Clade

A clade (/kleɪd/;[1][2] from Ancient Greek:

κλάδος, klados, "branch"), also known as a
monophyletic group or natural group,[3] is
a group of organisms that are
monophyletic—that is, composed of a
common ancestor and all its lineal
descendants.[4] Rather than the English
term, the equivalent Latin term cladus
(plural cladi) is often used in taxonomical
literature.
Cladogram (family tree) of a biological group. The last
common ancestor is the vertical line stem at the
bottom. The blue and red subgroups are clades; each
shows its common ancestor stem at the bottom of
the subgroup branch. The green subgroup is not a
clade; it is a paraphyletic group, because it excludes
the blue branch, even though it has also descended
from a common ancestor. The green subgroup
together with the blue one forms a clade again.

The common ancestor may be an

individual, a population, a species (extinct
or extant), and so on right up to a kingdom
and further. Clades are nested, one in
another, as each branch in turn splits into
smaller branches. These splits reflect
evolutionary history as populations
diverged and evolved independently.
Clades are termed monophyletic (Greek:
"one clan") groups.

Over the last few decades, the cladistic

approach has revolutionized biological
classification and revealed surprising
evolutionary relationships among
organisms.[5] Increasingly, taxonomists try
to avoid naming taxa that are not clades;
that is, taxa that are not monophyletic.
Some of the relationships between
organisms that the molecular biology arm
of cladistics has revealed are that fungi
are closer relatives to animals than they
are to plants, archaea are now considered
different from bacteria, and multicellular
organisms may have evolved from
archaea.[6]

The term "clade" is also used with a similar

meaning in other fields besides biology,
such as historical linguistics; see
Cladistics § In disciplines other than
biology.

Etymology
The term "clade" was coined in 1957 by
the biologist Julian Huxley to refer to the
result of cladogenesis, the evolutionary
splitting of a parent species into two
distinct species, a concept Huxley
borrowed from Bernhard Rensch.[7][8]

Many commonly named groups, rodents

and insects for example, are clades
because, in each case, the group consists
of a common ancestor with all its
descendant branches. Rodents, for
example, are a branch of mammals that
split off after the end of the period when
the clade Dinosauria stopped being the
dominant terrestrial vertebrates 66 million
years ago. The original population and all
its descendants are a clade. The rodent
clade corresponds to the order Rodentia,
and insects to the class Insecta. These
clades include smaller clades, such as
chipmunk or ant, each of which consists
of even smaller clades. The clade "rodent"
is in turn included in the mammal,
vertebrate and animal clades.

History of nomenclature and

taxonomy
Early phylogenetic tree by Haeckel, 1866. Groups once
thought to be more advanced, such as birds ("Aves"),
are placed at the top.

The idea of a clade did not exist in pre-

Darwinian Linnaean taxonomy, which was
based by necessity only on internal or
external morphological similarities
between organisms – although as it
happens, many of the better known animal
groups in Linnaeus' original Systema
Naturae (notably among the vertebrate
groups) do represent clades. The
phenomenon of convergent evolution is,
however, responsible for many cases
where there are misleading similarities in
the morphology of groups that evolved
from different lineages.

With the increasing realization in the first

half of the 19th century that species had
changed and split through the ages,
classification increasingly came to be
seen as branches on the evolutionary tree
of life. The publication of Darwin's theory
of evolution in 1859 gave this view
increasing weight. Thomas Henry Huxley,
an early advocate of evolutionary theory,
proposed a revised taxonomy based on a
concept strongly resembling clades,[9]
although the term clade itself would not be
coined until 1957 by his grandson, Julian
Huxley. For example, the elder Huxley
grouped birds with reptiles, based on fossil
evidence.[9]

German biologist Emil Hans Willi Hennig

(1913 – 1976) is considered to be the
founder of cladistics.[10] He proposed a
classification system that represented
repeated branchings of the family tree, as
opposed to the previous systems, which
put organisms on a "ladder", with
supposedly more "advanced" organisms at
the top.[5][11]

Taxonomists have increasingly worked to

make the taxonomic system reflect
evolution.[11] When it comes to naming,
however, this principle is not always
compatible with the traditional rank-based
nomenclature (in which only taxa
associated with a rank can be named)
because there are not enough ranks to
name a long series of nested clades. For
these and other reasons, phylogenetic
nomenclature has been developed; it is
still controversial.

As an example, the full current

classification of Anas platyrhynchos (the
mallard duck) has 40 clades from
Eukaryota down: see
https://species.wikimedia.org/wiki/Anas_p
latyrhynchos and click on "Expand".

Definitions

Gavialidae, Crocodylidae and Alligatoridae are clade

names that are here applied to a phylogenetic tree of
crocodylians.

A clade is by definition monophyletic,

meaning that it contains one ancestor
(which can be an organism, a population,
or a species) and all its
descendants.[note 1][12][13] The ancestor can
be known or unknown; any and all
members of a clade can be extant or
extinct.

Clades and phylogenetic trees

The science that tries to reconstruct
phylogenetic trees and thus discover
clades is called phylogenetics or
cladistics, the latter term coined by Ernst
Mayr (1965), derived from "clade". The
results of phylogenetic/cladistic analyses
are tree-shaped diagrams called
cladograms; they, and all their branches,
are phylogenetic hypotheses.[14]

Three methods of defining clades are

featured in phylogenetic nomenclature:
node-, stem-, and apomorphy-based (see
Phylogenetic nomenclature§Phylogenetic
definitions of clade names for detailed
definitions).

