Megaloceros: Difference between revisions

Megaloceros; Temporal range: late Early Pleistocene to Middle Holocene ~1.2–0.0077 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Skeleton of Megaloceros giganteus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
Family:	Cervidae
Tribe:	Cervini
Genus:	†Megaloceros; Brookes, 1828
Species
	†M. giganteus (type); †M. matritensis; †M. novocarthaginiensis; †M. savini;
Synonyms
	Megaceros (Owen, 1844);

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Megaloceros (from Greek: μεγαλος megalos κερας keras, 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.^[1]^[2]^[3] Megaloceros has been suggested to be closely related to other genera of "giant deer", like the East Asian genus Sinomegaceros, and the European Praemegaceros.

Nomenclatural history

Megaloceros giganteus was originally described in 1899 as Alce gigantea based on specimens found in Ireland. With Alce being a variant of the genus Alces used for elk/moose.^[4]

In 1827 Joshua Brookes, in a listing of his zoological collection, named the Megaloceros (spelled Megalocerus in the earlier editions) in the following passage:^[5]^[6]

Amongst other Fossil Bones, there [are] ... two uncommonly fine Crania of the Megalocerus antiquorum (Mihi). (Irish), with unusually fine horns, (in part restored)
— Joshua Brookes, Brookesian Museum. The Museum of Joshua Brookes, Esq. Anatomical and Zoological Preparations, p 20.

The etymology being from Greek: μεγαλος megalos "great" κερας keras "horn, antler".^[7] 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.^[8] 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" 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.^[8] 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."^[8] The original spelling of Megalocerus was never used after its original publication.^[6]

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.^[8] Despite being a junior synonym, Megaloceros remained in obscurity and Megaceros became the common genus name for the taxon.^[6] The combination "Megaceros giganteus" was in use by 1871.^[9] 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.^[6]^[10]

Taxonomic composition

Other than the type species Megaloceros giganteus, the composition of the genus is contested.^[11]^[12]^[13] While considered to be part of the genus Megaloceros by many authors,^[14] M. savini and related taxa (novocarthaginiensis and matritensis) are split into the separate genus Praedama by some scholars.^[11]^[12]^[15]

The genus Megaloceros is widely agreed to belong the subfamily Cervinae.^[12] 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.^[11] 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.^[16]

Relationships of Megaloceros mitochondrial genomes, after Xiao et al. 2023.^[16]

Dama

Sinomegaceros Megaloceros

Sinomegaceros pachyosteus (China)

Megaloceros giganteus (Russia, Belgium)

Sinomegaceros ordosianus (China, Russia)

	Sinomegaceros pachyosteus (China)

	Megaloceros giganteus (Europe, Russia)

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^[13]
M. savini: Middle Pleistocene European species^[11], with a temporal range spanning approximately 750-450,000 years ago,^[14] slightly larger than a caribou/reindeer, first fossils found near Sainte Savine, France and near Soria, Spain. Its antlers were straight, with thorn-like 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.^[17]
M. matritensis^[14]: Mid-Pleistocene species, lived around 300-400 ka 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 2 m (6.6 ft) 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.^[18]

"M". stravpolensis from the Early Pleistocene of Southwestern Russia^[19] has been subsequently suggested to belong to Arvernoceros.^[12]^[14]

