Ungulates are members of the diverse clade Euungulata, which primarily consists of large mammals with hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined to be a polyphyletic and thereby invalid clade based on molecular data. As a result, true ungulates had since been reclassified to the newer clade Euungulata in 2001 within the clade Laurasiatheria while Paenungulata has been reclassified to a distant clade Afrotheria. Living ungulates are divided into two orders: Perissodactyla including equines, rhinoceroses, and tapirs; and Artiodactyla including cattle, antelope, pigs, giraffes, camels, sheep, deer, and hippopotamuses, among others. Cetaceans such as whales, dolphins, and porpoises are also classified as artiodactyls, although they do not have hooves. Most terrestrial ungulates use the hoofed tips of their toes to support their body weight while standing or moving. Two other orders of ungulates, Notoungulata and Litopterna, both native to South America, became extinct at the end of the Pleistocene, around 12,000 years ago.
Artiodactyls are placental mammals belonging to the order Artiodactyla. Typically, they are ungulates which bear weight equally on two of their five toes. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, most perissodactyls bear weight on an odd number of the five toes. Another difference between the two orders is that many artiodactyls digest plant cellulose in one or more stomach chambers rather than in their intestine. Molecular biology, along with new fossil discoveries, has found that cetaceans fall within this taxonomic branch, being most closely related to hippopotamuses. Some modern taxonomists thus apply the name Cetartiodactyla to this group, while others opt to include cetaceans within the existing name of Artiodactyla. Some researchers use "even-toed ungulates" to exclude cetaceans and only include terrestrial artiodactyls, making the term paraphyletic in nature.
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.
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.
Pakicetus is an extinct genus of amphibious cetacean of the family Pakicetidae, which was endemic to Indian Subcontinent during the Ypresian period, about 50 million years ago. It was a wolf-like mammal, about 1–2 m long, and lived in and around water where it ate fish and other animals. The name Pakicetus comes from the fact that the first fossils of this extinct amphibious whale were discovered in Pakistan. The vast majority of paleontologists regard it as the most basal whale, representing a transitional stage between land mammals and whales. It belongs to the even-toed ungulates with the closest living non-cetacean relative being the hippopotamus.
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.
Sirenia is the order of placental mammals which comprises modern "sea cows" and their extinct relatives. They are the only extant herbivorous marine mammals and the only group of herbivorous mammals to have become completely aquatic. Sirenians are thought to have a 50-million-year-old fossil record. They attained modest diversity during the Oligocene and Miocene, but have since declined as a result of climatic cooling, oceanographic changes, and human interference. Two genera and four species are extant: Trichechus, which includes the three species of manatee that live along the Atlantic coasts and in rivers and coastlines of the Americas and western Africa, and Dugong, which is found in the Indian and Pacific oceans.
Zygophyseter varolai is an extinct sperm whale that lived during the Tortonian age of the Late Miocene 11.2 to 7.6 million years ago. It is known from a single specimen from the Pietra Leccese Formation in Italy. It was a member of a stem group of fossil macroraptorial sperm whales also including Brygmophyseter, Acrophyseter, and Livyatan. It probably grew to be around 6.5 to 7 meters in length and shared some characteristics with other raptorials, such as large teeth with tooth enamel that were functional in both the upper and lower jaws which the modern sperm whale lacks. It also had a beak, the ability to echolocate prey, and could have probably swum faster than the modern-day sperm whale which can reach 4 kilometers per hour (2.5 mph). These were probably used in the capture of large prey, such as large fish, seals, and whales. In fact, its common name, the killer sperm whale, refers to its feeding habits that would have had a resemblance to the modern-day killer whale.
Remingtonocetus is an extinct genus of early cetacean freshwater aquatic mammals of the family Remingtonocetidae endemic to the coastline of the ancient Tethys Ocean during the Eocene. It was named after naturalist Remington Kellogg.
Brygmophyseter, known as the biting sperm whale, is an extinct genus of toothed whale in the sperm whale family with one species, B. shigensis. When it was first described in 1994, the species was placed in the genus Scaldicetus based on tooth morphology, but this was later revised in 1995. In 2006, it was classified into the genus Naganocetus, which is considered to be a junior synonym. The only known specimen, a nearly complete skeleton, was dated to be around 16–15 million years old. Brygmophyseter is thought to have been 6.5–7 meters (21–23 ft) long, and it probably had 11 or 12 teeth in the upper and lower jaws. Brygmophyseter is part of a group of macroraptorial sperm whales which tended to be apex predators using their large teeth to catch struggling prey such as whales. It had a spermaceti organ which was probably used for biosonar like in the modern sperm whale. The whale has made an appearance on The History Channel's TV series Jurassic Fight Club.
Maiacetus is a genus of early cetacean from the Habib Rahi Formation of Pakistan.
Protocetidae, the protocetids, form a paraphyletic, diverse and heterogeneous group of extinct cetaceans known from Asia, Europe, Africa, South America, and North America.
Livyatan is an extinct genus of macroraptorial sperm whale containing one known species: L. melvillei. The genus name was inspired by the biblical sea monster Leviathan, and the species name by Herman Melville, the author of the famous novel Moby-Dick about a white bull sperm whale. Herman Melville often referred to whales as "Leviathans" in his book. It is mainly known from the Pisco Formation of Peru during the Tortonian stage of the Miocene epoch, about 9.9–8.9 million years ago (mya); however, finds of isolated teeth from other locations such as Chile, Argentina, the United States (California), South Africa and Australia imply that either it or a close relative survived into the Pliocene, around 5 mya, and may have had a global presence. It was a member of a group of macroraptorial sperm whales and was probably an apex predator, preying on whales, seals and so forth. Characteristically of raptorial sperm whales, Livyatan had functional, enamel-coated teeth on the upper and lower jaws, as well as several features suitable for hunting large prey.
Huaridelphis is an extinct genus of river dolphins from the Early Miocene. The type species is H. raimondii, found in the Chilcatay Formation of the Pisco Basin.
Aetiocetidae is an extinct family of toothed baleen whales known from the Oligocene and latest Eocene, so far only from rocks deposited in the North Pacific Ocean. The whales ranged in size from 3 to 8 metres long. Many of the described specimens were discovered from the Upper Oligocene of the Japanese Morawan Formation, the largest known one from the Morawan's Upper tuffaceous siltstone. Other formally described extinct toothed mysticetis from this time are smaller, from 3 to 4 metres in length. Mysticeti with true baleen are seen in fossils from the Upper Oligocene. The monophyly of the family is still uncertain, as are the evolutionary relationship between the early toothed baleen whales and the early and extant edentulous baleen whales. However, the cladistic analyses of Coronodon and Mystacodon seem to indicate that Aetiocetidae and Llanocetidae are more closely related to crown Mysticeti than to Mammalodontidae, Coronodon, and Mystacodon.
Mystacodon is a genus of toothed baleen whale from the Late Eocene Yumaque Member of Paracas Formation of the Pisco Basin in southwestern Peru. It is the oldest known baleen whale, and was probably a suction feeder of small prey on the seafloor.
Pisco Basin is a sedimentary basin extending over 300 kilometres (190 mi) in southwestern Peru. The basin has a 2 kilometres (6,600 ft) thick sedimentary fill, which is about half the thickness of more northern foreland basins in Peru.
