| Togocetus Temporal range: Late Eocene, | |
|---|---|
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Artiodactyla |
| Infraorder: | Cetacea |
| Family: | † Protocetidae |
| Genus: | † Togocetus Gingerich & Cappetta 2014 |
| Species: | †T. traversei |
| Binomial name | |
| †Togocetus traversei | |
Togocetus (“Togo whale”) is a genus of extinct cetacean from the Lutetian (lower Eocene) of Togo, known from a fossilized skeleton discovered a few kilometers north-east of Lomé.
The skeleton was found in a phosphate mining area, Kpogamé-Hahotoé, which is located just north of Lake Togo. It was embedded in a phospharenite bone bed dating back to 46 – 44 million years ago, and overlying an older rock unit, the Tabligbo Group. [3] The remains were described in 2014 by Philip D. Gingerich and Henri Cappetta, who established for it the new monotypic genus Togocetus and the new species T. traversei, dedicated to Michel Traverse. [1]
According to the two authors, Togocetus was a semiaquatic animal which must have weighed around 300–400 kilograms (660–880 lb). It was a protocetid with rather primitive traits such as a still quite long neck, a digitigrade manus and a swim-specialized pes. [4] It shared many similarities with the protocetid genera Protocetus and Pappocetus , the main differences being a smaller mandibular canal, the loss of the fovea capitis femoris (hence of the ligament of head of femur) and some traits related to the molar trigonids. [5] Later analysis of ear structure evidence originally attributed to Togocetus raised some question regarding established understanding of protocetid evolution. [6] The original analysis revealed a contradiction in assumed relationship between ear ossicle structure and mandibular canal size; later research and additional evidence revealed multiple places where more modern, specialized traits were blended unexpectedly with archaic ones, causing some controversy. Despite the fragmentary nature of the fossil record and the subtleties differentiating protocetids, T. traversei has consistently been shown to have unique features defining it, including a completely novel entoglenoid shape and several fossae otherwise not seen in contemporary cetaceans. [7]
Rodhocetus is an extinct genus of protocetid early whale known from the Lutetian of Pakistan. The best-known protocetid, Rodhocetus is known from two partial skeletons that taken together give a complete image of an Eocene whale that had short limbs with long hands and feet that were probably webbed and a sacrum that was immobile with four partially fused sacral vertebrae. It is one of several extinct whale genera that possess land mammal characteristics, thus demonstrating the evolutionary transition from land to sea.
Pakicetidae is an extinct family of Archaeoceti that lived during the Early Eocene in Pakistan. Unlike modern cetaceans, they had well developed limbs and were capable of walking.
The evolution of cetaceans is thought to have begun in the Indian subcontinent from even-toed ungulates (Artiodactyla) 50 million years ago (mya) and to have proceeded over a period of at least 15 million years. Cetaceans are fully aquatic mammals belonging to the order Artiodactyla and branched off from other artiodactyls around 50 mya. Cetaceans are thought to have evolved during the Eocene, the second epoch of the present-extending Cenozoic Era. Molecular and morphological analyses suggest Cetacea share a relatively recent closest common ancestor with hippopotami and that they are sister groups. Being mammals, they surface to breathe air; they have five finger bones (even-toed) in their fins; they nurse their young; and, despite their fully aquatic life style, they retain many skeletal features from their terrestrial ancestors. Research conducted in the late 1970s in Pakistan revealed several stages in the transition of cetaceans from land to sea.
Andrewsarchus, meaning "Andrews' ruler", is an extinct genus of artiodactyl that lived during the Middle Eocene in what is now China. The genus was first described by Henry Fairfield Osborn in 1924 with the type species A. mongoliensis based on a largely complete cranium. A second species, A. crassum, was described in 1977 based on teeth. A mandible, formerly described as Paratriisodon, does probably belong to Andrewsarchus as well. The genus has been historically placed in the families Mesonychidae or Arctocyonidae, or was considered to be a close relative of whales. It is now regarded as the sole member of its own family, Andrewsarchidae, and may have been related to entelodonts. Fossils of Andrewsarchus have been recovered from the Middle Eocene Irdin Manha, Lushi, and Dongjun Formations of Inner Mongolia, each dated to the Irdinmanhan Asian land mammal age.
Archaeoceti, or Zeuglodontes in older literature, is a paraphyletic group of primitive cetaceans that lived from the Early Eocene to the late Oligocene. Representing the earliest cetacean radiation, they include the initial amphibious stages in cetacean evolution, thus are the ancestors of both modern cetacean suborders, Mysticeti and Odontoceti. This initial diversification occurred in the shallow waters that separated India and Asia 53 to 45 mya, resulting in some 30 species adapted to a fully oceanic life. Echolocation and filter-feeding evolved during a second radiation 36 to 35 mya.
Protocetus atavus is an extinct species of primitive cetacean from Egypt. It lived during the middle Eocene period 45 million years ago. The first discovered protocetid, Protocetus atavus was described by Fraas 1904 based on a cranium and a number of associated vertebrae and ribs found in middle Lutetian Tethyan marine limestone from Gebel Mokattam near Cairo, Egypt.
