Dagbatitherium
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''Dagbatitherium'' is an extinct genus of
proboscidean The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. From ...
s. So far a single molar from the phosphate basins of
Togo Togo (), officially the Togolese Republic (french: République togolaise), is a country in West Africa. It is bordered by Ghana to the west, Benin to the east and Burkina Faso to the north. It extends south to the Gulf of Guinea, where its c ...
in West Africa has been found. The fossil dates to the Middle Eocene, around 47 million years ago. A striking feature of the tooth are the three pairs of cusps oriented transversely to the longitudinal axis of the tooth. This feature is found in more derived proboscideans, which are grouped in the
Elephantiformes Elephantiformes is a suborder within the order Proboscidea that contains the elephants as well as their extinct relatives. Elephant relatives such as the mammoths and the mastodons are included in this designation. References Mammal s ...
. For its age, ''Dagbatitherium'' is the earliest member of Elephantiformes to date. Furthermore, it is characterized by a low tooth crown and a humped occlusal pattern. The genus was described in 2021.


Discovery and naming

The only tooth of ''Dagbatitherium'' found so far is from West Africa and was recovered in the Hahotoé-Kpogamé phosphate complex. These are phosphate-bearing deposits formed in a coastal basin. They extend over a length of 30 to 35 km and a width of 2 to 3 km in a northeast-southwest direction between the towns of Dagbati and Aveta in Togo. First explored by drilling in the 1950s, they have been mined economically in large open-pit operations since the 1960s. This requires the removal of up to 30 m thick overlying sediments. The phosphate deposits were formed in a shallow marine environment. In principle, three units can be distinguished, consisting first of 10 to 15 m thick phosphate-bearing marks (Couche 3 and Couche 2), followed by 1 to 8 m thick phospharenites (Couche 1) and about 1 m thick phosphate-bearing clays (Couche 0). Especially the two lower depositional units are very rich in fossils, dominated by remains of sharks and rays. In the phospharenites, however, several bone deposits are also formed with, among others, fossils of whales and manatees. The tooth of ''Dagbatitherium'' was found in the basal parts of the phospharenites in the mining area of Dagbati. Numerous minerals of terrestrial origin and pollen indicate a once coastal situation. Accompanying foraminifera were also found. Based on these, an age dating to the Middle Eocene about 46.5 to 44 million years ago could be determined for the deposits. The genus name ''Dagbatitherium'' is composed of the name of the locality and the Greek word θηρίον (thērion) for "beast". Together with the genus, the species ''Dagbatitherium tassyi'' was erected. The species epithet honors the researcher Pascal Tassy, who devoted his scientific work to the study of proboscideans.


Description

''Dagbatitherium'' is the smallest and oldest representative of the Elephantiformes to date, a group of proboscideans that led to today's elephants. However, only one lower molar is currently known, possibly representing the first or second of the molar row. Its length is 37.5 mm, its width 29.2 mm. The tooth was characterized by its low (brachyodont) dental crown, which is a difference from the higher ones in other early Elephantiformes such as Phiomia or ''
Palaeomastodon ''Palaeomastodon'' an extinct genus of Proboscidea. ''Palaeomastodon'' fossils have been found in Africa, where they lived some 36-35 million years ago. They are believed to be the ancestors of elephants or mastodons. ''Palaeomastodon'' lived in ...
''. Two distinct pairs of cusps existed on the masticatory surface, oriented transversely to the longitudinal tooth axis. The two lip-sided cusps (protoconid and hypoconid) appeared broader and lower than the tongue-sided ones (metaconid and entoconid). The respective pairs of cusps were divided by a deep pit in the central axis of the tooth. The broad lip-sided cusps caused the tooth crown wall to be more inclined, while the tongue-sided one was steeper. Sharper ridges were sometimes formed on the cusps, causing the bunodont (humped) occlusal surface pattern to tend more toward a bunolophodont (with ridges) one. In addition to the main cusps, individual secondary cusps also occurred. A third pair of cusps joined at the posterior end of the tooth. It thus gave the molar a trilophodont (with three pairs of cusps/groins) appearance. This is a typical characteristic of the Elephantiformes and separates the group from the more primitive proboscideans with bilophodont teeth (with two cusp pairs/groins). In the
Deinotheriidae Deinotheriidae ("terrible beasts") is a family of prehistoric elephant-like proboscideans that lived during the Cenozoic era, first appearing in Africa, then spreading across southern Asia (Indo-Pakistan) and Europe. During that time, they cha ...
, which also possessed a trilophodont anterior molar, the individual cusp pairs were much more clearly fused into ridges. As another distinctive feature, the tooth in ''Dagbatitherium'' ended behind the third cusp pair in a cingulum, a low enamel bulge. In addition, it already exhibited a three-layered enamel pattern, again typical of more derived proboscideans, but absent in some earlier forms.


