A clade (from Ancient Greek κλάδος, klados, "branch") or monophylum (see monophyletic) is a group consisting of an ancestor and all its descendants, a single "branch" on the "tree of life". The ancestor may be an individual, a breeding pair, a population or even a species (extinct or extant). Many familiar groups, the rodents and the insects for example, are clades; others, like the lizards and the monkeys, are not.
Increasingly, taxonomists try to name preferably or only taxa that are clades.
Clade and ancestor
A clade is by definition monophyletic, meaning it contains one ancestor (which can be an organism, a population, or a species) and all its descendants.[note 1] 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 being derived from "clade" by Ernst Mayr (1965). The results of phylogenetic/cladistic analyses are tree-shaped diagrams called cladograms; they, and all their branches, are phylogenetic hypotheses.
Three methods of defining clades are featured in phylogenetic nomenclature: node-, stem-, and apomorphy-based:
- In node-based definition, clade name A refers to the least inclusive clade containing taxa (or specimens) X, Y, etc. A is the clade consisting of the last common ancestor of these taxa together with all the descendents of that ancestor. The ancestor is the branch point, or node.
- In stem-based definition, A refers to the most inclusive clade containing X, Y, etc., and their common ancestor, down to where Z branches off below A. Taxa are included between the node of A and down to (but not including) the branching point to Z; that is, the stem of A.
- In apomorphy-based definition, A refers to the clade identified by an apomorphy (a trait) found in X, Y, etc., and their common ancestor.
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, 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.
- A clade A is basal to a clade B if A branches off the lineage leading to B before B 's first member. In the diagram to the right, the strepsirrhine clade, including the lemurs and lorises, is basal to the hominoids, the apes and humans. Some authors have used "basal" differently to mean a clade that is "more primitive" or less species-rich than its sister clade; others consider this usage to be incorrect.
Nomenclature and taxonomy
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 publication of Darwin's theory of evolution in 1859, the concept of a "natural system" of taxonomy gained a theoretical basis, and the idea was born that groups used in a system of classification should represent branches on the evolutionary tree of life. In the century and a half since then, taxonomists have increasingly worked to make the taxonomic system reflect evolution. When it comes to naming, however, this principle is not always compatible with the traditional rank-based nomenclature. In the latter, only taxa associated with a rank can be named, yet there are not enough ranks to name a long series of nested clades; also, taxon names cannot be defined in a way that guarantees them to refer to clades. For these and other reasons, phylogenetic nomenclature has been developed; it is still controversial.
- Binomial nomenclature
- Biological classification
- Crown group
- Phylogenetic nomenclature
- 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
- Dupuis, Claude (1984). "Willi Hennig's impact on taxonomic thought". Annual Review of Ecology and Systematics 15: 1–24. ISSN 0066-4162.
- "International Code of Phylogenetic Nomenclature. Version 4c. Chapter I. Taxa.". 2010. Retrieved 22 September 2012.
- 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.
- 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.
- 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
- 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