Afrotheria

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Afrotheria
Temporal range: Paleocene - Holocene, 65–0 Ma
Afrotheria.jpg
1. Aardvark 2. Dugong 3. Black and rufous elephant shrew 4. West Indian manatee 5. Golden mole 6. Rock hyrax 7. African bush elephant 8. Tailless tenrec
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Atlantogenata
Superorder: Afrotheria
Stanhope et al., 1998
Orders

See below

Afrotheria is a clade of mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephant shrews (also known as sengis), tenrecs, aardvarks, hyraxes, elephants, sea cows, and several extinct clades. They share few anatomical features but many are partly or entirely African in their distribution. This probably reflects the fact that Africa was an island continent through the early Cenozoic. Because the continent was isolated by water, Laurasian groups such as insectivores, rabbits, carnivorans and ungulates could not become established. Instead, the niches occupied by those groups were filled by tenrecs, hyraxes and elephants that evolved from the ancestral afrothere. This adaptive radiation may have occurred in response to the Cretaceous–Paleogene mass extinction.

The common ancestry of these animals was not recognized until the late 1990s.[1] Historically, the Paenungulata had been linked to other ungulates; the golden mole, tenrecs, and elephant shrews with the traditional (and polyphyletic) Insectivora; and the aardvarks with the pangolins and the xenarthrans within the invalid taxon Edentata. Continuing work on the molecular[2][3] and morphological[4][5][6][7] diversity of afrotherian mammals has provided ever increasing support for their common ancestry.

Evolutionary relationships[edit]

The afrotherian clade was originally proposed in 1998[1] based on analyses of DNA sequence data. However, previous studies had hinted at the close interrelationships among subsets of endemic African mammals, some of these studies date to the 1920s;[8] there were also sporadic papers in the 1980s[9] and 1990s.[10][11] The core of the Afrotheria consists of the Paenungulata, i.e., elephants, sea cows, and hyraxes, a group with a long history among comparative anatomists.[12][13] Hence, while DNA sequence data have proven essential to infer the existence of the Afrotheria as a whole, and while the Afroinsectiphilia (insectivoran-grade afrotheres including tenrecs, golden moles, sengis, and aardvarks) were not recognized as part of Afrotheria without DNA data, some precedent is found in the comparative anatomical literature for the idea that at least part of this group forms a clade. The Paleocene genus Ocepeia, which is the most completely-known Paleocene African mammal and the oldest afrotherian known from a complete skull, shares similarities with both Paenungulata and Afroinsectiphilia, and may help to characterize the ancestral body type of afrotherians.[14]

Since the 1990s, increasing molecular and anatomical data have been applied to the classification of animals; both types of data support the idea that afrotherian mammals are descended from a single common ancestor to the exclusion of other mammals. On the anatomical side, features shared by most, if not all, afrotheres include high vertebral counts,[7] aspects of placental membrane formation,[15] the shape of the ankle bones,[5][6] and the relatively late eruption of the permanent dentition.[16] The snout is unusually long and mobile in several Afrotherian species.[17] Studies of genomic data, including millions of aligned nucleotides sampled for a growing number of placental mammals, also support Afrotheria as a clade.[18][19] Additionally, there might be some dental synapomorphies uniting afroinsectiphilians, if not afrotheres as a whole: p4 talonid and trigonid of similar breadth, a prominent p4 hypoconid, presence of a P4 metacone and absence of parastyles on M1–2.[6][20]

Afrotheria is now recognized as one of the three major groups within the Eutheria (containing placental mammals).[21] Relations within the three cohorts, Afrotheria, Xenarthra, Boreoeutheria, and the identity of the placental root, remain somewhat controversial.[4]

Afrotheria as a clade has usually been discussed without a Linnaean rank, but has been assigned the rank of cohort, magnorder, and superorder. One reconstruction, which applies the molecular clock, proposes that the oldest split occurred between Afrotheria and the other three some 105 million years ago in the mid-Cretaceous, when the African continent was separated from other major land masses.[22] This idea is consistent with the fossil record of Xenarthra, which is restricted to South America (following recent consensus that Eurotamandua is not a xenarthran[23]).

