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Cladus: Eukaryota
Supergroup: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Classis: Mammalia
Subclassis: Theria
Infraclassis: Placentalia
Superordo: †Meridiungulata
Ordo: †Notoungulata
Subordo: †Typotheria
Familis: Mesotheriidae


Mesotheriidae ("Middle Beasts") is an extinct family of notoungulate mammals known from the Eocene through the Pleistocene of South America. Mesotheriids were small to medium-sized herbivorous mammals adapted for digging.


Mesotheriids were small to medium sized notoungulates; larger forms were approximately the size of a sheep (Shockey et al., 2007). Additionally, the family is characterized by specializations of the teeth and skeleton. In the dentition, all mesotheriids have ever-growing incisors with enamel restricted to the anterior surface, a condition termed gliriform, as it also occurs in Glires (rodents and lagomorphs). The cheek teeth (premolars and molars) of mesotheriids are high-crowned (hypsodont) and in advanced members of the family, the cheek teeth are also ever-growing (Shockey et al., 2007). Mesotheriid skeletons are heavily built and show features associated with digging in living mammals. In particular, fossorial characteristics of mesotheriids include deeply fissured claws, presence of a sesamoid bone in the elbow and reinforcement of the pelvic girdle by addition of vertebrae to the sacrum and fusion of the sacrum and innominate (Shockey et al., 2007).


A biomechanical study of the skeleton of three mesotheriid genera (Trachytherus, Plesiotypotherium, and Mesotherium) spanning the temporal range of the family indicates that most or all mesotheriids were adapted for digging (Shockey et al., 2007). Shockey et al. concluded that mesotheriids likely dug for roots and tubers and were most similar in their diet and behavior to living wombats, although no living group is perfectly analogous. Extensive burrowing was considered possible but unlikely given the relatively large size of most mesotheriids.
[edit] Geographic and Temporal Distribution

As with almost all other notoungulates, mesotheriids are known only from the Cenozoic of South America (McKenna and Bell, 1997). Unlike some other families, mesotheriid fossils are not found across the continent. Instead, mesotheriids are most abundant and diverse in faunas from middle latitudes in Bolivia and Chile, particularly the Altiplano (Flynn et al., 2005). Mesotheriids fossils are rare in high latitude Patagonian faunas and absent entirely from tropical faunas in northern South America.

The earliest potential record of a mesotheriid is ?Trachytherus mendocensis from the late Eocene[3] or early Oligocene[4] Divisadero Largo Formation of Mendoza Province, Argentina (Shockey et al., 2007), but Cerdeño et al. (2006) have suggested that this specimen may actually derive from early Miocene rocks that overlie the Divisadero Largo Formation. The earliest secure records of the family come from the late Oligocene, when the family is represented by the genera Anatrachytherus and Trachytherus from Argentina and Bolivia (Reguero and Castro, 2004). The family reached its greatest diversity in the Miocene (Flynn et al., 2005), and mesotheriids persisted into the middle Pleistocene, in the form of the type genus, Mesotherium (McKenna and Bell, 1997). Mesotheriidae was one of only three notoungulate families to persist into the Quaternary, the others being Hegetotheriidae and Toxodontidae.


Within the order Notoungulata, Mesotheriidae is placed in the suborder Typotheria (Cifelli, 1993). In fact, Typotheria is named for the genus Typotherium, a synonym of Mesotherium (Shockey et al., 2007). In addition to Mesotheriidae, Typotheria traditionally includes other small bodied notoungulates in the families Oldfieldthomasiidae, Interatheriidae, and Archaeopithecidae (Simpson, 1967; McKenna and Bell, 1997). Recent opinion, however, favors inclusion of two additional families in Typotheria, Archaeohyracidae and Hegetotheriidae (Croft and Anaya, 2006). These families have traditionally been placed in a separate suborder, Hegetotheria, but phylogenetic studies indicate that their exclusion would render Typotheria paraphyletic (Cifelli, 1993; Billet et al., 2007). Within Typotheria, both Cifelli and Billet et al. indicate that mesotheriids are more closely related to archaeohyracids and hegetotheriids than to the remaining typotherian families. In fact, Billet et al.'s analysis indicates that both Mesotheriidae and Hegetotheriidae originated from within Archaeohyracidae.

McKenna and Bell (1997) recognized three subfamilies within Mesotheriidae: Fiandraiinae, Mesotheriinae, and Trachytheriinae. However, Flynn et al. (2005) have suggested that Fiandraia, the only known fiandraiine, is not a mesotheriid and may represent a toxodontid instead. Of the remaining subfamilies, Trachytheriinae includes earlier (Eocene and Oligocene) forms and may be paraphyletic with respect to Mesotheriinae, which includes more derived genera from the Miocene and later (Reguero and Castro, 2004).
Classification of Mesotheriidae:[5]

