Brontotheriidae







































Brontotheres
Temporal range: 56–34 Ma

PreЄ

Є

O

S

D

C

P

T

J

K

Pg

N








Eocene


Brontotherium hatcheri.jpg

Brontotherium hatcheri skeleton at the
National Museum of Natural History, Washington, DC

Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Order:
Perissodactyla
Suborder:
Hippomorpha
Family:
†Brontotheriidae



Marsh, 1873
Genera

See text



Synonyms


  • Menodontidae

  • Titanotheriidae



Brontotheriidae, also called Titanotheriidae, is a family of extinct mammals belonging to the order Perissodactyla, the order that includes horses, rhinoceroses, and tapirs. Superficially, they looked rather like rhinos, although they were actually more closely related to horses; Equidae and Brontotheriidae make up the suborder Hippomorpha. They lived around 56–34 million years ago, until the very close of the Eocene.




Contents






  • 1 Characteristics and evolution


  • 2 Classification of Brontotheres


  • 3 Notes


  • 4 References


  • 5 External links





Characteristics and evolution





Megacerops




Skull of Rhinotitan


Brontotheres retain four toes on their front feet and three toes on their hind feet. Their teeth are adapted to shearing (cutting) relatively nonabrasive vegetation. Their molars have a characteristic W-shaped ectoloph (outer shearing blade).


The evolutionary history of this group is well known, due to an excellent fossil record in North America.[1] The earliest brontotheres, such as Eotitanops, were rather small, no more than a meter in height, and were hornless.


Brontotheres, over time, evolved massive body sizes, although some small species, such as Nanotitanops, did persist through the Eocene. Some genera, such as Dolichorhinus, evolved highly elongated skulls. Later brontotheres were massive in size, up to 2.5 m (8.2 ft) in height with bizarre horn-like skull appendages. For instance the North American brontothere Megacerops evolved large sexually dimorphic paired horns above their noses. The sexually dimorphic horns suggest that brontotheres were highly gregarious (social) and males may have performed some sort of head-clashing behavior in competition for mates. However, unlike rhinos, the horns of brontotheres are composed of bone, the frontal bone and nasal bone, and were placed side-to-side rather than front-to-back.


Brontotheres probably became extinct due to an inability to adapt to drier conditions and tougher vegetation (such as grasses) that spread during the Oligocene.[1]



Classification of Brontotheres





Classification for Brontotheriidae after Mihlbachler 2008[2] And Mader 2010[3]

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 Brontotheriidae 












 

 Eotitanopinae
 
 












 

 Palaeosyopinae
 
 












 

 Brontotheriinae
 
 












 

 Brontotheriini
 
 












 

 Rhadinorhinina
 
 












 

 Telmatheriina
 
 












 

 Brontotheriina
 
 












 

 Brontotheriita
 
 

 Embolotheriita
 
 
 
 
 
 
 
 
 

Two classification systems for Brontotheriidae are presented below. The first contains 43 genera and 8 subfamilies, and although it is based on a 1997 publication by McKenna and Bell, it summarizes research that was conducted before 1920 and is badly outdated. The second classification is based on 2004 and 2005 research by Mihlbachler et al., which indicates that many of the previous subfamily names are invalid. Several more recently discovered brontotheres are included in the newer classification.




Life reconstruction of Protitanops curryi


Although Lambdotherium and Xenicohippus were previously included in Brontotheriidae, they are no longer considered members of this family. Lambdotherium, though excluded, may be the closest known relative to brontotheres. Xenicohippus is now thought to be an early member of the horse family, Equidae.












