Basal (phylogenetics)
In phylogenetics, basal is the direction of the base (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to the root. Note that extant taxa that lie on branches connecting directly to the root are not more closely related to the root than any other extant taxa.[1][2][3]
While there must always be two or more equally "basal" clades sprouting from the root of every cladogram, those clades may differ widely in taxonomic rank,[n 1] species diversity, or both.[n 2] If C is a basal clade within D that has the lowest rank of all basal clades within D,[n 3] C may be described as the basal taxon of that rank within D.[n 4] The concept of a 'key innovation' implies some degree of correlation between evolutionary innovation and diversification.[4][5][6][n 5] However, such a correlation does not make a given case predicable, so ancestral characters should not be imputed to the members of a less species-rich basal clade without additional evidence.[1][2][7][8][n 6]
In general, clade A is more basal than clade B if B is a subgroup of the
Usage
A basal group in the stricter sense forms a sister group to the rest of the larger clade,[citation needed] as in the following case:
|
Basal clade #2 |
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.[citation needed]
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
Despite the ubiquity of the usage of basal,
Examples
Flowering plants
The
|
' Basal angiosperms ' |
Great apes
This section includes a list of general references, but it lacks sufficient corresponding inline citations. (November 2021) |
Within the
Homininae |
| ||||||||||||||||||||||||
Moreover, orangutans are a sister group to Homininae and are the basal genus in the great ape family Hominidae as a whole.
Hominidae |
| ||||||||||||||||||
Subfamilies Homininae and Ponginae are both basal within Hominidae, but given that there are no nonbasal subfamilies in the cladogram it is unlikely the term would be applied to either. In general, a statement to the effect that one group (e.g., orangutans) is basal, or branches off first, within another group (e.g., Hominidae) may not make sense unless the appropriate taxonomic level(s) (genus, in this case) is specified. If that level cannot be specified (i.e., if the clade in question is unranked) a more detailed description of the relevant sister groups may be needed. As can be seen, the term is not reflective of ancestral states or proximity to the common ancestor of extant species.
In this example, orangutans differ from the other genera in their Asian range. This fact plus their basal status provides a hint that the most recent common ancestor of extant great apes may have been Eurasian (see below), a suggestion that is consistent with other evidence.
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.[22][23] 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.[23]
- rafting event.[26][27] 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.[28]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.[29]
- Extant australidelphian marsupials constitute about 240 species in Australasia and one species (the
- While the bat superfamily two in Madagascar, the basal position of the Malagasy family[36] 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.[37]
- The genus endemicto North America).
Notes
- ^ Meaning the lowest taxonomic ranks of the respective clades; the highest ranks should be the same (assuming they are ranked).
- Amborellales). Meanwhile, the other (unranked) sister basal clade has about 250,000 species.
- ^ For example, C might be a genus and the other basal clade(s) might have the higher ranks of subfamily or family.
- ^ In the great apes example, Gorilla is the basal genus of subfamily Homininae, while Pongo is the basal genus of family Hominidae. The two basal clades of the latter both have the highest rank of subfamily, i.e. Homininae and Ponginae.
- ^ Greater diversification of a clade may also be associated with colonization of a new land mass, especially if larger or less competitive than the ancestral land mass; see the coral snake, marsupial and noctilionoid bat examples.
- ^ That is, in the diagram below, both basal clades #1 and #2 are more basal than non-basal clade #1, which in turn is more basal than non-basal clades #2 and #3.
- origin of life. Thus, from a phylogenetic standpoint, the notion of a lineage being basal is nonsensical. However, in genetics, basal lineage refers to a lineage connecting a common ancestor with a single variant alleleto a branch ancestor with two descendant variants.
- ^ New Caledonia is viewed as a refugium; i.e., in this case the geographic location of the basal clade is not thought to provide evidence for the locale in which angiosperms originated.
- ^ These conclusions have been supported by the finding of Eocene fossil remains of the microbiotherian Woodburnodon casei in Antarctica,[30] which is presumed to have served as a way station on the migration route to Australia before the final breakup of Gondwana.
- Valdivian forest as the monito del monte), is basal to the Australasian families Limnodynastidae and Myobatrachidae,[31] with about 120 extant species, suggesting a South American origin for the group.[32] This is consistent with the finding of a fossil from the South American family in Antarctica.[33]
- ^ Ratites may have similarly traveled overland from South America to colonize Australia;[34] a fossil ratite is known from Antarctica,[35] and South American rheas are more basal within the group than Australo-Pacific ratites.[34]
References
- ^ a b c d Smith, Stacey (2016-09-19). "For the love of trees: The ancestors are not among us". For the love of trees. Retrieved 2022-03-07.
- ^ S2CID 82371239.
- ^ S2CID 12285373.
- .
- S2CID 4431205.
- S2CID 58567195.
- OCLC 861200134.
- ^ S2CID 23140594.
- PMID 18662376.
- JSTOR 3872687.
- PMID 16291999.
- .
- S2CID 146118264.
- ^ a b Essig, F. B. (2014-07-01). "What's so primitive about Amborella?". Botany Professor. Retrieved 2014-10-04.
- S2CID 220375828.
- ^ PMID 21652302.
- ^ .
- ^ PMID 20668664.
- PMID 19487676..
- PMID 10968927.
- PMID 16617429.
- ^ Kukso, F. (2016-11-08). "Seafaring Spiders Made It around the World—in 8 Million Years". Scientific American. Retrieved 2016-11-10.
- ^ PMID 27732621.
- PMID 18598742.
- S2CID 56054202.
- JSTOR 1563429.
- PMID 18782726.
- S2CID 243821392.
- JSTOR 3893186.
- ^ Goin, F. J.; Zimicz, N.; Reguero, M. A.; Santillana, S. N.; Marenssi, S. A.; Moly, J. J. (2007). "New marsupial (Mammalia) from the Eocene of Antarctica, and the origins and affinities of the Microbiotheria". Revista de la Asociación Geológica Argentina. 62 (4): 597–603. Retrieved 2016-07-17.
- PMID 21723399.
- PMID 28673970.
- PMID 32327670.
- ^ PMID 27989673.
- ^ Tambussi, C.P.; Noriega, J.I.; Gazdzicki, A.; Tatur, A.; Reguero, M.A.; Vizcaino, S.F. (1994). "Ratite bird from the Paleogene La Meseta Formation, Seymour Island, Antarctica" (PDF). Polish Polar Research. 15 (1–2): 15–20. Archived from the original (PDF) on 2019-12-28. Retrieved 28 December 2019.
- S2CID 25912333.
- PMID 24504061.
- PMID 16341006.
- hdl:2246/1588.
- ISBN 978-0-231-13528-3.