Classification

''Dagbatitherium'' is an extinct genus of the order Proboscidea. Its trilophodont-built molar places it in the group Elephantiformes, which unites the more modern evolutionary lineages of the proboscideans. In contrast, the
Plesielephantiformes Plesielephantiformes is an extinct clade of large herbivorous mammals and one of two suborders of the Proboscidea, a group containing elephant Elephants are the largest existing land animals. Three living species are currently recognise ...
(which are paraphyletic) are the most basal representatives of the order, which are characterized by a bilophodont dental structure, i.e. they have only two pairs of cusps or ridges.Jeheskel Shoshani, W. J. Sanders und Pascal Tassy: ''Elephants and other Proboscideans: a summary of recent findings and new taxonomic suggestions.'' In: G. Cavarretta et al. (Hrsg.): ''The World of Elephants – International Congress. Consiglio Nazionale delle Ricerche.'' Rom, 2001, S. 676–679 ''Dagbatitherium is'' considered to be the oldest member of the Elephantiformes known to date. Other forms like ''Phiomia'' and ''Palaeomastodon'' are much younger and were mostly described from the
Late Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "daw ...
and
Early Oligocene The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chatti ...
of the Fayyum. Compared to these two, ''Dagbatitherium'' was strikingly smaller, had lower tooth crowns, and a more complex enamel pattern. However, a trait that all three forms have in common is a striking bunodont (humped) occlusal surface design. The latter feature contrasts with the sometimes more distinctly lophodont teeth of earlier or contemporaneous members of the Plesielephantiformes, such as ''
Phosphatherium ''Phosphatherium escuillei'' is a basal proboscidean that lived from the Late Paleocene to the early stages of the Ypresian age until the early Thanetian some 56 million years ago in North Africa. Research has suggested that ''Phosphatherium'' ex ...
'', ''
Numidotherium ''Numidotherium'' ("Numidia beast") is an extinct genus of early proboscideans, discovered in 1984, that lived during the middle Eocene of North Africa some 46 million years ago. It was about 90-100 cm tall at the shoulder and weighed about 250-3 ...
'', or ''
Barytherium ''Barytherium'' (meaning "heavy beast") is a genus of an extinct family (Barytheriidae) of primitive proboscideans that lived during the late Eocene and early Oligocene in North Africa. The type species is ''Barytherium grave'', found at the begi ...
'', in which the individual cusps of a pair are connected with ridges. In dental structure, ''Dagbatitherium'' and other early elephantiforms thus more closely resemble ''
Moeritherium ''Moeritherium'' ("the beast from Lake Moeris") is an extinct genus of primitive proboscideans. These prehistoric mammals are related to the elephant and, more distantly, sea cows and hyraxes Hyraxes (), also called dassies, are small, thick ...
'' and ''
Saloumia ''Saloumia'' is an extinct genus of the order Proboscidea. It is one of the oldest members of the order and lived in the middle Eocene of Senegal. It is known only from a single molar, whose pronounced bumpy chewing surface indicates it is probab ...
'', which form an archaic group of proboscideans that was sometimes considered a lateral branch. Further similarities are found in ''
Eritherium ''Eritherium'' is an extinct genus of early Proboscidea found in the Ouled Abdoun basin (early Thanetian age), Morocco. It lived about 60 million years ago. It was first named by Emmanuel Gheerbrant in 2009 and the type species is ''Eritherium ...
'', the most primitive member of the order known to date, which also possessed bunodont grinders. Altogether, however, the relationships of the early proboscideans of the Paleogene are still poorly known. Within the Elephantiformes, ''Dagbatitherium'' has not yet been assigned to a specific family. Below is a phylogenetic tree of early Proboscidea, based on the work of Hautier and colleagues (2021).


References

{{Taxonbar, from=Q109233780 Elephantiformes Prehistoric placental genera Eocene mammals of Africa Fossils of Togo Eocene proboscideans Fossil taxa described in 2021