However, Afrotheria itself does not have a fossil record restricted to Africa,[24] and appear in fact to be have evolved in the continent's isolation.[25] More recent, genomic-scale phylogenies favor the hypothesis that Afrotheria and Xenarthra comprise sister taxa at the base of the placental mammal radiation, suggesting an ancient gondwannan clade of placental mammals[26]

Relations between the various afrotherian orders are still being studied. On the basis of molecular studies, elephants and manatees appear to be related, and likewise elephant shrews and aardvarks.[27] These findings are compatible with the work of earlier anatomists.[12][13]

Phylogeny[edit]

Afrotheria
Afroinsectiphilia
Tubulidentata

OrycteropodidaeAardvark2 (PSF) colourised.png


Afroinsectivora
Macroscelidea

MacroscelididaeRhynchocyon chrysopygus-J Smit white background.jpg


Afrosoricida

ChrysochloridaeThe animal kingdom, arranged according to its organization, serving as a foundation for the natural history of animals (Pl. 18) (Chrysochloris asiatica).jpg



TenrecidaeBrehms Thierleben - Allgemeine Kunde des Thierreichs (1876) (Tenrec ecaudatus).jpg





Paenungulata
Hyracoidea

ProcaviidaeDendrohyraxEminiSmit white background.jpg


Tethytheria
Proboscidea

ElephantidaeElephant white background.png


Sirenia

DugongidaeDugong dugon Hardwicke white background.jpg



TrichechidaeManatee white background.jpg






Current status and distribution[edit]

Many extant members of Afrotheria appear to have a high risk of extinction (perhaps related to the large size of many). Species loss within this already small group would comprise a particularly great loss of genetic and evolutionary diversity. The IUCN Afrotheria Specialist Group notes that Afrotheria, as currently reconstructed, includes nearly a third of all mammalian orders currently found in Africa and Madagascar, but only 75 of more than 1,200 mammalian species in those areas.[28]

While most extant species assigned to Afrotheria live in Africa, some (such as the Indian elephant and three of the four sirenian species) occur elsewhere; many of these are also endangered. Prior to the Quaternary extinction event, proboscideans were present on every continent of the world except Australia and Antarctica. Hyraxes lived in much of Eurasia as recently as the end of the Pliocene; the extinct afrotherian orders of embrithopods, and desmostylians were also once widely distributed. (However, the desmostylians have recently been viewed as possible perissodactyls, rather than afrotheres,[29] though this is still controversial.[25])

Classification[edit]

Afrotheria is a clade of placental mammals, the stem designation for which is Eutheria. Based on precedent, some clades are junior synonyms and arguably should be replaced.[30][31]

See also[edit]

Notes[edit]

