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Molar (tooth)

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
tooth", from mola, millstone and dens,
tooth. Molars show a great deal of
diversity in size and shape across
mammal groups.

A lower wisdom tooth after extraction.


Artery Posterior superior

alveolar artery
Permanent teeth of right half of lower dental arch,
seen from above: In this diagram, a healthy
Latin dentes
wisdom tooth (third, rearmost molares
molar) is included.
MeSH D008963

TA A05.1.03.007
FMA 55638
Anatomical terminology

Image showing molar teeth and their arrangement in

the mouth of an adult human being

Human anatomy
In humans, the molar teeth have either
four or five cusps. Adult humans have 12
molars, in four groups of three at the back
of the mouth. The third, rearmost molar in
each group is called a wisdom tooth. It is
the last tooth to appear, breaking through
the front of the gum at about the age of
20, although this varies from individual to
individual. Race can also affect the age at
which this occurs, with statistical
variations between groups.[1] In some
cases, it may not even erupt at all.

The human mouth contains upper

(maxillary) and lower (mandibular) molars.
They are: maxillary first molar, maxillary
second molar, maxillary third molar,
mandibular first molar, mandibular second
molar, and mandibular third molar.

Mammal evolution
In mammals, the crown of the molars and
premolars is folded into a wide range of
complex shapes. The basic elements of
the crown are the more or less conical
projections called cusps and the valleys
that separate them. The cusps contain
both dentine and enamel, whereas minor
projections on the crown, called
crenulations, are the result of different
enamel thickness. Cusps are occasionally
joined to form ridges and expanded to
form crests. Cingula are often incomplete
ridges that pass around the base of the

Mammalian, multicusped cheek teeth

probably evolved from single-cusped teeth
in reptilians, although the diversity of
therapsid molar patterns and the
complexity in the molars of the earliest
mammals make determining how this
happened impossible. According to the
widely accepted "differentiation theory",
additional cusps have arisen by budding or
outgrowth from the crown, while the
rivalling "concrescence theory" instead
proposes that complex teeth evolved by
the clustering of originally separate
conical teeth. Therian mammals
(placentals and marsupials) are generally
agreed to have evolved from an ancestor
with tribosphenic cheek teeth, with three
main cusps arranged in a triangle.[2]

Comparison of cheek teeth in various taxa: 1, reptile;

2, Dromatherium (a Triassic cynodont); 3,
Microconodon (a Triassic eucynodont); 4,
Spalacotherium (a Cretaceous symmetrodont); 5,
Amphitherium (a Jurassic mammal)

Generalized cusp of a mammalian molar: ant, anterior;

pos, posterior; ci, lingual cingulum; pa, paraconid; pr,
protoconid; me, metaconid; hy, hypoconid; hl,
hypoconulid; ec, entocristid; tb, talonid basin

Each major cusp on an upper molar is

called a cone and is identified by a prefix
dependent on its relative location on the
tooth: proto-, para-, meta-, hypo-, and ento-.
Suffixes are added to these names: -id is
added to cusps on a lower molar (e.g.,
protoconid); -ule to a minor cusp (e.g.,
protoconulid). A shelf-like ridge on the
lower part of the crown (on an upper
molar) is called a cingulum; the same
feature on the lower molar a cingulid, and
a minor cusp on these, for example, a
cingular cuspule or conulid.[3]

Tribosphenic …
Generalized tribosphenic molar: The protocone is on
the lingual (tongue) side, while the anterior paracone
and posterior metacone are on the buccal (cheek)
side of the jaw).

The design that is considered one of the

most important characteristics of
mammals is a three-cusped shape called a
tribosphenic molar. This molar design has
two important features: the trigonid, or
shearing end, and the talonid, or crushing
heel. In modern tribosphenic molars, the
trigonid is towards the front of the jaw and
the talonid is towards the rear.

