Megaceroides
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''Megaceroides algericus'' is an extinct species of deer known from the Late Pleistocene to the
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
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 subspecies of
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or hart, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of we ...
. It is considered to be closely related to the giant deer species of Eurasia.


Taxonomy

The species was first described by
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
as ''Cervus algericus'' in 1890 from a maxilla with teeth found near Hammam Maskhoutine in Algeria. The species ''Cervus pachygenys'' was erected for a pachyostotic mandible and an isolated molar found in Algeria by Auguste Pomel in 1892. Léonce Joleaud in two publications in 1914 and 1916 synonymised the two species, and suggested affinities with the giant deer of Europe, and placed it in the newly erected subgenus ''Megaceroides'' within the genus ''Megaceros'' (junior synonym of ''
Megaloceros ''Megaloceros'' (from Greek: + , literally "Great Horn"; see also Lister (1987)) is an extinct genus of deer whose members lived throughout Eurasia from the early Pleistocene to the beginning of the Holocene and were important herbivores durin ...
'') as the
type species In 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 species that contains the biological type specime ...
. Camille Arambourg in publications in 1932 and 1938 raised ''Megaceroides'' to the full genus rank, and described additional cranial material, which were figured but not described in detail. In 1953 Augusto Azzaroli published a systematic taxonomy for ''Megaloceros'', he avoided using the name ''Megaceroides'', and suggested affinities to his proposed "''verticornis'' group" of ''Megaloceros'' species, and noted its similarities to '' Sinomegaceros pachyosteus.'' The taxonomic relationships of giant deer and their smaller insular relatives (often referred to collectively as members of the
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...
Megacerini, though it is not known whether the grouping is monophyletic) are unresolved, with a long and convoluted taxonomic history. The genus ''Megaceroides'' is also used as a synonym of '' Praemegaceros'' by some authors, reflecting to its unresolved taxonomic position with respect to other giant deer. It was considered to belong to ''Megaloceros'' by Hadjouis in 1990 with ''Megaceroides'' at subgenus rank. Affinities with ''Megaloceros'' have been suggested based on several morphological grounds. A comprehensive description of the taxon by Roman Croitor published in 2016 suggested that it originated from ''Megaloceros mugharensis'' from the Middle Pleistocene of the Levant, with Middle Pleistocene cervid remains from North Africa possibly belonging to an ancestor of the species, and retained ''Megaceroides'' at generic rank. The craniodental morphology of ''Megaceroides algericus'' suggests its phylogenetic relationship with Eurasian giant deer ''Megaloceros giganteus''.


Distribution

The species was found within the Mediterranean region of northwest Africa north of the Atlas Mountains, with 26 known localities within Algeria and Morocco, extending from Bizmoune cave near Essaouira in the west to Hammam Maskhoutine and Puits des Chaachas in the East. The oldest known remains of the species are around 24,000 years
Before Present Before Present (BP) years, or "years before present", is a time scale used mainly in archaeology, geology and other scientific disciplines to specify when events occurred relative to the origin of practical radiocarbon dating in the 1950s. Becau ...
. No other deer are known to have been native to the African continent aside from ''M. algericus'' and the Barbary stag, an extant subspecies of
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or hart, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of we ...
, which is also native to the same area of northwest Africa.


Description

The species is known from limited material, and knowledge of post-cranial remains and antlers are poor. The estimated size of the animal is smaller than a red deer but slightly larger than a
fallow deer ''Dama'' is a genus of deer in the subfamily Cervinae, commonly referred to as fallow deer. Name The name fallow is derived from the deer's pale brown colour. The Latin word ''dāma'' or ''damma'', used for roe deer, gazelles, and antelopes ...
. It is known from a mostly complete skull of an aged individual with worn teeth found near Aïn Bénian in Algeria, alongside other fragmentary crania. The skull is broad, but the length of the skull, specifically the splanchnocranium (the region of the skull behind the dental region) is relatively short. This combination of a wide but proportionally short skull is a morphology that is unknown in any other deer species. The two halves of the mandible are estimated to have a contact angle of 60 degrees, which is extremely wide in comparison to other deer, to compensate for the wide skull. The parietals are flat, with a straight facial profile. The eyes face further outward and less forward than ''Sinomegaceros pachyosteus.'' The skull and dentary exhibit extreme thickening (
pachyostosis Pachyostosis is a non-pathological condition in vertebrate animals in which the bones experience a thickening, generally caused by extra layers of lamellar bone. It often occurs together with bone densification (osteosclerosis), reducing inner ca ...
) somewhat similar to that in ''Megaloceros,'' unlike ''Megaloceros'', the vomer is largely unaffected. The pachyostosis is among the most extreme of any known mammal. The upper canines are absent (there are no sockets for them present on the Aïn Bénian skull) and the lower fourth premolar (P4) is molarised. The preserved proximal portion of the antler is straight and cylindrical in cross section, and orientated anteriorly, laterally and slightly dorsally, the antler becomes flattened distally. An isolated radius suggested to belong to the taxon is known from
Berrouaghia Berrouaghia is a town and commune in Médéa Province, Algeria. According to the 1998 census it has a population of 58,780. Notable people * Benyoucef Benkhedda - Algerian politician History Berrouaghia was during the Roman Empire called Tanara ...
, which is robust and proportionally short in comparison to other cervids, and the mid shaft measurement of 40mm is proportionally wider than that of '' M. giganteus.'' Croitor suggests that several rock art drawings from the Altas and the Sahara depict the taxon, which show horned animals, some with antler like
tines Tines (; also spelled tynes), prongs or teeth are parallel or branching spikes forming parts of a tool or natural object. They are used to spear, hook, move or otherwise act on other objects. They may be made of metal, wood, bone or other hard, ...
including proportionally long tails. This interpretation, which had previously been suggested by other authors, has been criticised, noting that there are no known remains for the taxon from the Sahara, and that previous interpretations of the rock art representing deer had been based on faulty fossil identification of deer in Mali.


Ecology

The ecology of the species is unclear owing to the lack of living analogues for its unique morphology. On several morphological grounds, Croitor proposed that its habits were peri- or
semiaquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semi aquatic animals include: * Ve ...
, with the weak mastication ability and polished cheek teeth by attrition suggesting a preference for soft water plants, with taking of non-aquatic forage during the dry seasons. The extreme pachyostosis was suggested to have been a protection against attacks by crocodiles.


Extinction

Remains from the Tamar Hat and Taza I archaeological sites suggests that the species may have been hunted by people. The latest known date for the species from Bizmoune has an age estimated between 6641 and 6009 cal BP (4691 to 4059
BCE Common Era (CE) and Before the Common Era (BCE) are year notations for the Gregorian calendar (and its predecessor, the Julian calendar), the world's most widely used calendar era. Common Era and Before the Common Era are alternatives to the or ...
) at the end of the Epipaleolithic. The Holocene transition in North Africa also saw other extinctions of ungulates, including ''
Gazella A gazelle is one of many antelope species in the genus ''Gazella'' . This article also deals with the seven species included in two further genera, '' Eudorcas'' and '' Nanger'', which were formerly considered subgenera of ''Gazella''. A third ...
, Equus,'' '' Camelus'', the African subspecies of the aurochs and ''Syncerus'' ''antiquus''.


References

{{Taxonbar, from=Q96392525 Cervinae Prehistoric deer Cenozoic mammals of Africa Pleistocene even-toed ungulates Holocene extinctions Fossil taxa described in 1890 Taxa named by Richard Lydekker Prehistoric monotypic mammal genera