree with the fossil record. Although mammary glands are a signature feature of modern mammals, little is known about the evolution 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 ...
as these soft tissues are not often preserved in the fossil record. Most research concerning the evolution of mammals centers on the shapes of the teeth, the hardest parts of the
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 t ...
body. Other important research characteristics include the evolution of the middle ear bones, erect limb posture, 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 separ ...
, fur, hair, and
warm-blooded Warm-blooded is an informal term referring to animal species which can maintain a body temperature higher than their environment. In particular, homeothermic species maintain a stable body temperature by regulating metabolic processes. The on ...
ness.


Definition of "mammal"

While living mammal species can be identified by the presence of milk-producing
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 ...
s in the females, other features are required when classifying
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
, because mammary glands and other soft-tissue features are not visible in fossils. One such feature available for
paleontology 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 ...
, shared by all living mammals (including monotremes), but not present in any of the early
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 per ...
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, is shown in Figure 1 (on the right), namely: mammals use
two bones 2 (two) is a number, numeral and digit. It is the natural number following 1 and preceding 3. It is the smallest and only even prime number. Because it forms the basis of a duality, it has religious and spiritual significance in many cultur ...
for hearing that all other
amniotes 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 dis ...
use for eating. The earliest amniotes had a jaw joint composed 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 the quadrate (a small bone at the back of the upper jaw). All non-mammalian
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 t ...
s use this system including
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
s,
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 t ...
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 altho ...
s,
snake Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more ...
s,
crocodilia 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 livi ...
ns,
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 the
birds 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 ...
),
ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, alt ...
s,
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 ...
s and therapsids. But mammals have a different jaw joint, composed only of 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
(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 bones form the cheek series of the skull. The bone forms an ancestral co ...
(another small skull bone). In the Jurassic, their quadrate and articular bones evolved into the incus and malleus bones in 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 ...
.Mammalia: Overview – Palaeos
Mammals also have a double
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 ...
; they have two knobs at the base of the skull that fit into the topmost neck vertebra, while other tetrapods have a single occipital condyle. In a 1981 article, Kenneth A. Kermack and his co-authors argued for drawing the line between mammals and earlier synapsids at the point where the mammalian pattern of molar occlusion was being acquired and the dentary-squamosal joint had appeared. The criterion chosen, they noted, is merely a matter of convenience; their choice was based on the fact that "the lower jaw is the most likely skeletal element of a Mesozoic mammal to be preserved." Today, most paleontologists consider that animals are mammals if they satisfy this criterion.


The ancestry of mammals


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, with ...
s were
amniotes 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 dis ...
— their eggs had internal membranes that allowed 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 kept water in. This allowed amniotes to lay eggs on dry land, while amphibians generally need to lay their eggs in water (a few amphibians, such as the common Suriname toad, have
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...
other ways of getting around this limitation). The first amniotes apparently arose in the middle
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 ''Carboniferou ...
from the ancestral reptiliomorphs. Carroll R.L. (1991): The origin of reptiles. In: Schultze H.-P., Trueb L., (ed) ''Origins of the higher groups of tetrapods — controversy and consensus''. Ithaca: Cornell University Press, pp 331-353. Within a few million years, two important amniote lineages became distinct:
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, from which mammals are descended, and
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 syn ...
s, from which
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 altho ...
s, snakes, turtles/tortoises, crocodilians, dinosaurs, and birds are descended. The earliest known fossils of synapsids and sauropsids (such as '' Archaeothyris'' and ''
Hylonomus ''Hylonomus'' (; ''hylo-'' "forest" + ''nomos'' "dweller") is an extinct genus of reptile that lived 312 million years ago during the Late Carboniferous period. It is the earliest unquestionable reptile (''Westlothiana'' is older, but in fact it ...
'', respectively) date from about 320 to 315 million years ago. The times of origin are difficult to know, because vertebrate fossils from the late Carboniferous are very rare, and therefore the actual first occurrences of each of these types of animal might have been considerably earlier than the first fossil.


Synapsids

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 ...
skulls are identified by the distinctive pattern of the holes behind each eye, which served the following purposes: *made the skull lighter without sacrificing strength. *saved energy by using less bone. *probably provided attachment points for jaw muscles. Having attachment points further away from the jaw made it possible for the muscles to be longer and therefore to exert a strong pull over a wide range of jaw movement without being stretched or contracted beyond their optimum range. A number of creatures often - and incorrectly - believed to be
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, hence part of the reptile lineage and sauropsids, were in fact synapsids. This includes the well-known
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 Sphenacodont ...
.


Terms used for discussing non-mammalian synapsids

When referring to the ancestors and close relatives of mammals, paleontologists also use the following terms of convenience: *
Pelycosaur 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 ...
s — all synapsids, and all of their descendants, except for
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 - the eventual ancestor of mammals. The pelycosaurs included the largest land vertebrates of the
Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * 01'' (Richard Müller album), 2001 * ''01'' (Son of Dave album), 2000 * ''01'' (Urban Zakapa album), 2011 * ''O1'' (Hiroyuki Sawan ...
, such as the 6 m (20 ft) long '' Cotylorhynchus hancocki''. Among the other large pelycosaurs were '' Dimetrodon grandis'' and '' Edaphosaurus cruciger''. * Stem mammals (sometimes called protomammals or paramammals, and previously incorrectly called mammal-like reptiles) — all synapsids, and all of their descendants, except for mammals themselves. Stem mammals therefore include all pelycosaurs, and also all non-mammalian therapsids. Traditionally, these were known as "mammal-like reptiles", but this is incorrect and outdated; terms such as "stem mammal" are preferred instead, because these synapsids were neither reptiles, nor even part of reptile lineage. Pelycosaur and "mammal-like reptile" are both
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
terms (and in the latter case also based on incorrect historical belief that mammals evolved from reptiles rather than in parallel to them), and for that reason are disfavored and outdated terms rarely used in modern literature.


Therapsids

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 descended from
sphenacodonts Sphenacodontia is a stem-based clade of derived synapsids. It was defined by Amson and Laurin (2011) as "the largest clade that includes ''Haptodus baylei'', ''Haptodus garnettensis'' and '' Sphenacodon ferox'', but not ''Edaphosaurus pogonias' ...
, a primitive synapsid, 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 ± ...
, and took over from them as the dominant land vertebrates. They differ from earlier synapsids in several features of the skull and jaws, including larger
temporal fenestrae The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
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, wher ...
s that are equal in size. The therapsid lineage then went through several stages, leading to the evolution of
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 varie ...
s in the late Permian, some of which had begun to resemble early mammals: *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 separ ...
. Most books and articles interpret this as a prerequisite for the evolution of mammals' 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 ...
, because it enabled these animals to eat and breathe at the same time. But some scientists point out that some modern
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 ...
s use a fleshy secondary palate to separate the mouth from the airway, and that a ''bony'' palate provides a surface on which the tongue can manipulate food, facilitating chewing rather than breathing. The interpretation of the bony secondary palate as an aid to chewing also suggests the development of a faster metabolism, because chewing reduces the size of food particles delivered to the stomach and can therefore speed their digestion. In mammals, the palate is formed by two specific bones, but various Permian therapsids had other combinations of bones in the right places to function as a palate. *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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
gradually becomes the main bone of the lower jaw. *progress 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 erratic and very slow — for example: all herbivorous therapsids retained sprawling limbs (some late forms may have had semi-erect hind limbs); Permian carnivorous therapsids had sprawling forelimbs, and some late Permian ones also had semi-sprawling hindlimbs. In fact, modern
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 brai ...
s still have semi-sprawling forelimbs.


Therapsid family tree

(Simplified from Palaeos.com. Only those that are most relevant to the evolution of mammals are described below.) Only the dicynodonts, therocephalians, and cynodonts survived into the Triassic.


Biarmosuchia

The
Biarmosuchia Biarmosuchians are an extinct clade of non-mammalian synapsids from the Permian. They are the most basal group of the therapsids. All of them were moderately-sized, lightly-built carnivores, intermediate in form between basal sphenacodont " pelyc ...
were the most primitive and pelycosaur-like of the therapsids.


