Protungulatum Temporal range: Maastrichtian to Puercan late | |
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lower jaw of Protungulatum donnae | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Grandorder: | Ferungulata |
Clade: | Pan-Euungulata |
Family: | † Protungulatidae Chatterjee, Scotese & Bajpai, 2017 [1] |
Genus: | † Protungulatum Sloan & Van Valen, 1965 [2] |
Type species | |
†Protungulatum donnae Sloan & Van Valen, 1965 | |
Species | |
Protungulatum ('first ungulate') is an extinct genus of eutherian mammals within extinct family Protungulatidae, [1] and is possibly one of the earliest known placental mammals in the fossil record, that lived in North America from the Late Cretaceous to early Paleocene. [6] [3] [7]
Fossils of this genus were first found in the Bug Creek Anthills in northeastern Montana. The Bug Creek Anthills were initially believed to be Late Cretaceous (latest Maastrichtian) because of the presence of the remains of non-avian dinosaurs and common Cretaceous mammals, [8] but these were later shown to have been reworked [lower-alpha 1] from Late Cretaceous strata, and consequently the Bug Creek Anthills are currently believed to be Early Paleocene (Puercan) in age. [9] Remains from the Ravenscrag Formation of Saskatchewan, Canada have been assigned to Protungulatum donnae. These remains may also be Cretaceous in age, but the age of the Ravenscrag Formation is not entirely certain. In 2011, remains of a new species in this genus, Protungulatum coombsi from the Hell Creek Formation suggested that Protungulatum was present in both the Cretaceous and the Paleocene. [3] However, some later studies questioned this Late Cretaceous age for the genus, considering Protungulatum to be exclusively Paleocene in age. [10]
Studies differ on the placement of Protungulatum. This genus was initially assigned to the family Arctocyonidae, [11] in order Condylarthra, [12] [13] (a group of archaic "ungulates", that is now known to be polyphyletic). Some found that it is not a true placental mammal and it is eutherian found to be close to Placentalia. [14] [3] [15] [16] However, a number of studies consider it to be a placental mammal and stem relative to true ungulates in clade Pan-Euungulata, [17] [18] [19] [20] [7] [21] [22] [10] based on anatomy of inner ear. [7]
The phylogenetic relationships of genus Protungulatum are shown in the following cladogram. [17] [20]
Scrotifera |
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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.
Multituberculata is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, and reached a peak diversity during the Late Cretaceous and Paleocene. They eventually declined from the mid-Paleocene onwards, disappearing from the known fossil record in the late Eocene. They are the most diverse order of Mesozoic mammals with more than 200 species known, ranging from mouse-sized to beaver-sized. These species occupied a diversity of ecological niches, ranging from burrow-dwelling to squirrel-like arborealism to jerboa-like hoppers. Multituberculates are usually placed as crown mammals outside either of the two main groups of living mammals—Theria, including placentals and marsupials, and Monotremata—but usually as closer to Theria than to monotremes. They are considered to be closely related to Euharamiyida and Gondwanatheria as part of Allotheria.
Placental mammals are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguished from monotremes and marsupials in that the fetus is carried in the uterus of its mother to a relatively late stage of development. The name is something of a misnomer considering that marsupials also nourish their fetuses via a placenta, though for a relatively briefer period, giving birth to less developed young which are then nurtured for a period inside the mother's pouch. Placentalia represents the only living group within Eutheria, which contains all mammals more closely related to placentals than to marsupials.
Lambdopsalis is an extinct multituberculate mammal from the Late Paleocene of China and Mongolia. It is placed within the suborder Cimolodonta and is a member of the superfamily Taeniolabidoidea. Fossil remains have been found in the Late Paleocene Nomogen and Khashat Formations in Nao-mugen and Bayn Ulan of China and Mongolia, dated to 59-55 million years ago from the Thanetian age. Lambdopsalis bulla is the type species of this genus. The genus and species were named by Chow and Tao Qi in 1978.
Meniscoessus is a genus of extinct multituberculates from the Upper Cretaceous Period that lived in North America.
Cimexomys is an extinct North American mammal that lived from the Upper Cretaceous to the Paleocene. For a while, it shared the world with dinosaurs, but outlived them. It was a member of the extinct order Multituberculata and lies within the suborder Cimolodonta. It is perhaps a member of the Paracimexomys group, though it is not certain.