The tribosphenic tooth is found in

insectivores and young platypuses (adults
have no teeth). Upper molars look like
three-pointed mountain ranges; lowers
look like two peaks and a third off to the
The tribosphenic design appears
primitively in all groups of mammals.
Some paleontologists believe that it
developed independently in monotremes
(or australosphenidans), rather than being
inherited from an ancestor that they share
with marsupials and placentals (or
boreosphenidans); but this idea has critics
and the debate is still going on.[4] For
example, the dentition of the Early
Cretaceous monotreme Steropodon is
similar to those of Peramus and
dryolestoids, which suggests that
monotremes are related to some pre-
tribosphenic therian mammals,[5] but, on
the other hand, the status of neither of
these two groups is well-established.

Some Jurassic mammals, such as

Shuotherium and Pseudotribos, have
"reversed tribosphenic" molars, in which
the talonid is towards the front. This
variant is regarded as an example of
convergent evolution.[6]
From the primitive tribosphenic tooth,
molars have diversified into several unique
morphologies. In many groups, a fourth
cusp, the hypocone (hypoconid),
subsequently evolved (see below).

Quadrate …

Pig tooth
Quadrate (also called quadritubercular or
euthemorphic) molars have an additional
fourth cusp on the lingual (tongue) side
called the hypocone, located posterior to
the protocone. Quadrate molars appeared
early in mammal evolution and are present
in many species, including hedgehogs,
raccoons, and many primates, including
humans.[7] There may be a fifth cusp.

In many mammals, additional smaller

cusps called conules appear between the
larger cusps. They are named after their
locations, e.g. a paraconule is located
between a paracone and a metacone, a
hypoconulid is located between a
hypoconid and an entoconid.[7]

Bunodont …
Upper and lower dentition of a chimpanzee

In bunodont molars, the cusps are low and

rounded hills rather than sharp peaks.
They are most common among omnivores
such as pigs, bears, and humans.[7]
Bunodont molars are effective crushing
devices and often basically quadrate in

Hypsodont …
Hypsodont dentition is characterized by
high-crowned teeth and enamel that
extends far past the gum line, which
provides extra material for wear and tear.[9]
Some examples of animals with
hypsodont dentition are cattle and horses,
all animals that feed on gritty, fibrous
material. Hypsodont molars can continue
to grow throughout life, for example in
some species of Arvicolinae (herbivorous
Hypsodont molars lack both a crown and a
neck. The occlusal surface is rough and
mostly flat, adapted for crushing and
grinding plant material. The body is
covered with cementum both above and
below the gingival line, below which is a
layer of enamel covering the entire length
of the body. The cementum and the
enamel invaginate into the thick layer of

Brachydont …
The opposite condition to hypsodont is
called brachydont or brachyodont (from
brachys, "short"). It is a type of dentition
characterized by low-crowned teeth.
Human teeth are brachydont.[7]

A brachydont tooth has a crown above the

gingival line and a neck just below it, and
at least one root. A cap of enamel covers
the crown and extends down to the neck.
Cementum is only found below the
gingival line. The occlusal surfaces tend to
be pointed, well-suited for holding prey and
tearing and shredding.[10]

Zalambdodont …

Zalambdodont molars have three cusps,

one larger on the lingual side and two
smaller on the labial side, joined by two
crests that form a V- or λ-shape. The larger
inner cusp might be homologous with the
paracone in a tribosphenic molar, but can
also be fused with the metacone. The
protocone is typically missing. The two
smaller labial cusps are located on an
expanded shelf called the stylar shelf.
Zalambdodont molars are found in, for
example, golden moles and solenodons.[7]

Dilambdodont …

Like zalambdodont molars, dilambdodont

molars have a distinct ectoloph, but are
shaped like two lambdas or a W. On the
lingual side, at the bottom of the W, are the
metacone and paracone, and the stylar
shelf is on the labial side. A protocone is
present lingual to the ectoloph.
Dilambdodont molars are present in
shrews, moles, and some insectivorous

Lophodont …
Lophodont molars of Elephas (left) and Loxodonta
(center), compared to the nonlophodont mastodon

Lophodont teeth are easily identified by

the differentiating patterns of ridges or
lophs of enamel interconnecting the cusps
on the crowns. Present in most herbivores,
these patterns of lophs can be a simple,
ring-like edge, as in mole rats, or a
complex arrangement of series of ridges
and cross-ridges, as those in odd-toed
ungulates, such as equids.[8]
Lophodont molars have hard and
elongated enamel ridges called lophs
oriented either along or perpendicular to
the dental row. Lophodont molars are
common in herbivores that grind their food
thoroughly. Examples include tapirs,
manatees, and many rodents.[7]