Dinocephalians

Dinocephalia Dinocephalians (terrible heads) are a clade of large-bodied early therapsids that flourished in the Early and Middle Permian between 279.5 and 260 million years ago (Ma), but became extinct during the Capitanian mass extinction event. Dinocephal ...
ns ("terrible heads") included both carnivores and herbivores. They were large; '' Anteosaurus'' was up to 6 m (20 ft) long. Some of the carnivores had semi-erect hindlimbs, but all dinocephalians had sprawling forelimbs. In many ways they were very primitive therapsids; for example, they had no secondary palate and their jaws were rather "reptilian".


Anomodonts

The
anomodonts Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores.Chinsamy-Turan, A. (2011) ''Forerunners of Mammals: R ...
("anomalous teeth") were among the most successful of the herbivorous therapsids — one sub-group, the
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, typic ...
s, survived almost to the end of the Triassic. But anomodonts were very different from modern herbivorous mammals, as their only teeth were a pair of fangs in the upper jaw and it is generally agreed that they had beaks like those of birds or
ceratopsian Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period (geology), Period, although anc ...
s.


Theriodonts

The
theriodonts The theriodonts ( clade Theriodontia) are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts. Naming In 1876, Ric ...
("beast teeth") and their descendants had jaw joints in which 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 ...
bone of the lower jaw tightly gripped the very small quadrate bone of the skull. This allowed a much wider gape and allowed one group, the carnivorous
gorgonopsia Gorgonopsia (from the Greek Gorgon, a mythological beast, and 'aspect') is an extinct clade of sabre-toothed therapsids from the Middle to Upper Permian roughly 265 to 252 million years ago. They are characterised by a long and narrow skull, a ...
ns ("gorgon faces"), to develop "sabre teeth". However, the jaw hinge of the theriodont had a longer term significance — the much reduced size of the quadrate bone was an important step in the development of the mammalian jaw joint and middle ear. The gorgonopsians still had some primitive features: no bony secondary palate (other bones in the right places perform the same functions); sprawling forelimbs; hindlimbs that could operate in both sprawling and erect postures. The
therocephalia Therocephalia is an extinct suborder of eutheriodont therapsids (mammals and their close relatives) from the Permian and Triassic. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of thei ...
ns ("beast heads"), which appear to have arisen at about the same time as the gorgonopsians, had additional mammal-like features, e.g. their finger and toe bones had the same number of phalanges (segments) as in early mammals (and the same number that
primates 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 ...
have, including humans).


Cynodonts

The cynodonts, a theriodont group that also arose in the late
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 Paleo ...
, include the ancestors of all mammals. Cynodonts' mammal-like features include further reduction in the number of bones in the lower jaw, a secondary bony palate, cheek teeth with a complex pattern in the crowns, and a brain which filled the endocranial cavity. Multi-chambered burrows have been found, containing as many as 20 skeletons of the Early Triassic cynodont ''
Trirachodon ''Trirachodon'' (Greek: "three ridge tooth") is an extinct genus of cynodonts. Fossils have been found in the ''Cynognathus'' Assemblage Zone of the Beaufort Group in South Africa and the Omingonde Formation of Namibia, dating back to the Early ...
''; the animals are thought to have been drowned by a
flash flood A flash flood is a rapid flooding of low-lying areas: washes, rivers, dry lakes and depressions. It may be caused by heavy rain associated with a severe thunderstorm, hurricane, or tropical storm, or by meltwater from ice or snow flowing ...
. The extensive shared burrows indicate that these animals were capable of complex social behaviors. Their primitive synapsid and therapsid ancestors were very large (between ) but cynodonts gradually decreased in size (to ) even before the Permian-Triassic extinction event, probably due to competition with other therapsids. After the extinction event, the
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 m ...
n cynodont group rapidly decreased in size (to 4 in–1.5 ft (10–50 cm)) due to new competition with archosaurs and transitioned to
nocturnality Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
, evolving nocturnal features,
pulmonary alveoli A pulmonary alveolus (plural: alveoli, from Latin ''alveolus'', "little cavity"), also known as an air sac or air space, is one of millions of hollow, distensible cup-shaped cavities in the lungs where oxygen is exchanged for carbon dioxide. Al ...
,
bronchioles The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start ...
and a developed
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 ...
for a larger surface area for breathing, enucleated
erythrocytes Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek ''erythros'' for "red" and ''kytos'' for "holl ...
, a
large intestine The large intestine, also known as the large bowel, is the last part of the gastrointestinal tract and of the digestive system in tetrapods. Water is absorbed here and the remaining waste material is stored in the rectum as feces before bein ...
which bears a true colon after the
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, to which it is joined). The wo ...
,
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 ...
, a hairy, glandular and thermoregulatory skin (which releases
sebum A sebaceous gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in the greatest nu ...
and
sweat Perspiration, also known as sweating, is the production of fluids secreted by the sweat glands in the skin of mammals. Two types of sweat glands can be found in humans: eccrine glands and apocrine glands. The eccrine sweat glands are distrib ...
), and a 4-chambered
heart The heart is a muscular Organ (biology), 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 ca ...
to maintain their high metabolism, larger brains, and fully upright hindlimb (forelimbs remained semi sprawling, and became like that only later, in therians). Some skin glands may have evolved into mammary glands in females for fulfilling the metabolic demands of their offspring (which increased 10 times). Many skeletal changes occurred also: 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
bone became stronger and held differentiated teeth, the pair of nasal openings in the skull became fused. These evolutionary changes lead to the first
mammals 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 o ...
(size around ). They also adapted to a
burrowing An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of s ...
lifestyle, losing their big tail-based leg muscles which allowed dinosaurs to become bipedal, and explains why bipedal mammals are so rare.


Triassic takeover

The catastrophic mass extinction at the end of the Permian, around 252 million years ago, killed off about 70 percent of 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, with ...
species and the majority of land plants. As a result,
ecosystems An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
and food chains collapsed, and the establishment of new stable ecosystems took about 30 million years. With the disappearance of the gorgonopsians, which were dominant predators in the late Permian, the cynodonts' principal competitors for dominance of the carnivorous niches were a previously obscure sauropsid group, the
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-avia ...
s, which includes the ancestors of crocodilians and dinosaurs. The archosaurs quickly became the dominant carnivores, a development often called the "Triassic takeover". Their success may have been due to the fact that the early Triassic was predominantly arid and therefore archosaurs' superior water conservation gave them a decisive advantage. All known archosaurs have glandless skins and eliminate
nitrogenous waste 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, ...
in a
uric acid Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates, such as ammonium acid urate. Uric acid is a product of the metabolic breakdown ...
paste containing little water, while the cynodonts probably excreted most such waste in a solution of
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 ...
, as mammals do today; considerable water is required to keep urea dissolved. However, this theory has been questioned, since it implies synapsids were necessarily less advantaged in water retention, that synapsid decline coincides with climate changes or archosaur diversity (neither of which has been tested) and the fact that desert-dwelling mammals are as well adapted in this department as archosaurs, and some cynodonts like '' Trucidocynodon'' were large-sized predators. The Triassic takeover was probably a vital factor in the evolution of the mammals. Two groups stemming from the early cynodonts were successful in niches that had minimal competition from the archosaurs: the
tritylodonts 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 details o ...
, which were
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 mouthpar ...
s, and the mammals, most of which were small 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 were ...
s (although some, like '' Sinoconodon'', were carnivores that fed on vertebrate prey, while still others were herbivores or
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 nu ...
s). As a result: *The therapsid trend towards differentiated teeth with precise occlusion accelerated, because of the need to hold captured
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s and crush their
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
s. *As the body length of the mammals' ancestors fell below 10.5 cm (4 inches), advances in
thermal insulation Thermal insulation is the reduction of heat transfer (i.e., the transfer of thermal energy between objects of differing temperature) between objects in thermal contact or in range of radiative influence. Thermal insulation can be achieved with ...
and temperature regulation would have become necessary for nocturnal life. *Acute senses of hearing and smell became vital. **This accelerated the development of the mammalian middle ear (though the complete detachment of the middle ear bones from the jaw happened independently in monotremes and therians). **The increase in the size of the olfactory lobes of the brain increased brain weight as a percentage of total body weight. Brain tissue requires a disproportionate amount of energy. The need for more food to support the enlarged brains increased the pressures for improvements in insulation, temperature regulation and feeding. *Probably as a side-effect of the nocturnal life, mammals lost two of the four cone
opsins Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become Retinylidene proteins, but are usually still called opsins regardless. Most pro ...
, photoreceptors in the
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which the ...
, present in the eyes of the earliest amniotes. Paradoxically, this might have improved their ability to discriminate colors in dim light. This retreat to a nocturnal role is called a nocturnal bottleneck, and is thought to explain many of the features of mammals.


