''Ceratotherium neumayri'' is an extinct species of rhinoceros from the

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

epoch (specifically the

Vallesian The Vallesian age is a period of geologic time (11.6–9.0 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Turolian age and follows the Astaracian age. The so-called Vallesian Crisis resulted in th ...

and

Turolian The Turolian age is a period of geologic time (9.0–5.3 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Ruscinian age and follows the Vallesian age. The Turolian overlaps the Tortonian and Messinia ...

European land mammal ages) of the

Balkans The Balkans ( , ), corresponding partially with the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throug ...

(including GreeceGiaourtsakis, I.X. (2003). Late Neogene Rhinocerotidae of Greece: distribution, diversity and stratigraphical range. ''Deinsea'', ''10''(1), 235–254. and BulgariaGeraads D, Spassov N. (2009). Rhinocerotidae (Mammalia) from the Late Miocene of Bulgaria. Palaeontographica A. 287:99–122.) and

Western Asia West Asia (also called Western Asia or Southwest Asia) is the westernmost region of Asia. As defined by most academics, UN bodies and other institutions, the subregion consists of Anatolia, the Arabian Peninsula, Iran, Mesopotamia, the Armenian ...

(including Iran and

Anatolia Anatolia (), also known as Asia Minor, is a peninsula in West Asia that makes up the majority of the land area of Turkey. It is the westernmost protrusion of Asia and is geographically bounded by the Mediterranean Sea to the south, the Aegean ...

in Turkey).

Taxonomy

The species was originally named ''Atelodus neumayri'' by

Henry Fairfield Osborn Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was professor of anatomy at Columbia University, president of the American Museum of Natural History for 25 y ...

in 1900.Osborn HF (1900) Phylogeny of the rhinoceroses of Europe. Bull Am Mus Natur Hist 12:229–267. It is considered part of the tribe Dicerotini (also spelled Diceroti) or subtribe Dicerotina, indicating a close relationship to the extant African rhinoceroses, the

black rhinoceros The black rhinoceros (''Diceros bicornis''), also called the black rhino or the hooked-lip rhinoceros, is a species of rhinoceros native to East Africa, East and Southern Africa, including Angola, Botswana, Eswatini, Kenya, Lesotho, Malawi, Moza ...

(''Diceros bicornis'') and the

white rhinoceros The white rhinoceros, also known as the white rhino or square-lipped rhinoceros (''Ceratotherium simum''), is the largest extant species of rhinoceros and the most Sociality, social of all rhino species, characterized by its wide mouth adapted f ...

(''Ceratotherium simum''). The generic assignment of ''neumayri'' is disputed, having been placed in both ''

Ceratotherium ''Ceratotherium'' (from Greek: ''keras'' κέρας "horn" and ''thērion'' θηρίον "beast") is a genus within the family Rhinocerotidae. It comprises one living species, the white rhinoceros (''Ceratotherium simum''), and several extinct fo ...

'' and ''

Diceros ''Diceros'' (Greek: "two" (dio), "horn" (keratos)) is a genus of rhinoceros containing the extant black rhinoceros ''(Diceros bicornis)'' and several extinct species. Taxonomy ''Diceros'' is more closely related to the genus ''Ceratotherium'' ( ...

'' by various authors. Some paleontologists have proposed it as a common ancestor to both ''Ceratotherium'' and ''Diceros'', while others suggest it represents an early, distinct evolutionary branch not directly ancestral to the modern African rhinos. A 2022 study placed the species in the separate

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...

genus ''Miodiceros''.

Description

The species was a large sized rhinceros, and had two horns, a nasal and a frontal horn. The

nasal septum The nasal septum () separates the left and right airways of the Human nose, nasal cavity, dividing the two nostrils. It is Depression (kinesiology), depressed by the depressor septi nasi muscle. Structure The fleshy external end of the nasal s ...

was not ossified.

Ecology

Analysis of

dental microwear Dental microwear analysis is a method to infer diet and behavior in extinct animals, especially in fossil specimens. It has been used on a variety of taxa, including hominids, victoriapithecids, amphicyonids, canids, ursids, hyaenids, hyaenodont ...

patterns on the teeth of ''C. neumayri'' suggests that it was a mixed feeder. This diet means it was adaptable, capable of both

grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to free range (roam around) and consume wild vegetations in order to feed conversion ratio, convert the otherwise indigestible (by human diges ...

on grasses and

browsing Browsing is a kind of orienting strategy. It is supposed to identify something of relevance for the browsing organism. In context of humans, it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing open sh ...

on leaves and twigs from shrubs and trees.

Discoveries

Fossils attributed to ''C. neumayri'' have been found across southeastern Europe and Western Asia. Fossils of the species have been found in the

, including

Bulgaria Bulgaria, officially the Republic of Bulgaria, is a country in Southeast Europe. It is situated on the eastern portion of the Balkans directly south of the Danube river and west of the Black Sea. Bulgaria is bordered by Greece and Turkey t ...

and

Greece Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...

(such as

Pikermi Pikermi () is a suburb of Athens and a former community of East Attica regional unit, Greece. Since the 2011 local government reform it is part of the municipality Rafina-Pikermi, of which it is a municipal unit. The municipal unit has an area of ...

Samos Samos (, also ; , ) is a Greek island in the eastern Aegean Sea, south of Chios, north of Patmos and the Dodecanese archipelago, and off the coast of western Turkey, from which it is separated by the Mycale Strait. It is also a separate reg ...

, and Axios Valley). Discoveries also extend into

and

northern Iran Northern Iran (), is a geographical term that refers to a relatively large and fertile area, consisting of the southern border of the Caspian Sea and the Alborz mountains. It includes the provinces of Gilan, Mazandaran, and Golestan (ancie ...

, with occurrences in the

southern Caucasus The South Caucasus, also known as Transcaucasia or the Transcaucasus, is a geographical region on the border of Eastern Europe and West Asia, straddling the southern Caucasus Mountains. The South Caucasus roughly corresponds to modern Armenia, ...

. In 2012, a well-preserved skull from Gülşehir, dating to around 9.2 million years ago was found. This individual is believed to have died due to extreme temperatures from a

pyroclastic flow A pyroclastic flow (also known as a pyroclastic density current or a pyroclastic cloud) is a fast-moving current of hot gas and volcanic matter (collectively known as tephra) that flows along the ground away from a volcano at average speeds of b ...

(ignimbrite) associated with a volcanic eruption. Some authors have suggested that the species was also present in Africa, based on

remains found in

Tunisia Tunisia, officially the Republic of Tunisia, is a country in the Maghreb region of North Africa. It is bordered by Algeria to the west and southwest, Libya to the southeast, and the Mediterranean Sea to the north and east. Tunisia also shares m ...

originally attributed to ''C. douariense''.Pandolfi (2018)
Evolutionary history of Rhinocerotina (Mammalia, Perissodactyla).
Fossilia, Volume 2018

References

Dicerotini Miocene rhinoceroses Miocene mammals of Asia Miocene mammals of Europe Fossil taxa described in 1900 Taxa named by Henry Fairfield Osborn {{paleo-oddtoedungulate-stub