Draft:Haplogroup O-K18

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Haplogroup O-K18 also known as O-F2320 and (as of 2017) Haplogroup O1b1, is a human Y-chromosome DNA haplogroup. Haplogroup O-K18 is a descendant branch of Haplogroup O-P31. Based on its disjunct distribution, O-K18 can be further divided into south subclade O1b1a1-PK4 and north subclade O1b1a2-CTS4040. O-CTS4040 is widely distributed in East Asia, whereas O-PK4 is more frequent in South China and Southeast Asia. O-PK4 is best known for the high frequency of its O-M95 subclade among populations of Southeast Asia and among speakers of Austroasiatic languages in South Asia.

Origin
In a paper published in 2011 by a group of Chinese researchers affiliated with Fudan University, it has been suggested that China is the origin of the expansion of haplogroup O-M268, the parent haplogroup of O-F2320.

Distribution
Haplogroup O-K18 is distributed widely in Asia, from southern India to the Altai Mountains and Central Asia in the west, and from Indonesia to northern China and Japan in the east. According to its distribution, O-K18 can be roughly divided into north subclade O-CTS4040 and south subclade O-PK4. O-CTS4040 is overall uncommon, but it is relatively abundant in Northern and Eastern parts of China (about 5%). It is also found at low frequencies of approximately 1% or less at the periphery of its distribution in other Indo-Pacific area like Vietnamese, Koreans, Japanese, West Kalimantan, Hazaras, and Arabs (Qatar). The other haplogroup O-PK4 consists of O-F838 and O-M95. O-F838 are more frequent in the South Han in China, showing the same trend with the its parallel branch O-M95 in China. The other branch, O-M95, is the best known subclade for the whole Y Haplogroup O-K18. O-M95 is found only at marginally low frequencies of approximately 1% at the periphery of its distribution in southern India, Central Asia, northern China, and Japan, but many populations within the vast intervening territory in South Asia, Southeast Asia, and southern China display a greatly elevated frequency of Haplogroup O-M95 Y-chromosomes. Haplogroup O-M122, which attains its peak frequency among speakers of Sino-Tibetan and Hmong–Mien languages in China and Southeast Asia, and Haplogroup O-M119, which predominates among Taiwanese aborigines and many populations of the Philippines, also generally occur among speakers of Austroasiatic languages in South China and the Indochinese Peninsula, but usually at much lower frequencies than Haplogroup O-M95.

Modern northern Han Chinese Y haplogroups and mtdna match those of ancient northern Han Chinese ancestors 3,000 years ago from the Hengbei archeological site. 89 ancient samples were taken. Y haplogroups O3a, O3a3, M, O2a, Q1a1, and O* were all found in Hengbei samples.

According to the National Geographic project regarding O-M95: The Austro-Asiatic language family developed in groups containing men from this lineage. As these groups spread across Southeast Asia in successive waves, they spread their language. Today, the distribution of men from this lineage matches the pattern of these waves of migration. It is 42 percent of male lineages in Java, 40 percent of male lineages in Vietnam, and 38 percent of male lineages in Borneo. It accounts for 28 percent of the male population in Malaysia. It is present in Sumatra in about 14 percent of the male population. In mainland China, it is, on average, about 3 percent of the male population but a lot higher in ethnic minorities of South China. In South Asia, it is 9 percent of the Pardhan, between 1 and 2 percent of the Andh, and 10 percent of the Naikpod. It is around 59 percent of Balinese male lineages.

Haplogroup O-M95 is generally found in high percentages in most Austro-Asiatic ethnic groups but also found high in the Tai-Kradai people of South China and Southeast Asia, and Javanese, Sundanese, and Balinese of Indonesia. It is also widespread in ethnic minorities of South China related to language families of East Asian origin and Southeast Asian origin.

O-CTS4040
O-CTS4040 is relatively rare and is usually marked as O1B*/O2*-M268(PK4-, M176-) in the past academic report. It shares a common ancestor with its nearest outgroup, O-PK4, approximately 24,405 (95% CI 17,810 <-> 27,604) ybp according to Karmin et al. 2022, approximately 23,410 years before present according to 23mofang, or approximately 22,100 (95% CI 20,400 <-> 23,900) years before present according to YFull.

It is mainly distributed in East Asia and is mainly found in Han Chinese and occasionally found in Taiwan plains tribes, Vietnamese, Dai, Filipinos, Koreans, Japanese, West Kalimantan, Hazaras, and Arabs (Qatar). TMRCA of Han Chinese, Dai, Vietnamese, and Japanese members estimated to be 15,900 [95% CI 13,300 <-> 16,400] ybp. Relative paper illustrates O-P31/M268(PK4-, M176-) is found in North China (6.2%), East China (4.8%) and South China (3.1%).

