Splitting up the Ungulates

Get ready for a good dose of heresy
brought to you by the large reptile tree (LRT 1003 taxa) a fully resolved online morphological study.

A little backstory
According to Wikipedia: “Ungulates are any members of a diverse group of primarily large mammals that includes odd-toed ungulates such as horses and rhinoceroses, and even-toed ungulates such as cattle, pigs, giraffes, camels, deer, and hippopotami.”

“However, in 2009 morphological and molecular^[ work has found that aardvarks, hyraxes, sea cows, and elephants are more closely related to sengis, tenrecs, and golden moles than to the perissodactyls and artiodactyls, and form Afrotheria. Elephants, sea cows, and hyraxes are grouped together in the clade Paenungulata, while the aardvark has been considered as either a close relative to them or a close relative to sengis in the clade Afroinsectiphilia. This is a striking example of convergent evolution.

“There is now some dispute as to whether this smaller Ungulata is a cladistic (evolution-based) group, or merely a pheneticgroup (form taxon) or folk taxon (similar, but not necessarily related). Some studies have indeed found the mesaxonianungulates and paraxonian ungulates to form a monophyletic lineage, closely related to either the Ferae (the carnivorans and the pangolins) in the clade Fereuungulata or to the bats. Other studies found the two orders not that closely related, as some place the perissodactyls as close relatives to bats and Ferae in Pegasoferae and others place the artiodactyls as close relatives to bats.” 

About that last hypothesis ~
If I had recovered that relationship I would expect pitchforks and torches.

Today
the LRT presents a hypothesis of mammal relationships in which hippos nest apart from other artiodactyls and perissodactyls nest apart from those two, separated by the elephant/hyrax/manatee clade with the hyrax-like Ectocion at it base (Fig. 1).

Note:
Hippos do indeed nest with whales, but only the mysticete whales. And desmostylians separate hippos from mysticetes. Odontocetes, like Orcinus, had a separate ancestry as we learned earlier here. The even-toed artiodactyls nest together with pig-types toward the base and deer-types a little later. The odd-toed perissodactyls nest together.

There are no
tenrecs, aardvarks or golden moles in this sector of the LRT.

Ungulata is not a monophyletic cladistic group,
unless it also includes the elephants and whales with Mesonyx as a basal member, (a taxon close to the last common ancestor). I’m not the first one to come to this conclusion. Probably best just to drop this name from usage and realize that hooves appeared several times in mammal evolution.

A little extra study
cleared up the earlier Pappaceras issue. Now the nesting is more in accord with consensus with regard to the indricotheres, but they’re still closer to 3-toed horses than to 3-toed rhinos.

Also note:
the <50 bootstrap scores surrounding Hyracotherum, a basal horse/rhino. I could only separate sister taxa by 2 steps. It takes at least 3 steps to bounce the Boostrap score over 50. Small steps. Small steps.