  1. ^ a b Stanhope, M. J.; Waddell, V. G.; Madsen, O.; de Jong, W.; Hedges, S. B.; Cleven, G. C.; Kao, D.; Springer, M. S. (1998). "Molecular evidence for multiple origins of Insectivora and for a new order of endemic African insectivore mammals". Proceedings of the National Academy of Sciences. 95 (17): 9967–9972. Bibcode:1998PNAS...95.9967S. doi:10.1073/pnas.95.17.9967. PMC 21445Freely accessible. PMID 9707584. 
  2. ^ Springer, Mark S.; Michael J. Stanhope; Ole Madsen; Wilfried W. de Jong (2004). "Molecules consolidate the placental mammal tree" (PDF). Trends in Ecology & Evolution. 19 (8): 430–438. doi:10.1016/j.tree.2004.05.006. PMID 16701301. 
  3. ^ Robinson, T. J.; Fu, B.; Ferguson-Smith, M. A.; Yang, F. (2004). "Cross-species chromosome painting in the golden mole and elephant-shrew: support for the mammalian clades Afrotheria and Afroinsectiphillia but not Afroinsectivora". Proceedings of the Royal Society B: Biological Sciences. 271 (1547): 1477–1484. doi:10.1098/rspb.2004.2754. 
  4. ^ a b Asher RJ, Bennett N, Lehmann T (2009). "The new framework for understanding placental mammal evolution". BioEssays. 31 (8): 853–864. doi:10.1002/bies.200900053. PMID 19582725. 
  5. ^ a b Tabuce, R.; Marivaux, L.; Adaci, M.; Bensalah, M.; Hartenberger, J.-L.; Mahboubi, M.; Mebrouk, F.; Tafforeau, P.; Jaeger, J.-J. (2007). "Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade". Proceedings of the Royal Society B: Biological Sciences. 274 (1614): 1159–1166. doi:10.1098/rspb.2006.0229. 
  6. ^ a b c d Seiffert, Erik R (2007). "A new estimate of afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence". BMC Evolutionary Biology. 7 (1): 224. doi:10.1186/1471-2148-7-224. PMC 2248600Freely accessible. PMID 17999766. 
  7. ^ a b Sánchez‐Villagra, Marcelo R.; Narita, Yuichi; Kuratani, Shigeru (2007). "Thoracolumbar vertebral number: The first skeletal synapomorphy for afrotherian mammals". Systematics and Biodiversity. 5 (1): 1–7. doi:10.1017/S1477200006002258. 
  8. ^ Le Gros Clark, W.E. & C.F. Sonntag (1926). "A monograph of Orycteropus afer III, the skull, the skeleton of the trunk, and limbs". Proceedings of the Zoological Society London. 30: 445–485. 
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  11. ^ Springer, M. S.; Cleven, Gregory C.; Madsen, Ole; De Jong, Wilfried W.; Waddell, Victor G.; Amrine, Heather M.; Stanhope, Michael J. (1997). "Endemic African mammals shake the phylogenetic tree". Nature. 388 (6637): 61–64. doi:10.1038/40386. PMID 9214502. 
  12. ^ a b Simpson, G. G. (1945). "The principles of classification and a classification of mammals". Bulletin of the American Museum of Natural History. 85: 1–350. 
  13. ^ a b Tabuce, Rodolphe; Asher, Robert J.; Lehmann, Thomas (2008). "Afrotherian mammals: a review of current data" (PDF). Mammalia. 72 (1): 2–14. doi:10.1515/MAMM.2008.004. 
  14. ^ a b Gheerbrant, Emmanuel; Amaghzaz, Mbarek; Bouya, Baadi; Goussard, Florent; Letenneur, Charlène (2014). "Ocepeia (Middle Paleocene of Morocco): The Oldest Skull of an Afrotherian Mammal". PLoS ONE. 9 (2): e89739. Bibcode:2014PLoSO...989739G. doi:10.1371/journal.pone.0089739. PMC 3935939Freely accessible. PMID 24587000. 
  15. ^ Mess, Andrea; Carter, Anthony M. (2006). "Evolutionary transformations of fetal membrane characters in Eutheria with special reference to Afrotheria". Journal of Experimental Zoology Part B: Molecular and Developmental Evolution. 306B (2): 140–163. doi:10.1002/jez.b.21079. 
  16. ^ Asher, Robert J; Lehmann, Thomas (2008). "Dental eruption in afrotherian mammals". BMC Biology. 6 (1): 14. doi:10.1186/1741-7007-6-14. PMC 2292681Freely accessible. PMID 18366669. 
  17. ^ Christine M. Dengler-Crish; et al. (2006). "Organization of the somatosensory cortex in elephant shrews (E. edwardii)". Anat. Rec. 288A (8): 859–866. doi:10.1002/ar.a.20357. 
