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Crown group

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Crown group

In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. It is thus a way of defining a clade, a group consisting of a species and all its extant or extinct descendants. For example, Neornithes (birds) can be defined as a crown group, which includes the most recent common ancestor of all modern birds, and all of its extant or extinct descendants.

The concept was developed by Willi Hennig, the formulator of phylogenetic systematics, as a way of classifying living organisms relative to their extinct relatives in his "Die Stammesgeschichte der Insekten", and the "crown" and "stem" group terminology was coined by R. P. S. Jefferies in 1979. Though formulated in the 1970s, the term was not commonly used until its reintroduction in 2000 by Graham Budd and Sören Jensen.

It is not necessary for a species to have living descendants in order for it to be included in the crown group. Extinct side branches on the family tree that are descended from the most recent common ancestor of living members will still be part of a crown group. For example, if we consider the crown-birds (i.e. all extant birds and the rest of the family tree back to their most recent common ancestor), extinct side branches like the dodo or great auk are still descended from the most recent common ancestor of all living birds, so fall within the bird crown group. One very simplified cladogram for birds is shown below:

In this diagram, the clade labelled "Neornithes" is the crown group of birds: it includes the most recent common ancestor of all living birds and its descendants, living or not. Although considered to be birds (i.e. members of the clade Aves), Archaeopteryx and other extinct groups are not included in the crown group, as they fall outside the Neornithes clade, being descended from an earlier ancestor.

An alternative definition does not require any members of a crown group to be extant, only to have resulted from a "major cladogenesis event". The first definition forms the basis of this article.

Often, the crown group is given the designation "crown-", to separate it from the group as commonly defined. Both birds and mammals are traditionally defined by their traits, and contain fossil members that lived before the last common ancestors of the living groups or, like the mammal Haldanodon, were not descended from that ancestor although they lived later. Crown-Aves and Crown-Mammalia therefore differ slightly in content from the common definition of Aves and Mammalia. This has caused some confusion in the literature.

The cladistic idea of strictly using the topology of the phylogenetic tree to define groups necessitates other definitions than crown groups to adequately define commonly discussed fossil groups. Thus, a host of prefixes have been defined to describe various branches of the phylogenetic tree relative to extant organisms.

A pan-group or total group is the crown group and all organisms more closely related to it than to any other extant organisms. In a tree analogy, it is the crown group and all branches back to (but not including) the split with the closest branch to have living members. The Pan-Aves thus contain the living birds and all (fossil) organisms more closely related to birds than to crocodilians (their closest living relatives). The phylogenetic lineage leading back from Neornithes to the point where it merges with the crocodilian lineage, along with all side branches, constitutes pan-birds. In addition to non-crown group primitive birds like Archaeopteryx, Hesperornis and Confuciusornis, therefore, pan-group birds would include all dinosaurs and pterosaurs as well as an assortment of non-crocodilian animals like Marasuchus.

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