* Family †Mesotheriidae

o Subfamily †Fiandraiinae[6]
+ †Fiandraia (Miocene)
# †F. romeroi
o Subfamily †Trachytheriinae (paraphyletic)
+ †Trachytherus (?l. Eocene-l. Oligocene[7])
# †T. alloxus
# †"T. mendocensis"
# †T. spegazzinianus
# †T. subandinus
o Subfamily †Mesotheriinae
+ †Eotypotherium (e. Miocene[8])
# †E. chico
+ †Altitypotherium (e. Miocene[8])
# †A. chucalensis
# †A. paucidens
+ †Microtypotherium (m./l. Miocene)
# †M. choquecotense

o Subfamily †Mesotheriinae (continued)
+ †Eutypotherium (m. Miocene)
# †E. lehmannnitschei
# †E. superans
+ †Caraguatypotherium (m.-l. Miocene[9])
# †C. munozi
+ †Plesiotypotherium (l. Miocene)
# †P. achirense
+ †Typotheriopsis (l. Miocene)
# †T. chasicoensis
# †T. silveyrai
+ †Pseudotypotherium (l. Miocene-?m. Pleistocene)
# †P. exiguum
# †P. subinsigne
+ †Mesotherium (e.-m. Pleistocene)
# †M. cristatum
# †M. hystatum
# †M. maendrum
# †M. pachygnathum
+ †Hypsitherium[10] (e. Pliocene)
# †H. bolivianum


1. ^ First appearance from Reguero and Castro (2004) with age of fauna following Shockey and Flynn (2007); last appearance from McKenna & Bell (1997).
2. ^ Compiled from McKenna & Bell (1997), Reguero and Castro (2004), and Shockey et al. (2007).
3. ^ Shockey and Flynn, 2007, p.24
4. ^ Reguero and Castro, 2004
5. ^ Trachytheriinae following Billet et al. (2008); Mesotheriinae following Anaya and MacFadden (1995) and Shockey et al. (2007). Temporal ranges following McKenna and Bell (1997) except where noted.
6. ^ Inclusion tentative following Flynn et al. (2005).
7. ^ FAD following Cerdeño et al. (2006) and Shockey and Flynn (2007).
8. ^ a b Following Croft et al. (2004).
9. ^ Following Flynn et al. (2005).
10. ^ Recognized as valid by McKenna and Bell (1997), this genus has not been evaluated in recent studies.


* Billet, G.A., Muizon, C. de, and Quispe, B.M. 2008. Late Oligocene mesotheriids (Mammalia, Notoungulata) from Salla and Lacayani (Bolivia): implications for basal mesotheriid phylogeny and distribution. Zoological Journal of the Linnean Society 152:153-200.
* Billet, G.A., Patterson, B., and Muizon, C. de. 2007. The latest archaeohyracids representatives (Mammalia, Notoungulata) from the Deseadan of Bolivia and Argentina; pp. 39–43 in E. Díaz-Martínez and I. Rábano (eds.), 4th European Meeting on the Palaeontology and Stratigraphy of Latin America. Instituto Geológico y Minero de España, Madrid. ISBN 978-84-7840-707-1 [1]
* Cerdeño, E., González Riga, B., and Bordonaro, O. 2006. Primer hallazgo de mamíferos en la Formación Mariño (Mioceno) en Divisadero Largo (Mendoza, Argentina). Ameghiniana 43:205-214. [2]
* Cifelli, R. L. 1993. The phylogeny of the native South American ungulates. pp. 195–216 in F. S. Szalay, M. J. Novacek and M. C. McKenna (eds.) Mammal Phylogeny, Volume 2, Placentals. Springer-Verlag, New York. ISBN 0-387-97853-4
* Croft, D.A., and Anaya, F. 2006. A new middle Miocene hegetotheriid (Notoungulata: Typotheria) and a phylogeny of Hegetotheriidae. Journal of Vertebrate Paleontology 26:387-399.
* Croft, D.A., Flynn, J.J. and Wyss, A.R. 2004. Notoungulata and Litopterna of the Early Miocene Chucal Fauna, Northern Chile. Fieldiana Geology 50(1):1-52. [3]
* Flynn, J. J., Croft, D.A., Charrier, R., Wyss, A.R., Hérail, G., and García, M. 2005. New Mesotheriidae (Mammalia, Notoungulata, Typotheria), geochronology and tectonics of the Caragua area, northernmost Chile. Journal of South American Earth Sciences 19:55-74.
* McKenna, Malcolm C., and Bell, Susan K. 1997. Classification of Mammals Above the Species Level. Columbia University Press, New York, 631 pp. ISBN 0-231-11013-8
* Reguero, M.A., and Castro, P.V. 2004. Un nuevo Trachytheriinae (Mammalia, †Notoungulata) del Deseadense (Oligoceno tardío) de Patagonia, Argentina: implicancias en la filogenia, biogeografía y bioestratigrafía de los Mesotheriidae. Revista Geológica de Chile 31:45–64. [4]
* Shockey, B.J., Croft, D.A., and Anaya, F. 2007. Analysis of function in the absence of extant functional homologues: a case study using mesotheriid notoungulates (Mammalia). Paleobiology 33:227-247.
* Shockey, B.J., and Flynn, J.J. 2007. Morphological diversity in the postcranial skeleton of Casamayoran (?middle to late Eocene) Notoungulata and foot posture in notoungulates. American Museum Novitates 3601:1-26. [5]
* Simpson, G.G. 1967. The beginning of the age of mammals in South America. Part 2, Systematics : Notoungulata, concluded (Typotheria, Hegetotheria, Toxodonta, Notoungulata incertae sedis), Astrapotheria, Trigonostylopoidea, Pyrotheria, Xenungulata, Mammalia incertae sedis. Bulletin of the American Museum of Natural History 137:1-259. [6]

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