References

^ 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.
^ 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.
^ Gonzalez, S., Kitchener, A. C. & Lister, A. M. 2000. Survival of the Irish elk into the Holocene. Nature 405, 753-754.
^ Blumenbach J. 1799. Handbuch der Naturgeschichte (6th Ed.) 16: 697
^ Joshua Brookes (1827) "Brookesian Museum. The Museum of Joshua Brookes, Esq. Anatomical and Zoological Preparations" London Gold and Walton
^ ^a ^b ^c ^d Lister, A M, 1987 Megaloceros Brookes 1828 Mammalia Artiodactyla Proposed Emendation Of The Original Spelling The Bulletin of zoological nomenclature. 44 255–256
^ "Megaloceros". Oxford Dictionary. Archived from the original on 12 July 2020. Retrieved 2020-07-12 – via Lexico.
^ ^a ^b ^c ^d Lister, A.M. (1987). "Megaceros or Megaloceros? The nomenclature of the giant deer". Quaternary Newsletter. 52: 14–16.
^ Vogt, Carl (1871). Lehrbuch der Geologie und Petrefactenkunde : zum Gebrauche bei Vorlesungen und zum Selbstunterrichte. p. 7. OCLC 162473843.
^ International Commission on Zoological Nomenclature 1989. Opinion 1566. Megaloceros Brookes, 1828 (Mammalia, Artiodactyla): original spelling emended. Bulletin of zoological nomenclature 46: 219–220.
^ ^a ^b ^c ^d Vislobokova, I. A. (December 2013). "Morphology, taxonomy, and phylogeny of megacerines (Megacerini, Cervidae, Artiodactyla)". Paleontological Journal. 47 (8): 833–950. doi:10.1134/S0031030113080017. ISSN 0031-0301.
^ ^a ^b ^c ^d Croitor, Roman (2018). Plio-Pleistocene deer of Western Palearctic : taxonomy, systematics, phylogeny. pp. 72 (stavropolensis) 93–94 (Praedama). ISBN 978-9975-66-609-1. OCLC 1057238213.
^ ^a ^b Made, J. van der (2015). "The latest Early Pleistocene giant deer Megaloceros novocarthaginiensis n. sp. and the fallow deer "Dama df. vallonnetensis" from Cueva Victoria (Murcia, Spain)". Mastia. 11–13: 269–323. S2CID 44675184.
^ ^a ^b ^c ^d Van der Made, Jan (2019). "The dwarfed 'giant deer' Megaloceros matritensis n.sp. from the Middle Pleistocene of Madrid - A descendant of M. savini and contemporary to M. giganteus". Quaternary International. 520: 110–139. Bibcode:2019QuInt.520..110V. doi:10.1016/j.quaint.2018.06.006. S2CID 133792579. Retrieved 2019-02-04.
^ Athanassiou, Athanassios (2022), Vlachos, Evangelos (ed.), "The Fossil Record of Continental Fossil Deer (Mammalia: Artiodactyla: Cervidae) in Greece", Fossil Vertebrates of Greece Vol. 2, Cham: Springer International Publishing, pp. 205–247, doi:10.1007/978-3-030-68442-6_6, ISBN 978-3-030-68441-9, retrieved 2023-11-15
^ ^a ^b Xiao, Bo; Rey-lglesia, Alba; Yuan, Junxia; Hu, Jiaming; Song, Shiwen; Hou, Yamei; Chen, Xi; Germonpré, Mietje; Bao, Lei; Wang, Siren; Taogetongqimuge; Valentinovna, Lbova Liudmila; Lister, Adrian M.; Lai, Xulong; Sheng, Guilian (November 2023). "Relationships of Late Pleistocene Giant Deer as Revealed by Sinomegaceros Mitogenomes from East Asia". iScience: 108406. doi:10.1016/j.isci.2023.108406. PMC 10690636.
^ Strani, Flavia; DeMiguel, Daniel; Alba, David M.; Moyà-Solà, Salvador; Bellucci, Luca; Sardella, Raffaele; Madurell-Malapeira, Joan (April 2019). "The effects of the "0.9 Ma event" on the Mediterranean ecosystems during the Early-Middle Pleistocene transition as revealed by dental wear patterns of fossil ungulates". Quaternary Science Reviews. 210: 80–89. doi:10.1016/j.quascirev.2019.02.027.
^ Croitor, Roman (July–September 2016). "Systematical position and paleoecology of the endemic deer Megaceroides algericus Lydekker, 1890 (Cervidae, Mammalia) from the late Pleistocene-early Holocene of North Africa". Geobios. 49 (4): 265–283. doi:10.1016/j.geobios.2016.05.002.
^ Titov, V. V.; Shvyreva, A. K. (January 2016). "Deer of the genus Megaloceros (Mammalia, Cervidae) from the Early Pleistocene of Ciscaucasia". Paleontological Journal. 50 (1): 87–95. doi:10.1134/S0031030116010111. ISSN 0031-0301. S2CID 131336166.

