Paraphyly

In taxonomy, a group is paraphyletic if it

consists of the group's last common
ancestor and all descendants of that
ancestor excluding a few—typically only
one or two—monophyletic subgroups.
The group is said to be paraphyletic with
respect to the excluded subgroups. A
paraphyletic group cannot be a clade, or
monophyletic group, which is any group
of species that includes a common
ancestor and all of its descendants. One
or more members of a paraphyletic
group is more closely related to the
excluded group(s) than it is to other
members of the paraphyletic group. The
term is commonly used in phylogenetics
(a subfield of biology) and in linguistics.
Paraphyletic groups are identified by a
combination of synapomorphies and
symplesiomorphies.

In this phylogenetic tree, the green group is

paraphyletic; it is composed of a common ancestor
(the lowest green vertical stem) and its
descendants, but it excludes the blue group (a
monophyletic group) which diverged from the green
group.
The term was coined by Willi Hennig to
apply to well-known taxa like Reptilia
(reptiles) which, as commonly named
and traditionally defined, is paraphyletic
with respect to mammals and birds.
Reptilia contains the last common
ancestor of reptiles and all descendants
of that ancestor, including all extant
reptiles as well as the extinct synapsids,
except for mammals and birds. Other
commonly recognized paraphyletic
groups include fish, monkeys, and
lizards.[1]

If many subgroups are missing from the

named group, it is said to be
polyparaphyletic.
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
Reptilia (green field) is a paraphyletic group
comprising all amniotes (Amniota) except for two
subgroups: Mammalia (mammals) and Aves (birds);
therefore, Reptilia is not a clade. In contrast,

Amniota itself is a clade, which is a monophyletic

group.
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 …

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.

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), 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, as
traditionally defined, is paraphyletic
because it excludes birds (class Aves)
and mammals. Under the ranks of the
ICZN Code, these three taxa are separate
classes. However, mammals hail from
the synapsids (which were once
described as "mammal-like reptiles") and
birds are sister taxon to a group of
dinosaurs (part of Diapsida), both of
which are "reptiles".[13] Alternatively,
reptiles are paraphyletic because they
gave rise to (only) birds. Birds and
reptiles together make Sauropsids, a
clade of Amniota that is the sister group
of the clade that includes mammals.

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 here from formal
classifications. [19][20]
Paraphyly in species …

Species have a special status in

systematics as being an observable
feature of nature itself and as the basic
unit of classification.[21] Some
articulations of the phylogenetic species
concept require species to be
monophyletic, but paraphyletic species
are common in nature, to the extent that
they do not have a single common
ancestor. Indeed, for sexually
reproducing taxa, no species has a
"single common ancestor" organism.
Paraphyly is common in speciation,
whereby a mother species (a
paraspecies) gives rise to a daughter
species without itself becoming
extinct.[22] Research indicates as many
as 20 percent of all animal species and
between 20 and 50 percent of plant
species are paraphyletic.[23][24]
Accounting for these facts, some
taxonomists argue that paraphyly is a
trait of nature that should be
acknowledged at higher taxonomic
levels.[25][26]

Cladists advocate a phylogenetic species

concept [27] that does not consider
species to exhibit the properties of
monophyly or paraphyly, concepts under
that perspective which apply only to
groups of species. [28] They consider
Zander's extension of the "paraphyletic
species" argument to higher taxa to
represent a category error [29]

Uses for paraphyletic groups …

When the appearance of significant traits

has led a subclade on an evolutionary
path very divergent from that of a more
inclusive clade, it often makes sense to
study the paraphyletic group that
remains without considering the larger
clade. For example, the Neogene
evolution of the Artiodactyla (even-toed
ungulates, like deer, cows, pigs and
hippopotamuses - note that Cervidae,
Bovidae, Suidae and Hippopotamidae,
the families that contain these various
artiodactyls, are all monophyletic groups)
has taken place in environments so
different from that of the Cetacea
(whales, dolphins, and porpoises) that
the Artiodactyla are often studied in
isolation even though the cetaceans are
a descendant group. The prokaryote
group is another example; it is
paraphyletic because it is composed of
two Domains (Eubacteria and Archaea)
and excludes (the eukaryotes). It is very
useful because it has a clearly defined
and significant distinction (absence of a
cell nucleus, a plesiomorphy) from its
excluded descendants.
Also, some systematists recognize
paraphyletic groups as being involved in
evolutionary transitions, the development
of the first tetrapods from their ancestors
for example. Any name given to these
hypothetical ancestors to distinguish
them from tetrapods—"fish", for example
—necessarily picks out a paraphyletic
group, because the descendant
tetrapods are not included.[30] Other
systematists consider reification of
paraphyletic groups to obscure inferred
patterns of evolutionary history.[31]

The term "evolutionary grade" is

sometimes used for paraphyletic
groups.[32] Moreover, the concepts of
monophyly, paraphyly, and polyphyly
have been used in deducing key genes
for barcoding of diverse group of
species.[33]

Independently evolved traits …

Current phylogenetic hypotheses of

tetrapod relationships imply that
viviparity, the production of offspring
without the external laying of a fertilized
egg, developed independently in the
lineages that led to humans (Homo
sapiens) and southern water skinks
(Eulampus tympanum, a kind of lizard).
Put another way, viviparity is a
synapomorphy for Theria within
mammals, and an autapomorphy for
Eulamprus tympanum (or perhaps a
synapomorphy, if other Eulamprus
species are also viviparous).