Analysis of DNA extracted from a tooth from what are believed to be the remains of Cao Ding shows that he belonged to this clade. The researchers also found that the Y-chromosome of Cao Ding matches those of self-proclaimed living descendants of Cao Cao who hold lineage records dating back to more than 100 generations ago. Cao Cao laid the foundation of Cao Wei, one of three major states that succeeded the Han Dynasty of China.

In Yangshao culture (around 5000 BC), there is an ancient male who belongs to haplogroup O-PAGE59 in WangGou site (Zhengzhou, Henan, China). This is currently the oldest discovered ancient DNA that has been confirmed to be derived from O-CTS4040.

O-PK4
The coalescence age of O-PK4 is 13,911 (95% CI 11,147 <-> 15,915) ybp according to Karmin et al. 2022, 13,060 ybp according to 23mofang, or 12,900 (95% CI 11,700 <-> 14,200) years before present according to YFull. It mainly consists of two subclades: O-F838 and O-M95. It is best known for the high frequency of its O-M95 subclade among populations of Southeast Asia and among speakers of Austroasiatic languages in South Asia.

O-F838
This lineage has been relocated upstream of M95 following a paper published on the subject in 2011. Found in three samples of Han Chinese: 3/65 = 4.6% South China, 1/129 = 0.8% North China, 1/167 = 0.6% East China.

According to 23mofang, O-F838 (TMRCA 10,730 ybp) currently accounts for the Y-DNA of approximately 1.40% of all males in China, with its distribution being densest in the South Central Region of China.

Peng et al. (2013) found O-PK4(xM95), which probably should belong to O-F838 according to the phylogenetic tree of human Y-DNA as it is currently resolved, in a Bamar individual in Ayeyarwady Region, Myanmar.

Trejaut et al. (2014) found O-PK4(xM95) in one of 18 individuals sampled on Ambon Island, Indonesia, one of 24 individuals sampled in Hanoi, Vietnam, six of 258 miscellaneous Han volunteers in Taiwan, one of 60 Minnan in Taiwan, and one of 85 Siraya in Pingtung, Taiwan.

Wang et al. (2014) found O-PK4(xM95) in two of a sample of 46 Khams Tibetans from Xinlong County, Sichuan.

O-M95
This subclade is downstream from O-PK4. It reaches high frequencies among the populations of the islands of Sumatra, Java, Bali, and Borneo in western Indonesia (Karafet 2010). It has been found to be by far the most common Y-chromosome haplogroup among the Balinese, occurring in approximately 58.6% (323/551) of a sample of Balinese men. It is found around 70% frequency among Bhumij of East India. It has been found in 17.1% (6/35) of a sample of Malagasy in Madagascar (Hurles 2005) and in 1.7% (1/60) of a sample of Swahili people in Kilifi, Kenya. It is one of the most frequently occurring Y-DNA haplogroups among men in Malaysia, Thailand, Laos, Cambodia, Vietnam, and Myanmar. It is also very common among minority ethnic groups in India and China, especially those who have ethnolinguistic connections with populations in Southeast Asia (e.g. Munda peoples, Khasi people, and Nicobarese people in India and Kra–Dai peoples, Blang people, and Mang people in China).

O-M95(xM88) is relatively infrequent in other populations, but a study published in 2006 has found it in samples of Daurs (6/39 = 15.4%), Qiang people (3/33 = 9.1%), She people (3/34 = 8.8%), Hani people (2/34 = 5.9%), Yao people in Liannan, Guangdong (2/35 = 5.7%), Japanese people (2/47 = 4.3%), Evenks in China (1/26 = 3.8%), Han Chinese in Lanzhou, Gansu (1/30 = 3.3%), Han Chinese in Yili, Xinjiang (1/32 = 3.1%), Han Chinese in Chengdu, Sichuan (1/34 = 2.9%), and Yao people in Bama, Guangxi (1/35 = 2.9%).