From cynodonts to crown mammals


Fossil record

Mesozoic synapsids that had evolved to the point of having a jaw joint composed of the dentary and squamosal bones are preserved in few good fossils, mainly because they were mostly smaller than rats: *They were largely restricted to environments that are less likely to provide good
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
.
Floodplains A floodplain or flood plain or bottomlands is an area of land adjacent to a river which stretches from the banks of its channel to the base of the enclosing valley walls, and which experiences flooding during periods of high discharge.Goudi ...
as the best terrestrial environments for fossilization provide few mammal fossils, because they are dominated by medium to large animals, and the mammals could not compete with archosaurs in the medium to large size range. *Their delicate bones were vulnerable to being destroyed before they could be fossilized — by scavengers (including
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 fr ...
and
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 am ...
) and by being trodden on. *Small fossils are harder to spot and more vulnerable to being destroyed by weathering and other natural stresses before they are discovered. In the past years, however, the number of Mesozoic fossil mammals has increased decisively; only 116 genera were known in 1979, for example, but about 310 in 2007, with an increase in quality such that "at least 18 Mesozoic mammals are represented by nearly complete skeletons".


Mammals or mammaliaforms

Some writers restrict the term "mammal" to 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 ...
mammals, the group consisting of the most recent common ancestor of the monotremes,
marsupials 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 ...
, and placentals, together with all the descendants of that ancestor. In an influential 1988 paper, Timothy Rowe advocated this restriction, arguing that "ancestry... provides the only means of properly defining taxa" and, in particular, that the divergence of the monotremes from the animals more closely related to marsupials and placentals "is of central interest to any study of Mammalia as a whole." To accommodate some related taxa falling outside the crown group, he defined the
Mammaliaformes 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 c ...
as comprising "the last common ancestor of Morganucodontidae and Mammalia s he had defined the latter termand all its descendants." Besides Morganucodontidae, the newly defined taxon includes Docodonta and Kuehneotheriidae. Though haramiyids have been referred to the mammals since the 1860s, Rowe excluded them from the Mammaliaformes as falling outside his definition, putting them in a larger clade, the Mammaliamorpha. Some writers have adopted this terminology noting, to avoid misunderstanding, that they have done so. Most paleontologists, however, still think that animals with the dentary-squamosal jaw joint and the sort of molars characteristic of modern mammals should formally be members of Mammalia. Where the ambiguity in the term "mammal" may be confusing, this article uses "mammaliaform" and "crown mammal".


Family tree – cynodonts to crown group mammals

(based o
Cynodontia:Dendrogram – Palaeos


Morganucodontidae

The
Morganucodon ''Morganucodon'' ("Glamorgan tooth") is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, ''Morganucodon'' is well represent ...
tidae first appeared in the late Triassic, about 205 million years ago. They are an excellent example of transitional fossils, since they have both the dentary-squamosal and articular-quadrate jaw joints. They were also one of the first discovered and most thoroughly studied of the mammaliaforms outside of the crown-group mammals, since an unusually large number of morganucodont fossils have been found.


Docodonts

Docodonts, among the most common Jurassic mammaliaforms, are noted for the sophistication of their molars. They are thought to have had general semi-aquatic tendencies, with the fish-eating ''
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 ...
'' ("beaver tail"), which lived in the mid-
Jurassic The Jurassic ( ) is a geologic period and 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 Jurassic constitutes the middle period of ...
about 164M years ago and was first discovered in 2004 and described in 2006, being the most well-understood example. ''Castorocauda'' was not a crown group mammal, but it is extremely important in the study of the evolution of mammals because the first find was an almost complete skeleton (a real luxury in paleontology) and it breaks the "small nocturnal insectivore" stereotype: See also the news item at *It was noticeably larger than most Mesozoic mammaliaform fossils — about from its nose to the tip of its tail, and may have weighed . *It provides the earliest absolutely certain evidence of hair and fur. Previously the earliest was '' Eomaia'', a crown group mammal from about 125M years ago. *It had aquatic adaptations including flattened tail bones and remnants of soft tissue between the toes of the back feet, suggesting that they were webbed. Previously the earliest known semi-aquatic mammaliaforms were from the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
, about 110M years later. *''Castorocaudas powerful forelimbs look adapted for digging. This feature and the spurs on its ankles make it resemble the
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or mono ...
, which also swims and digs. *Its teeth look adapted for eating fish: the first two molars had cusps in a straight row, which made them more suitable for gripping and slicing than for grinding; and these molars are curved backwards, to help in grasping slippery prey.


''Hadrocodium''

The family tree above shows ''
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 (, paleocoordina ...
'' as an "aunt" of crown mammals. This mammaliaform, dated about 195M years ago in the very early Jurassic, exhibits some important features: *The jaw joint consists only of the squamosal and dentary bones, and the jaw contains no smaller bones to the rear of the dentary, unlike the therapsid design. *In
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 mo ...
and early mammaliaforms 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 from the middle ear. Its function is to transmit sound from the ...
may have stretched over a trough at the rear of the lower jaw. But ''Hadrocodium'' had no such trough, which suggests its ear was part of the
cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, as it is in crown-group mammals — and hence that the former
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 ...
and quadrate had migrated to the middle ear and become the malleus and incus. On the other hand, the dentary has a "bay" at the rear that mammals lack. This suggests that ''Hadrocodium's'' dentary bone retained the same shape that it would have had if the articular and quadrate had remained part of the jaw joint, and therefore that ''Hadrocodium'' or a very close ancestor may have been the first to have a fully mammalian middle ear. *Therapsids and earlier mammaliaforms had their jaw joints very far back in the skull, partly because the ear was at the rear end of the jaw but also had to be close to the brain. This arrangement limited the size of the braincase, because it forced the jaw muscles to run round and over it. ''Hadrocodium's'' braincase and jaws were no longer bound to each other by the need to support the ear, and its jaw joint was further forward. In its descendants or those of animals with a similar arrangement, the brain case was free to expand without being constrained by the jaw and the jaw was free to change without being constrained by the need to keep the ear near the brain — in other words it now became possible for mammaliaforms both to develop large brains and to adapt their jaws and teeth in ways that were purely specialized for eating.


Earliest crown mammals

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 ...
mammals, sometimes called 'true mammals', are the
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, ...
mammals and their relatives back to their last common ancestor. Since this group has living members, DNA analysis can be applied in an attempt to explain the evolution of features that do not appear in fossils. This endeavor often involves
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, a technique that has become popular since the mid-1980s.


Family tree of early crown mammals

Cladogram after Z.-X Luo. († marks extinct groups)


Color vision

Early amniotes had four
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become Retinylidene proteins, but are usually still called opsins regardless. Most ...
s in the cones of their retinas to use for distinguishing colours: one sensitive to red, one to green, and two corresponding to different shades of blue. The green opsin was not inherited by any crown mammals, but all normal individuals did inherit the red one. Early crown mammals thus had three cone opsins, the red one and both of the blues. All their extant descendants have lost one of the blue-sensitive opsins but not always the same one: monotremes retain one blue-sensitive opsin, while marsupials and placentals retain the other (except
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 th ...
ns, which later lost the other blue opsin as well). Some placentals and marsupials, including higher primates, subsequently evolved green-sensitive opsins; like early crown mammals, therefore, their vision is trichromatic.