"Groups" based on independently-

developed traits such as these examples
of viviparity represent examples of
polyphyly, not paraphyly.

Not paraphyly …

Amphibious fish are polyphyletic, not

paraphyletic. Although they appear
similar, several different groups of
amphibious fishes such as
mudskippers and lungfishes evolved
independently in a process of
convergent evolution in distant
relatives faced with similar ecological
circumstances.[34]
Flightless birds are polyphyletic
because they independently (in
parallel) lost the ability to fly.[35]
Animals with a dorsal fin are not
paraphyletic, even though their last
common ancestor may have had such
a fin, because the Mesozoic ancestors
of porpoises did not have such a fin,
whereas pre-Mesozoic fish did have
one.
Quadrupedal archosaurs are not a
paraphyletic group. Bipedal dinosaurs
 like Eoraptor, ancestral to quadrupedal
ones, were descendants of the last
common ancestor of quadrupedal
dinosaurs and other quadrupedal
archosaurs like the crocodilians.

Non-exhaustive list of paraphyletic

groups
…

The following list recapitulates a number

of paraphyletic groups proposed in the
literature, and provides the
corresponding monophyletic taxa.
 Paraphyletic Corresponding References and
Excluded clades
taxon monophyletic taxon notes
[36]
Prokaryotes Eukaryota Cellular organisms

Animalia, Plantae, and [37]

Protista Eukaryota
Fungi
[38]
Invertebrates Vertebrata Animalia

Sponge Eumetazoa Animalia [39] [40]

[41][42]
Worm Multiple groups Nephrozoa
[43]
Radiata Bilateria Eumetazoa

Lophotrochozoa, [44]
Platyzoa Spiralia
Mesozoa
[45]
Fish Tetrapoda Vertebrate

Reptiles Birds Sauropsida [46]

[47]
Lizard Snakes, Amphisbaenia Squamates

Cimolodonta, [48]
Plagiaulacidans Multituberculata
Arginbaataridae
[49]
Pelycosaurs Therapsida Synapsida

Even-toed [50][51]
Cetacea Cetartiodactyla
ungulates
[52]
Archaeoceti Neoceti Cetacea
[53]
Prosimians Simiiformes Primates

Crustaceans Hexapoda Tetraconata [17][18]

[54]
Wasps Ants, Bees Apocrita

Sawfly Apocrita Hymenoptera [14]

[55]
Vespoidea Apoidea, Ants Euaculeata
[56]
Parasitica Aculeata Apocrita
[57]
Nautiloidea Ammonoidea, Coleoidea Cephalopoda

Embryophyte (Land [58]

Charophyte Streptophyta
plants)

Bryophyte Tracheophyte Embryophyte [59][58]

[60]
Gymnosperm Angiosperm Spermatophyte

Dicotyledon Monocotyledon Angiosperm [11]

 [61]
Moth Butterfly Lepidoptera

Coral Medusozoa, Myxozoa Cnidaria [62][63]

[64][65][66]
Jellyfish Hydroidolina Medusozoa
[67][68]
Cycloneuralia Panarthropoda Ecdysozoa

Rotifera Acanthocephala Syndermata [69][70]

Mecoptera Siphonaptera Mecopteroidea

Leptothecata,
Anthoathecata Hydroidolina
Siphonophorae

Monkey Hominoidea Simiiformes

Linguistics
The concept of paraphyly has also been
applied to historical linguistics, where the
methods of cladistics have found some
utility in comparing languages. For
instance, the Formosan languages form
a paraphyletic group of the Austronesian
languages because they consist of the
nine branches of the Austronesian family
that are not Malayo-Polynesian and are
restricted to the island of Taiwan.[71]
See also
Glossary of scientific naming

Notes
1. The history of flowering plant
classification can be found under
History of the classification of
flowering plants.

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Paraphyletic groups as natural units of
biological classification

External links

Look up paraphyletic in Wiktionary,

the free dictionary.

Funk, D. J.; Omland, K. E. (2003).

"Species-level paraphyly and polyphyly:
Frequency, cause and consequences,
with insights from animal
 mitochondrial DNA" (PDF). Annual
Review of Ecology, Evolution, and
Systematics. 34: 397–423.
doi:10.1146/annurev.ecolsys.34.01180
2.132421 .

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