Editing Nyala (section)

==Taxonomy and naming==
{{cladogram|align=left|title=
|caption=Phylogenetic relationships of the mountain nyala from combined analysis of all molecular data (Willows-Munro et.al. 2005)
|cladogram={{clade | style=font-size:90%;line-height:100%;width:300px;
|1={{clade
|1={{clade |label1=
 |1= {{clade|label1=
           |1=
         {{clade
         |label1=
         |1={{clade
            |1={{clade
                |1={{clade
                   |1=[[Giant eland]]
                   |2=[[Common eland]]}}
                |2={{clade
                   |1=[[Greater kudu]]
                   |2={{clade
                       |1=[[Mountain nyala]]
                       |2={{Clade
                           |1={{clade
                              |1={{clade
                                  |1=[[Bongo (antelope)|Bongo]]
                                  |2=[[Sitatunga]]}}
                           |2={{Clade
                           |1={{clade
                              |1={{clade
                                  |1=[[Cape bushbuck]]
|2=[[Harnessed bushbuck]] }} }} }} }} }} }} }} }} 
         |2='''Lowland nyala'''}}
|2=[[Lesser kudu]] }} }} }} }} }} }}
The nyala was first [[species description|described]] by [[George French Angas]], an English naturalist, in 1849. The [[scientific name]] of nyala is ''Tragelaphus angasii''. The name ''angasii'' is attributed to Angas, who said that [[John Edward Gray]] had named this species after Angas' father, [[George Fife Angas]] of South Australia. According to Article 50.1.1 of the [[International Code of Zoological Nomenclature]] and [[International Commission on Zoological Nomenclature]], though, this is insufficient to state Gray as the author.<ref name=MSW3/> The name "nyala" is the [[Tsonga language|Tsonga]] name for this antelope, which is likely the source of the English, along with [[Zulu language|Zulu]] {{Lang|zu|inyala}}.<ref>OED2</ref> Its first known use was in 1899. The word has a [[Bantu languages|Bantu]] origin, similar to the [[Venda language|Venda]] word {{Lang|ve|dzì-nyálà}} (nyala buck).<ref>{{Merriam-Webster|Nyala|access-date=30 January 2016}}</ref>

The nyala is the second [[taxon]] to branch off from the tragelaphine family tree just after the [[lesser kudu]]. As the nyala line has remained separate for a considerable time (over 5&nbsp;million years), some authorities have placed it in its own monotypic genus ''Nyala''. ''Nyala'' was proposed in 1912 by American zoologist [[Edmund Heller]], who also proposed ''Ammelaphus'' for the lesser kudu,<ref name=heller>{{cite book| author=Heller, E.| author-link=Edmund Heller| title=New Genera and Races of African Ungulates| publisher=Smithsonian Institution| location=Washington D. C.| url=https://www.biodiversitylibrary.org/pdf3/005508900035814.pdf| page=15| date= 2 November 1912}}</ref> but it was not widely recognized. It was re-erected as a valid genus in 2011 under the classification of [[Peter Grubb (zoologist)|Peter Grubb]] and [[Colin Groves]],<ref>{{cite book| last1=Groves|first1=C.| last2=Grubb|first2=P.| title=Ungulate Taxonomy| year=2011}}</ref> but has not been embraced by taxonomic authorities such as the Mammal Diversity Database.<ref name=burton>{{cite book| last=Burton| first=M.| title=International wildlife encyclopedia| year=2002| publisher=Marshall Cavendish| location=New York| isbn=978-0-7614-7269-8| author2=Burton, R.| edition=3rd| pages=1765–6| chapter=Nyala| chapter-url=https://archive.org/details/internationalwil03burt0}}</ref><ref>{{Cite web |title=Nyala |url=https://www.mammaldiversity.org/explore.html#genus=Tragelaphus&species=angasii&id=1006275 |access-date=7 August 2022 |website=Mammal Diversity Database}}</ref> Among its closest extant relatives are the forest-inhabiting [[Bongo (antelope)|bongo]], the [[Cape bushbuck|bushbuck]], the large [[Common eland|common]] and [[Giant eland|giant elands]], the [[Greater kudu|greater]] and [[Lesser kudu|lesser kudus]], the [[mountain nyala]] (of [[Ethiopia]]) and the swamp-dwelling [[sitatunga]], all of which share similar characteristics, such as hornless females, vertical side-stripes and other unique white markings, and the spiraling horns and distinct "beard" or [[dewlap]] of males, often with a raised trail of fur extending down the back and underside.{{cn|date=May 2024}}
[[File:ZOO Hellabrunn, München - kudui.jpg|thumb|Nyala family in captivity]]

