Basal (phylogenetics)

A basal group in the stricter sense forms a sister group to the rest of the larger clade, as in the following case: While it is easy to identify a basal clade in such a cladogram, the appropriateness of such an identification is dependent on the accuracy and completeness of the diagram. It is often assumed in this example that the terminal branches of the cladogram depict all the extant taxa of a given rank within the clade; this is one reason the term basal is highly deceptive, as the lack of additional species in one clade is taken as evidence of morphological affinity with ancestral taxa. Additionally, this qualification does not ensure that the diversity of extinct taxa (which may be poorly known) is represented. In phylogenetics, the term basal cannot be objectively applied to clades of organisms, but tends to be applied selectively and more controversially to groups or lineages thought to possess ancestral characters, or to such presumed ancestral traits themselves. In describing characters, "ancestral" or "plesiomorphic" are preferred to "basal" or "primitive", the latter of which may carry false connotations of inferiority or a lack of complexity. An extant basal group may or may not resemble the last common ancestor of a larger clade to a greater degree than other groups, and is separated from that ancestor by the same amount of time as all other extant groups. However, there are cases where the unusually small size of a sister group does indeed correlate with an unusual number of ancestral traits, as in Amborella (see below). This is likely a source of the mis-use of the term. Other famous examples of this phenomenon are the oviparous reproduction and nipple-less lactation of monotremes, a clade of mammals with just five species, and the archaic anatomy of the tuatara, a basal clade of lepidosaurian with a single species. ==Examples==

Flowering plants '', the most basal extant angiosperm The flowering plant family Amborellaceae, restricted to New Caledonia in the southwestern Pacific, is a basal clade of extant angiosperms, consisting of the most species, genus, family and order within the group that are sister to all other angiosperms (out of a total of about 250,000 angiosperm species). The traits of Amborella trichopoda are regarded as providing significant insight into the evolution of flowering plants; for example, it has "the most primitive wood (consisting only of tracheids), of any living angiosperm" as well as "simple, separate flower parts of indefinite numbers, and unsealed carpels". The cladogram below is based on Ramírez-Barahona et al. (2020), with species counts taken from the source indicated. }}}}}} Great apes Within the great apes, gorillas (eastern and western) are a sister group to chimpanzees, bonobos and humans. These five species form a clade, the subfamily Homininae (African apes), of which Gorilla has been termed the basal genus. However, if the analysis is not restricted to genera, the Homo plus Pan clade is also basal. of marsupials derived from retroposon data shows the basal position of South American Didelphimorphia within Marsupialia, and the basal position of South American Dromiciops within otherwise Australasian Australidelphia. (Of course, lesser apes are entirely Asiatic.) However, orangutans also differ from African apes in their more highly arboreal lifestyle, a trait generally viewed as ancestral among the apes. ==Relevance to biogeographic history==

Given that the deepest phylogenetic split in a group is likely to have occurred early in its history, identification of the most basal subclade(s) in a widely dispersed taxon or clade can provide valuable insight into its region of origin; however, the lack of additional species in a clade is not evidence that it carries the ancestral state for most traits. Most deceptively, people often believe that the direction of migration away from the area of origin can also be inferred (as in the Amaurobioides and Noctilionoidea cases below). As with all other traits, the phylogeographic location of one clade that connects to the root does not provide information about the ancestral state. Examples where such unjustified inferences may have been made include: • Spiders of the genus Amaurobioides are present in South Africa, Australia, New Zealand and Chile. The most basal clade is South African; DNA sequence evidence indicates that after their South American ancestors reached South Africa, they dispersed eastward all the way back to South America over an interval of about 8 million years. This suggested that iguanids once had a widespread Gondwanan distribution; after the Malagasy and New World representatives were separated by vicariance, less isolated Old World iguanids became extinct through competition with other lizard groups (e.g., agamids). In contrast, western Pacific iguanids are nested deeply within American iguanids, having apparently colonized their isolated range after an epic 10,000 km rafting event. However, a 2022 study found oplurids to be closely allied with the American iguanians Leiosauridae, having only diverged 60 million years ago following a likely rafting event of their own. Due to this, neither of the Old World "iguanids" are thought to represent basal lineages. • Coral snakes comprise about 16 species in Asia and over 65 species in the Americas. However, none of the American clades are basal, implying that the group's ancestry was in the Old World. • Extant australidelphian marsupials constitute about 240 species in Australasia and one species (the monito del monte) in South America. The fact that the monito del monte occupies a basal position (the most basal species, genus, family and order) in the superorder Australidelphia is an important clue that its origin was in South America. This conclusion is consistent with the fact that the South American order Didelphimorphia is basal within infraclass Marsupialia; i.e., extant marsupials as a whole also appear to have originated in South America. • While the bat superfamily Noctilionoidea has over 200 species in the Neotropics, two in New Zealand, and two in Madagascar, the basal position of the Malagasy family suggests, in combination with the fossil record and the next-most-basal placement of the New Zealand family, that the superfamily originated in Africa and then migrated eastward to South America, proliferating there but surviving in the Old World only in refugia. • The genus Urocyon (gray and island foxes) is basal in the canine subfamily, suggesting a North American origin of the nearly worldwide group. This is consistent with fossil evidence indicating a North American origin for the canid family as a whole (the other two canid subfamilies, the extinct Borophaginae and Hesperocyoninae, the latter being basal in Canidae, were both endemic to North America). == See also ==