Basal (phylogenetics)

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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

core eudicots
. No extant taxon is closer to the root than any other.

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:

Root 

 Basal clade #1

 Non-basal clade #1

 Non-basal clade #2  

 Non-basal clade #3

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

plesiomorphic" are preferred to "basal" or "primitive", the latter of which may carry false connotations of inferiority or a lack of complexity.[1] The terms ''deep-branching'' or ''early-branching'' are similar in meaning, and equally may misrepresent extant taxa that lie on branches connecting directly to the root node as having more ancestral character states.[1][2]

Despite the ubiquity of the usage of basal,

oviparous reproduction and nipple-less lactation of monotremes, a clade of mammals[11] with just five species, and the archaic anatomy of the tuatara,[12] a basal clade of lepidosaurian
with a single species.

Examples

Flowering plants

Amborella trichopoda
, the most basal extant angiosperm

The

fossil record could potentially also be helpful in this respect, but is absent in this case).[14] The cladogram below is based on Ramírez-Barahona et al. (2020),[15]
with species counts taken from the source indicated.

 
Angiosperms
 

Amborellales (1 species)[16]

Nymphaeales (about 90 species)[17]

Austrobaileyales (about 95 species)[17]

Magnoliids (about 9,000 species)[16]

Chloranthales (about 80 species)[17]

Monocots (about 70,000 species)[16]

Ceratophyllales (about 6 species)[16]

Eudicots (about 175,000 species)[16]

 '
Basal angiosperms
'

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.

Dromiciops within otherwise Australasian Australidelphia.[18]
Myzopodidae
. Locations with only fossil members are indicated by red stars.
 Homininae 

Humans (Homo sapiens)

Bonobos (Pan paniscus)

Chimpanzees (Pan troglodytes)

Eastern gorillas (Gorilla beringei)

Western gorillas (Gorilla gorilla)

Moreover, orangutans are a sister group to Homininae and are the basal genus in the great ape family Hominidae as a whole.

 Hominidae 
 Ponginae 

Orangutans (Pongo spp.)

 Homininae 

Humans (Homo sapiens)

Chimpanzees (Pan spp.)

Gorillas (Gorilla spp.)

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.

arboreal lifestyle, a trait generally viewed as ancestral among the apes.[20][21]

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:

Notes

  1. ^ Meaning the lowest taxonomic ranks of the respective clades; the highest ranks should be the same (assuming they are ranked).
  2. Amborellales
    ). Meanwhile, the other (unranked) sister basal clade has about 250,000 species.
  3. ^ For example, C might be a genus and the other basal clade(s) might have the higher ranks of subfamily or family.
  4. ^ 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.
  5. ^ 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.
  6. Ursidae,[9] but its specializations for a bamboo diet are not ancestral ursid characters.[10]
  7. ^ 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.
  8. 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 allele
    to a branch ancestor with two descendant variants.
  9. ^ 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.
  10. ^ 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.
  11. 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]
  12. ^ 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

  1. ^ 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.
  2. ^
    S2CID 82371239
    .
  3. ^ .
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  7. .
  8. ^ .
  9. .
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  14. ^ a b Essig, F. B. (2014-07-01). "What's so primitive about Amborella?". Botany Professor. Retrieved 2014-10-04.
  15. S2CID 220375828
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  16. ^ .
  17. ^ .
  18. ^ .
  19. ..
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  22. ^ Kukso, F. (2016-11-08). "Seafaring Spiders Made It around the World—in 8 Million Years". Scientific American. Retrieved 2016-11-10.
  23. ^
    PMID 27732621
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  29. .
  30. ^ 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.
  31. PMID 21723399
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  33. .
  34. ^ .
  35. ^ 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.
  36. S2CID 25912333
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External links