Australosphenida and Ausktribosphenidae

Ausktribosphenidae is a group name that has been given to some rather puzzling finds that: *appear to have
tribosphenic molar The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
s, a type of tooth that is otherwise known only in placentals and marsupials. *come from mid-
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 ...
deposits in Australia — but Australia was connected only to Antarctica, and placentals originated in the Northern Hemisphere and were confined to it until
continental drift Continental drift is the hypothesis that the Earth's continents have moved over geologic time relative to each other, thus appearing to have "drifted" across the ocean bed. The idea of continental drift has been subsumed into the science of pl ...
formed land connections from North America to South America, from Asia to Africa and from Asia to India (the late Cretaceous ma
here
shows how the southern continents are separated). *are represented only by teeth and jaw fragments, which is not very helpful. Australosphenida is a group that has been defined in order to include the Ausktribosphenidae and monotremes. ''Asfaltomylos'' (mid- to late Jurassic, from
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and g ...
) has been interpreted as a basal australosphenid (animal that has features shared with both Ausktribosphenidae and monotremes; lacks features that are peculiar to Ausktribosphenidae or monotremes; also lacks features that are absent in Ausktribosphenidae and monotremes) and as showing that australosphenids were widespread throughout Gondwanaland (the old Southern Hemisphere super-continent). Recent analysis of ''
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 somewhere between 121 and 112.5 million years ago, suggests that it was a "crown group" (advanced and relatively specialised) monotreme. This was taken as evidence that the basal (most primitive) monotremes must have appeared considerably earlier, but this has been disputed (see the following section). The study also indicated that some alleged Australosphenids were also "crown group" monotremes (e.g. '' Steropodon'') and that other alleged Australosphenids (e.g. ''Ausktribosphenos'', ''Bishops'', ''Ambondro'', ''Asfaltomylos'') are more closely related to and possibly members of the Therian mammals (group that includes marsupials and placentals, see below).


Monotremes

''
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 ...
'', from Australia, is the earliest known monotreme. A 2007 study (published 2008) suggests that it was not a basal (primitive, ancestral) monotreme but a full-fledged
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or mono ...
, and therefore that the platypus and
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 ...
lineages diverged considerably earlier. A more recent study (2009), however, has suggested that, while ''Teinolophos'' was a type of platypus, it was also a basal monotreme and predated the radiation of modern monotremes. The semi-aquatic lifestyle of platypuses prevented them from being outcompeted by the marsupials that migrated to Australia millions of years ago, since joeys need to remain attached to their mothers and would drown if their mothers ventured into water (though there are exceptions like the water opossum and the
lutrine opossum ''Lutreolina'' is a genus of opossum found in South America. Both extant species in this genus are known as lutrine opossums. They have an otter-like body plan and occasionally semiaquatic tendencies, hence the genus name ''Lutreolina'', which i ...
; however, they both live in South America and thus don't come into contact with monotremes). Genetic evidence has determined that echidnas diverged from the platypus lineage as recently as 19-48M, when they made their transition from semi-aquatic to terrestrial lifestyle. Monotremes have some features that may be inherited from the
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 varie ...
ancestors: *like lizards and birds, they use the same orifice to urinate, defecate and reproduce ("monotreme" means "one hole"). *they lay eggs that are leathery and uncalcified, like those of lizards, turtles and crocodilians. Unlike other mammals, female monotremes do not have nipples and feed their young by "sweating" milk from patches on their bellies. These features are not visible in fossils, and the main characteristics from paleontologists' point of view are: *a slender
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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
bone in which the coronoid process is small or non-existent. *the external opening of the ear lies at the posterior base of the jaw. *the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
bone is small or non-existent. *a primitive
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of ...
with strong
ventral 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 prov ...
elements: coracoids, clavicles and
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals (marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In t ...
. Note:
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. ...
n mammals have no interclavicle. *sprawling or semi-sprawling forelimbs.


Multituberculates

Multituberculates (named for the multiple
tubercles In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection, ...
on their "
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
") are often called the "rodents of the Mesozoic", but this is an example of
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 ...
rather than meaning that they are closely related to the
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 n ...
ia. They existed for approximately 120 million years—the longest fossil history of any mammal lineage—but were eventually outcompeted by rodents, becoming extinct during the early
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but t ...
. Some authors have challenged the phylogeny represented by the cladogram above. They exclude the multituberculates from the mammalian crown group, holding that multituberculates are more distantly related to extant mammals than even the Morganucodontidae. Multituberculates are like undisputed crown mammals in that their jaw joints consist of only 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
and
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 bones form the cheek series of the skull. The bone forms an ancestral co ...
bones-whereas the quadrate and
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 ...
bones are part of the middle ear; their teeth are differentiated, occlude, and have mammal-like
cusp A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifurc ...
s; they have a
zygomatic arch In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
; and the structure of the
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
suggests that they gave birth to tiny helpless young, like modern marsupials. On the other hand, they differ from modern mammals: *Their "molars" have two parallel rows of tubercles, unlike the tribosphenic (three-peaked) molars of uncontested early crown mammals. *The chewing action differs in that undisputed crown mammals chew with a side-to-side grinding action, which means that the molars usually occlude on only one side at a time, while multituberculates' jaws were incapable of side-to-side movement—they chewed, rather, by dragging the lower teeth backwards against the upper ones as the jaw closed. *The anterior (forward) part of the zygomatic arch mostly consists of 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. T ...
(upper jawbone) rather than the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
, a small bone in a little slot in the maxillary process (extension). *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 bones form the cheek series of the skull. The bone forms an ancestral co ...
does not form part of the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
. *The rostrum (snout) is unlike that of undisputed crown mammals; in fact it looks more like that of a
pelycosaur 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 ...
, 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 Sphenacodont ...
''. The multituberculate rostrum is box-like, with the large flat maxillae forming the sides, the
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 * ...
the top, and the tall
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 ...
at the front.


Theria

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. ...
("beasts") is 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 ter ...
originating with the last common ancestor of 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 ...
(including placentals) and
Metatheria 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 w ...
(including marsupials). Common features include: *no
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals (marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In t ...
. * coracoid bones non-existent or fused with the
shoulder blades The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
to form
coracoid process The coracoid process (from Greek κόραξ, raven) is a small hook-like structure on the lateral edge of the superior anterior portion of the scapula (hence: coracoid, or "like a raven's beak"). Pointing laterally forward, it, together with t ...
es. *a type of crurotarsal ankle joint in which: the main joint is between the
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
and
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
; the
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. S ...
has no contact with the tibia but forms a heel to which muscles can attach. (The other well-known type of crurotarsal ankle is seen in crocodilians and works differently — most of the bending at the ankle is between the calcaneum and astragalus). *tribosphenic molars.