  18. ^ Murphy, W. J.; Pringle, T. H.; Crider, T. A.; Springer, M. S.; Miller, W. (2007). "Using genomic data to unravel the root of the placental mammal phylogeny". Genome Research. 17 (4): 413–421. doi:10.1101/gr.5918807. PMC 1832088Freely accessible. PMID 17322288. 
  19. ^ Nikolaev, Sergey; Montoya-Burgos, Juan I.; Margulies, Elliott H.; NISC Comparative Sequencing Program; Rougemont, Jacques; Nyffeler, Bruno; Antonarakis, Stylianos E. (2007). "Early History of Mammals Is Elucidated with the ENCODE Multiple Species Sequencing Data". PLoS Genetics. 3 (1): e2. doi:10.1371/journal.pgen.0030002. PMC 1761045Freely accessible. PMID 17206863. 
  20. ^ Cote S, Werdelin L, Seiffert ER, Barry JC (March 2007). "Additional material of the enigmatic Early Miocene mammal Kelba and its relationship to the order Ptolemaiida". Proc Natl Acad Sci USA. 104 (13): 5510–5. Bibcode:2007PNAS..104.5510C. doi:10.1073/pnas.0700441104. PMC 1838468Freely accessible. PMID 17372202. 
  21. ^ William J. Murphy; Eduardo Eizirik; Mark S. Springer; et al. (14 December 2001). "Resolution of the Early Placental Mammal Radiation Using Bayesian Phylogenetics" (PDF). Science. 294 (5550): 2348–2351. Bibcode:2001Sci...294.2348M. doi:10.1126/science.1067179. PMID 11743200. 
  22. ^ Springer, M. S.; Murphy, W. J.; Eizirik, E.; O'Brien, S. J. (2003). "Placental mammal diversification and the Cretaceous-Tertiary boundary". Proceedings of the National Academy of Sciences. 100 (3): 1056–1061. Bibcode:2003PNAS..100.1056S. doi:10.1073/pnas.0334222100. PMC 298725Freely accessible. PMID 12552136. 
  23. ^ Rose KD, Emry RJ, Gaudin TJ, Storch G (2005). "Xenarthra and Pholidota.". In Rose KD, Archibald JD. The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades. Baltimore, MD: Johns Hopkins University Press. 
  24. ^ Zack S.P.; Penkrot T.A.; Bloch J.I.; Rose K.D. (2005). "Affinities of 'hyopsodontids' to elephant shrews and a Holarctic origin of Afrotheria". Nature. 434 (7032): 497–501. Bibcode:2005Natur.434..497Z. doi:10.1038/nature03351. PMID 15791254. 
  25. ^ a b Gheerbrant, Emmanuel; Filippo, Andrea; Schmitt, Arnaud (2016). "Convergence of Afrotherian and Laurasiatherian Ungulate-Like Mammals: First Morphological Evidence from the Paleocene of Morocco". PLOS ONE. 11 (7): e0157556. doi:10.1371/journal.pone.0157556. 
  26. ^ Prasad, A. B.; Allard, M. W.; Green, E. D. (2008). "Confirming the Phylogeny of Mammals by Use of Large Comparative Sequence Data Sets". Molecular Biology and Evolution. 25 (9): 1795–1808. doi:10.1093/molbev/msn104. PMC 2515873Freely accessible. PMID 18453548. 
  27. ^ Svartman, M.; Stanyon, R. (2012). "The Chromosomes of Afrotheria and Their Bearing on Mammalian Genome Evolution" (PDF). Cytogenetic and Genome Research. 137 (2–4): 144–153. doi:10.1159/000341387. PMID 22868637. 
  28. ^ "What is Afrotheria?". IUCN Afrotheria Specialist Group. Retrieved 10 December 2013. 
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  30. ^ "Afrosoricida". Mammal Species of the World, 3rd edition. Retrieved 17 March 2014. 
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  32. ^ Rodolphe Tabuce et al. (2007) Early Tertiary mammals from North Africa reinforce the molecular Afrotheria clade. Proc. R. Soc. B 2007 274, doi:10.1098/rspb.2006.0229
  33. ^ A Anthony Ravel; Maeva Orliac (2014). "The inner ear morphology of the 'condylarthran' Hyopsodus lepidus". Historical Biology. 27 (8): 957–969. doi:10.1080/08912963.2014.915823. 
  34. ^ Horovitz, Inés; Gerhard Storch & Thomas Martin (2005). "Ankle structure in Eocene pholidotan mammal Eomanis krebsi and its taxonomic implications" (PDF). Acta Palaeontol. Pol. 50 (3): 545–548. 

References[edit]

External links[edit]