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Eutriconodonta

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Eutriconodonta

Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia (including pre-contact India), 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.

Traditionally seen as the classical Mesozoic small mammalian insectivores, discoveries over the years have shown them to be among the best examples of the diversity of mammals in this time period, including a vast variety of bodyplans, ecological niches and locomotion methods.

"Triconodonta" had long been used as the name for an order of early mammals which were close relatives of the ancestors of all present-day mammals, characterized by molar teeth with three main cusps on a crown that were arranged in a row. The group originally included only the family Triconodontidae and taxa that were later assigned to the separate family Amphilestidae, but was later expanded to include other taxa such as Morganucodon or Sinoconodon. The phylogenetic analyses found that all these taxa did not form a natural group, and that some traditional "triconodonts" were more closely related to therian mammals than others. Some traditional "triconodonts" do seem to form a natural group (or "clade"), and this was given the name Eutriconodonta (or "true triconodonts").

Most analyses use only dental and mandibular characters. Gao et al. (2010) conducted a second analysis as well, using a modified version of the matrix from the analysis of Luo et al. (2007); this analysis involved a broader range of Mesozoic mammaliaforms and more characters, including postcranial ones. Both Luo et al. (2007) and the second analysis of Gao et al. (2010) recovered a more inclusive monophyletic Eutriconodonta that also contained gobiconodontids and Amphilestes; in the second analysis of Gao et al. it also contained Juchilestes (recovered as amphidontid in their first analysis, the only amphidontid included in their second analysis). However, Gao et al. (2010) stressed that jeholodentids and gobiconodontids are the only eutriconodonts with known postcranial skeletons; according to the authors, it remains uncertain whether the results of their second analysis represent true phylogeny or are merely "a by-product of long branch attraction of jeholodentids and gobiconodontids". Phylogenetic studies conducted by Zheng et al. (2013), Zhou et al. (2013) and Yuan et al. (2013) recovered monophyletic Eutriconodonta containing triconodontids, gobiconodontids, Amphilestes, Jeholodens and Yanoconodon.

The exact phylogenetic placement of eutriconodonts within Mammaliaformes is also uncertain. Zhe-Xi Luo, Zofia Kielan-Jaworowska and Richard Cifelli (2002) conducted an analysis that recovered eutriconodonts within the crown group of Mammalia, i.e. the least inclusive clade containing monotremes and therian mammals. The analysis found eutriconodonts to be more closely related to therian mammals than monotremes were, but more distantly than (paraphyletic) amphitheriids, dryolestids, spalacotheriid "symmetrodonts" and multituberculates were. This result was mostly confirmed by Luo et al. (2007), the second analysis of Gao et al. (2010), Zheng et al. (2013), Zhou et al. (2013) and Yuan et al. (2013), although in the phylogenies of Luo et al. (2007) and Yuan et al. (2013) eutriconodonts were in unresolved polytomy with multituberculates and trechnotherians. If confirmed this would make eutriconodonts one of the groups that can be classified as mammals by any definition. Several other extinct groups of Mesozoic animals that are traditionally considered to be mammals (such as Morganucodonta and Docodonta) are now placed just outside Mammalia by those who advocate a 'crown-group' definition of the word "mammal". However, Luo, Kielan-Jaworowska and Cifelli (2002) tested alternative possible phylogenies as well, and found that recovering eutriconodonts outside the crown group of Mammalia required only five additional steps compared to the most parsimonious solution. The authors stated that such placement of eutriconodonts is less likely than their placement within the mammalian crown group, but it cannot be rejected on a statistical basis.

The most recent cladogram is by Thomas Martin et al. 2015, in their description of Spinolestes. Eutriconodonts are recovered as a largely monophyletic group within Theriimorpha.

A 2020 study found them paraphyletic in regards to crown group Mammalia.

When eutriconodonts first appeared is unclear. The earliest remains come from the late Early Jurassic (Toarcian), but they already represent a variety of groups: the volaticotherian Argentoconodon, the alticonodontine Victoriaconodon and the gobiconodontid Huasteconodon, as well as the putative eutriconodont "Dyskritodon" indicus. They achieve their peak diversity across the Early Cretaceous, before largely disappearing from the fossil record in the early Late Cretaceous outside of North America. The Maastrichtian genus Indotriconodon is the youngest representative of the group, hailing from the intertrappean beds of India; the Campanian/Maastrichtian Austrotriconodon was originally referred to as a late surviving member of the clade, but has since been moved to Dryolestoidea.

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