Metatheria

The living
Metatheria 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 w ...
are all marsupials (animals with pouches). A few fossil
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 ...
, such as the Mongolian late Cretaceous ''Asiatherium'', may be marsupials or members of some other metatherian group(s). The oldest known metatherian is '' Sinodelphys'', found in 125M-year-old early Cretaceous
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, especiall ...
in China's northeastern
Liaoning Province Liaoning () is a coastal 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 is the northernmost ...
. The fossil is nearly complete and includes tufts of fur and imprints of soft tissues. Didelphimorphia (common opossums of the
Western Hemisphere The Western Hemisphere is the half of the planet Earth that lies west of the prime meridian (which crosses Greenwich, London, United Kingdom) and east of the antimeridian. The other half is called the Eastern Hemisphere. Politically, the te ...
) first appeared in the late Cretaceous and still have living representatives, probably because they are mostly semi-
arboreal Arboreal locomotion is the 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. The habitats pose nu ...
unspecialized
omnivores 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 nut ...
. Tracks from the Early Cretaceous of Angola show the existence of raccoon-size mammals 118 million years ago. The best-known feature of marsupials is their method of reproduction: *The mother develops a kind of
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example ...
sack in her womb that delivers nutrients to the
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 ...
. Embryos of
bandicoots Bandicoots are a group of more than 20 species of small to medium-sized, terrestrial, largely nocturnal marsupial omnivores in the order Peramelemorphia. They are endemic to the Australia–New Guinea region, including the Bismarck Archipelago ...
, koalas and
wombats Wombats are short-legged, muscular quadrupedal marsupials that are native to Australia. They are about in length with small, stubby tails and weigh between . All three of the extant species are members of the family Vombatidae. They are adap ...
additionally form placenta-like organs that connect them to the
uterine 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 ...
wall, although the placenta-like organs are smaller than in placental mammals and it is not certain that they transfer nutrients from the mother to the embryo. *Pregnancy is very short, typically four to five weeks. The embryo is born at a very early stage of development, and is usually less than long at birth. It has been suggested that the short pregnancy is necessary to reduce the risk that the mother's
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
will attack the embryo. *The newborn marsupial uses its forelimbs (with relatively strong hands) to climb to a
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 ...
, which is usually in a pouch on the mother's belly. The mother feeds the baby by contracting muscles over her mammary glands, as the baby is too weak to suck. The newborn marsupial's need to use its forelimbs in climbing to the nipple was historically thought to have restricted metatherian evolution, as it was assumed that the forelimb couldn't become specialised intro structures like wings, hooves or flippers. However, several
bandicoot Bandicoots are a group of more than 20 species of small to medium-sized, terrestrial, largely nocturnal marsupial omnivores in the order Peramelemorphia. They are endemic to the Australia–New Guinea region, including the Bismarck Archipelago t ...
s, most notably the
pig-footed bandicoot ''Chaeropus'', known as the pig-footed bandicoots, is a genus of small mammals that became extinct during the 20th century. They were unique marsupials, of the order Peramelemorphia (bandicoots and bilbies), with unusually thin legs, yet were abl ...
, have true hooves similar to those of placental ungulates, and several marsupial gliders have evolved. Although some marsupials look very like some placentals (the
thylacine The thylacine ( , or , also ) (''Thylacinus cynocephalus'') is an extinct carnivorous marsupial that was native to the Australian mainland and the islands of Tasmania and New Guinea. The last known live animal was captured in 1930 in Tasma ...
, "marsupial tiger" or "marsupial wolf" is a good example), marsupial skeletons have some features that distinguish them from placentals: *Some, including the thylacine, have four molars; whereas no known placental has more than three. *All have a pair of palatal fenestrae, window-like openings on the bottom of the skull (in addition to the smaller nostril openings). Marsupials also have a pair of marsupial bones (sometimes called " epipubic bones"), which support the pouch in females. But these are not unique to marsupials, since they have been found in fossils of multituberculates, monotremes, and even eutherians — so they are probably a common ancestral feature that disappeared at some point after the ancestry of living placental mammals diverged from that of marsupials. Some researchers think the epipubic bones' original function was to assist locomotion by supporting some of the muscles that pull the thigh forwards.


Eutheria

The time of appearance of the earliest eutherians has been a matter of controversy. On one hand, recently discovered fossils of '' Juramaia'' have been dated to 160 million years ago and classified as eutherian. Fossils of '' Eomaia'' from 125 million years ago in the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145& ...
have also been classified as eutherian. A recent analysis of phenomic characters, however, classified ''Eomaia'' as pre-eutherian and reported that the earliest clearly eutherian specimens came from '' Maelestes'', dated to 91 million years ago. That study also reported that eutherians did not significantly diversify until after the catastrophic extinction at the
Cretaceous–Paleogene boundary The Cretaceous–Paleogene (K–Pg) boundary, formerly known as the Cretaceous–Tertiary (K–T) boundary, is a geological signature, usually a thin band of rock containing much more iridium than other bands. The K–Pg boundary marks the end ...
, about 66 million years ago. ''Eomaia'' was found to have some features that are more like those of marsupials and earlier metatherians: * Epipubic bones extending forwards from the pelvis, which are not found in any modern placental, but are found in all other mammals — early mammaliaforms, non-placental eutherians, marsupials, and monotremes — as well as in the
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 varie ...
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 that are closest to mammals. Their function is to stiffen the body during locomotion. This stiffening would be harmful in pregnant placentals, whose abdomens need to expand. * A narrow pelvic outlet, which indicates that the young were very small at birth and therefore pregnancy was short, as in modern marsupials. This suggests that the placenta was a later development. * Five incisors in each side of the upper jaw. This number is typical of metatherians, and the maximum number in modern placentals is three, except for
homodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example, ...
s, such as the
armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, alo ...
. But ''Eomaia's'' molar to
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
ratio (it has more pre-molars than molars) is typical of eutherians, including placentals, and not normal in marsupials. ''Eomaia'' also has a Meckelian groove, a primitive feature of the lower jaw that is not found in modern placental mammals. These intermediate features are consistent with
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
estimates that the placentals diversified about 110M years ago, 15M years after the date of the ''Eomaia'' fossil. ''Eomaia'' also has many features that strongly suggest it was a climber, including several features of the feet and toes; well-developed attachment points for muscles that are used a lot in climbing; and a tail that is twice as long as the rest of the spine. Placentals' best-known feature is their method of reproduction: * The embryo attaches itself to the
uterus 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 ...
via a large
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 mate ...
via which the mother supplies food and oxygen and removes waste products. * Pregnancy is relatively long and the young are fairly well-developed at birth. In some species (especially herbivores living on plains) the young can walk and even run within an hour of birth. It has been suggested that the evolution of placental reproduction was made possible by
retrovirus A retrovirus is a type of virus that inserts a DNA copy of its RNA genome into the DNA of a host cell that it invades, thus changing the genome of that cell. Once inside the host cell's cytoplasm, the virus uses its own reverse transcriptas ...
es that: * make the interface between the placenta and uterus into a
syncytium A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleu ...
, i.e. a thin layer of cells with a shared external membrane. This allows the passage of oxygen, nutrients and waste products, but prevents the passage of blood and other cells that would cause the mother's
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
to attack the
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 dev ...
. * reduce the aggressiveness of the mother's immune system, which is good for the foetus but makes the mother more vulnerable to infections. From a paleontologist's point of view, eutherians are mainly distinguished by various features of their teeth, ankles and feet.


