Praemegaceros cazioti
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''Praemegaceros'' 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
deer A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) ...
, known from the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
and
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
of Western Eurasia. ''Praemegaceros'' is considered to be a genus of "giant deer", with many species having an estimated body mass of around , considerably larger than most living deer. The genus contains the
subgenera In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
''Praemegaceros,'' ''Orthogonoceros'' and ''Nesoleipoceros''. It has sometimes been synonymised with ''
Megaloceros ''Megaloceros'' (from Greek: + , literally "Great Horn"; see also Lister (1987)) is an extinct genus of deer whose members lived throughout Eurasia from the Pleistocene to the early Holocene. The type and only undisputed member of the genus ...
'' and ''
Megaceroides ''Megaceroides algericus'' is an extinct species of deer known from the Late Pleistocene to the Holocene of North Africa. It is one of only two species of deer known to have been native to the African continent, alongside the Barbary stag, a sub ...
'', but is regarded as a distinct genus by most studies. Some authors have considered the genus closely related to ''Megaloceros'', but this has been disputed by others. The earliest species like ''P. obscurus'' and ''P. verticornis'' appeared in Europe between 2 and 1.5 million years ago. The genus was widely distributed across Europe, West and Central Asia during the Early-
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
, with fossils having been discovered in France, Georgia, Germany, England, Greece, Israel, Italy, Romania, Russia Spain, Syria, and Tajikistan. The genus was extinct in mainland Europe and Asia by end of the Middle Pleistocene. An insular species, ''P. cazioti'' survived into the Late Pleistocene and Holocene in isolation on the
Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...
-
Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...
archipelago until around 5500 BCE.


Taxonomy

''Praemegaceros'' was first described as a subgenus of ''Cervus'' by Portis in 1920, to include ''Cervus'' (''Praemegaceros'') ''dawkinsi.'' Kalkhe in 1956 named ''Orthogonoceros'' with the type species of ''Orthogonoceros'' (previously ''Cervus'') ''verticornis.'' Kalkhe in 1965 recognised that these genera were synonyms, with ''Praemegaceros'' having priority. Radulesco & Samson in 1967 designated ''P.'' ''dawkinsi'' as the type species of the genus, while also naming the new genera ''Allocaenelaphus, Psecupsoceros,'' and ''Nesoleipoceros'' which are now recognised as synonyms of the genus. Some authors have used the genus "''Megaceroides''" to include the species usually placed in ''Praemegaceros'', but the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of ''Megaceroides'', the North African ''
Megaceroides algericus ''Megaceroides algericus'' is an extinct species of deer known from the Late Pleistocene to the Holocene of North Africa. It is one of only two species of deer known to have been native to the African continent, alongside the Barbary stag, a subs ...
'' is morphologically distinct from ''Praemegaceros''. ''Praemegaceros'' is widely agreed to be belong to the subfamily
Cervinae The Cervinae or the Old World deer, are a subfamily of deer. Alternatively, they are known as the plesiometacarpal deer, due to having lost the parts of the second and fifth metacarpal bones closest to the foot (though retaining the parts away fr ...
. ''Praemegaceros'' is considered a genus of "giant deer", and has often been placed in the tribe Megacerini, alongside genera like ''
Megaloceros ''Megaloceros'' (from Greek: + , literally "Great Horn"; see also Lister (1987)) is an extinct genus of deer whose members lived throughout Eurasia from the Pleistocene to the early Holocene. The type and only undisputed member of the genus ...
'' and ''
Sinomegaceros ''Sinomegaceros'' is an extinct genus of deer known from the Late Pliocene/Early Pleistocene to Late Pleistocene of Central and East Asia. It is considered to be part of the group of "giant deer" (often referred to collectively as members of the ...
,'' though other authors have disputed the close relationship between ''Praemegaceros'' and ''Megaloceros'', alternatively suggesting that the two genera obtained their large size independently, and instead propose a close relationship of ''Praemegaceros'' with the genus ''
Eucladoceros ''Eucladoceros'' (Greek language, Greek for "well-branched antler") is an extinct genus of large deer whose fossils have been discovered across Eurasia, from Europe to China, spanning from the Late Pliocene-Early Pleistocene. It is noted for i ...
''. Classification according to Croitor, 2018.


