Formed circa 45,400–45,840 years BP (NO/NO1, based on YFull 2017,[1] and previous estimates of: 41,500 [95% CI 37,400 <-> 45,600] years BP,[2] 48,871 [95% CI 37,095 <-> 58,831] years BP,[3] and 50,800 or 43,500 years BP[4])
Coalescence age
Circa 41,500 – 40,067 years BP (NO/NO1, based on YFull 2017,[1] and previous estimates of: 36,800 [95% CI 34,300 <-> 39,300] years BP,[2] 41,900 [31,294 <-> 51,202] years BP,[3] and 44,700 or 38,300 years BP depending on mutation rate[4])
The location of NO1 at the SNP M214 follows the taxonomy set out by Karmin et al. 2022,[9] and conforms to a structure shown by ISOGG (2022).[8] (However neither Karmin nor ISOGG has integrated one of the findings of Poznik et al. 2016:[6] that there was a subclade generation both above haplogroup NO1 (M214) and beneath K-M2308. That is, both ISOGG and Karmin et al. continued to equate K2a with haplogroup NO.)
Before 2016, the subclades compromising both NO and NO1 were not recognised, and were regarded as synonymous with K2a.[6] Researchers such as David Poznik (Poznik et al. 2016) documented examples of previously unknown subclades of haplogroup K2, in both ancient remains and living individuals, which: (firstly) had several, varying suites of the SNPs regarded previously as uniquely defining K2a and NO, but also (secondly) lacked any of the SNPs specifically identifying haplogroups Haplogroup N (M231) and Haplogroup O (M175).[6] This demonstrated conclusively that multiple stages of development separated K2a from NO, which therefore constituted "grandparent" and "grandchild" clades. Poznik et al. 2016 used the name "K2a1" (a.k.a. K-M2313) for the Y-DNA of some of the individuals who belonged to K2a(xK2a*,NO), while also mentioning that K-M2313 did not include all examples of K2a(xK2a*,NO).
As of 2022, the International Society of Genetic Genealogy (ISOGG) refers to NO-M214 as "NO1", and to K2a (M2308)/K-M2313 as "NO".[8] There may be at least one other primary branch of NO: the ISOGG official Y-DNA haplogroup tree lists a haplogroup known as "NO1~" [sic] (CTS707/M2306) alongside NO-M214 (which ISOGG refers to as "NO1").[8] The tilde (~) indicates that its exact position of NO1~ in the phylogeny is unknown. It may be a primary branch or sibling of NO, it may be a primary branch or sibling of K2a, K-M2313, or it may instead be a primary branch of K2a.
Based on the projected origins of K2a, K-M2313, and the basal haplogroups N* and O* respectively, NO* probably originated in East Asia.[5][6]
Distribution
While there is some evidence of NO* being found in living individuals, these examples are not well-researched. Further research may instead identify them as belonging to N* (M231), N1, or the provisional subclade N2 (F3373/M2283/Page56/S323).[11][12] These cases include:
5.7% (2/35) of Bouyei males, in China or Vietnam;[12]
a pool of four samples of Japanese males at 2.9% (6/210), particularly in Tokushima Prefecture at a rate of 5.7% (4/70),[11] and;
Members of Haplogroup NO* include a Telugu of Indian origin sampled in the United Kingdom and a Malay sampled in Singapore.[6][1]
Two sets of ancient remains previously considered as possibly belonging to NO have since been reclassified upstream to K2a.[citation needed]
Ust'-Ishim man dates from approximately 45,000 BP and was found in Omsk Oblast, Russia.[13] (Until 2016 these remains were erroneously classified as K2*.)
Oase 1: the remains found in Romania of a male who lived 37,000-42,000 years BP.[14]
Likewise, cases previously regarded as possible examples of NO* or NO1*, and since ruled out, include:
two Han Chinese males previously found to be negative for M175 (i.e. Haplogroup O) and LLY22g (an obsolete, possibly inaccurate marker for N1), have subsequently have been found to belong to N* (N-M231),[15] and;
a clade first identified in South India, defined by the SNP M147 and labelled "pre-NO" and "Haplogroup X", among other names, was found to be a sibling of NO (K2a) within Haplogroup K2 (K-M526); the new clade was renamed K2e.[citation needed]
Subclades
Phylogenetic tree
This phylogeny of haplogroups K2a, K2a1, and NO is based on YFull 2018,[1] Poznik 2016, ISOGG 2018, Karafet 2008.[6][8][16]
K2aK-M2308 (M2308) Found only in the ancient remains "Ust'-Ishim man" (c. 45,000 BP) and "Oase 1" (c. 39,500 BP).[6]
K2a1K-M2313 (M2313/Z4858) [6] Named by Poznik 2016; previously not distinguished from K2a.
NOK-M214 (M214/Page39; F176/M2314; CTS5858/M2325/F346; CTS11572) [1][6][8] Poznik 2016 and ISOGG 2018 distinguish between NO1 (M214), N and O.
^Van Oven M, Van Geystelen A, Kayser M, Decorte R, Larmuseau HD (2014). "Seeing the wood for the trees: a minimal reference phylogeny for the human Y chromosome". Human Mutation. 35 (2): 187–91. doi:10.1002/humu.22468. PMID24166809. S2CID23291764.
^K-M2313*, which as yet has no phylogenetic name, has been documented in two living individuals, who have ethnic ties to India and South East Asia. In addition, K-Y28299, which appears to be a primary branch of K-M2313, has been found in three living individuals from India. See: Poznik op. cit.; YFull YTree v5.08, 2017, "K-M2335", and; PhyloTree, 2017, "Details of the Y-SNP markers included in the minimal Y tree" (Access date of these pages: 9 December 2017)
^ Haplogroup S, as of 2017, is also known as K2b1a. (Previously the name Haplogroup S was assigned to K2b1a4.)
^ Haplogroup M, as of 2017, is also known as K2b1b. (Previously the name Haplogroup M was assigned to K2b1d.)
^ abG. David Poznik, Yali Xue, Fernando L. Mendez, et al., 2016, "Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences." Nature Genetics vol. 48, no. 6 (June): pp. 593–599. doi:10.1038/ng.3559.
^ abRootsi, Siiri; Zhivotovsky, Lev A; Baldovič, Marian; Kayser, Manfred; Kutuev, Ildus A; Khusainova, Rita; Bermisheva, Marina A; Gubina, Marina; Fedorova, Sardana A; Ilumäe, Anne-Mai; Khusnutdinova, Elza K; Voevoda, Mikhail I; Osipova, Ludmila P; Stoneking, Mark; Lin, Alice A; Ferak, Vladimir; Parik, Jüri; Kivisild, Toomas; Underhill, Peter A; Villems, Richard; et al. (2007). "A counter-clockwise northern route of the Y-chromosome haplogroup N from Southeast Asia towards Europe". European Journal of Human Genetics. 15 (2): 204–211. doi:10.1038/sj.ejhg.5201748. PMID17149388.
^Tatiana M. Karafet, Brian Hallmark, Murray P. Cox et al., "Major East-West Division Underlies Y Chromosome Stratification Across Indonesia," MBE Advance Access published March 5, 2010.