Expansion of ecological niches in the Mesozoic

Generally speaking, most species of mammaliaforms did occupy the niche of small, nocturnal insectivores, but recent finds, mainly in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, show that some species and especially crown group mammals were larger and that there was a larger variety of lifestyles than previously thought. For example: * '' Adalatherium hui'' is a large sized, erect limbed herbivore from 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 ...
of
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
. *''
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 ...
'', a member of Docodonta which lived in the middle Jurassic about 164 million years, was about long, weighed , had a
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 a ...
-like tail that was adapted for swimming, limbs adapted for swimming and digging, and teeth adapted for eating fish. Another docodont, '' Haldanodon'', also had semi-aquatic habits, and indeed aquatic tendencies were probably common among docodonts based on their prevalence in wetland environments. The eutriconodonts ''
Liaoconodon ''Liaoconodon'' is an extinct genus of early mammal from the early Cretaceous (early Aptian stage, approximately 120 Ma). It is a eutriconodont which lived in what is now the Jianchang of Liaoning Province, eastern China. It is known from ...
'' and ''
Yanoconodon ''Yanoconodon'' is a monotypic genus of extinct early mammal whose representative species ''Yanoconodon allini'' lived during the Mesozoic in what is now China. The holotype fossil of ''Yanoconodon'' was excavated in the Yan Mountains about ...
'' have more recently also have been suggested to be freshwater swimmers, lacking ''Castorocaudas powerful tail but possessing paddle-like limbs; the eutriconodont '' Astroconodon'' has similarly been suggested as being semi-aquatic in the past, albeit to less convincing evidence. * Multituberculates are allotherians that survived for over 125 million years (from mid-Jurassic, about 160M years ago, to late
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
, about 35M years ago) are often called the "rodents of the Mesozoic". As noted above, they may have given birth to tiny live neonates rather than laying eggs. * ''
Fruitafossor ''Fruitafossor'' was a termite-eating mammal endemic to North America during the Late Jurassic epoch (around 150 mya). The description is based on a complete skeleton of a chipmunk-sized animal. It was discovered on March 31, 2005, in Fruita, ...
'', from the late Jurassic period about 150 million years ago, was about the size of a
chipmunk Chipmunks are small, striped rodents of the family Sciuridae. Chipmunks are found in North America, with the exception of the Siberian chipmunk which is found primarily in Asia. Taxonomy and systematics Chipmunks may be classified either as ...
and its teeth, forelimbs and back suggest that it broke open the nest of social insects to prey on them (probably
termites 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 Blat ...
, as ants had not yet appeared). * Similarly, the gobiconodontid '' Spinolestes'' possessed adaptations for fossoriality and convergent traits with placental
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. ...
ns like scutes and xenarthrous vertebrae, so it too might have had anteater like habits. It is also notable for the presence of quills akin to those of modern
spiny mice The term spiny mouse refers to any species of rodent within the genus ''Acomys''. Similar in appearance to mice of the genus ''Mus'', spiny mice are small mammals with bare, scaled tails. However, their coats are endowed with unusually stiff ...
. * '' Volaticotherium'', from the boundary the early Cretaceous about 125M years ago, is the earliest-known gliding mammal and had a gliding membrane that stretched out between its limbs, rather like that of a modern
flying squirrel Flying squirrels (scientifically known as Pteromyini or Petauristini) are a tribe of 50 species of squirrels in the family Sciuridae. Despite their name, they are not in fact capable of full flight in the same way as birds or bats, but they a ...
. This also suggests it was active mainly during the day. The closely related '' Argentoconodon'' also shows similar adaptations that may also suggest aerial locomotion. * '' Repenomamus'', a eutriconodont from the early Cretaceous 130 million years ago, was a stocky,
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 by ...
-like predator that sometimes preyed on young dinosaurs. Two species have been recognized, one more than long and weighing about , the other less than long and weighing . * ''
Schowalteria ''Schowalteria'' is a genus of extinct mammal from the Cretaceous of Canada. It is the earliest known representative of Taeniodonta, a specialised lineage of non-placental eutherian mammals otherwise found in Paleocene and Eocene deposits. It is ...
'' is a Late Cretaceous species almost as large if not larger than ''R. giganticus'' that shows speciations towards herbivory, comparable to those of modern ungulates. * Zhelestidae is a lineage of Late Cretaceous herbivorous
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 ...
ns, to the point of being mistaken for stem-
ungulate 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, ...
s. * Similarly, mesungulatids are also fairly large sized herbivorous mammals from the Late Cretaceous *
Deltatheroida Deltatheroida is an extinct group of basal metatherians that were distantly related to modern marsupials. The majority of known members of the group lived in the Cretaceous; one species, '' Gurbanodelta kara'', is known from the late Paleocene ( ...
ns were
metatheria 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 w ...
ns that were specialised towards carnivorous habits, and possible forms like '' Oxlestes'' and '' Khudulestes'' might have been among the largest Mesozoic mammals, though their status as deltatheroidans is questionable. * '' Ichthyoconodon'', a eutriconodont from the
Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ag ...
of Morocco, is currently known from molariforms found in marine deposits. These teeth are sharp-cusped and similar in shape to those of piscivorous mammals, and unlike the teeth of contemporary mammals they do not show degradation, so rather than being carried down by river deposits the animal died ''in situ'' or close. This has been taken to mean that it was a
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 ...
, likely one of the few examples known from the Mesozoic. Alternatively, its close relations to '' Volaticotherium'' and '' Argentoconodon'' might suggest that it was a flying mammal. * '' Didelphodon'' is a Late Cretaceous riverine species of stagodontid marsupialiform with a
durophagous Durophagy is the eating behavior of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled mollusks, or crabs. It is mostly used to describe fish, but is also used when describing reptiles, including fossil tu ...
dentition, robust jaws similar to a modern
Tasmanian devil The Tasmanian devil (''Sarcophilus harrisii'') ( palawa kani: purinina) is a carnivorous marsupial of the family Dasyuridae. Until recently, it was only found on the island state of Tasmania, but it has been reintroduced to New South Wales ...
, and a postcranial skeleton very similar in size and shape to an
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 we ...
. This animal has been lauded as the strongest bite of all Mesozoic mammals. It possibly specialized on eating
freshwater crab Around 1,300 species of freshwater crabs are distributed throughout the tropics and subtropics, divided among eight families. They show direct development and maternal care of a small number of offspring, in contrast to marine crabs, which relea ...
s and
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is est ...
s. * Tracks of a raccoon-sized mammaliaform representing the morphofamily Ameghinichnidae are described from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145& ...
(late
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...
)
Calonda Formation Corund ( hu, Korond, Hungarian pronunciation: ) is a commune in Harghita County, Romania. It lies in the Székely Land, an ethno-cultural region in eastern Transylvania, in the "Salt Region" (''Ținutul Sării'' or ''Sóvidék''). Corund is fa ...
(
Angola , national_anthem = "Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordinat ...
) by Mateus ''et al.'' (2017), who name a new ichnotaxon '' Catocapes angolanus''. A study on Mesozoic mammaliaforms suggests that they were a primary factor in constraining mammalian body size, rather than solely competition from dinosaurs.


Evolution of major groups of living mammals

There are currently vigorous debates between traditional
paleontologists 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 foss ...
and molecular phylogeneticists about how and when the modern groups of mammals diversified, especially the placentals. Generally, the traditional paleontologists date the appearance of a particular group by the earliest known fossil whose features make it likely to be a member of that group, while the molecular phylogeneticists suggest that each lineage diverged earlier (usually in the Cretaceous) and that the earliest members of each group were anatomically very similar to early members of other groups and differed only in their
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
. These debates extend to the definition of and relationships between the major groups of placentals.


Molecular phylogenetics-based family tree of placental mammals

Molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
uses features of organisms'
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s to work out family trees in much the same way as paleontologists do with features of fossils — if two organisms' genes are more similar to each other than to those of a third organism, the two organisms are more closely related to each other than to the third. Molecular phylogeneticists have proposed a family tree that is both broadly similar to but has notable differences from that of the paleontologists. Like paleontologists, molecular phylogeneticists have differing ideas about various details, but here is a typical family tree according to molecular phylogenetics: Note that the diagram shown here omits extinct groups, as one cannot extract DNA from fossils. Here are the most significant of the differences between this family tree and the one familiar to paleontologists: *The top-level division is between Atlantogenata and Boreoeutheria, instead of between Xenarthra and the rest. However, analysis of
transposable element A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Transp ...
insertions supports a three-way top-level split between Xenarthra, Afrotheria and Boreoeutheria and the Atlantogenata clade does not receive significant support in recent distance-based molecular phylogenetics. *Afrotheria contains several groups that are only distantly related according to the paleontologists' version: Afroinsectiphilia ("African insectivores"), Tubulidentata (aardvarks, which paleontologists regard as much closer to odd-toed ungulates than to other members of Afrotheria), Macroscelidea (elephant shrews, usually regarded as close to rabbits and rodents). The only members of Afrotheria that paleontologists would regard as closely related are Hyracoidea (hyraxes), Proboscidea (elephants) and Sirenia (manatees, dugongs). *Insectivores are split into three groups: one is part of Afrotheria and the other two are distinct sub-groups within Boreoeutheria. *Bats are closer to Carnivora and odd-toed ungulates than to Primates and Dermoptera (colugos). *Perissodactyla (odd-toed ungulates) are closer to Carnivora and bats than to Artiodactyla (even-toed ungulates). The grouping together of the Afrotheria has some geological justification. All surviving members of the Afrotheria originate from South American or (mainly) African lineages — even the
Indian elephant The Indian elephant (''Elephas maximus indicus'') is one of four extant recognised subspecies of the Asian elephant and native to mainland Asia. Since 1986, the Asian elephant has been listed as Endangered on the IUCN Red List as the wild pop ...
, which diverged from an African lineage about . As
Pangaea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
broke up, Africa and South America separated from the other continents less than 150M years ago, and from each other between 100M and 80M years ago. So it would not be surprising if the earliest eutherian immigrants into Africa and South America were isolated there and radiated into all the available
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s. Nevertheless, these proposals have been controversial. Paleontologists naturally insist that fossil evidence must take priority over deductions from samples of the DNA of modern animals. More surprisingly, these new family trees have been criticised by other molecular phylogeneticists, sometimes quite harshly: *
Mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
's mutation rate in mammals varies from region to region — some parts hardly ever change and some change extremely quickly and even show large variations between individuals within the same species. *Mammalian mitochondrial DNA mutates so fast that it causes a problem called "saturation", where random noise drowns out any information that may be present. If a particular piece of mitochondrial DNA mutates randomly every few million years, it will have changed several times in the 60 to 75M years since the major groups of placental mammals diverged.