Subgenus ''Orthogonoceros''


''Praemegaceros pliotarandoides''

Known from the late Early Pleistocene and Middle Pleistocene of Italy, North Greece, Moldova, South Ukraine and the Azov Sea Region. It was named by De Alessandri in 1903 for remains found in North Italy. ''Psekupsoceros orientalis'' is a junior synonym of the taxon. Estimated body mass of around .


''Praemegaceros verticornis''

Named by William Boyd Dawkins in 1872 on the basis of a partial antler from the Cromer Forest Bed. Croitor suggests that ''Cervus belgrandi'' is a junior synonym. Known from late Early Pleistocene to Middle Pleistocene localities in England, Spain, France, Germany and Italy. Estimated body mass of around . Isotopic analysis of specimens from the Early Pleistocene (about 1.6 million years ago) Venta Micena locality of southern Spain, suggests that at this locality the species was almost exclusively
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 ...
(feeding on leaves and twigs of high growing plants like shrubs and trees) in forested environments.


Subgenus ''Nesoleipoceros''


''Praemegaceros solilhacus''

The species was named by Robert in 1930 for remains from Soleilhac, an early Middle Pleistocene site located in the Massif Central, France. The neotype specimen consists of a partial left frontal with an attached partial antler. The morphology of the preserved antler strongly resembles that of ''P. cazioti,'' which suggests a close relationship. Other known sites are from the early Middle Pleistocene of France, Germany, Italy, South Russia and Moldova. This taxon is suggested to be the largest species of ''Praemegaceros'', with an estimated mass of . The taxon appears to have been extinct by the late Middle Pleistocene.


''Praemegaceros sardous–Praemegaceros cazioti''

Endemic to the
Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...
-
Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...
archipelago from the late Middle Pleistocene to the Early Holocene, the ancestor of ''P. cazioti'' appears to have dispersed to the islands during the Middle Pleistocene, with the earliest well dated records of ''Praemegaceros'' at Su Fossu de Cannas in Sardinia being over 450,000 years in age. Other early remains of the genus in Sardinia are referred to the species ''P. sardus/sardous''. The oldest remains assigned to ''P. cazioti'' date to approximately 300,000 years ago. ''P. cazioti'' is smaller than ''P. sardus'', being slightly larger than a
fallow deer Fallow deer is the common name for species of deer in the genus ''Dama'' of subfamily Cervinae. There are two living species, the European fallow deer (''Dama dama''), native to Europe and Anatolia, and the Persian fallow deer (''Dama mesopotamic ...
, with an estimated body mass of around , exhibiting an over 75% size reduction from its presumed mainland ancestor ''Praemegaceros solilhacus,'' an example of
insular dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
. The cranial morphology appears to be unaffected by the insular dwarfism. Two chronologically separated subspecies are known, which are suggested to be
chronospecies A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...
. ''P. cazioti cazioti'' dates to the late Middle Pleistocene and Late Pleistocene and is characterised by smaller brachyodont teeth and ''P. cazioti algarensis'' from the end of the Late Pleistocene is larger than ''P. cazioti cazioti'' and is characterised by large mesodont upper cheek teeth, and a long premolar series, which are suggested to be adaptions for a grazing diet. The youngest date for ''P. cazioti'' is around 5500 BCE, making it the latest surviving ''Praemegaceros'' species.


