Tutcetus | |
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Holotype specimen block | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Artiodactyla |
Infraorder: | Cetacea |
Family: | † Basilosauridae |
Genus: | † Tutcetus Antar et al., 2023 |
Type species | |
†Tutcetus rayanensis Antar et al., 2023 |
Tutcetus is an extinct genus of diminutive basilosaurid cetacean from the Bartonian of Egypt. Tutcetus, named after the child pharaoh Tutankhamun, is both one of the oldest known basilosaurids from Africa and the smallest member of the family. It is suggested that the type specimen, a subadult close to maturity, only measured approximately 2.5 m (8 ft 2 in) long. The genus is monotypic, only including the species T. rayanensis.
Tutcetus was first described in August 2023 on the basis of a single specimen, MUVP 501. [1] The specimen consists of an incomplete skull found alongside both mandibles, the hyoid, and some of the first vertebrae of the neck. [1] The remains stem from the Fayum Depression of Egypt, which is well known for its rich record of early whales. [2] More specifically, the bones of Tutcetus stem from the Sath El-Hadid Formation, which dates to the early Bartonian (ca. 41 mya). [1]
The name Tutcetus is a reference to the pharaoh Tutankhamun, also known as King Tut, a child king who died when he was only 18 years old. The name was chosen to reflect the small size and young age of Tutcetus. The second part of the generic name is the Ancient Greek word for whale, "cetus". The species name meanwhile means "from the Wadi El-Rayan Area" in reference to the type locality. [1]
Most of the sides of the nasal bones is in contact with the maxillae, with only a small section towards the front of the nasals contacting the premaxillae. [1]
The upper and lower premolars of Tutcetus differ from those of all other basilosaurids in the number of accessory tooth cusps. Tutcetus features two mesial accessory cusps (directed towards the front of the jaw) and three distal accessory cusps (directed towards the back of the jaw). The premolars also differ in some other aspects. They are notably more gracile than those of other basilosaurids and the tooth enamel is much smoother. The fourth premolar appears to have been the largest tooth in both the upper and lower jaw. The first premolar does not appear to have had a replacement, which suggests one of two things. Either the deciduous first premolar is retained even into adulthood, or the first premolar is developed without having a temporary precursor. [1]
Tutcetus may have been the smallest known basilosaurid with an estimated length of 2.51–2.55 m (8 ft 3 in – 8 ft 4 in) and an estimated weight of 180.4–187.1 kg (398–412 lb). Although maturity is difficult to determine in fossil, Antar and colleagues note several factors observed in Tutcetus that give clues to the animal's age at the time of its death . These factors include both the stage of the tooth eruption and the fusion of the bones, which are all more advanced than those of the oldest known Dorudon juvenile and suggest that the holotype of Tutcetus was an advanced subadult at the verge of adulthood. [1]
According to the Bayesian tip-dating analysis conducted for Tutcetus, Basilosauridae was a paraphyletic clade that included both traditional basilosaurids as well as all modern whale groups (Mysticeti and Odontoceti). [2] Among core basilosaurids, Tutcetus was recovered within a weakly-supported group of early diverging, middle Eocene whales that also included Chrysocetus from North America and Africa as well as Ocucajea from South America. Within this group, termed the Tutcetus-clade by Antar et al., Tutcetus was found to be most closely related to Ocucajea, with Chrysocetus having diverged before the split between the other two. [1]
Pelagiceti | |
Mohamed Antar, Abdullah Gohar, Hesham Sallam and colleagues note that, beyond determining the age at which the animal died, the pattern of tooth replacement seen in Tutcetus may also give additional clues to how the animal lived. Tutcetus had molar teeth that erupted rather early, suggesting that the switch from deciduous to permanent teeth occurred rapidly in members of this species. According to Antar et al., rapid onset of tooth replacement is associated with animals that mature quickly and don't grow exceptionally old. They argue that the inferred low lifespan of Tutcetus, combined with its diminutive size, indicate that it was a precocial animal that grew quicker and died younger than the larger basilosaurids of its time. [1]
The deposits in which Tutcetus was found in match the type of warm, shallow, tropical waters that whales will seek out as calving grounds, with authors suggesting that the locality represents just that. Antar and colleagues note that the stage of tooth replacement during which the holotype specimen of Tutcetus died might suggest that the species had a relatively younger infant mortality rate compared to the much better sampled Dorudon. They further highlight how such patterns would be expected in species that give birth every year to a single young. However, they also point out how this hypothesis cannot be tested until more material of Tutcetus is found and described. [1]
Another aspect of Tutcetus' biology briefly explored in the type description is the relation between its environment and its small body size. While the small size of Tutcetus could be simply a left over of its smaller protocetid ancestry, it is also possible that the small size of it could be tied directly during the period of global warming known as the Lutetian thermal maximum. As aging and mortality are more prominent in warmer conditions, the small size and early maturing of Tutcetus could have been adaptations to these warmer conditions, allowing for the animal to reproduce more quickly. On the flipside, the large sizes of later basilosaurids such as Basilosaurus itself may have been driven by the middle Eocene climatic optimum or the brief cooling period between the Lutetian thermal maximum and the middle Eocene climatic optimum. Antar and colleagues conclude that body size in these early whales may have been primarily driven by climate, whereas body shape would have been mostly changed in response to competition. [1]
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.