Timing of placental evolution

Recent molecular phylogenetic studies suggest that most placental
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 ...
diverged late in the Cretaceous period, about 100 to 85 million years ago, but that modern families first appeared later, in the late Eocene and early
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 ...
epochs 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 configu ...
period. Fossil-based analyses, on the contrary, limit the placentals to the Cenozoic. Many Cretaceous fossil sites contain well-preserved lizards, salamanders, birds, and mammals, but not the modern forms of mammals. It is possible that they simply did not exist, and that the
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...
runs fast during major evolutionary radiations. On the other hand, there is fossil evidence from of
hoof The hoof (plural: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits, yet the rum ...
ed mammals that may be ancestors of modern
ungulate 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, ...
s. Fossils of the earliest members of most modern groups date from the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
, a few date from later and very few from the Cretaceous, before the extinction of the dinosaurs. But some paleontologists, influenced by molecular phylogenetic studies, have used statistical methods to
extrapolate In mathematics, extrapolation is a type of estimation, beyond the original observation range, of the value of a variable on the basis of its relationship with another variable. It is similar to interpolation, which produces estimates between kn ...
''backwards'' from fossils of members of modern groups and concluded that
primates 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 ...
arose in the late Cretaceous. However, statistical studies of the fossil record confirm that mammals were restricted in size and diversity right to the end of the Cretaceous, and rapidly grew in size and diversity during the Early Paleocene.


Evolution of mammalian features


Jaws and middle ears

''
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 (, paleocoordina ...
'', whose fossils date from the early Jurassic, provides the first clear evidence of fully mammalian jaw joints and middle ears, in which the jaw joint is formed by 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
and
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 bones form the cheek series of the skull. The bone forms an ancestral co ...
bones while 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 ...
and quadrate move to the middle ear, where they are known as the incus and malleus. One analysis of the monotreme ''
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 ...
'' suggested that this animal had a pre-mammalian jaw joint formed by the angular and quadrate bones and that the definitive mammalian middle ear evolved twice independently, in monotremes and in
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. ...
n mammals, but this idea has been disputed. For other opinions see "Technical comments" linked from same Web page In fact, two of the suggestion's authors co-authored a later paper that reinterpreted the same features as evidence that ''Teinolophos'' was a full-fledged
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or mono ...
, which means it would have had a mammalian jaw joint and middle ear.


Lactation

It has been suggested that lactation's original function was to keep eggs moist. Much of the argument is based on monotremes (egg-laying mammals): *While the amniote egg is usually described as able to evolve away from water, most reptile eggs actually need moisture if they are not to dry out. *Monotremes do not have nipples, but secrete milk from a hairy patch on their bellies. *During incubation, monotreme eggs are covered in a sticky substance whose origin is not known. Before the eggs are laid, their shells have only three layers. Afterwards, a fourth layer appears with a composition different from that of the original three. The sticky substance and the fourth layer may be produced by the mammary glands. *If so, that may explain why the patches from which monotremes secrete milk are hairy. It is easier to spread moisture and other substances over the egg from a broad, hairy area than from a small, bare nipple. Later research demonstrated that
casein Casein ( , from Latin ''caseus'' "cheese") is a family of related phosphoproteins ( αS1, aS2, β, κ) that are commonly found in mammalian milk, comprising about 80% of the proteins in cow's milk and between 20% and 60% of the proteins in hum ...
s already appeared in the common mammalian ancestor approximately 200–310 million years ago. The question of whether secretions of a substance to keep eggs moist translated into actual lactation in therapsids is open. A small mammaliomorph called '' Sinocodon'', generally assumed to be the sister group of all later mammals, had front teeth in even the smallest individuals. Combined with a poorly ossified jaw, they very probably did not suckle. Thus suckling may have evolved right at the pre-mammal/mammal transition. However, tritylodontids, generally assumed to be more basal, show evidence of suckling.
Morganucodonta Morganucodonta (" Glamorgan teeth") is an extinct order of basal Mammaliaformes, a group including crown-group mammals (Mammalia) and their close relatives. Their remains have been found in Southern Africa, Western Europe, North America, India a ...
ns, also assumed to be basal
Mammaliaformes 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 c ...
, also show evidence of lactation.


Digestive system

The evolution of the digestive system has formed a significant influence in mammal evolution. With the emergence of mammals, the digestive system was modified in a variety of ways depending on the animal's diet.  For example, cats and most carnivores have simple large intestines, while the horse as a herbivore has a voluminous large intestine. An ancestral feature of
ruminant Ruminants (suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The ...
s is their multi-chambered (usually four-chambered) stomach, which evolved about 50 million years ago. Along with morphology of the gut,
gastric acid Gastric acid, gastric juice, or stomach acid is a digestive fluid formed within the stomach lining. With a pH between 1 and 3, gastric acid plays a key role in digestion of proteins by activating digestive enzymes, which together break down the ...
ity has been proposed as a key factor shaping the diversity and composition of microbial communities found in the vertebrate gut. Comparisons of stomach acidity across trophic groups in mammal and bird taxa show that scavengers and carnivores have significantly higher stomach acidities compared to herbivores or carnivores feeding on phylogenetically distant prey such as insects or fish. Despite the lack of fossilization of the gut, microbial evolution of the gut can be inferred from the interrelationships of existing animals, microbes and probable foodstuffs. Mammals are metagenomic, in that they are composed of not only their own genes, but also those of all of their associated microbes.
Gut microbiota 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 ...
has co-diversified as mammalian species have evolved. Recent studies indicate that adaptive divergence between mammalian species is shaped in part by changes in the gut microbiota. The house mouse may have evolved not only with, but also in response to, the unique bacteria inhabiting its gut.


Hair and fur

The first 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. ''Me ...
'', from 164M years ago in the mid-Jurassic. As both mammals ''Megaconus'' and ''Castorocauda'' have a double coat of hair, with both guard hairs and an undercoat, it may be assumed that their last common ancestor did as well. This animal must have been Triassic as it was an ancestor of the Triassic ''
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 was ...
''. More recently, the discovery of hair remnants in Permian coprolites pushes back the origin of mammalian hair much further back in the synapsid line to
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ...
therapsids. In the mid-1950s, some scientists interpreted 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. T ...
e (upper jaws) 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 ...
e (small bones in front of the maxillae) of
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 varie ...
s as channels that 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 sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are coarse ...
) and suggested that this was evidence of hair or fur. It was soon pointed out, however, that foramina do not necessarily show that an animal had vibrissae; the modern lizard '' Tupinambis'' has foramina that are almost identical to those found in the non-mammalian 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''. A
trace fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or animal itself. Trace fossils contrast with body fossils, ...
from the Lower Triassic had been erroneously regarded as a cynodont footprint showing hair, but this interpretation has been refuted. A study of cranial openings for facial nerves connected whiskers in extant mammals indicate the Prozostrodontia, small immediate ancestors of mammals, presented whiskers similar to mammals, but that less advanced therapsids would either have immobile whiskers or no whisker at all. Fur may have evolved from whiskers. Whiskers themselves may have evolved as a response to nocturnal and/or burrowing lifestyle. Ruben & Jones (2000) note that the Harderian glands, which secrete
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 in ...
s for coating the fur, were present in the earliest mammals like ''
Morganucodon ''Morganucodon'' ("Glamorgan tooth") is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, ''Morganucodon'' is well represent ...
'', but were absent in near-mammalian therapsids like ''Thrinaxodon''. The Msx2 gene associated with hair follicle maintenance is also linked to the closure of the
parietal eye A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhyth ...
in mammals, indicating that fur and lack of pineal eye is linked. The pineal eye is present in ''Thrinaxodon'', but absent in more advanced cynognaths (the
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 m ...
). Insulation is the "cheapest" way to maintain a fairly constant body temperature, without consuming energy to produce more body heat. Therefore, the possession of hair or fur would be good evidence of homeothermy, but would not be such strong evidence of a high metabolic rate.


