Megaloceros

Megaloceros (from Greek: Template:Lang Template:Transliteration + Template:Lang Template:Transliteration, 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, Megaloceros giganteus, vernacularly known as the "Irish elk" or "giant deer", is also the best known. Fallow deer are thought to be their closest living relatives.<ref>Lister, A. M., Edwards, C. J., Nock, D. A. W., Bunce, M., van Pijlen, I. A., Bradley, D. G., Thomas, M. G. & Barnes, I. 2005. The phylogenetic position of the 'giant deer' Megaloceros giganteus. Nature 438, 850-853.</ref><ref>Mennecart, B., deMiguel, D., Bibi, F., Rössner, G. E., Métais, G., Neenan, J. M., Wang, S., Schulz, G., Müller, B. & Costeur, L. 2017. Bony labyrinth morphology clarifies the origin and evolution of deer. Scientific Reports 7: 13176.</ref><ref>Gonzalez, S., Kitchener, A. C. & Lister, A. M. 2000. Survival of the Irish elk into the Holocene. Nature 405, 753-754.</ref> Megaloceros has been suggested to be closely related to other genera of "giant deer", like the East Asian genus Sinomegaceros (whose species are sometimes included in Megaloceros), and the European Praemegaceros.

Nomenclatural history

Megaloceros giganteus was originally described in 1799 as Alce gigantea by Johann Friedrich Blumenbach based on specimens found in Ireland. With Alce being a variant of the genus Alces used for elk/moose.<ref name="Handbuch der Naturgeschichte">Blumenbach J. 1799. Handbuch der Naturgeschichte (6th Ed.) 16: 697</ref>

In 1827 Joshua Brookes, in a listing of his zoological collection, named the Megaloceros (spelled Megalocerus in the earlier editions) in the following passage:<ref>Joshua Brookes (1827) "Brookesian Museum. The Museum of Joshua Brookes, Esq. Anatomical and Zoological Preparations" London Gold and Walton</ref><ref name="Lister 1987 spelling">Lister, A M, 1987 Megaloceros Brookes 1828 Mammalia Artiodactyla Proposed Emendation Of The Original Spelling The Bulletin of zoological nomenclature. 44 255–256</ref>Template:BlockquoteThe etymology being from Greek: Template:Lang Template:Transliteration "great" + Template:Lang Template:Transliteration "horn, antler".<ref>Template:Cite web</ref> The type and only species named in the description being Megaloceros antiquorum, based on Irish remains now considered to belong to M. giganteus, making the former a junior synonym. The original description was considered by Adrian Lister in 1987 to be inadequate for a taxonomic definition.<ref name="Lister 1987 nomenclature">Template:Cite journal</ref> In 1828 Brookes published an expanded list in the form of a catalogue for an upcoming auction, which included the Latin phrase "Cornibus deciduis palmatis" ("palmate deciduous horns") as a description of the remains. The 1828 publication was approved by International Commission on Zoological Nomenclature (ICZN) in 1977 as an available publication for the basis of zoological nomenclature.<ref name="Lister 1987 nomenclature" /> Adrian Lister in 1987 judged that "the phase "Cornibus deciduis palmatis" constitutes a definition sufficient under the [International Code of Zoological Nomenclature] (article 12) to validate Megalocerus."<ref name="Lister 1987 nomenclature" /> The original spelling of Megalocerus was never used after its original publication.<ref name="Lister 1987 spelling" />

In 1844 Richard Owen named another synonym of the Irish elk, including it within the newly named subgenus Megaceros, Cervus (Megaceros) hibernicus. This has been suggested to be derived from another junior synonym of the Irish elk described by J. Hart in 1825, Cervus megaceros.<ref name="Lister 1987 nomenclature" /> Despite being a junior synonym, Megaloceros remained in obscurity and Megaceros became the common genus name for the taxon.<ref name="Lister 1987 spelling" /> The combination "Megaceros giganteus" was in use by 1871.<ref>Template:Cite book</ref> George Gaylord Simpson in 1945 revived the original Megaloceros name, which became progressively more widely used, until a taxonomic decision in 1989 by the ICZN confirmed the priority of Megaloceros over Megaceros, and Megaloceros to be the correct spelling.<ref name="Lister 1987 spelling" /><ref>International Commission on Zoological Nomenclature 1989. Opinion 1566. Megaloceros Brookes, 1828 (Mammalia, Artiodactyla): original spelling emended. Bulletin of zoological nomenclature 46: 219–220.</ref>