Subgenus ''Praemegaceros''


''Praemegaceros obscurus''

Known from the late
Villafranchian Villafranchian age ( ) is a period of geologic time (3.5–1.0 Ma) spanning the Late Pliocene and Early Pleistocene used more specifically with European Land Mammal Ages. Named by Italian geologist Lorenzo Pareto for a sequence of terrestrial se ...
of Europe (including Central Italy, Central Romania and Moldova) and the Near East, extending from the
Cromer Forest Bed The Cromer Forest-bed Formation, sometimes known as the Cromer Forest Bed, is a geological formation in Norfolk, England. It consists of river gravels, estuary and floodplain sediments predominantly silt, sand, and muds as well as peat along the ...
in England to Ubeidiya, Israel,
Dmanisi Dmanisi ( ka, დმანისი, tr, , ) is a town and archaeological site in the Kvemo Kartli region of Georgia (country), Georgia approximately 93 km southwest of the nation’s capital Tbilisi in the river valley of Mashavera. Abandoned i ...
, Georgia and the Azov region, Russia. Croitor suggests an origin in South Asia for this species, descended from taxa possibly referrable to '' Panolia sp.'' from the
Siwaliks The Sivalik Hills, also known as Churia Hills, are a mountain range of the outer Himalayas. The literal translation of "Sivalik" is 'tresses of Shiva'. The hills are known for their numerous fossils, and are also home to the Soanian Middle Pale ...
. Estimated body mass of around .


''Praemegaceros dawkinsi''

Named in 1882 from remains from the
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
Cromer Forest Bed in
Norfolk Norfolk ( ) is a Ceremonial counties of England, ceremonial county in England, located in East Anglia and officially part of the East of England region. It borders Lincolnshire and The Wash to the north-west, the North Sea to the north and eas ...
by
Edwin Tulley Newton Edwin Tulley Newton (4 May 1840 – 28 January 1930) was a British paleontologist. Newton originally worked at handicrafts, but was able to attend Thomas Henry Huxley's lectures and by 1865, was appointed as his assistant. In 1882, he beca ...
which were originally described in 1872 as belonging to ''P. verticornis'' by
William Boyd Dawkins Sir William Boyd Dawkins (26 December 183715 January 1929) was a British geologist and archaeologist. He was a member of the Geological Survey of Great Britain, Curator of the Manchester Museum and Professor of Geology at Owens College, M ...
. The body mass is estimated to be around , with a mesodont dentition. Croitor suggests that because the pedicles (base of the antlers) are robust and similar to those of giant deer, that ''P. dawkinsi'' represents a dwarfed form. The species is considered to be endemic to Britain, with the antler morphology simplified relative to other ''Praemegaceros'' species.


''Praemegaceros mosbachensis''

Named by Wolfgang Soergel in 1927 for the species found in the lowest level of the Middle Pleistocene Mosbach locality in Germany. It was included in the “verticornis” group of Azzaroli's 1953 classification. It has been suggested to a be synonym of ''P. verticornis.'' However, Croitor suggests that these similarities are the result of
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and pa ...
, and proposes that ''P. mosbachensis'' represents an intermediate form between ''P. obscurus'' and ''P. dawkinsi.''


Ecology

During the Early-Middle Pleistocene, mainland species of ''Praemegaceros'' are suggested to have been preyed upon by the sabertooth cats ''
Megantereon ''Megantereon'' is an extinct genus of prehistoric machairodontine saber-toothed cat that lived in Eurasia, Africa and possibly North America from the late Pliocene to the Middle Pleistocene. It is a member of the tribe Smilodontini, and close ...
'' and ''
Homotherium ''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
'', and the "European jaguar" ''
Panthera gombaszoegensis ''Panthera gombaszoegensis'', also known as the European jaguar, is a ''Panthera'' species that lived from about 2.0 to 0.3 million years ago in Europe, as well as likely elsewhere in Eurasia. The first fossils were Excavation (archaeology), exca ...
''.''''


References

{{Taxonbar, from=Q25095511 Prehistoric deer Pleistocene Artiodactyla Pleistocene genus extinctions Piacenzian first appearances Pliocene Artiodactyla Prehistoric Artiodactyla genera