In 2005, Sandi Willows-Munro (of the [[University of KwaZulu-Natal]]) and colleagues carried out a [[mitochondrial DNA]] analysis of the nine ''Tragelaphus'' species. Mitochondrial DNA and [[nDNA|nuclear DNA]] data were compared. The results showed the tribe Tragelaphini to be [[monophyletic]], with the [[lesser kudu]] (''T.&nbsp; imberbis'') [[Basal (phylogenetics)|basal]] in the phylogeny, followed by the nyala.<ref name=cladogram>{{cite journal|last1=Willows-Munro|first1=S.|last2=Robinson|first2=T. J.|last3=Matthee|first3=C. A.|title=Utility of nuclear DNA intron markers at lower taxonomic levels: Phylogenetic resolution among nine ''Tragelaphus'' spp.|journal=Molecular Phylogenetics and Evolution|date=June 2005|volume=35|issue=3|pages=624–36|doi=10.1016/j.ympev.2005.01.018|pmid=15878131|bibcode=2005MolPE..35..624W }}</ref><ref name="groves2014">{{cite journal|last1=Groves|first1=C.|title=Current taxonomy and diversity of crown ruminants above the species level|journal=Zitteliana|date=2014|volume=32|issue=B|pages=5–14|url=http://www.palmuc.de/bspg/images/pdf/zitteliana32/1_groves.pdf|issn=1612-4138|access-date=2016-03-22|archive-url=https://web.archive.org/web/20160204110133/http://www.palmuc.de/bspg/images/pdf/zitteliana32/1_groves.pdf|archive-date=2016-02-04|url-status=dead}}</ref> On the basis of mitochondrial data, studies have estimated that the lesser kudu separated from its [[sister clade]] around 13.7&nbsp;million years ago. However, nuclear DNA data shows lesser kudu and nyala forming a clade, which collectively separated from the sister clade 13.8&nbsp;million years ago.<ref name="ropiquet">{{cite journal|last1=Ropiquet|first1=A.|title=Etude des radiations adaptatives au sein des Antilopinae (Mammalia, Bovidae)|journal=Ph.D. Thesis, Université Paris|date=2006|volume=6|issue=1–247}}</ref><ref name="hassanin">{{cite journal|last1=Hassanin|first1=A.|last2=Delsuc|first2=F.|last3=Ropiquet|first3=A.|last4=Hammer|first4=C.|last5=Jansen van Vuuren|first5=B.|last6=Matthee|first6=C.|last7=Ruiz-Garcia|first7=M.|last8=Catzeflis|first8=F.|last9=Areskoug|first9=V.|last10=Nguyen|first10=T.T.|last11=Couloux|first11=A.|title=Pattern and timing of diversification of Cetartiodactyla (Mammalia, Laurasiatheria), as revealed by a comprehensive analysis of mitochondrial genomes|journal=Comptes Rendus Biologies|date=2012|volume=335|issue=1|pages=32–50|doi=10.1016/j.crvi.2011.11.002|pmid=22226162|doi-access=free}}</ref>

===Genetics and evolution===
The nyala has 55 male [[chromosome]]s and 56 female chromosomes.<ref name=Rubes>{{cite journal| last=Rubes| first=J.|author2=Kubickova, S. |author3=Pagacova, E. |author4=Cernohorska, H. |author5=Berardino, D. |author6=Antoninova, M. |author7=Vahala, J. |author8= Robinson, T. J. | title=Phylogenomic study of spiral-horned antelope by cross-species chromosome painting| journal=Chromosome Research|year=2008| volume=16| issue=7| pages=935–47| doi=10.1007/s10577-008-1250-6| pmid=18704723| s2cid=23066105}}</ref> The [[Y chromosome]] has been [[Chromosomal translocation|translocated]] onto the 14th chromosome, as in other [[Tragelaphus|tragelaphids]], but no [[Chromosomal inversion|inversion]] of the Y chromosome occurs.<ref name=MSW3>{{MSW3 Artiodactyla | id = 14200727 | page = 126}}</ref> Cranial studies have shown that the [[mountain nyala]] and nyala, though sharing a common name, are actually distant relatives.<ref name=antelope/>

Fossil evidence suggests that the nyala has been a separate species since the end of the [[Miocene]] (5.8&nbsp;million years ago). Genetic evidence suggests that the proto-nyala had some early hybridization with the proto-lesser kudu, but the two have remained separate long after this crossing.<ref name=MSW3/>