Prototheria
Encyclopedia
Prototheria is a taxonomic group, or taxon
, to which the order Monotremata
belongs. It is conventionally ranked as a subclass within the mammal
s.
Most of the animals in this group are extinct. The egg-laying monotremes are known from fossils of the Cretaceous
and Cenozoic
periods; they are represented today by the platypus
and several species of echidna
.
The names Prototheria, Metatheria
and Eutheria
(meaning "first beasts", "changed beasts", and "true beasts") refer to the three mammalian groupings which have living representatives. Each of the three may be defined as a total clade
containing a living crown-group (respectively the Monotremata, Marsupialia and Placentalia) plus any fossil species which are more closely related to that crown-group than to any other living animals.
The threefold division of living mammals into monotremes, marsupials and placentals was already well established when Thomas Huxley
proposed the names Metatheria and Eutheria to incorporate the two latter groups in 1880. Initially treated as subclasses, Metatheria and Eutheria are by convention now grouped as infraclasses of the subclass Theria
, and in more recent proposals have been demoted further (to cohorts or even magnorders), as cladistic reappraisals of the relationships between living and fossil mammals have suggested that the Theria itself should be reduced in rank.
Prototheria, on the other hand, was generally recognised as a subclass until quite recently, on the basis of an hypothesis which defined the group by two supposed synapomorphies: (1) formation of the side wall of the braincase from a bone called the anterior lamina, contrasting with the alisphenoid in therians; and (2) a linear alignment of molar cusps, contrasting with a triangular arrangement in therians. These characters appeared to unite monotremes with a range of Mesozoic
fossil orders (Morganucodon
ta, Triconodonta
, Docodonta
and Multituberculata
) in a broader clade for which the name Prototheria was retained, and of which monotremes were thought to be only the last surviving branch (Benton 2005: 300, 306).
The evidence which was held to support this grouping is now universally discounted. In the first place, examination of embryos has revealed that the development of the braincase wall is essentially identical in therians and in 'prototherians': the anterior lamina simply fuses with the alisphenoid in therians, and therefore the 'prototherian' condition of the braincase wall is primitive for all mammals while the therian condition can be derived from it. Additionally, the linear alignment of molar cusps is also primitive for all mammals. Therefore, neither of these states can supply a uniquely shared derived character which would support a 'prototherian' grouping of orders in contradistinction to Theria (Kemp 1983).
In a further reappraisal, the molars of embryonic and fossil monotremes (living monotreme adults are toothless) appear to demonstrate an ancestral pattern of cusps which is similar to the triangular arrangement observed in therians. Some peculiarities of this dentition support an alternative grouping of monotremes with certain recently-discovered fossil forms into a proposed new clade known as the Australosphenida
, and also suggest that the triangular array of cusps may have evolved independently in australosphenidans and therians (Luo et al. 2001, 2002).
The Australosphenida hypothesis remains controversial, and some taxonomists (e.g. McKenna & Bell 1997) prefer to maintain the name Prototheria as a fitting contrast to the other group of living mammals, the Theria. In theory, the Prototheria is taxonomically redundant, since Monotremata is currently the only order which can still be confidently included, but its retention might be justified if new fossil evidence, or a re-examination of known fossils, enables extinct relatives of the monotremes to be identified and placed within a wider grouping.
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
, to which the order Monotremata
Monotreme
Monotremes are mammals that lay eggs instead of giving birth to live young like marsupials and placental mammals...
belongs. It is conventionally ranked as a subclass within the mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
s.
Most of the animals in this group are extinct. The egg-laying monotremes are known from fossils of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
and Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...
periods; they are represented today by the platypus
Platypus
The platypus is a semi-aquatic mammal endemic to eastern Australia, including Tasmania. Together with the four species of echidna, it is one of the five extant species of monotremes, the only mammals that lay eggs instead of giving birth to live young...
and several species of echidna
Echidna
Echidnas , also known as spiny anteaters, belong to the family Tachyglossidae in the monotreme order of egg-laying mammals. There are four extant species, which, together with the platypus, are the only surviving members of that order and are the only extant mammals that lay eggs...
.
The names Prototheria, Metatheria
Metatheria
Metatheria is a grouping within the animal class Mammalia. First proposed by Thomas Henry Huxley in 1880, it is nearly synonymous with the earlier taxon Marsupialia though it is slightly wider since it also contains the nearest fossil relatives of marsupial mammals.The earliest known...
and Eutheria
Eutheria
Eutheria is a group of mammals consisting of placental mammals plus all extinct mammals that are more closely related to living placentals than to living marsupials . They are distinguished from noneutherians by various features of the feet, ankles, jaws and teeth...