Erect limbs

Understanding of the evolution of erect limbs in mammals is incomplete — living and fossil
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 brai ...
s have sprawling limbs. Some scientists think that the parasagittal (non-sprawling) limb posture is limited to the
Boreosphenida Tribosphenida is a group (infralegion) of mammals that includes the ancestor of ''Hypomylos'', Aegialodontia and Theria (the last common ancestor of marsupials and placentals plus all of its descendants). Its current definition is more or less s ...
, a group that contains the therians but not, for example, the multituberculates. In particular, they attribute a parasagittal stance to the therians ''Sinodelphys'' and ''Eomaia'', which means that the stance had arisen by 125 million years ago, in the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145& ...
. However, they also discuss that earlier mammals had more erect forelimbs as opposed to the more sprawling hindlimbs, a trend still continued to some extent in modern placentals and marsupials.


Warm-bloodedness

"
Warm-blooded Warm-blooded is an informal term referring to animal species which can maintain a body temperature higher than their environment. In particular, homeothermic species maintain a stable body temperature by regulating metabolic processes. The on ...
ness" is a complex and rather ambiguous term, because it includes some or all of the following: *
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 ...
, the ability to generate heat internally rather than via behaviors such as basking or muscular activity. * Homeothermy, maintaining a fairly constant body temperature. Most
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s have an optimum operating temperature; efficiency drops rapidly outside the preferred range. A homeothermic organism needs only to possess enzymes that function well in a small range of temperatures. * Tachymetabolism, maintaining a high metabolic rate, particularly when at rest. This requires a fairly high and stable body temperature because of the Q10 effect: biochemical processes run about half as fast if an animal's temperature drops by 10 °C. Since scientists cannot know much about the internal mechanisms of extinct creatures, most discussion focuses on homeothermy and tachymetabolism. However, it is generally agreed that endothermy first evolved in non-mammalian synapsids such 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, typic ...
s, which possess body proportions associated with heat retention, high vascularised bones with
Haversian canal Haversian canals (sometimes canals of Havers) are a series of microscopic tubes in the outermost region of bone called cortical bone. They allow blood vessels and nerves to travel through them to supply the osteocytes. Structure Each Haversia ...
s, and possibly hair. More recently, it has been suggested that endothermy evolved as far back as '' Ophiacodon''. Modern monotremes have a low body temperature compared to marsupials and placental mammals, around . Phylogenetic bracketing suggests that the body temperatures of early crown-group mammals were not less than that of extant monotremes. There is cytological evidence that the low metabolism of monotremes is a secondarily evolved trait.


Respiratory turbinates

Modern mammals have respiratory turbinates, convoluted structures of thin bone in the nasal cavity. These are lined with
mucous membrane A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It i ...
s that warm and moisten inhaled air and extract heat and moisture from exhaled air. An animal with respiratory turbinates can maintain a high rate of breathing without the danger of drying its lungs out, and therefore may have a fast metabolism. Unfortunately these bones are very delicate and therefore have not yet been found in fossils. But rudimentary ridges like those that support respiratory turbinates have been found in advanced Triassic
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 varie ...
s, such as ''
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 ...
'' and '' Diademodon'', which suggests that they may have had fairly high metabolic rates.


Bony secondary palate

Mammals have a secondary bony palate, which separates the respiratory passage from the mouth, allowing them to eat and breathe at the same time. Secondary bony palates have been found in the more advanced cynodonts and have been used as evidence of high metabolic rates. But some cold-blooded vertebrates have secondary bony palates (crocodilians and some lizards), while birds, which are warm-blooded, do not.


Diaphragm

A muscular
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 ...
helps mammals to breathe, especially during strenuous activity. For a diaphragm to work, the ribs must not restrict the abdomen, so that expansion of the chest can be compensated for by reduction in the volume of the abdomen and ''vice versa''. Diaphragms are known in caseid pelycosaurs, indicating an early origin within synapsids, though they were still fairly inefficient and likely required support from other muscle groups and limb motion. The advanced cynodonts have very mammal-like rib cages, with greatly reduced lumbar ribs. This suggests that these animals had more developed diaphragms, were capable of strenuous activity for fairly long periods and therefore had high metabolic rates. On the other hand, these mammal-like rib cages may have evolved to increase agility. However, the movement of even advanced therapsids was "like a wheelbarrow", with the hindlimbs providing all the thrust while the forelimbs only steered the animal, in other words advanced therapsids were not as agile as either modern mammals or the early dinosaurs. So the idea that the main function of these mammal-like rib cages was to increase agility is doubtful.


Limb posture

The
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 mo ...
had sprawling forelimbs and semi-erect hindlimbs. This suggests that
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 ...
would have made it rather difficult for them to move and breathe at the same time, but not as difficult as it is for animals such as lizards, which have completely sprawling limbs. Advanced therapsids may therefore have been significantly less active than modern mammals of similar size and so may have had slower metabolisms overall or else been bradymetabolic (lower metabolism when at rest).


Brain

Mammals are noted for their large brain size relative to body size, compared to other animal groups. Recent findings suggest that the first brain area to expand was that involved in smell. Scientists scanned the skulls of early mammal species dating back to 190–200 million years ago and compared the brain case shapes to earlier pre-mammal species; they found that the brain area involved in the sense of smell was the first to enlarge. This change may have allowed these early mammals to hunt insects at night when dinosaurs were not active. After the extinction of the dinosaurs 66 million years ago, mammals began to increase in body size as new niches became available, but their brain lagged behind their bodies for the first ten million years. Relative to body size the brain of Paleocene mammal was relatively smaller than that of Mesozoic mammals. It wasn’t until the Eocene that the mammalian brains began to catch up with their bodies, particularly in certain areas associated with their senses.Mammals’ bodies outpaced their brains right after the dinosaurs died
/ref>


See also

*
History of life The history of life on Earth traces the processes by which living and fossil organisms evolved, from the earliest emergence of life to present day. Earth formed about 4.5 billion years ago (abbreviated as ''Ga'', for ''gigaannum'') and evide ...
* Evolution of primates * Evolution of ungulates ** Evolution of even-toed ungulates ** Evolution of odd-toed ungulates * Genome diversity and karyotype evolution of mammals * List of examples of convergent evolution in mammals * '' Juramaia''


References


Further reading

* Robert L. Carroll, ''Vertebrate Paleontology and Evolution'', W. H. Freeman and Company, New York, 1988 . Chapters XVII through XXI * Nicholas Hotton III, Paul D. MacLean, Jan J. Roth, and E. Carol Roth, editors, ''The Ecology and Biology of Mammal-like Reptiles'', Smithsonian Institution Press, Washington and London, 1986 * T. S. Kemp, ''The Origin and Evolution of Mammals'', Oxford University Press, New York, 2005 * * *


External links


The Cynodontia
covers several aspects of the evolution of cynodonts into mammals, with plenty of references.
Mammals
BBC Radio 4 discussion with Richard Corfield, Steve Jones & Jane Francis (''In Our Time'', Oct. 13, 2005) {{DEFAULTSORT:Evolution Of Mammals