Paraphyly

Article
Talk
Read
Edit
View history

Tools
From Wikipedia, the free encyclopedia

In this phylogenetic tree, the green group is paraphyletic; it is composed of a

common ancestor (the lowest green vertical stem) and some of its descendants, but
it excludes the blue group (a monophyletic group) which diverged from the green
group.
In taxonomy, a grouping is paraphyletic if it consists of the grouping's last
common ancestor and most of its descendants, but excludes a few monophyletic
subgroups. The grouping is said to be paraphyletic with respect to the excluded
subgroups. In contrast, a monophyletic grouping (a clade) includes a common
ancestor and all of its descendants. The terms are commonly used in phylogenetics
(a subfield of biology) and in the tree model of historical linguistics.
Paraphyletic groups are identified by a combination of synapomorphies and
symplesiomorphies. If many subgroups are missing from the named group, it is said
to be polyparaphyletic.

The term was coined by Willi Hennig to apply to well-known taxa like Reptilia
(reptiles) which is paraphyletic with respect to birds. Reptilia contains the last
common ancestor of reptiles and all descendants of that ancestor except for birds.
Other commonly recognized paraphyletic groups include fish, monkeys, and lizards.
[1][page needed]

Etymology
The term paraphyly, or paraphyletic, derives from the two Ancient Greek words παρά
(pará), meaning "beside, near", and φῦλον (phûlon), meaning "genus, species",[2][3]
and refers to the situation in which one or several monophyletic subgroups of
organisms (e.g., genera, species) are left apart from all other descendants of a
unique common ancestor.

Conversely, the term monophyly, or monophyletic, builds on the Ancient Greek prefix
μόνος (mónos), meaning "alone, only, unique",[2][3] and refers to the fact that a
monophyletic group includes organisms consisting of all the descendants of a unique
common ancestor.

By comparison, the term polyphyly, or polyphyletic, uses the Ancient Greek prefix
πολύς (polús), meaning "many, a lot of",[2][3] and refers to the fact that a
polyphyletic group includes organisms arising from multiple ancestral sources.

Phylogenetics

Cladogram of the primates, showing a monophyly (the simians, in yellow), a

paraphyly (the prosimians, in blue, including the red patch), and a polyphyly (the
night-active primates, the lorises and the tarsiers, in red).
In cladistics
Further information: Cladistics
Groups that include all the descendants of a common ancestor are said to be
monophyletic. A paraphyletic group is a monophyletic group from which one or more
subsidiary clades (monophyletic groups) are excluded to form a separate group.
Philosopher of science Marc Ereshefsky has argued that paraphyletic taxa are the
result of anagenesis in the excluded group or groups.[4] Cladists do not grant
paraphyletic assemblages the status of "groups" or reify them with explanations,
because they represent evolutionary non-events [5]

A group whose identifying features evolved convergently in two or more lineages is

polyphyletic (Greek πολύς [polys], "many"). More broadly, any taxon that is not
paraphyletic or monophyletic can be called polyphyletic. Empirically, the
distinction between polyphyletic groups and paraphyletic groups is rather
arbitrary, since the character states of common ancestors are inferences, not
observations.[citation needed]

These terms were developed during the debates of the 1960s and 1970s accompanying
the rise of cladistics.

Paraphyletic groupings are considered problematic by many taxonomists, as it is not

possible to talk precisely about their phylogenetic relationships, their
characteristic traits and literal extinction.[6][7] Related terms are stem group,
chronospecies, budding cladogenesis, anagenesis, or 'grade' groupings. Paraphyletic
groups are often relics from outdated hypotheses of phylogenic relationships from
before the rise of cladistics.[8]

Examples

Wasps are paraphyletic, consisting of the clade Apocrita without ants and bees,
which are not usually considered to be wasps; the sawflies ("Symphyta") too are
paraphyletic, as the Apocrita are nested inside the Symphytan clades.
The prokaryotes (single-celled life forms without cell nuclei) are a paraphyletic
grouping, because they exclude the eukaryotes, a descendant group. Bacteria and
Archaea are prokaryotes, but archaea and eukaryotes share a common ancestor that is
not ancestral to the bacteria. The prokaryote/eukaryote distinction was proposed by
Edouard Chatton in 1937[9] and was generally accepted after being adopted by Roger
Stanier and C.B. van Niel in 1962. The botanical code (the ICBN, now the ICN)
abandoned consideration of bacterial nomenclature in 1975; currently, prokaryotic
nomenclature is regulated under the ICNB with a starting date of 1 January 1980 (in
contrast to a 1753 start date under the ICBN/ICN).[10]

Among plants, dicotyledons (in the traditional sense) are paraphyletic because the
group excludes monocotyledons. "Dicotyledon" has not been used as a botanic
classification for decades, but is allowed as a synonym of Magnoliopsida.[note 1]
Phylogenetic analysis indicates that the monocots are a development from a dicot
ancestor. Excluding monocots from the dicots makes the latter a paraphyletic group.
[11]

Among animals, several familiar groups are not, in fact, clades. The order
Artiodactyla (even-toed ungulates) as traditionally defined is paraphyletic because
it excludes Cetaceans (whales, dolphins, etc.). Under the ranks of the ICZN Code,
the two taxa are separate orders. Molecular studies, however, have shown that the
Cetacea descend from artiodactyl ancestors, although the precise phylogeny within
the order remains uncertain. Without the Cetaceans the Artiodactyls are
paraphyletic.[12] The class Reptilia is paraphyletic because it excludes birds
(class Aves). Under the ranks of the ICZN Code, these two taxa are separate
classes. However birds are sister taxon to a group of dinosaurs (part of Diapsida),
both of which are "reptiles".[13] .

Osteichthyes, bony fish, are paraphyletic when circumscribed to include only

Actinopterygii (ray-finned fish) and Sarcopterygii (lungfish, etc.), and to exclude
tetrapods; more recently, Osteichthyes is treated as a clade, including the
tetrapods.[14][15]

The "wasps" are paraphyletic, consisting of the narrow-waisted Apocrita without the
ants and bees.[16] The sawflies (Symphyta) are similarly paraphyletic, forming all
of the Hymenoptera except for the Apocrita, a clade deep within the sawfly tree.
[14] Crustaceans are not a clade because the Hexapoda (insects) are excluded. The
modern clade that spans all of them is the Tetraconata.[17][18]

One of the goals of modern taxonomy over the past fifty years has been to eliminate
paraphyletic "groups", such as the examples given here, from formal
classifications.[19][20]