Resolution of the long-standing controversy over the type species of the genus Pseudotypotherium (Notoungulata,Typotheria, Mesotheriidae)

Mesotheres (Notoungulata: Typotheria) are among the most common mammals found in upper Miocene to Pliocene deposits of central Argentina, including the classic type Monte Hermoso locality, which defines the Montehermosan South American Land Mammal “Age”. Nevertheless, the correct name for the mesothere species from this site has been shrouded in uncertainty for well over a century due to questions of taxonomic priority, specimen provenance, and ontogenetic changes in dental formula. Since the mesotheres from Monte Hermoso were named, three distinct species have been formally considered as the type species of the genus: (1) Pseudotypotherium bravardi; (2) “Pseudotypotherium” maendrum; and (3) Pseudotypotherium exiguum. However, none of these species is a nominal species of the Pseudotypotherium genus; all three were originally referred to Typotherium. Article 67.2 of the International Code of Zoological Nomenclature (ICZN, 1999) indicates that only species considered as nominal species are eligible to set the type; in the case of Pseudotypotherium, these include: P. pulchrum, P. carlesi, P. hystatum, and P. carhuense. We conclude that Pseudotypotherium pulchrum F. Ameghino, 1904 (holotype MACN A 10299, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Ameghino Collection), is the type species of the mesotheriid notoungulate genus from Monte Hermoso. According to Article 68.2, F. Ameghino fixed the type by original designation in 1904 when he described P. pulchrum and included “n. g., n. sp.”. Two of the other species previously considered species P. (= T.) bravardi and P. (= T.) exiguum are invalid as type species according to Article 70.2, since their designations overlooked the previous type fixation. The third species (M. (= T.) maendrum) represents a different mesothere genus (Mesotherium) that only occurs in younger (Pleistocene) deposits. Our analysis puts an end to a historical debate that has been ongoing for more than a century regarding the identity of this well-represented late Miocene–Pliocene mesotheriine genus (Pseudotypotherium). This study provides a solid taxonomic foundation for future studies on intraspecific and ontogenetic variation of Pseudotypotherium pulchrum.