MammalsFossil range:
Late Triassic - Recent |
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Madras Treeshrew
(Anathana ellioti) |
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Scientific classification |
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Subclasses |
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The mammals are the
class of vertebrate animals characterized by the production of milk in females
for the nourishment of young, from mammary glands present on most species and
specialized skin glands in monotremes that seep or ooze milk; the presence of
hair or fur; specialized teeth; three minute bones within the ear; the presence
of a neocortex region in the brain; and endothermic or "warm-blooded"
bodies, and, in most cases, the existence of a placenta in the ontogeny. The
brain regulates endothermic and circulatory systems, including a four-chambered
heart. Mammals encompass some 5,500 species (including humans), distributed in
about 1,200 genera, 152 families and up to 46 orders, though this varies with
the classification scheme.
Phylogenetically, Mammalia
is defined as all descendants of the most recent common ancestor of monotremes
(e.g., echidnas and platypuses) and therian mammals (marsupials and
placentals).
Living mammal species are
easily identified by the fact that the females have mammary glands which
produce milk.
But other features are
required when classsifying fossils, because mammary glands and most of the
other features listed above are not visible in fossils. The evolution of
mammals from reptiles was a gradual process which took about 70 million years,
from the beginning of the mid-Permian to the mid-Jurassic, and by the beginning
of the mid-Triassic there were many species which looked very like mammals.
Paleontologists therefore
use as the distinguishing feature one that is shared by all living mammals
(including monotremes) but is not present in any of the early Triassic
mammal-like reptiles - mammals use for hearing two bones which all reptiles use
for eating. The earliest reptiles had a jaw joint composed of the articular (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 reptiles use this system, including
lizards, crocodilians, dinosaurs (and their descendants the birds) and
mammal-like reptiles. But mammals have a different jaw joint, composed only
of the dentary (the lower jaw bone which carries the teeth) and the squamosal
(another small skull bone) - the word "only" is important here, as
some late mammal-like reptiles had both types of jaw joint. And in mammals the
old quadrate and articular bones have become the incus and malleus bones in the
middle ear. Note: "non-mammalian reptiles" above implies that mammals
are a sub-group of reptiles, and that is exactly what cladistics says they are.
Mammals also have a double
occipital condyle, i.e. they have two knobs at the base of the skull which fit
into the topmost neck vertebra, and other vertebrates have a single occipital
condyle. But paleontologists use only the jaw joint and middle ear as criteria
for identifying fossil mammals, because it would be very confusing if they
found a fossil which had one feature but not the other.
The vast majority of
mammals have seven cervical vertebrae (bones in the neck); this includes bats,
giraffes, whales, and humans. The few exceptions include the manatee and the
two-toed sloth, which each have only six cervical vertebrae, and the three-toed
sloth with nine cervical vertebrae.
See the section Mammalian
lungs in the Lung article.
The mammalian heart has
four chambers: the right atrium, right ventricle, left atrium, and left
ventricle. Atria are for receiving blood; ventricles are for pumping blood to
the lungs and body. The ventricles are larger than the atria and their walls
are thick, because muscular walls are needed to forcefully pump the blood from
the heart to the body and lungs. Deoxygenated blood from the body enters the
right atrium, which pumps it to the right ventricle. The right ventricle pumps
blood to the lungs, where carbon dioxide diffuses out, and oxygen diffuses in.
From the lungs, oxygenated blood enters the left atrium, where it is pumped to
the left ventricle (the largest and strongest of the 4 chambers), which pumps
it out to the rest of the body, including the heart's own blood supply.
All mammalian brains
possess a neocortex which is a brain region that is unique to mammals.
Mammals have integumentary
systems made up of three layers: the outermost epidermis, the dermis, and the
hypodermis. This characteristic is not unique to mammals, since it is found in
all vertebrates.
The epidermis is typically
ten to thirty cells thick, its main function being to provide a waterproof
layer. Its outermost cells are constantly lost; its bottommost cells are
constantly dividing and pushing upward. The middle layer, the dermis, is
fifteen to forty times thicker than the epidermis. The dermis is made up of
many components such as bony structures and blood vessels. The hypodermis is
made up of adipose tissue. Its job is to store lipids, and to provide
cushioning and insulation. The thickness of this layer varies widely from
species to species.