(meaning "first beasts", "changed beasts", and "true beasts") refer to the three mammalian groupings which have living representatives. Each of the three may be defined as a total clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
containing a living crown-group (respectively the Monotremata, Marsupialia and Placentalia) plus any fossil species which are more closely related to that crown-group than to any other living animals.
The threefold division of living mammals into monotremes, marsupials and placentals was already well established when Thomas Huxley
Thomas Huxley
Thomas Henry Huxley PC FRS was an English biologist, known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution....
proposed the names Metatheria and Eutheria to incorporate the two latter groups in 1880. Initially treated as subclasses, Metatheria and Eutheria are by convention now grouped as infraclasses of the subclass Theria
Theria
Theria is a subclass of mammals that give birth to live young without using a shelled egg, including both eutherians and metatherians . The only omitted extant mammal group is the egg-laying monotremes....
, and in more recent proposals have been demoted further (to cohorts or even magnorders), as cladistic reappraisals of the relationships between living and fossil mammals have suggested that the Theria itself should be reduced in rank.
Prototheria, on the other hand, was generally recognised as a subclass until quite recently, on the basis of an hypothesis which defined the group by two supposed synapomorphies: (1) formation of the side wall of the braincase from a bone called the anterior lamina, contrasting with the alisphenoid in therians; and (2) a linear alignment of molar cusps, contrasting with a triangular arrangement in therians. These characters appeared to unite monotremes with a range of Mesozoic
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...
fossil orders (Morganucodon
Morganucodon
Morganucodon is an early mammalian genus which lived during the Late Triassic. It first appeared about 205 million years ago. This has also been identified with Eozostrodon. Unlike many other early mammals, Morganucodon is well represented by abundant and well preserved, though in the vast...
ta, Triconodonta
Triconodonta
Triconodonta is the generic name for a group of early mammals which were close relatives of the ancestors of all present-day mammals. Triconodonts lived between the Triassic and the Cretaceous. They are one of the groups that can be classified as mammals by any definition...
, Docodonta
Docodonta
Docodonta is an order of extinct proto-mammals that lived during the mid- to late-Mesozoic era. Their most distinguishing physical features were their relatively sophisticated set of molars, from which the order gets its name. In the fossil record, Docodonta is represented primarily by isolated...
and Multituberculata
Multituberculata
The Multituberculata were a group of rodent-like mammals that existed for approximately one hundred and twenty million years—the longest fossil history of any mammal lineage—but were eventually outcompeted by rodents, becoming extinct during the early Oligocene. At least 200 species are...
) in a broader clade for which the name Prototheria was retained, and of which monotremes were thought to be only the last surviving branch (Benton 2005: 300, 306).
The evidence which was held to support this grouping is now universally discounted. In the first place, examination of embryos has revealed that the development of the braincase wall is essentially identical in therians and in 'prototherians': the anterior lamina simply fuses with the alisphenoid in therians, and therefore the 'prototherian' condition of the braincase wall is primitive for all mammals while the therian condition can be derived from it. Additionally, the linear alignment of molar cusps is also primitive for all mammals. Therefore, neither of these states can supply a uniquely shared derived character which would support a 'prototherian' grouping of orders in contradistinction to Theria (Kemp 1983).
In a further reappraisal, the molars of embryonic and fossil monotremes (living monotreme adults are toothless) appear to demonstrate an ancestral pattern of cusps which is similar to the triangular arrangement observed in therians. Some peculiarities of this dentition support an alternative grouping of monotremes with certain recently-discovered fossil forms into a proposed new clade known as the Australosphenida
Australosphenida
The Australosphenida are a clade of mammals. Today, living specimens exist only in Australia and New Guinea with only five surviving species, but fossils have been found in Madagascar and Argentina...
, and also suggest that the triangular array of cusps may have evolved independently in australosphenidans and therians (Luo et al. 2001, 2002).
The Australosphenida hypothesis remains controversial, and some taxonomists (e.g. McKenna & Bell 1997) prefer to maintain the name Prototheria as a fitting contrast to the other group of living mammals, the Theria. In theory, the Prototheria is taxonomically redundant, since Monotremata is currently the only order which can still be confidently included, but its retention might be justified if new fossil evidence, or a re-examination of known fossils, enables extinct relatives of the monotremes to be identified and placed within a wider grouping.