Terminology

Cladogram of modern primate groups; all tarsiers are

C adog a o ode p ate g oups; a ta s e s a e
haplorhines, but not all haplorhines are tarsiers; all
apes are catarrhines, but not all catarrhines are apes;
etc.

The relationship between clades can be

described in several ways:

A clade located within a clade is said to

be nested within that clade. In the
diagram, the hominoid clade, i.e. the
apes and humans, is nested within the
primate clade.
Two clades are sisters if they have an
immediate common ancestor. In the
diagram, lemurs and lorises are sister
clades, while humans and tarsiers are
not.
A clade A is basal to a clade B if A
branches off the lineage leading to B
before the first branch leading only to
members of B. In the adjacent diagram,
the strepsirrhine/prosimian clade, is
basal to the hominoids/ape clade.
However, in this example, both
Haplorrhine as prosimians should be
considered as most basal groupings. It
is better to say that the prosimians are
the sister group to the rest of the
primates.[15] This way one also avoids
unintended and misconceived
connotations about evolutionary
 advancement, complexity, diversity,
ancestor status, and ancienity e.g. due
to impact of sampling diversity and
extinction.[15][16] Basal clades should not
be confused with stem groupings, as the
latter is associated with paraphyletic or
unresolved groupings.

In popular culture
Clade is the title of a novel by James
Bradley, who chose it both because of its
biological meaning and also because of
the larger implications of the word.[17]

An episode of Elementary is titled "Dead

Clade Walking" and deals with a case
involving a rare fossil.

See also
Adaptive radiation
Binomial nomenclature
Biological classification
Cladistics
Crown group
Monophyly
Paraphyly
Phylogenetic network
Phylogenetic nomenclature
Phylogenetics
Polyphyly
Notes
1. A semantic case has been made that
the name should be "holophyletic", but
this term has not acquired widespread
use. For more information, see
holophyly.

References
1. Wells, John C. (2008). Longman
Pronunciation Dictionary (3rd ed.).
Longman. ISBN 978-1-4058-8118-0.
2. "clade" . Merriam-Webster Dictionary.
Retrieved 19 April 2020.
3. Martin, Elizabeth; Hin, Robert (2008). A
Dictionary of Biology. Oxford
University Press.
4. Cracraft, Joel; Donoghue, Michael J.,
eds. (2004). "Introduction".
Assembling the Tree of Life . Oxford
University Press. p. 1 . ISBN 978-0-19-
972960-9.
5. Palmer, Douglas (2009). Evolution: The
Story of Life. Berkeley: University of
California Press. p. 13.
 . Pace, Norman R. (18 May 2006). "Time
for a change". Nature. 441 (7091):
289. Bibcode:2006Natur.441..289P .
doi:10.1038/441289a . ISSN 1476-
4687 . PMID 16710401 .
S2CID 4431143 .
7. Dupuis, Claude (1984). "Willi Hennig's
impact on taxonomic thought" .
Annual Review of Ecology and
Systematics. 15: 1–24.
doi:10.1146/annurev.es.15.110184.00
0245 .
 . Huxley, J. S. (1957). "The three types
of evolutionary process". Nature. 180
(4584): 454–455.
Bibcode:1957Natur.180..454H .
doi:10.1038/180454a0 .
S2CID 4174182 .
9. Huxley, T.H. (1876): Lectures on
Evolution. New York Tribune. Extra. no
36. In Collected Essays IV: pp 46-138
original text w/ figures
10. Brower, Andrew V. Z. (2013). "Willi
Hennig at 100". Cladistics. 30 (2):
224–225. doi:10.1111/cla.12057 .
11. ”Evolution 101". page 10 .
Understanding Evolution website.
University of California, Berkeley.
Retrieved 26 February 2016.
12. "International Code of Phylogenetic
Nomenclature. Version 4c. Chapter I.
Taxa" . 2010. Retrieved 22 September
2012.
13. Envall, Mats (2008). "On the difference
between mono-, holo-, and
paraphyletic groups: a consistent
distinction of process and pattern" .
Biological Journal of the Linnean
Society. 94: 217. doi:10.1111/j.1095-
8312.2008.00984.x .
14. Nixon, Kevin C.; Carpenter, James M.
(1 September 2000). "On the Other
"Phylogenetic Systematics" ".
Cladistics. 16 (3): 298–318.
doi:10.1111/j.1096-
0031.2000.tb00285.x .
S2CID 73530548 .
15. Krell, F.-T. & Cranston, P. (2004). "Which
side of the tree is more basal?".
Systematic Entomology. 29 (3): 279–
281. doi:10.1111/j.0307-
6970.2004.00262.x .
S2CID 82371239 .
1 . Smith, Stacey (19 September 2016).
"For the love of trees: The ancestors
are not among us" . For the love of
trees. Retrieved 23 March 2019.
17. "Choosing the Book title 'Clade' " .
Penguin Group Australia. 2015.
Retrieved 20 January 2015.

External links

Look up clade in Wiktionary, the free

dictionary.

Evolving Thoughts: "Clade"

DM Hillis, D Zwickl & R Gutell. "Tree of
life ". An unrooted cladogram depicting
 around 3000 species.
"Phylogenetic systematics, an
introductory slide-show on evolutionary
trees" —University of California, Berkeley

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