No mammals are known to
have hair that is naturally blue or green in color. Some cetaceans, along with
the mandrills appear to have shades of blue skin. Many mammals are indicated as
having blue hair or fur, but in all known cases, it has been found to be a
shade of grey. The two-toed sloth can seem to have green fur, but this color is
caused by algae growths.
Most mammals give birth to
live young (vivipary), but a few (the monotremes) lay eggs. Live birth also
occurs in some non-mammalian species, such as guppies and hammerhead sharks;
thus it is not a distinguishing characteristic of mammals. Although all mammals
are endothermic, so are birds, and so this too is not a defining feature.
A characteristic of mammals
is that they have mammary glands, a defining feature present only in mammals.
The monotremes branched from other mammals early on, and do not have nipples,
but they do have mammary glands. Most mammals are terrestrial, but some are
aquatic, including sirenia (manatees and dugongs) and the cetaceans (dolphins
and whales). Whales are the largest of all animals. There are semi-aquatic
species such as seals which come to land to breed but spend most of the time in
water.
The only mammals for which
true flight has been observed are bats; mammals such as flying squirrels and
flying lemurs are more accurately classified as gliding mammals.
Mammals belong to a group
of amniotes called the synapsids, which are distinguished by having a single
hole (temporal fenestra) low on each side of the skull on each side where jaw
muscles attach. In comparison, dinosaurs, birds, and most living reptiles are
diapsids, with two temporal fenestrae on each side of the skull; and turtles,
with no temporal fenestra, are anapsids. The synapsids diverged from the other
reptile lineages very early, in the late Carboniferous, and one of the earliest
examples is Archaeothyris.
The synapsid pelycosaurs
were the dominant land vertebrates of the early Permian. The therapsids
probably evolved from pelycosaur ancestors, and consist of a sequence of groups
which became increasingly mammal-like, especially the Triassic cynodonts
("dog-teeth"). The main article on synapsids presents a probable
phylogeny ("family tree") which shows how mammals evolved from early
synapsids.
Some cynodonts such as the
early to mid Triassic Cynognathus had erect limbs and possibly hair or
fur, and therefore may have been at least fairly warm-blooded.
Several groups of
mammaliformes ("almost mammals") arose from the mid Triassic onwards
- these had non-mammalian jaw joints but almost certainly had hair and were
warm-blooded. Megazostrodon is a good example. Some of them were very
successful in their own right, for example the multituberculates appeared in
the mid-Jurassic and became extinct about 100M years later, in the Oligocene.
The first true mammals
appeared in the early Jurassic, over 70M years after the first therapsids and
about 30M years after the first mammaliformes. Hadrocodium is an example of the
transition to true mammal status - it had a mammalian jaw joint but there is
some debate about whether its middle ear was fully mammalian. The triconodonts
may have been the earliest true mammals known so far.
Like their predecessors,
and like modern monotremes, the first mammals laid eggs. The earliest known
fossils of marsupial and placental mammals date from the mid to late
Cretaceous, about 170M years after the first therapsids.
Lactation is the
distinguishing feature of living mammals, but no-one is sure about when it
evolved. Since all living mammals (including monotremes) produce milk, we can
be confident that their last common ancestor (whatever and whenever that was)
produced milk. Some scientists have suggested that the cynodonts produced milk,
but this idea is a plausible speculation with little hard evidence to support
it.
Throughout the Mesozoic
therapsids, mammaliformes and mammals lived in the shadow of archosaurs - first
the "thecodonts" and following, the dinosaurs were the dominant land
vertebrates.
From the mid-Triassic
onwards, mammaliformes and mammals were mostly about between shrew-sized and
rat-sized. But in January 2005, the journal Nature reported the
discovery of two 130 million year old fossils of Repenomamus, one more than a
meter in length, the other having remains of a baby dinosaur in its stomach (Nature,
Jan. 15, 2005 [1]). And the 2004 discovery in China of a 164 million year old
50 cm long aquatic mammal-like fossil of a thus far unknown species, dubbed
Castorocauda, by a team led by Dr. Ji Qiang of Nanjing University and the
Chinese Academy of Geological Sciences, was reported in February 2006 in the
journal Science (Science, Feb. 24, 2006 [2]).