Mesodma is an extinct genus of mammal, a member of the extinct order Multituberculata within the suborder Cimolodonta, family Neoplagiaulacidae. It lived during the upper Cretaceous and Paleocene Periods of what is now North America. The earliest definitive record is from the late Santonian stage strata of the Straight Cliffs Formation. A single premolar tooth from the lower Cenomanian stage strata of the Cedar Mountain Formation has been tentatively assigned to this genus based on its similarity, but its describers noted that it is unlikely that Mesodma lived during that time.
Eutheria, also called Pan-Placentalia, is the clade consisting of placental mammals and all therian mammals that are more closely related to placentals than to marsupials.
Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wastebasket taxon, having served as a dumping ground for classifying ungulates which had not been clearly established as part of either Perissodactyla or Artiodactyla, being composed thus of several unrelated lineages.
Purgatorius is a genus of seven extinct eutherian species typically believed to be the earliest example of a primate or a proto-primate, a primatomorph precursor to the Plesiadapiformes, dating to as old as 66 million years ago. The first remains were reported in 1965, from what is now eastern Montana's Tullock Formation, specifically at Purgatory Hill in deposits believed to be about 63 million years old, and at Harbicht Hill in the lower Paleocene section of the Hell Creek Formation. Both locations are in McCone County, Montana.
Cimolesta is an extinct order of non-placental eutherian mammals. Cimolestans had a wide variety of body shapes, dentition and lifestyles, though the majority of them were small to medium-sized general mammals that bore superficial resemblances to rodents, lagomorphs, mustelids, and marsupials.
Dinocerata or Uintatheria, also known as uintatheres, is an extinct order of large herbivorous hoofed mammals with horns and protuberant canine teeth, known from the Paleocene and Eocene of Asia and North America. With body masses ranging up to 4,500 kilograms (9,900 lb) they represent some of the earliest known large mammals.
Ferungulata is a grandorder of placental mammals that groups together mirorder Ferae and clade Pan-Euungulata. It has existed in two guises, a traditional one based on morphological analysis and a revised one taking into account more recent molecular analyses. The Fereungulata is a sister group to the order Chiroptera (bats) and together they make clade Scrotifera.
Cimolestes is a genus of early eutherians with a full complement of teeth adapted for eating insects and other small animals. Paleontologists have disagreed on its relationship to other mammals, in part because quite different animals were assigned to the genus, making Cimolestes a grade taxon of animals with similar features rather than a genus of closely related ones. Fossils have been found in North America, South America, Europe and Africa. Cimolestes first appeared during the Late Cretaceous of North America. According to some paleontologists, Cimolestes died out at the start of the Paleocene, while others report the genus from the early Eocene.
Deltatheroida is an extinct group of basal metatherians that were distantly related to modern marsupials. The majority of known members of the group lived in the Cretaceous; one species, Gurbanodelta kara, is known from the late Paleocene (Gashatan) of China. Their fossils are restricted to Central Asia and North America. This order can be defined as all metatherians closer to Deltatheridium than to Marsupialia.
Leptictida is a possibly paraphyletic extinct order of eutherian mammals. Their classification is contentious: according to cladistic studies, they may be (distantly) related to Euarchontoglires, although they are more recently regarded as the first branch to split from basal eutherians. One recent large-scale cladistic analysis of eutherian mammals favored lepictidans as close to the placental crown-clade; and several other recent analyses that included data from Cretaceous non-eutherian mammals found Leptictis to belong to the superorder Afrotheria.
Phenacodontidae is an extinct family of large herbivorous mammals traditionally placed in the “wastebasket taxon” Condylarthra, which may instead represent early-stage perissodactyls. They lived from the late early Paleocene to early middle Eocene and their fossil remains have been found in North America and Europe. The only unequivocal Asian phenacodontid is Lophocion asiaticus.
Zhelestidae is a lineage of extinct eutherian mammals. Occurring in the Late Cretaceous from the Turonian to the Maastrichtian, they were an extremely successful group, with representatives present in Europe, Asia, India, Africa and North America, ostensibly rendering them a cosmopolitan clade. They were specialised towards an herbivorous lifestyle and were in fact initially considered stem-ungulates, but the presence of epipubics and "archaic" dental characters render them as non-placental eutherians.
Gypsonictops is an extinct genus of leptictidan mammals of the family Gypsonictopidae, which was described in 1927 by George Gaylord Simpson. Species in this genus were small mammals and the first representatives of the order Leptictida, that appeared during the Upper Cretaceous.
Pan-Euungulata is a clade of placental mammals from grandorder Ferungulata, consisting of the taxon Euungulata [sensu stricto] and all taxa (species) more closely related to it than to any other living species.