Old Classification (summarized by McKenna and Bell, 1997)
 New classification (Mihlbachler et al., 2004a, 2004b; Mihlbachler, 2005)


  • Family Brontotheriidae


    • Pakotitanops incertae sedis, from Pakistan


    • Nanotitanops incertae sedis, from Asia

      • Subfamily Lambdotheriinae


        • Lambdotherium, from North America


        • Xenicohippus, from North America



      • Subfamily Palaeosyopinae


        • Palaeosyops (including Eotitanops), from North America, 0.5 m tall


        • Mulkrajanops, from Pakistan, 1.25 m tall



      • Subfamily Dolichorhininae


        • Metarhinus, from North America, 1 m tall


        • Sphenocoelus, from North America, 1.25 m tall


        • Mesatirhinus, from North America, 1 m tall



      • Subfamily Brontotheriinae


        • Duchesneodus, from North America


        • Brontotherium, from North America


        • Megacerops, from North America, 2.5 m tall



      • Subfamily Embolotheriinae


        • Titanodectes, from Asia


        • Embolotherium, from Mongolia, 2.5 m tall


        • Protembolotherium, from Outer Mongolia, 2 m tall



      • Subfamily Brontopinae


        • Brachydiastematherium, from Eastern Europe, 2 m tall


        • Pachytitan, from Inner Mongolia, 2 m tall


        • Dianotitan, from China, 2 m tall


        • Gnathotitan, from Inner Mongolia, 2.5 m tall


        • Microtitan, from Inner Mongolia, 0.75 m tall


        • Epimanteoceras, from Inner Mongolia, 2 m tall


        • Protitan, from Inner Mongolia, 2 m tall


        • Rhinotitan, from Inner Mongolia, 2.5 m tall


        • Metatitan, from Mongolia, 1.5 m tall


        • Protitanotherium, from North America, 2 m tall


        • Parabrontops, from Mongolia, 2 m tall


        • Oreinotherium, from North America


        • Brontops, from North America


        • Protitanops, from North America, 2 m tall


        • Pygmaetitan, from China, 0.5 m tall



      • Subfamily Telmatheriinae


        • Acrotitan, from Inner Mongolia, 0.3 m tall


        • Desmatotitan, from Inner Mongolia, 1.25 m tall


        • Arctotitan, from China


        • Hyotitan, from Inner Mongolia, 2.2 m tall


        • Sthenodectes, from North America, 1.25 m tall


        • Telmatherium (including Metatelmatherium), from North America and Inner Mongolia, 1.5 m tall


