Prototheria (; from Greek πρώτος, ''prōtos'', first, + θήρ, ''thēr'', wild animal) is a subclass to which the orders Monotremata,

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 ...

Docodonta Docodonta is an order of extinct mammaliaforms that lived during the Mesozoic, from the Middle Jurassic to Early Cretaceous. They are distinguished from other early mammaliaforms by their relatively complex molar teeth, from which the order ...

, Triconodonta and

Multituberculata Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...

have been assigned, although the validity of the subclass has been questioned. Most of the animals in this group are extinct. The egg-laying monotremes are known from fossils of 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 ...

and

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 ...

periods; they are represented today by 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 ...

and several species of

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 ...

. The names Prototheria,

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 ...

, and

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 ...

(meaning "first beasts", "changed beasts", and "true beasts", respectively) refer to the three mammalian groupings of which we have living representatives. Each of the three may be defined as a total

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 ...

containing a living crown-group (respectively the Monotremata,

Marsupialia Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...

and

Placentalia Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...

) 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 Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The stori ...

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

infraclass In biological classification, class ( la, classis) is a taxonomic rank, as well as a taxonomic unit, a taxon, in that rank. It is a group of related taxonomic orders. Other well-known ranks in descending order of size are life, domain, king ...

es of the subclass

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. ...

, and in more recent proposals have been demoted further (to cohorts or even

magnorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...

s), as

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

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 a hypothesis which defined the group by two supposed

synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ha ...

: (1) formation of the side wall of the braincase from a bone called the anterior lamina, contrasting with the

alisphenoid The greater wing of the sphenoid bone, or alisphenoid, is a bony process of the sphenoid bone; there is one on each side, extending from the side of the body of the sphenoid and curving upward, laterally, and backward. Structure The greater win ...

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 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 ...

fossil orders (

, Triconodonta and

) in a broader clade for which the name Prototheria was retained, and of which monotremes were thought to be only the last surviving branch. 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. 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 The Australosphenida are a clade of mammals, containing mammals with tribosphenic molars, known from the Jurassic to Mid-Cretaceous of Gondwana. They are thought to have acquired their tribosphenic molars independently from those of Tribosphenid ...

, and also suggest that the triangular array of cusps may have evolved independently in australosphenidans and therians. 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.

Notes

{{Taxonbar, from=Q715778 Mammal taxonomy Taxa named by Theodore Gill