Ichthyolestes is an extinct genus of archaic cetacean that was endemic to Indo-Pakistan during the Lutetian stage. To date, this monotypic genus is only represented by Ichthyolestes pinfoldi.
Artiocetus is an extinct genus of early whales belonging to the family Protocetidae. It was a close relative to Rodhocetus and its tarsals indicate it resembled an artiodactyl.
Remingtonocetidae is a diverse family of early aquatic mammals of the order Cetacea. The family is named after paleocetologist Remington Kellogg.
Basiloterus is an extinct genus of late-Eocene archaeocete whale from the Drazinda Formation in southwestern Punjab, Pakistan and possibly also the Barton Group of England. Known from two isolated lumbar vertebrae, the elongated nature of these elements has been taken as possible evidence that Basiloterus was a close relative of the better-known Basilosaurus. This was also the reasoning behind its name, which roughly translates to "another king". However, publications since then not only lead to some major changes of the internal relationships within Basilosauridae but have also called into question how diagnostic elongated vertebrae are for members of this group, as other early whales have developed similar anatomy independently. Though the identity of Basiloterus as a basilosaurid is generally maintained, its exact position within more recent interpretations of the family is unclear.
The Qasr el Sagha Formation is a geological formation located in Egypt. The formation is part of the Wadi El Hitan World Heritage Site. The Qasr el Sagha Formation overlies the Birket Qarun Formation and is overlain by the Gebel Qatrani Formation. The sandstones and shales of the formation were deposited in a deltaic to shallow marine environment. It dates to the Late Eocene.
Protocetidae, the protocetids, form a diverse and heterogeneous group of extinct cetaceans known from Asia, Europe, Africa, South America, and North America.
Aegyptocetus is an extinct genus of protocetid archaeocete whale known from Egypt.
Pappocetus is an extinct protocetid cetacean known from the Eocene of southern Nigeria's Ameki Formation and Togo. More recently, fossil teeth and femurs have also been discovered in the Aridal Formation of the Sahara Desert in southwestern Morocco.
Carolinacetus is an extinct protocetid early whale found in the Bartonian Tupelo Bay Formation in Berkeley County, South Carolina.
Eocetus is an extinct protocetid early whale known from the early-late Eocene Giushi Formation in Gebel Mokattam, outside Cairo, Egypt. Fossil remains have also been discovered in the Aridal Formation of the Sahara Desert in southwestern Morocco.
Elephantiformes is a suborder within the order Proboscidea. Members of this group are primitively characterised by the possession of upper tusks, an elongated mandibular symphysis and lower tusks, and the retraction of the facial region of the skull indicative of the development of a trunk. The earliest known member of the group, Dagbatitherium is known from the Eocene (Lutetian) of Togo, which is only known from isolated teeth, while other primitive elephantiforms like Phiomia and Palaeomastodon are known from the Early Oligocene onwards. Phiomia and Palaeomastodon are often collectively referred to as "palaeomastodonts" and assigned to the family Palaeomastodontidae. Most diversity of the group is placed in the subclade Elephantimorpha, which includes mastodons, as well as modern elephants and gomphotheres (Elephantida). It is disputed as to whether Phiomia is closely related to both Mammutidae and Elephantida with Palaeomastodon being more basal, or if Palaeomastodon is closely related to Mammutidae and Phiomia more closely related to Elephantida.
Dagbatitherium is an extinct genus of proboscideans. So far a single molar from the phosphate basins of Togo 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. 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.
Tupelocetus is an extinct genus of early cetacean found in the Bartonian Middle Eocene Tupelo Bay Formation, in Berkeley County, South Carolina.
Gingerich, P. D.; Cappetta, H. (2014). "A new archaeocete and other marine mammals (Cetacea and Sirenia) from lower middle Eocene phosphate deposits of Togo". Journal of Paleontology . 88 (1): 109–129. Bibcode:2014JPal...88..109G. doi:10.1666/13-040. S2CID 85915213.
MKassegne, K. E (11 June 2021). "First partial cranium of Togocetus from Kpogamé (Togo) and the protocetid diversity in the Togolese phosphate basin". Annales de Paléontologie. 107 (2): 102488. Bibcode:2021AnPal.10702488K. doi: 10.1016/j.annpal.2021.102488 . S2CID 236757400.
Mourlam, M.J.; Orliac, M.J. (2017). "Protocetid (cetacea, artiodactyla) bullae and petrosals from the Middle Eocene locality of KPOGAMÉ, Togo: New insights into the early history of cetacean hearing". Journal of Systematic Palaeontology. 16 (8): 621–644. Bibcode:2018JSPal..16..621M. doi:10.1080/14772019.2017.1328378. S2CID 89774296 – via EBSCO.
Archaeocete genera by family | |||||||
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| Pakicetidae | |||||||
| Ambulocetidae | |||||||
| Remingtonocetidae | |||||||
| Protocetidae |
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| Basilosauridae |
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