Brontotheriidae

Brontotheres Temporal range: Eocene
Megacerops skeleton at the National Museum of Natural History, Washington, DC
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Perissodactyla
Family:	†Brontotheriidae Marsh, 1873
Genera
See text
Synonyms
Menodontidae Titanotheriidae

Brontotheriidae 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 with some developing bony nose horns, and were some of the earliest mammals to have evolved large body sizes of several tonnes. They lived around 56–34 million years ago, until the very close of the Eocene. Brontotheres had a Holarctic distribution, with the exception of Western Europe: they occupied North America, Asia, and Eastern Europe. They were the first fossilized mammals to be discovered west of the Mississippi, and were first discovered in South Dakota.

Characteristics and evolution

Life reconstruction of Megacerops

Life reconstruction of Protitanops curryi

Skull of Rhinotitan

This group has also been referred to as "Titanotheres." "Titan" refers to the mythological Greek gods who were symbols of strength and large size, and "theros" is Greek for "wild animal." "Bronto" is Greek for "thunder," which may be how this group got the nickname "thunder beasts."

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 wear patterns observed on brontothere teeth suggests a folivorous diet. Early Brontotheres had brachydont teeth with thick enamel, while later forms evolved a more hypsodont style tooth with thinner enamel.

Brontotheres also shared an elongated postorbital cranium, meaning that their skulls are lengthened between their eyes and ears. They also had anteroposteriorly abbreviated (shortened) faces.

The evolutionary history of this group is well known due to an excellent fossil record in North America. The earliest stem-brontotheres, had an estimated body mass of only 18 kilograms (40 lb)

The earliest brontotheres, such as Eotitanops, were rather small, no more than a meter in height, and hornless.

Brontotheres evolved massive bodies, with some species standing over 2.5 meters (7 feet) tall, with body masses of over a tonne, perhaps exceeding 4,000 kilograms (8,800 lb), in large individuals of Megacerops, although some small species such as Nanotitanops did persist through the Eocene. Some genera, such as Dolichorhinus, evolved highly elongated skulls. Some later brontotheres developed horn-like bony projections of the skull. The North American brontothere Megacerops, for example, evolved large sexually dimorphic paired horns above their noses. The sexually dimorphic horns, along with highly developed neck musculature, suggest that brontotheres were highly gregarious (social) and males may have performed some sort of head-clashing behavior in competition for mates. Females had smaller appendages, which may have been used to ward off predators and protect young. In Asia, another species of brontothere, Embolotherium, evolved a similarly gigantic body size; however, instead of the slingshot-like horns of the Megacerops, they evolved a single elongated bony process that was composed of both nasal and frontal bones. Embolotherium may have used its large nasal cavity to make vocalizations in order to communicate with others of its species. Unlike rhinoceros, in which the horns are made of keratin, however, 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. Similarly to Giraffes, their horns were covered in skin and did not have grooves for nutrient blood vessels. There is some evidence of secondary bone growth, likely due to impact from head clashing.

Brontotheres had likely adapted to the warmer and more humid climates of the Eocene, and probably became extinct because they could not adapt to the drier conditions and more open landscapes of the Oligocene.

Discovery

Brontothere model reconstruction (Parvicornis occidentalis) at the San Diego Natural History Museum

Brontotheres were one of the first fossilized mammals to be discovered west of the Mississippi, with the first fossil being found in 1846 in the Badlands, South Dakota. Joseph Leidy was the first researcher to scientifically describe brontothere fossils, followed by Cope and Marsh, who studied skulls and entire skeletons. Marsh came up with the term "Brontotheridae," identified them as odd-toed ungulates, and identified distinguishing characteristics of the group. Brontotheriidae fossils have been found in eastern Europe, eastern Russia, Kazakstan, Pakistan, southeast Asia, Korea, Japan, the southeastern U.S., and Canada.

Classification

Brontotheres are an early diverging clade within Perissodactyla. Although historically suggested to be closely related to horses, phylogenetic analyses have recovered them to lie outside the clade containing chalicotheres, rhinoceroses, tapirs and horses, or more closely related to chalicotheres, rhinoceroses and tapirs than to horses.

Classification for Brontotheriidae after Mihlbachler 2008 and Mader 2010

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.

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 India, 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 India, 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

Further reading

• 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. (eds.). Paleogene Mammals. Bulletin 26. New Mexico Museum of Natural History and Science. pp. 129–136. Retrieved 29 October 2013.
• 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 29 October 2013.

External links

• Brontotheroidea at Mikko's Phylogeny Archive