When two lophs form transverse, often

ring-shaped, ridges on a tooth, the
arrangement is called bilophodont. This
pattern is common in primates, but can
also be found in lagomorphs (hares,
rabbits, and pikas) and some rodents.[7][8]

Extreme forms of lophodonty in elephants

and some rodents (such as Otomys) is
known as loxodonty.[7] The African
elephant belongs to a genus called
Loxodonta because of this feature.

Selenodont …

In selenodont molars (so-named after

moon goddess Selene), the major cusp is
elongated into crescent-shaped ridge.
Examples include most even-toed
ungulates, such as cattle and deer.[7][8]

Secodont …

Carnassials of a Eurasian wolf

Many carnivorous mammals have
enlarged and blade-like teeth especially
adapted for slicing and chopping called
carnassials. A general term for such blade-
like teeth is secodont or plagiaulacoid.[7]

See also
Dental formula

1. Rozkovcová, E; Marková, M; Dolejsí, J
(1999). "Studies on agenesis of third
molars amongst populations of
different origin". Sbornik Lekarsky. 100
(2): 71–84. PMID 11220165 .
2. Zhao, Weiss & Stock 2000, Acquisition
of multi-cusped cheek teeth in
mammals, p. 154
3. Myers et al. 2013b
4. Stokstad 2001
5. Luo, Cifelli & Kielan-Jaworowska 2001
6. Luo, Ji & Yuan 2007
7. Myers et al. 2013a
8. Lawlor 1979, pp. 13–4
9. Flynn, Wyss & Charrier 2007
10. Kwan, Paul W.L. (2007). "Digestive
system I" (PDF). Tufts University.
Archived from the original (PDF) on
2012-09-13. Retrieved 2013-05-18.

Flynn, John J.; Wyss, André R.; Charrier,
Reynaldo (May 2007). "South America's
Missing Mammals" . Scientific American: 68–
75. OCLC 17500416 . Retrieved 2013-05-11.
Lawlor, T.E. (1979). "The Mammalian
Skeleton" (PDF). Handbook to the Orders and
Families of Living Mammals. Mad River Press.
ISBN 978-0-916422-16-5. OCLC 5763193 .
Archived from the original (PDF) on 2010-07-
01. Retrieved 2013-05-12.
Luo, Zhe-Xi; Cifelli, Richard L.; Kielan-
Jaworowska, Zofia (4 January 2001). "Dual
origin of tribosphenic mammals". Nature.
409 (6816): 53–7. doi:10.1038/35051023 .
PMID 11343108 .
Luo, Z.-X.; Ji, Q.; Yuan, C.-X. (November
2007). "Convergent dental adaptations in
pseudo-tribosphenic and tribosphenic
mammals". Nature. 450 (7166): 93–97.
doi:10.1038/nature06221 . PMID 17972884 .
Myers, P.; Espinosa, R.; Parr, C. S.; Jones, T.;
Hammond, G. S.; Dewey, T. A. (2013a). "The
Basic Structure of Cheek Teeth" . Animal
Diversity Web, University of Michigan.
Retrieved 2013-05-12.
Myers, P.; Espinosa, R.; Parr, C. S.; Jones, T.;
Hammond, G. S.; Dewey, T. A. (2013b). "The
Diversity of Cheek Teeth" . Animal Diversity
Web, University of Michigan. Archived from
the original on 2013-04-05. Retrieved
Stokstad, E. (January 2001). "Tooth Theory
Revises History of Mammals" . Science. 291
(5501): 26.
501.26 . PMID 11191993 .
Zhao, Z.; Weiss, K. M.; Stock, D. W. (2000).
"Development and evolution of dentition
patterns and their genetic basis". In Teaford,
Mark F; Smith, Moya Meredith; Ferguson,
Mark WJ (eds.). Development, Function and
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Press. pp. 152–72. ISBN 978-0-511-06568-2.

External links
Overview of molar morphology and
terminology -

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