Proterotheriidae is an extinct family of litoptern ungulates known from the Eocene-Late Pleistocene of South America. Members of the group were small to medium-sized cursorial herbivores with brachydont teeth, with their toes showing progressive reduction, with later members of the group bearing weight on a single large toe similar to living horses.

Description

Skeleton of Diadiaphorus

Two subfamilies and 18 genera of Proterotheriidae are known. All forms were small or medium-sized. Typical is a reduction of the number of toes and brachydont or mesodont teeth. The family is recorded since the late Palaeocene. Various fossils are known from many parts of the South American continent. The diversity decreased in the Miocene to Pliocene and it has been assumed for a long time that they entirely disappeared in the late Pliocene. However, fossils found in Argentina, Brazil and Uruguay show that one member of the group, Neolicaphrium recens survived into the Late Pleistocene.

Better known genera of the family include Diadiaphorus and Thoatherium from the Miocene.

Interpretation of Thoatherium

Taxonomy

Proterotheriidae is traditionally considered to include two subfamilies, Anisolambdinae and Proterotheriinae. Anisolambdinae (also called Anisolambdidae in some studies) was proposed to unite the primitive and earlier forms Anisolambda, Eolicaphrium, Paranisolambda, Protheosodon, Guilielmofloweria, Heteroglyphis, Lambdaconops, Wainka and Xesmodon. However, the phylogenetic analysis of McGrath and colleagues recovered the included genera to neither form their own clade, or to universally represent basal taxa outside the genera of Proterotheriinae per Soria, making Anisolambdinae a polyphyletic group of unrelated organisms.

Proterotheriidae was redefined by McGrath and colleagues in 2019 to be all taxa closer to Tetramerorhinus than Macrauchenia, Tricoelodus or Protolipterna. The cladogram below shows the modified results of their phylogenetic analysis, where incomplete taxa were placed based on morphology. The unrelated genera of the polyphyletic taxon Anisolambdinae or Anisolambdidae is highlighted in pink.

ProterotheriidaeMegadolodinae Megadolodus Bounodus Indaleciidae Indalecia Adiantoides Paramacrauchenia Anisolambda Lambdaconus suinus Lambdaconus inaequifacies Lambdaconus lacerum Paranisolambda Picturotherium Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Megadolodinae Megadolodus Bounodus
MegadolodinaeMegadolodus Bounodus
Megadolodus
Bounodus
Indaleciidae Indalecia Adiantoides Paramacrauchenia Anisolambda Lambdaconus suinus Lambdaconus inaequifacies Lambdaconus lacerum Paranisolambda Picturotherium Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Indaleciidae Indalecia Adiantoides
IndaleciidaeIndalecia Adiantoides
Indalecia
Adiantoides
Paramacrauchenia Anisolambda Lambdaconus suinus Lambdaconus inaequifacies Lambdaconus lacerum Paranisolambda Picturotherium Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Paramacrauchenia
Anisolambda Lambdaconus suinus Lambdaconus inaequifacies Lambdaconus lacerum Paranisolambda Picturotherium Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Anisolambda
Lambdaconus suinus Lambdaconus inaequifacies
Lambdaconus suinus
Lambdaconus inaequifacies
Lambdaconus lacerum Paranisolambda Picturotherium Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Lambdaconus lacerum
Paranisolambda Picturotherium Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Paranisolambda
Picturotherium Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Picturotherium
Neodolodus Protheosodon Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Neodolodus Protheosodon
Neodolodus
Protheosodon
Anisolophus floweri Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Anisolophus floweri
Anisolophus australis Anisolophus minusculus Prolicaphrium Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Anisolophus australis Anisolophus minusculus Prolicaphrium
Anisolophus australis
Anisolophus minusculus Prolicaphrium
Anisolophus minusculus
Prolicaphrium
Olisanophus Mesolicaphrium Diplasiotherium Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Olisanophus Mesolicaphrium Diplasiotherium
Olisanophus
Mesolicaphrium Diplasiotherium
Mesolicaphrium
Diplasiotherium
Villarroelia Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Villarroelia
Tetramerorhinus fleaglei Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Tetramerorhinus fleaglei
Tetramerorhinus prosistens Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Tetramerorhinus prosistens
Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Tetramerorhinus cingulatum Tetramerorhinus lucarius Tetramerorhinus mixtum
Tetramerorhinus cingulatum
Tetramerorhinus lucarius Tetramerorhinus mixtum
Tetramerorhinus lucarius
Tetramerorhinus mixtum
Diadiaphorus Thoatherium Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Diadiaphorus Thoatherium
Diadiaphorus
Thoatherium
Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Brachytherium Thoatheriopsis Proterotherium Neobrachytherium ameghinoi
Brachytherium
Thoatheriopsis Proterotherium Neobrachytherium ameghinoi
Thoatheriopsis
Proterotherium Neobrachytherium ameghinoi
Proterotherium
Neobrachytherium ameghinoi
Eoauchenia Neobrachytherium ullumense Neobrachytherium morenoi Neobrachytherium intermedium Neolicaphrium Epitherium Uruguayodon
Eoauchenia
Neobrachytherium ullumense
Neobrachytherium morenoi Neobrachytherium intermedium
Neobrachytherium morenoi
Neobrachytherium intermedium
Neolicaphrium Epitherium Uruguayodon
Neolicaphrium
Epitherium Uruguayodon
Epitherium
Uruguayodon

Palaeobiology

Locomotion

Based on the morphology of their hindlimbs, proterotheriids likely locomoted by making use of a rotary or half-bound gait, in contrast to the transverse gallop used by equids.