Ambulocetus is a genus of early amphibious cetacean from the Kuldana Formation in Pakistan, roughly 48 or 47 million years ago during the Early Eocene (Lutetian). It contains one species, Ambulocetus natans, known solely from a near-complete skeleton. Ambulocetus is among the best-studied of Eocene cetaceans, and serves as an instrumental find in the study of cetacean evolution and their transition from land to sea, as it was the first cetacean discovered to preserve a suite of adaptations consistent with an amphibious lifestyle. Ambulocetus is classified in the group Archaeoceti—the ancient forerunners of modern cetaceans whose members span the transition from land to sea—and in the family Ambulocetidae, which includes Himalayacetus and Gandakasia.
Basilosaurus is a genus of large, predatory, prehistoric archaeocete whale from the late Eocene, approximately 41.3 to 33.9 million years ago (mya). First described in 1834, it was the first archaeocete and prehistoric whale known to science. Fossils attributed to the type species B. cetoides were discovered in the United States. They were originally thought to be of a giant reptile, hence the suffix "-saurus", Ancient Greek for "lizard". The animal was later found to be an early marine mammal, prompting attempts at renaming the creature, which failed as the rules of zoological nomenclature dictate using the original name given. Fossils were later found of the second species, B. isis, in 1904 in Egypt, Western Sahara, Morocco, Jordan, Tunisia, and Pakistan. Fossils have also been unearthed in the southeastern United States and Peru.
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.
Basilosauridae is a family of extinct cetaceans. They lived during the middle to the early late Eocene and are known from all continents, including Antarctica. They were probably the first fully aquatic cetaceans. The group is noted to be a paraphyletic assemblage of stem group whales from which the monophyletic Neoceti are derived.
Dorudon ("spear-tooth") is a genus of extinct basilosaurid ancient whales that lived alongside Basilosaurus 40.4 to 33.9 million years ago in the Eocene. It was a small whale, with D. atrox measuring 5 metres (16 ft) long and weighing 1–2.2 metric tons. Dorudon lived in warm seas around the world and fed on small fish and mollusks. Fossils have been found along the former shorelines of the Tethys Sea in present-day Egypt and Pakistan, as well as in the United States, New Zealand and Western Sahara.
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.
Zygorhiza ("Yoke-Root") is an extinct genus of basilosaurid early whale known from the Late Eocene of Louisiana, Alabama, and Mississippi, United States, and the Bartonian to the late Eocene of New Zealand . Specimens reported from Europe are considered Dorudontinae incertae sedis.
Pachycetus is an extinct genus of pachycetine basilosaurid from Middle Eocene of the eastern United States and Europe. The best known remains generally suggest that Pachycetus lived during the Bartonian, however, fossil finds have also been recovered from sediments of less certain age that could suggest that it may have also lived during the Late Lutetian and Early Priabonian. Pachycetus is primarily known from vertebrae and ribs and is characterized by its highly osteosclerotic and pachyostotic skeleton. This means the bones not only featured thickened rings of cortical bone surrounding the internal cancellous bone, but the cortical bone was furthermore much denser than in other basilosaurids. Two species of Pachycetus are recognized: Pachycetus paulsonii from Europe and Pachycetus wardii from the United States. A third species might be represented by "Zeuglodon" wanklyni.
