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In the broad phylogenetic usage, a clade including Cetacea together with the traditional even-toed ungulates. Whale ancestry is therefore artiodactyl ancestry, despite the loss of the adult terrestrial hoofed body plan.

A paleontologist who used ancient protein sequences to test evolution relationships of extinct South American ungulates, illustrating palaeoproteomics when recoverable DNA is unavailable.^[ref]

An ungulate is conventionally a hoofed mammal. The name has also been applied broadly to extinct mammals with superficially similar anatomy, so a historical grouping by body form need not specify one exact modern evolutionary lineage. Convergent traits can make fossil relationships difficult to infer.

Ancient-protein analysis tested the relationships of two enigmatic South American forms. Collagen sequences recovered from fossils were compared with sequences from living mammals, placing the sampled extinct taxa near the odd-toed ungulate lineage rather than the alternative placements suggested by some anatomical comparisons. ^[ref] Molecular preservation thereby supplied evidence complementary to shape, even where recoverable ancient DNA had been lacking.