        • Sivatitanops, from Asia and Europe



      • Subfamily Menodontinae


        • Diplacodon, from North America, 2 m tall


        • Eotitanotherium, from North America


        • Notiotitanops, from North America, 2 m tall


        • Menodus, from Europe and North America


        • Ateleodon, from North America

  • Family Brontotheriidae


    • Pakotitanops incertae sedis, from Pakistan


    • Mulkrajanops incertae sedis, from Pakistan, 1.25 m tall


    • Eotitanops, from North America, 0.5 m tall


    • Palaeosyops, from North America, 1 m tall

    • Subfamily Brontotheriinae


      • Bunobrontops, from Asia


      • Mesatirhinus, from North America, 1 m tall


      • Dolichorhinus, from North America, 1.25 m tall


      • Sphenocoelus, from North America, 1.25 m tall


      • Desmatotitan, from Inner Mongolia, 1.25 m tall


      • Fossendorhinus, from North America


      • Metarhinus, from North America, 1 m tall


      • Microtitan, from Inner Mongolia, 0.75 m tall


      • Sthenodectes, from North America, 1.25 m tall


      • Telmatherium, from North America, 1.25 m tall


      • Metatelmatherium, from North America and Inner Mongolia, 1.25 m tall


      • Epimanteoceras, from Inner Mongolia, 2 m tall


      • Hyotitan incertae sedis, from Inner Mongolia, 2.2 m tall


      • Nanotitanops incertae sedis, from Asia


      • Pygmaetitan incertae sedis, from China, 0.5 m tall


      • Acrotitan incertae sedis, from Inner Mongolia, 0.3 m tall


      • Arctotitan incertae sedis, from China


      • Qufutitan incertae sedis, from China

      • Tribe Brontotheriini


        • Protitan, from Inner Mongolia, 2 m tall


        • Protitanotherium, from North America, 2 m tall


        • Rhinotitan, from Inner Mongolia, 2.5 m tall


        • Diplacodon (including Eotitanotherium), from North America, 2 m tall


        • Pachytitan, from Inner Mongolia, 2 m tall


        • Brachydiastematherium, from Eastern Europe, 2 m tall


        • Sivatitanops, from Asia and Europe

        • Subtribe Embolotheriina


          • Gnathotitan, from Inner Mongolia, 2.5 m tall


          • Aktautitan, from Kazakhstan, 2.5 m tall


          • Metatitan, from Mongolia, 1.5 m tall


          • Nasamplus, from Inner Mongolia


          • Protembolotherium, from Outer Mongolia, 2 m tall


          • Embolotherium (including Titanodectes), from Mongolia, 2.5 m tall



        • Subtribe Brontotheriina


          • Parabrontops, from Mongolia, 2 m tall


          • Protitanops, from North America, 2 m tall


          • Notiotitanops, from North America, 2 m tall


          • Dianotitan, from China, 2 m tall


          • Duchesneodus, from North America


          • Megacerops (including Menodus, Brontotherium, Brontops, Menops, Titanotherium, Ateleodon, and Oreinotherium), from North America, 2.5 m tall


Notes





  1. ^ ab Titanotherium


  2. ^ Mihlbachler, Matthew C. "Species taxonomy, phylogeny and biogeography of the Brontotheriidae (Mammalia, Perissodactyla) ; Bulletin of the American Museum of Natural History, no. 311". American Museum of Natural History. Retrieved 4 September 2013..mw-parser-output cite.citation{font-style:inherit}.mw-parser-output q{quotes:"""""""'""'"}.mw-parser-output code.cs1-code{color:inherit;background:inherit;border:inherit;padding:inherit}.mw-parser-output .cs1-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Lock-green.svg/9px-Lock-green.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-limited a,.mw-parser-output .cs1-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Lock-gray-alt-2.svg/9px-Lock-gray-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Lock-red-alt-2.svg/9px-Lock-red-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration{color:#555}.mw-parser-output .cs1-subscription span,.mw-parser-output .cs1-registration span{border-bottom:1px dotted;cursor:help}.mw-parser-output .cs1-hidden-error{display:none;font-size:100%}.mw-parser-output .cs1-visible-error{font-size:100%}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration,.mw-parser-output .cs1-format{font-size:95%}.mw-parser-output .cs1-kern-left,.mw-parser-output .cs1-kern-wl-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right,.mw-parser-output .cs1-kern-wl-right{padding-right:0.2em}


  3. ^ Mader, Bryn J. "A species-level revision of the North American brontotheres Eotitanops and Palaeosyops (Mammalia, Perissodactyla)" (PDF). Department of Biological Sciences and Geology, Queensborough Community College. Retrieved 4 September 2013.




References


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  • McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals Above the Species Level. New York: Columbia University Press. ISBN 0231110138. OCLC 37345734.


  • Mihlbachler, M.C. (2004). Phylogenetic Systematics of the Brontotheriidae (Mammalia, Perissodactyla) (Ph.D. thesis). Columbia University.


  • Mihlbachler, M.C.; Lucas, S.G.; Emry, R.J. (2004a). "The holotype specimen of Menodus giganteus, and the 'insoluble' problem of Chadronian brontothere taxonomy". In Lucas, S.G.; Zeigler, K.; Kondrashov, P.E. Paleogene Mammals. Bulletin 26. New Mexico Museum of Natural History and Science. pp. 129–136. Retrieved October 2013. Check date values in: |accessdate= (help)


  • Mihlbachler, M.C.; Lucas, S.G.; Emry, R.J.; Bayshashov, B. (2004b). "A new brontothere (Brontotheriidae, Perissodactyla, Mammalia) from the Eocene of the Ily Basin of Kazakhstan and a phylogeny of Asian "horned" brontotheres" (PDF). American Museum Novitates. 3439: 1–43. doi:10.1206/0003-0082(2004)439<0001:ANBBPM>2.0.CO;2. Retrieved October 2013. Check date values in: |accessdate= (help)




External links



  • Brontotheroidea at Mikko's Phylogeny Archive









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