Georgiacetus is an extinct genus of ancient whale known from the Eocene period of the United States. Fossils are known from Georgia, Alabama, and Mississippi and protocetid fossils from the right time frame, but not yet confirmed as Georgiacetus, have been found in Texas and South Carolina.
Cynthiacetus is an extinct genus of basilosaurid early whale that lived during the Late Eocene Specimens have been found in the southeastern United States and Peru.
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.
Chrysocetus is a genus of extinct early whale known from Late Eocene-aged fossils of the eastern United States and western Africa. It is the second smallest basilosaurid after Saghacetus.
Ocucajea is an extinct genus of basilosaurid cetacean from Middle Eocene deposits of southern Peru. Ocucajea is known from the holotype MUSM 1442, a partial skeleton. It was collected in the Archaeocete Valley site, from the Paracas Formation of the Pisco Basin about 40.4 to 37.2 million years ago.
Supayacetus is an extinct genus of basilosaurid cetacean from the Middle Eocene Paracas Formation of Peru. It has been noted for its relatively small size and basal morphology, with the sternum bearing close resemblance to those of protocetids. Due to this, it has been traditionally placed as one of the basalmost basilosaurids, except for a 2023 study that places it within the family Pachycetinae close to Neoceti. Supayacetus is monotypic, meaning the genus includes only a single species: S. muizoni.
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.
Pachycetinae is an extinct subfamily of basilosaurid cetaceans that lived during the middle Eocene. The best-dated remains stem from Bartonian strata, but some finds suggest that they could have first appeared during the Lutetian and may have survived until the Priabonian. Fossils of pachycetines are chiefly known from the southern United States, Ukraine, Morocco and Germany, among others. They differ from other basilosaurids in having pachyostotic and osteosclerotic vertebrae and ribs, making them denser and heavier by comparison. Based on this it has been suggested that these whales lived in shallow waters and that these thickened bones act as a buoyancy control as seen in sirenians. Analysis of the teeth suggests that pachycetines had a varying diet, with the robust teeth of the larger Pachycetus indicating that it possibly fed on sharks, whereas the more gracile teeth of Antaecetus suggest a diet of smaller prey items. The clade currently only includes two genera, Antaecetus and Pachycetus, but a 2023 study suggests that the Peruvian Supayacetus may at least be a close relative.
Antaecetus is an extinct genus of pachycetine basilosaurid from the middle Eocene Aridal Formation of Morocco as well as the Fayum, Egypt. Antaecetus, although known from fewer remains in total, is markedly more complete than the closely related Pachycetus, with one specimen preserving large parts of the vertebral column up to the lumbar vertebrae and a well preserved skull. Based on these remains Antaecetus appears to have been smaller than Pachycetus, with a proportionally smaller head and much more gracile teeth. Both genera however share a highly osteosclerotic and pachyostotic skeleton, greatly increasing their weight and possibly serving as additional ballast. In turn the elongation and thickening of the vertebrae severely impacts the animals movement, most likely causing it to have been much slower and far less mobile than other archaeocetes. It has been proposed that Antaecetus was a slow moving inhabitant of shallow coastal waters, where it would ambush fish and marine molluscs. Only a single species of Antaecetus is known, Antaecetus aithai, which was originally described as a species of Pachycetus.
Tupelocetus is an extinct genus of early cetacean found in the Bartonian Middle Eocene Tupelo Bay Formation, in Berkeley County, South Carolina.
The Sath El-Hadid Formation, translating to "Iron Surface" in Arabic "سطح الحديد", is a geological formation in Egypt characterized by a nummulitic limestone bank containing large and small Nummulites. Introduced into the stratigraphy of the south Fayum area by Iskander in 1943, this formation is significant in the middle Eocene (Bartonian).