Dryolestida | |
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Skeleton of Henkelotherium | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Clade: | Cladotheria |
Superorder: | † Dryolestoidea Butler, 1939 |
Order: | † Dryolestida Prothero, 1981 |
Families and genera | |
Dryolestida is an extinct order of mammals, known from the Jurassic and Cretaceous. They are considered basal members of the clade Cladotheria, close to the ancestry of therian mammals. It is also believed that they developed a fully mammalian jaw and also had the three middle ear bones. Most members of the group, as with most Mesozoic mammals, are only known from fragmentary tooth and jaw remains.
The group contains Dryolestidae and the possibly paraphyletic Paurodontidae, and some other unplaced genera, which were small insectivores, known from the Middle Jurassic to Early Cretaceous of Laurasia, primarily Europe and North America, with a single record from Asia. During the Late Jurassic in North America and from the Late Jurassic to Early Cretaceous in Europe, they were among the most diverse mammal groups. [1] They have sometimes been placed as part of a broader group, the "Dryolestoidea", which typically includes Dryolestida and Meridiolestida, a diverse group of mammals including both small insectivores and mid-large sized herbivores known from the Late Cretaceous to Miocene of South America and possibly Antarctica. However, in many phylogenetic analyses, Meridiolestida are recovered as an unrelated group of basal cladotherians, rendering "Dryolestoidea" paraphyletic. [2]
Dryolestids were formerly considered part of Pantotheria and/or Eupantotheria. The clade Quirogatheria, erected by José Bonaparte in 1992, is often used as a synonym for Dryolestida. Originally, Quirogatheria was meant to include Brandoniidae, but this family is now included with the dryolestids.
Dryolestids are mostly represented by teeth, fragmented dentaries and parts of the rostrum. The Jurassic forms retained a coronoid and splenial, but the Cretaceous forms lack these. Another primitive feature is the presence of a Meckelian groove (Meridiolestidans lost it altogether). [3] A fundamentally modern ear is known in at least Dryolestes and mesungulatids. [4] [5]
The basal non dryolestid dryolestidan Henkelotherium from the Late Jurassic of Portugal is known from a partial articulated skeleton, and is thought to have been arboreal, adapted to climbing and living in trees. [6]
Tooth enamel evolved differently in marsupials and eutherians. In a first phase, during the late Triassic and Jurassic, prisms separated from the interprismatic matrix, probably independently in several Mesozoic mammal lineages. More derived enamel types evolved in a second phase, during the Tertiary and Quaternary, but without replacing the old prismatic enamel, instead forming various combinations of three-dimensional structures (called schmelzmuster). Dryolestid dentition is thought to resemble the primitive mammalian dentition before the marsupial-eutherian differentiation and dryolestids are candidates to be the last common ancestor of the two mammalian subclasses. [7] In mesungulatids molar tooth eruption is delayed compared to other dryolestoids. [8]
Dryolestids are known from the Jurassic through Early Cretaceous of the Northern Hemisphere (North America, Eurasia, and North Africa) and from the Late Cretaceous through to the Miocene of South America. [3] Drylestoids are very rarely found in the Cenozoic, as are the few other Mesozoic mammals with later descendants, such as multituberculates, monotremes, and gondwanatheres. [9]
The oldest named member of Dryolestidae is Anthracolestes from the Middle Jurassic (Bathonian) aged Itat Formation in western Siberia. [10] Fragmentary remains attributable to dryolestidans are known from the equivalently aged Forest Marble Formation of England [11] and the Anoual Formation of Morocco. [12]
The youngests fossils of Dryolestidans in the Northern Hemisphere are the dryolestids Crusafontia cuencana from Uña and the Camarillas Formation, Spain and Minutolestes submersus and Beckumia sinemeckelia from Balve, Germany, which all date to the Barremian-Aptian stages of the Early Cretaceous, [13] though a fragmentary lower molar from the late Cretaceous Mesaverde Formation in Wyoming has been tentatively attributed to Dryolestidae. [14] In South America, by contrast, Meridiolestida thrived in the Late Cretaceous, diversifying in a myriad of forms such as the saber-toothed Cronopio and the herbivorous mesungulatids, becoming some of the most ecologically diverse Mesozoic South American mammals. [15] Groebertherium from the Late Cretaceous of South America has a more primitive morphology similar to Northern Hemisphere dryolestids and may be more closely related to the North Hemisphere dryloestidans than to Meridiolestida. [16]
With the advent of the Cenozoic, dryolestoids declined drastically in diversity, with only the large dog-sized herbivore Peligrotherium being known from the Palaeocene. The exact reasons for this decline are not clear; most likely they simply did not recover from the K-Pg event. Nonetheless, meridiolestidans would continue to survive until the Miocene, from when Necrolestes is known; a gap of 50 million years exists between it and Peligrotherium . [17] A tooth fragment, now lost, found in the Eocene aged La Meseta Formation of the Antarctic peninsula, is possibly a meridiolestidan. [18]