@@ Line 17: / Line 17: @@
 '''''Megaloceros''''' (from [[Ancient Greek language|Greek]]: {{lang|grc|μεγαλος}} {{transliteration|grc|megalos}}   {{lang|grc|κερας}} {{transliteration|grc|keras}}, 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 species|type]] and only undisputed member of the genus, ''[[Irish elk|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]],'' and the European ''[[Praemegaceros]]''.
-== Taxonomy ==
+== Nomenclatural history ==
+''[[Megaloceros giganteus]]'' was originally described in 1899 as ''Alce gigantea'' based on specimens found in Ireland. With ''Alce'' being a variant of the genus ''[[Alces]]'' used for [[Moose|elk/moose]].''<ref name="Handbuch der Naturgeschichte">Blumenbach J. 1799. ''[https://books.google.com/books?id=9pJSAAAAcAAJ&dq=alce gigantea&pg=PA697 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) [https://books.google.com/books?id=4DwAAAAAQAAJ&pg=PA20 "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 [https://www.biodiversitylibrary.org/part/348#/summary ''Megaloceros'' Brookes 1828 Mammalia Artiodactyla Proposed Emendation Of The Original Spelling] ''The Bulletin of zoological nomenclature''. '''44''' 255–256</ref>{{Blockquote|text=Amongst other Fossil Bones, there [are] ... two uncommonly fine Crania of the ''Megalocerus antiquorum'' (Mihi). (Irish), with unusually fine horns, (in part restored)|author=Joshua Brookes|title=Brookesian Museum. The Museum of Joshua Brookes, Esq. Anatomical and Zoological Preparations|source=p 20.}}The etymology being from [[Ancient Greek language|Greek]]: {{lang|grc|μεγαλος}} {{transliteration|grc|megalos}} "great"   {{lang|grc|κερας}} {{transliteration|grc|keras}} "horn, antler".<ref>{{Cite web |title=''Megaloceros'' |url=https://www.lexico.com/en/definition/megaloceros |url-status=dead |archive-url=https://web.archive.org/web/20200712195102/https://www.lexico.com/en/definition/megaloceros |archive-date=12 July 2020 |access-date=2020-07-12 |website=Oxford Dictionary |language=en |via=Lexico}}</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">{{cite journal |last1=Lister |first1=A.M. |year=1987 |title=''Megaceros'' or ''Megaloceros''? The nomenclature of the giant deer |url=https://www.researchgate.net/publication/264785006 |journal=Quaternary Newsletter |volume=52 |pages=14–16}}</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"'' 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 <nowiki>[</nowiki>[[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>{{Cite book |last=Vogt |first=Carl |author-link=Carl Vogt |url=https://books.google.com/books?id=m25CAAAAYAAJ&q="megaceros giganteus"&pg=PA96 |title=Lehrbuch der Geologie und Petrefactenkunde : zum Gebrauche bei Vorlesungen und zum Selbstunterrichte |year=1871 |pages=7 |oclc=162473843}}</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 [[Principle of Priority|priority]] of ''Megaloceros'' over ''Megaceros'', and ''Megaloceros'' to be the correct spelling.