''Pappaceras'' is an
extinct
Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of rhinocerotoids from the
Early Eocene
In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
of
Asia
Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
belonging to
Paraceratheriidae
Paraceratheriidae is an extinct family of long-limbed, hornless rhinocerotoids, native to Asia and Eastern Europe that originated in the Eocene epoch and lived until the end of the Oligocene. They represent some of the largest terrestrial mammals ...
.
Taxonomy
In
1963
Events January
* January 1 – Bogle–Chandler case: Commonwealth Scientific and Industrial Research Organisation scientist Dr. Gilbert Bogle and Mrs. Margaret Chandler are found dead (presumed poisoned), in bushland near the Lane Cove ...
, material including a partial skull containing cheek teeth was unearthed in Late Eocene deposits of Mongolia. These remains were identified as from a true rhinoceros by Wood, who found them an important discovery with the scant amount of previous cranial material of early rhinocerotids available. On July 25, the same year, a paper was published by Wood concerning the taxonomy and osteology of these remains, in which he named them a new genus and species (or binomial) as well as re-ranking a previously named family as a subfamily containing the new taxon. The binomial created was ''Pappaceras confluens'', classified as a close relative of ''Forstercooperia'' within Forstercooperiinae (before Forstercooperiidae, named in
1940
A calendar from 1940 according to the Gregorian calendar, factoring in the dates of Easter and related holidays, cannot be used again until the year 5280.
Events
Below, events related to World War II have the "WWII" prefix.
January
*Janu ...
by Kretzoi). Wood noted that the generic name is derived from the
Latin
Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word ''πaππos'', "grandfather", and the
Greek
Greek may refer to:
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group
*Greek language, a branch of the Indo-European language family
**Proto-Greek language, the assumed last common ancestor of all kno ...
words ''alpha'', "without", and ''keras'', "horn", translating as "Grandfather without horn". The species name is based on the confluent morphology of the teeth. The
catalogue number for the skull is AMNH 26660, and it specifically preserved a "front half of the skull and a complete lower jaw, with most of the teeth and remaining
alveoli
Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit.
Uses in anatomy and zoology
* Pulmonary alveolus, an air sac in the lungs
** Alveolar cell or pneumocyte
** Alveolar duct
** Alveolar macrophage
* M ...
, totaling a full placental
series
Series may refer to:
People with the name
* Caroline Series (born 1951), English mathematician, daughter of George Series
* George Series (1920–1995), English physicist
Arts, entertainment, and media
Music
* Series, the ordered sets used i ...
". Other remains included a portion of the mandible and a premolar. All of these specimens were from the lame locality, the Upper Gray Clays, of the Irdin Manha Formation in
Inner Mongolia
Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an Autonomous regions of China, autonomous region of China. Its border includes two-thirds of the length of China's China–Mongolia border, border with the country of Mongolia. ...
.
In the revision by Radnisky, it was found that this species was assignable to ''Forstercooperia'', and the new combination ''F. confluens'' was erected.
This species is well known, although in the 1981 review of ''Forstercooperia'', it was synonymized with ''F. grandis''.
In the 1960s, newly uncovered material from the Arshanto Formation was identified as belonging to a new species of rhinocerotoid. Originally, they were found to be from ''F. confluens'', as they were in the same location as that species holotype. They were later assigned to ''Forstercooperia'' sp., with no new name being given. The material included an almost complete skull, an almost complete lower jaw, an anterior portion of the skull, and an
astragalus
Astragalus may refer to:
* ''Astragalus'' (plant), a large genus of herbs and small shrubs
*Astragalus (bone)
The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
. These bones were first assigned a new species by Lucas ''et al.'', ''Forstercooperia minuta''. They were found to be a unique species based on their size and the anatomy of their teeth. The species has been retained in the species complex of ''Forstercooperia'' throughout major revisions, by Lucas ''et al.'' in 1981,
Lucas and Sobus in
1989
1989 was a turning point in political history with the "Revolutions of 1989" which ended communism in Eastern Bloc of Europe, starting in Poland and Hungary, with experiments in power-sharing coming to a head with the opening of the Berlin W ...
,
and Holbrook and Lucas in
1997
Events January
* January 1 – The Emergency Alert System is introduced in the United States.
* January 11 – Turkey threatens Cyprus on account of a deal to buy Russian S-300 missiles, prompting the Cypriot Missile Crisis.
* January 1 ...
.
In
1977
Events January
* January 8 – 1977 Moscow bombings, Three bombs explode in Moscow within 37 minutes, killing seven. The bombings are attributed to an Armenian separatist group.
* January 10 – Mount Nyiragongo erupts in eastern Zaire (no ...