A phylogenetic analysis conducted by Rougier et al. (2012) indicated that meridiolestidans might not be members of Dryolestoidea but instead slightly more closely related to the placental mammals, marsupials and amphitheriids. Paurodontids were also recovered as not belonging to Dryolestida, but instead as a sister group of Meridiolestida in this analysis. [19] An analysis conducted by Averianov, Martin and Lopatin (2013) did not recover meridiolestidans as members of Dryolestida as well, but it found them to be the sister group of spalacotheriid "symmetrodonts" instead. However, paurodontids were recovered as members of Dryolestida in this analysis. [20] On the other hand, an analysis conducted by Chimento, Agnolin and Novas (2012) did recover meridiolestidans as members of Dryolestoidea. [21]
Cladogram after Lasseron and colleagues (2022), which found Donodontidae and Meridiolestida unrelated to Dryolestida: [2]
Cladotheria |
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Docodonta is an order of extinct Mesozoic mammaliaforms. They were among the most common mammaliaforms of their time, persisting from the Middle Jurassic to the Early Cretaceous across the continent of Laurasia. They are distinguished from other early mammaliaforms by their relatively complex molar teeth. Docodont teeth have been described as "pseudotribosphenic": a cusp on the inner half of the upper molar grinds into a basin on the front half of the lower molar, like a mortar-and-pestle. This is a case of convergent evolution with the tribosphenic teeth of therian mammals. There is much uncertainty for how docodont teeth developed from their simpler ancestors. Their closest relatives may have been certain Triassic "symmetrodonts", namely Woutersia, Delsatia.
Volaticotherium antiquum is an extinct, gliding, insectivorous mammal that lived in Asia during the Jurassic period, around 164 mya. It is the only member of the genus Volaticotherium.
Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack et al. in 1973 as a replacement name for the paraphyletic Triconodonta.
Vincelestes is an extinct genus of mammal that lived in what is now South America during the Early Cretaceous. It is closely related to modern therian mammals as part of Cladotheria.
Gobiconodon is an extinct genus of carnivorous mammals belonging to the family Gobiconodontidae. Undisputed records of Gobiconodon are restricted to the Early Cretaceous of Asia and North America, but isolated teeth attributed to the genus have also been described from formations in England and Morocco dating as far back as the Middle Jurassic. Species of Gobiconodon varied considerably in size, with G. ostromi, one of the larger species, being around the size of a modern Virginia opossum. Like other gobiconodontids, it possessed several speciations towards carnivory, such as shearing molariform teeth, large canine-like incisors and powerful jaw and forelimb musculature, indicating that it probably fed on vertebrate prey. Unusually among predatory mammals and other eutriconodonts, the lower canines were vestigial, with the first lower incisor pair having become massive and canine-like. Like the larger Repenomamus there might be some evidence of scavenging.
Trechnotheria is a group of mammals that includes the therians and some fossil mammals from the Mesozoic Era. It includes both the extinct symmetrodonts and the living Cladotheria.
Dryolestidae is an extinct family of Mesozoic mammals, known from the Middle Jurassic to the Early Cretaceous of the North Hemisphere. The oldest known member, Anthracolestes, is known from the Middle Jurassic Itat Formation of Western Siberia, but most other representatives are known from the Late Jurassic of North America and the Late Jurassic and Early Cretaceous of Europe. Most members are only known from isolated teeth and jaw fragments. Like many other groups of early mammals, they are thought to have been insectivores. They are generally classified in Cladotheria, meaning that they are considered to be more closely related to marsupials and placentals than to monotremes. They are placed as part of the broader Dryolestida, which also includes the Paurodontidae, and also sometimes the South American-Antarctic Meridiolestida, which are often considered unrelated cladotherians. Dryolestidae taxon is not based on a phylogenetic definition, but instead on the possession of unequal roots for the molars of the lower jaw. Additionally, the clade is distinguished by hypsodonty in lower molars, and uneven labio-lingual height for the alveolar borders of the dentary.
Amphitheriidae is a family of Mesozoic mammals restricted to the Middle Jurassic of Britain, with indeterminate members also possibly known from the equivalently aged Itat Formation in Siberia and the Anoual Formation of Morocco. They were members of Cladotheria, more derived than members of Dryolestida, and forming a close relationship with Peramuridae. Amphitheriidae is the only family of the order Amphitheriida.
Cladotheria is a clade of mammals. It contains modern therian mammals and several extinct groups, such as the "dryolestoids", amphitheriids and peramurids. The clade was named in 1975 by Malcolm McKenna. In 2002, it was defined as a node-based taxon containing "the common ancestor of dryolestids and living therians, plus all its descendants". A different, stem-based definition was given in 2013, in which Cladotheria contains all taxa that are closer to Mus musculus than to the "symmetrodont" Spalacotherium tricuspidens.