<ref name="Lister 1987 spelling" /><ref>International Commission on Zoological Nomenclature 1989. [https://www.biodiversitylibrary.org/part/548#/summary 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">{{Cite book |last=Croitor |first=Roman |url=https://hal.archives-ouvertes.fr/hal-01737207/document |title=Plio-Pleistocene deer of Western Palearctic : taxonomy, systematics, phylogeny |year=2018 |isbn=978-9975-66-609-1 |pages=72 (''stavropolensis'') 93–94 (''Praedama'') |oclc=1057238213}}</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>{{Citation |last=Athanassiou |first=Athanassios |title=The Fossil Record of Continental Fossil Deer (Mammalia: Artiodactyla: Cervidae) in Greece |date=2022 |url=https://link.springer.com/10.1007/978-3-030-68442-6_6 |work=Fossil Vertebrates of Greece Vol. 2 |pages=205–247 |editor-last=Vlachos |editor-first=Evangelos |access-date=2023-11-15 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-68442-6_6 |isbn=978-3-030-68441-9}}</ref>
-''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">{{Cite journal |last=Vislobokova |first=I. A. |date=December 2013 |title=Morphology, taxonomy, and phylogeny of megacerines (Megacerini, Cervidae, Artiodactyla) |url=http://link.springer.com/10.1134/S0031030113080017 |journal=Paleontological Journal |language=en |volume=47 |issue=8 |pages=833–950 |doi=10.1134/S0031030113080017 |issn=0031-0301}}</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 (genus)|Dama]]''.<ref name=":3">{{Cite journal |last=Xiao |first=Bo |last2=Rey-lglesia |first2=Alba |last3=Yuan |first3=Junxia |last4=Hu |first4=Jiaming |last5=Song |first5=Shiwen |last6=Hou |first6=Yamei |last7=Chen |first7=Xi |last8=Germonpré |first8=Mietje |last9=Bao |first9=Lei |last10=Wang |first10=Siren |last11=Taogetongqimuge |last12=Valentinovna |first12=Lbova Liudmila |last13=Lister |first13=Adrian M. |last14=Lai |first14=Xulong |last15=Sheng |first15=Guilian |date=November 2023 |title=Relationships of Late Pleistocene Giant Deer as Revealed by Sinomegaceros Mitogenomes from East Asia |url=https://linkinghub.elsevier.com/retrieve/pii/S2589004223024835 |journal=iScience |language=en |pages=108406 |doi=10.1016/j.isci.2023.108406|doi-access=free |pmc=10690636 }}</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">{{Cite journal |last=Vislobokova |first=I. A. |date=December 2013 |title=Morphology, taxonomy, and phylogeny of megacerines (Megacerini, Cervidae, Artiodactyla) |url=http://link.springer.com/10.1134/S0031030113080017 |journal=Paleontological Journal |language=en |volume=47 |issue=8 |pages=833–950 |doi=10.1134/S0031030113080017 |issn=0031-0301}}</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 (genus)|Dama]]''.<ref name=":3">{{Cite journal |last=Xiao |first=Bo |last2=Rey-lglesia |first2=Alba |last3=Yuan |first3=Junxia |last4=Hu |first4=Jiaming |last5=Song |first5=Shiwen |last6=Hou |first6=Yamei |last7=Chen |first7=Xi |last8=Germonpré |first8=Mietje |last9=Bao |first9=Lei |last10=Wang |first10=Siren |last11=Taogetongqimuge |last12=Valentinovna |first12=Lbova Liudmila |last13=Lister |first13=Adrian M. |last14=Lai |first14=Xulong |last15=Sheng |first15=Guilian |date=November 2023 |title=Relationships of Late Pleistocene Giant Deer as Revealed by Sinomegaceros Mitogenomes from East Asia |url=https://linkinghub.elsevier.com/retrieve/pii/S2589004223024835 |journal=iScience |language=en |pages=108406 |doi=10.1016/j.isci.2023.108406|doi-access=free |pmc=10690636 }}</ref>
 Relationships of ''Megaloceros'' mitochondrial genomes, after Xiao et al. 2023.<ref name=":3" />{{clade|''[[Dama (genus)|Dama]]''|{{clade

Revision as of 12:36, 17 June 2024

Nomenclatural history

Taxonomic composition

Species

References

Further reading