, some the first description of a dentary from Kazakhstan's
Sargamys Formation was published. Authored by Gabunia, the paper figured the dentary, as well as some other material. In the images caption, the dentary was assigned to as ''Forstercooperia'' sp. although the text used a different name. In the text, the dentary and its teeth were assigned to ''Forstercooperia crudus'', although no size was mentioned. As the text did not have a description in it of ''F. crudus'', the name is now considered a ''
nomen nudum
In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
''. Its material is possibly assignable to ''F. minuta'', however.
In 1997, other material from Kazakhstan, specifically the
Kolpak Formation, was assigned to ''F. minuta'', meaning that it certainly lived in Kazakhstan at the same time as ''F. crudus''.
In a 1981 paper focusing purely on the genus ''Forstercooperia'', it was found that there was very little diversity in the species found valid by Radinsky. This paper, authored by
Spencer G. Lucas and
Robert Schoch and
Earl Manning and published in 1981 reviewed all currently-named species of ''Forstercooperia'', and named the new species ''F. minuta''. ''F. crudus'', named in 1977 by Gabunia, was found to be a ''nomen nudum'', and ''Pappaceras'' synonymized with ''Forsterocooperia''.
However, recent study has found ''Pappaceras'' to be generically distinct.
[Haibing Wang; Bin Bai; Jin Meng; Yuanqing Wang (2016). "Earliest known unequivocal rhinocerotoid sheds new light on the origin of Giant Rhinos and phylogeny of early rhinocerotoids". Scientific Reports. 6: Article number 39607. doi:10.1038/srep39607.]
Evolution
The
superfamily
SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Rhinocerotoidea
Rhinocerotoidea is a superfamily (taxonomy), superfamily of Perissodactyla, perissodactyls that appeared 56 million years ago in the Paleocene. They included four extinct families, the Amynodontidae, the Hyracodontidae, the Paraceratheriidae, an ...
can be traced back to the
early Eocene
In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
—about 50 million years ago—with early precursors such as ''Hyrachyus''. Rhinocerotoidea contains three families; the
Amynodontidae
Amynodontidae ("defensive tooth") is a family of extinct perissodactyls related to true rhinoceroses. They are commonly portrayed as semiaquatic hippo-like rhinos but this description only fits members of the Metamynodontini; other groups of ...
, the
Rhinocerotidae
A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
("true rhinoceroses"), and the Hyracodontidae. The diversity within the rhinoceros group was much larger in prehistoric times; sizes ranged from dog-sized to the size of Paraceratherium. There were long-legged,
cursorial
A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
forms and squat,
semi aquatic forms. Most species did not have horns. Rhinoceros fossils are identified as such mainly by characteristics of their teeth, which is the part of the animals most likely to be preserved. The upper molars of most rhinoceroses have a
pi (π) shaped pattern on the crown, and each lower molar has paired L-shapes. Various skull features are also used for identification of fossil rhinoceroses.
Description

''Pappaceras'' is known from a vast amount of cranial material, although only some scant postcranial remains. The largest species is ''P. confluens'', followed ''P. minuta''.
The average size of all species, is about equal with a large dog, even though later genera like ''Juxia'' reached the size of a modern horse and ''Paraceratherium'' exceeded the size of the largest African elephant.
Each species is distinguished by cheek tooth morphology, with the remaining skull quite similar.
Like primitive rhinocerotoids, ''Pappaceras'' possesses blunt ends on the tips of its nasals, above the nasal incision. Unlike all modern rhinoceroses, the nasals of ''Pappaceras'', as well as many related genera, lack rugosities, which suggests that they lacked any form of horn. The nasal incision extends fairly far into the upper jaw, ending just posterior to the canine. ''Pappaceras'' possesses a small post-incisor diastema, not as large as its descendants, and similar in size to that of ''Hyracodon''.
Distribution and habitat
Remains of ''Pappaceras'' have been found all across Asia. Most important remains are from the Early Eocene Arshanto Formation of Inner Mongolia (China).
In 1963, the species ''Pappaceras confluens'' was described from the same region, probably within the same formation.
''Pappaceras minuta'', is based on a maxilla from the Irdin Manha Formation as well.
''P. confluens'' is known from
Shara Murun Formation,
Ulan Shireh Formation and
Houldjin Formation, as well as the Arshanto Formation. ''P. minuta'' is known from specimens only from the Arshanto Formation, as is ''P. meiomenus'', and possibly the Shara Murun Formation of China,
and the Kolpak Formation of Kazakhstan,
References
{{Taxonbar, from=Q49002435
Paraceratheriidae
Eocene rhinoceroses
Prehistoric placental genera
Eocene mammals of Asia
Hyracodonts
Fossil taxa described in 1963