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I found this sentence in one paper. I think one, or even two commas should be there:

Their closest relatives, the zebras and horses are mammals.

or

Their closest relatives, the zebras and horses, are mammals.

"the zebras and horses" is the noun modifier, and should be delimited with commas. Am I right? Which variant is most correct? Do missing commas in general make sentences ungrammatical?

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I would call "zebras and horses" a parenthetical phrase (although I'm not sure of what type). You don't need the definite article (but you can use it if you like), and you are right about the commas:

Their closest relatives, zebras and horses, are mammals.

  • Alternatively, “their closest relatives” is a participial modifier with its copular participle implied, and the rest is a simple clause: “[Being] their closest relatives, zebras and horses are mammals”. That would make more sense with a disjunctive coordinator, though, like ‘though’. – Janus Bahs Jacquet Oct 26 '16 at 7:37

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