Triconodontidae is an extinct family of small, carnivorous mammals belonging to the order Eutriconodonta, endemic to what would become Asia, Europe, North America and probably also Africa and South America during the Jurassic through Cretaceous periods at least from 190–66 mya.
Paurodontidae is a family of Late Jurassic to Early Cretaceous mammals in the order Dryolestida. Remains of paurodontids have been found in the United States, Britain, Portugal, and Tanzania. The group likely represents a paraphyletic group of basal non dryolestid dryolestidans. Paurodon has been suggested to have been a specilast feeder on earthworms due to the morphology of its teeth closely resembling that of the golden mole genus Amblysomus.
Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.
Reigitherium was a mammal that lived during the Late Cretaceous, in the. Its fossils have been found in the Los Alamitos and the La Colonia Formations of Argentina.
Donodon is an extinct genus of mammal from the Ksar Metlili Formation of Talssint, Morocco, which has been dated to the Late Jurassic to Early Cretaceous epochs. The type species D. perscriptoris was described in 1991 by the palaeontologist Denise Sigogneau-Russell. A second species, D. minor, was named in 2022. Donodon was a member of Cladotheria, a group that includes therian mammals and some of their closest relatives. It differed from dryolestids in having upper molars that were not compressed mesiodistally. Some studies have suggested that it was closely related to various South American cladotherians in the clade Meridiolestida, with specific similarities to Mesungulatum, a herbivorous mesungulatid, being noted. On the other hand, a 2022 phylogenetic analysis found it to be only distantly related to meridiolestidans, and instead closer to crown group therians.
Donodontidae is an extinct family of cladotherian mammals known from the Late Jurassic and Early Cretaceous of North Africa. When originally named in 1991, Donodontidae was a monotypic family containing a single species: Donodon perscriptoris. In 2022, four more species were designated and placed within the family: Donodon minor, Stylodens amerrukensis, Anoualestes incidens, and Amazighodon orbis. All five species are endemic to the Ksar Metlili Formation of Morocco, which is dated to the Tithonian and Berriasian. Donodontid fossils are restricted to postcanine teeth and associated jaw fragments.
Cronopio is an extinct genus of small insectivorous mammal known from the early Late Cretaceous of the Río Negro region in Argentina. Its only species is Cronopio dentiacutus. It belongs to the Meridiolestida, an extinct group of mammals widespread in South America during the Late Cretaceous, which are more closely related to modern marsupials and placental mammals than to monotremes.
Necrolestes is an extinct genus of mammals, which lived during the Early Miocene in what is now Argentine Patagonia. It is the most recent known genus of Meridiolestida, an extinct group of mammals more closely related to therians than to monotremes, which were the dominant mammals in South America during the Late Cretaceous. It contains two species, N. patagonensis and N. mirabilis; the type species N. patagonensis was named by Florentino Ameghino in 1891 based on remains found by his brother, Carlos Ameghino in Patagonia. Fossils of Necrolestes have been found in the Sarmiento and Santa Cruz Formations. Its morphology suggests that it was a digging, subterranean-dwelling mole-like mammal that fed on invertebrates.
Leonardus is an extinct mammal genus from the Late Cretaceous of South America. It is a meridiolestidan, closely related to the also Late Cretaceous Cronopio and the Miocene Necrolestes.
Meridiolestida is an extinct clade of mammals known from the Cretaceous and Cenozoic of South America and possibly Antarctica. They represented the dominant group of mammals in South America during the Late Cretaceous. Meridiolestidans were morphologically diverse, containing both small insectivores such as the "sabretooth-squirrel" Cronopio, as well as the clade Mesungulatoidea/Mesungulatomorpha, which ranged in size from the shrew-sized Reigitherium to the dog-sized Peligrotherium. Mesungulatoideans had highly modified dentition with bunodont teeth, and were likely herbivores/omnivores. Meridiolestidans are generally classified within Cladotheria, more closely related to living marsupials and placental mammals (Theria) than to monotremes, barring one study recovering them as the sister taxa to spalacotheriid "symmetrodonts". However, more recent studies have stuck to the cladotherian interpretation. Within Cladotheria, they have often been placed in a group called Dryolestoidea together with Dryolestida, a group of mammals primarily known from the Jurassic and Early Cretaceous of the Northern Hemisphere. However, some analyses have found this group to be paraphyletic, with the meridiolestidans being more or less closely related to therian mammals than dryolestidans are. Meridiolestidans differ from dryolestidans in the absence of a parastylar hook on the molariform teeth and the lack of a Meckelian groove.
Groebertherium is a genus of dryolestoid mammal from the Late Cretaceous Los Alamitos and Allen Formations of Argentina. It is not closely related to other contemporary dryolestoids, all of which are part of the clade Meridiolestida.