Eutriconodonta

Eutriconodonts Temporal range: Middle Jurassic - Late Cretaceous, 167–70 Ma PreЄ Є O S D C P T J K Pg N
Life restoration of a Jeholodens
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	†Eutriconodonta Kermack et al., 1973
Subgroups
?†Amphidontidae ?†Amphilestidae ?†Gobiconodontidae ?†Klameliidae ?†Bocaconodon †Jeholodens †Liaoconodon †Yanoconodon †Triconodontidae

Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, 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.

Classification

Illustration of the lower jaw of Triconodon mordax, 1861

"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).

Cladogram after Averianov & Lopatin, 2011:

Holotheria	Gobiconodonta Gobiconodontidae Klameliidae Amphilestidae Amphilestes Phascolotherium Amphidontidae Eutriconodonta Bocaconodon Jeholodentidae Triconodontidae Tinodon Trechnotheria Kiyatherium Spalacotheriidae Zhangheotheriidae Cladotheria

Cladogram after Gao et al., 2010 and Meng, Wang & Li, 2011:

Holotheria	Amphidontidae Gobiconodontidae Eutriconodonta Jeholodens Yanoconodon Liaoconodon Triconodontidae Priacodon Triconodon Trioracodon Arundeloconodon Astroconodon Alticonodon Corvicondodon Phascolotherium Amphilestes Tinodon Trechnotheria Spalacotheriidae Zhangheotheriidae Cladotheria

The cladograms above resulted from the analyses using 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.

Biology

Like many other non-therian mammals, eutriconodonts retained classical mammalian synapomorphies like epipubic bones (and likely the associated reproductive constrictions), venomous spurs and sprawling limbs. However, the forelimb and shoulder anatomy of at least some species like Jeholodens are similar to those of therian mammals, though the hindlimbs remain more conservative.

Some eutriconodonts like Spinolestes and Volaticotherium were very well preserved, showing evidence of fur, internal organs and, in the latter, of patagia. Spinolestes shows hair similar to that of modern mammals, with compound hair follicles with primary and secondary hair, even preserving traces of a pore infection. It also possesses a clear thoracic diaphragm like modern mammals, as well as spines, dermal scutes and an ossified Meckel's cartilage. Furthermore, Spinolestes may also display signs of dermatophytosis, suggesting that gobiconodontids, like modern mammals, were vulnerable to this type of fungal infection.

The eutriconodont triconodont dentition has no analogue among living mammals, so comparisons are difficult. However, its clear that most if not all eutriconodonts were primarily carnivorous, given the presence of long, sharp canines, premolars with trenchant main cusps that were well suited to grasp and pierce prey, strong development of the madibular abductor musculature, bone crushing ability in at least some species and several other features.

Eutriconodonts are often among the largest mammals in Mesozoic faunal assemblages, displaying a broad size range from small shrew-like insectivores to large forms like Repenomamus, Gobiconodon and Jugulator. They were among the first mammals to be specialised for vertebrate prey, and likely occupied the highest trophic levels among mammals in their faunal communities. Several forms like Gobiconodon and Repenomamus show evidence of scavenging, being among the few Mesozoic mammals to have significantly exploited that.

At least in carnivorous niches, eutriconodonts were probably replaced by deltatheroidean metatherians, which are the dominant carnivorous mammals in Late Cretaceous faunal assemblages.

Some eutriconodonts were instead among the most specialised of Mesozoic mammals. Several taxa like Astroconodon, Dystritodon and Ichthyoconodon may show adaptations for piscivory and occur in aquatic settings, and though some caution is advised due to the lack of direct dental comparisons with modern mammals at least Ichthyoconodon's teeth shows no erosion associated with aquatic transportation, meaning that the animal died in situ or close.

Additionally, Volaticotherium and Argentoconodon show adaptations for aerial locomotion. Both genera are closely related, implying a long lived lineage of flying or gliding mammals. Ichthyoconodon is also part of this clade, and its appearance in marine deposits without tooth erosion could imply powered flight.

At least Spinolestes had xenarthrous vertebrae and osseous scutes, convergent to those of modern xenarthrans and to a lesser extent the hero shrew. This genus may have displayed an ecological role similar to that of modern anteaters, pangolins, echidnas, aardvark, aardwolf and numbat, being the second known Mesozoic mammal after Fruitafossor to have done so.