Taxonomic composition

Other than the type species Megaloceros giganteus, the composition of the genus is contested.<ref name=":2" /><ref name="Croitor 2018">Template:Cite book</ref><ref name=":1" /> While considered to be part of the genus Megaloceros by many authors,<ref name=":0" /> M. savini and related taxa (novocarthaginiensis and matritensis) are split into the separate genus Praedama by some scholars.<ref name=":2" /><ref name="Croitor 2018" /><ref>Template:Citation</ref>

The genus Megaloceros is widely agreed to belong the subfamily Cervinae.<ref name="Croitor 2018" /> Megaloceros has often been placed in the tribe Megacerini, alongside other "giant deer" genera like Sinomegaceros and Praemegaceros, though the taxonomy regarding giant deer as whole is uncertain and contested.<ref name=":2">Template:Cite journal</ref> A close relationship with Sinomegaceros has been supported by mitochondrial DNA, which found that the mitochondrial sequences of M. giganteus are nested within those of Sinomegaceros, suggesting that the two lineages interbred with each other after the initial split between them, with all mitochondrial genomes of Sinomegaceros more closely related to those of M. giganteus than to their closest living relative Dama.<ref name=":3">Template:Cite journal</ref>

Relationships of Megaloceros mitochondrial genomes, after Xiao et al. 2023.<ref name=":3" />Template:Clade

File:Megaloceros antler.png

Antlers of M. giganteus (bottom) and Praedama (often included in Megaloceros)

Species

Species ordered from oldest to youngest:

M. novocarthaginiensis

Described from the latest Early Pleistocene 0.9-0.8 Ma of Cueva Victoria in Spain. Known from antlers, teeth and postcranial material. Related and possibly ancestral to M. savini<ref name=":1">Template:Cite journal</ref>

M. savini

Middle Pleistocene European species<ref name=":2" />, with a temporal range spanning approximately 750-450,000 years ago,<ref name=":0" /> slightly larger than a caribou/reindeer, first fossils found near Sainte Savine, France and near Soria, Spain. Its antlers were straight, with thornlike prongs. The lowermost prongs near the base were palmate. Has been suggested to comprise the separate genus Praedama.

It is suggested to have been a mixed feeder or grazer.<ref>Template:Cite journal</ref>

M. matritensis<ref name="

0">Template:Cite journal</ref>

Mid-Pleistocene species, lived around 400-300,000 years ago near present-day Madrid, Spain, being contemporary with M. giganteus. The species had enlarged premolars, very thick molar enamel, and a low mandibular condyle. The species itself formed part of the diet of people which lived in the area. M. matritensis fossils are found associated to stone tools of late Acheulean and early Mousterian type. The species is thought to be descended from M. savini

M. giganteus

Largest, best known, and among the last species of the genus that stands about Template:Convert at the shoulders. Lived throughout Eurasia, from Ireland to Siberia during the late Middle Pleistocene to early Holocene.

Megaceroides algericus from the Late Pleistocene to Holocene of North Africa has been considered to be closely related and possibly derived from Megaloceros by some authors.<ref name=":02">Template:Cite journal</ref>

"M". stravpolensis from the Early Pleistocene of Southwestern Russia<ref>Template:Cite journal</ref> has been subsequently suggested to belong to Arvernoceros.<ref name="Croitor 2018" /><ref name=":0" /> The species "Megaloceros" cretensis from the island of Crete has been moved to the genus Candiacervus.<ref>Template:Citation</ref>

Further reading

• Template:Smallcaps (2006): Template:Cite journal
• Template:Smallcaps (1987): Megaceros or Megaloceros? The nomenclature of the giant deer. Quaternary Newsletter 52: 14–16.
• Template:Smallcaps (2005): Template:Cite journal