Some orders of placental
mammals appear to have arisen before the end of the Cretaceous - insectivores,
rodents, ungulates, and possibly primates. But they were not able to expand
into their modern ecological niches until after the extinction of the
dinosaurs.
Shortly after the start of
the Paleocene, mammals exploded into the ecological niches left by the
extinction of the dinosaurs and also found some which the dinosaurs apparently
never occupied:
Some orders of mammals only
appeared after the extinction of the dinosaurs:
Some Cenozoic mammals were
considerably larger than their nearest modern equivalents - but none approached
the size of the large dinosaurs.
George Gaylord Simpson's
"Principles of Classification and a Classification of Mammals" (AMNH Bulletin
v. 85, 1945) was the original source for the taxonomy listed here. Simpson laid
out a systematics of mammal origins and relationships that was universally
taught until the end of the 20th century. Since Simpson's classification, the
paleontological record has been recalibrated, and the intervening years have
seen much debate and progress concerning the theoretical underpinnings of
systematization itself, partly through the new concept of cladistics. Though
field work gradually made Simpson's classification outdated, it remained the
closest thing to an official classification of mammals.
A somewhat standardized
classification system has been adopted by most current mammalogy classroom
textbooks. The following taxonomy of extant and recently extinct mammals is
from Vaughan et al. (2000).
Class Mammalia
In 1997, the mammals were
comprehensively revised by Malcolm C. McKenna and Susan K. Bell, which has
resulted in the "McKenna/Bell classification".
McKenna and Bell, Classification
of Mammals: Above the species level, (1997) is the most comprehensive work
to date on the systematics, relationships, and occurrences of all mammal taxa,
living and extinct, down through the rank of genus. The new McKenna/Bell
classification was quickly accepted by paleontologists. The authors work
together as paleontologists at the American Museum of Natural History, New
York. McKenna inherited the project from Simpson and, with Bell, constructed a
completely updated hierarchical system, covering living and extinct taxa that
reflects the historical genealogy of Mammalia.
The McKenna/Bell
hierarchical listing of all of the terms used for mammal groups above the
species includes extinct mammals as well as modern groups, and introduces some
fine distinctions such as legions and sublegions (ranks which fall between
classes and orders) that are likely to be glossed over by the layman.
The published
re-classification forms both a comprehensive and authoritative record of
approved names and classifications and a list of invalid names.
Extinct groups are
represented by a dagger (†).
Class Mammalia
Molecular studies based on
DNA analysis have suggested new relationships among mammal families over the
last few years. Most of these findings have been independently validated by
Retrotransposon presence/absence data. The most recent classification systems
based on molecular studies have proposed four groups or lineages of placental
mammals. Molecular clocks suggest that these clades diverged from early common
ancestors in the Cretaceous, but fossils have not been found to corroborate
this hypothesis. These molecular findings are consistent with mammal zoogeography:
Following molecular DNA
sequence analyses, the first divergence was that of the Afrotheria 110–100 mya.
The Afrotheria proceeded to evolve and diversify in the isolation of the
African-Arabian continent. The Xenarthra, isolated in South America, diverged
from the Boreoeutheria approximately 100–95 mya. According to an alternative
view, the Xenarthra has the Afrotheria as closest allies, forming the
Atlantogenata as sistergroup to Boreoeutheria. The Boreoeutheria split into the
Laurasiatheria and Euarchontoglires between 95 and 85 mya; both of these groups
evolved on the northern continent of Laurasia. After tens of millions of years
of relative isolation, Africa-Arabia collided with Eurasia, exchanging
Afrotheria and Boreoeutheria. The formation of the Isthmus of Panama linked
South America and North America, which facilitated the exchange of mammal
species in the Great American Interchange. The traditional view that no
placental mammals reached Australasia until about 5 million years ago when bats
and murine rodents arrived has been challenged by recent evidence and may need
to be reassessed. These molecular results are still controversial because they
are not reflected by morphological data, and thus not accepted by many
systematists. Further there is some indication from Retrotransposon
presence/absence data that the traditional Epitheria hypothesis, suggesting
Xenarthra as the first divergence, might be true.
In light of all the options
available, the following classification system has been adopted for use in
related articles.
Class Mammalia
Wikipedia
http://en.wikipedia.org/w/index.php?title=Mammal&action=history