A study published in 2010 found O-M95(xM111) in 57.3% (367/641) Bali, 49.2% (30/61) Java, 31.3% (10/32) Malaysia, 20.9% (18/86) Borneo (Indonesia), 15.8% (6/38) Toba people in Sumatra, 13.0% (7/54) Mandar people in Sulawesi, 7.1% (5/70) Vietnam, 6.1% (10/165) Han Chinese, 4.6% (18/394) Flores, 3.4% (2/58) Miao in China, 2.1% (1/48) Philippines, 1.7% (1/60) Yao in China, and 0.3% (1/350) Sumba. (Karafet 2010)

Trejaut et al. (2014) found O-M95(xM88) in 36.2% (51/141) Java, 29.4% (5/17) Sulawesi, 25.3% (19/75) general population of Bangkok, 25% (2/8) Malaysia, 22.2% (4/18) Ambon, 19.2% (5/26) Sumatra, 12.0% (3/25) Kalimantan, 10.0% (3/30) Yami, 8.3% (2/24) Hanoi, Vietnam, 6.7% (4/60) Minnan in Taiwan, 5.9% (2/34) Hakka in Taiwan, 3.7% (1/27) Akka in Thailand, 3.5% (9/258) miscellaneous Han in Taiwan, 1.8% (1/55) Han in Fujian, 1.6% (6/370) Taiwan Plains Tribes. The authors did not find any cases of O-M95(xM88) among their samples from the Philippines (0/146) or Taiwan Highlands Tribes (0/325).

O-M88
This subclade is downstream from O-M95. The TMRCA of O-M88, which is also known as O-M111, is estimated to be 6,607 (95% CI 5,216 <-> 7,632) ybp according to Karmin et al. 2022, 5,950 ybp according to 23mofang, or 5,600 [95% CI 5,000 <-> 6,300] years before present according to YFull. The entire O-M88 clade is estimated to share a most recent common ancestor with O-CTS5854, most members of which have been found in southern China, Laos, and Thailand, but some also in northern China, Japan, Vietnam, and the Philippines, 10,071 (95% CI 7,821 <-> 11,536) ybp according to Karmin et al. 2022, 9,500 [95% CI 8,600 <-> 10,500] years before present according to YFull, or 8,980 ybp according to 23mofang.

O-M88 is frequently found among Tai peoples, Vietnamese people, Hani-Akha people, She people, and some tribal peoples in Laos (including Aheu people, Xinh Mul people, Alak people, Kuy people, and So people ), with a moderate distribution among Cambodians, Qiang people, Yi people, Tujia people, Hlai, Miao, Yao, Cham people, Taiwanese aborigines, populations of Borneo, the Philippines, and Malaysia (Karafet 2010), and Han Chinese of Sichuan, Hunan, Guangxi, Guangdong, Yunnan, and Taiwan.

Trejaut et al. (2014) found O-M88 in 37.5% (21/56) Bunun, 25.9% (7/27) Akka in Thailand, 25.0% (6/24) Hanoi, Vietnam, 17.3% (13/75) general population of Bangkok, Thailand, 5.0% (7/141) Java, 3.4% (5/146) Philippines, 3.3% (1/30) Yami, 2.9% (1/34) Hakka in Taiwan, 1.7% (1/60) Minnan in Taiwan, 1.55% (4/258) Han in Taiwan, and 0.54% (2/370) Taiwan Plains Tribes (including 1/18 Papora and 1/38 Siraya from the Tainan coast).

Macholdt et al. (2020) found Y-DNA that belongs to subclades of O-M88 (O-F2758, O-F1399, O-Z24091, O-F2890, and O-Z24014) in 69.4% (25/36) of a sample of Lolo, 32.4% (12/37) of a sample of Nung, 28.0% (14/50) of a sample of Kinh, 22.2% (8/36) of a sample of Lachi, 12.9% (4/31) of a sample of Lahu, 11.6% (5/43) of a sample of Dao, 10.6% (5/47) of a sample of Tày, 8.3% (3/36) of a sample of Pathen, 8.3% (2/24) of a sample of Ede, 6.1% (2/33) of a sample of Hanhi, 4.2% (1/24) of a sample of Thái, and 3.7% (1/27) of a sample of Giarai from Vietnam.

O-M297
More research is needed on this lineage. It is claimed to be downstream from M95 and parallel to M88.

Phylogenetic history
Prior to 2002, there were in academic literature at least seven naming systems for the Y-Chromosome Phylogenetic tree. This led to considerable confusion. In 2002, the major research groups came together and formed the Y-Chromosome Consortium (YCC). They published a joint paper that created a single new tree that all agreed to use. Later, a group of citizen scientists with an interest in population genetics and genetic genealogy formed a working group to create an amateur tree aiming at being above all timely. The table below brings together all of these works at the point of the landmark 2002 YCC Tree. This allows a researcher reviewing older published literature to quickly move between nomenclatures.

Research publications
The following research teams per their publications were represented in the creation of the YCC Tree. • α and

• β

• γ

• δ

• ε

• ζ

• η

Phylogenetic trees
This phylogenetic tree of haplogroup O subclades is based on the YCC 2008 tree and subsequent published research.


 * O-M95 (M95)
 * O-M88 (M88, M111)

Genetics
• Genetic genealogy

• Haplogroup

• Haplotype

• Human Y-chromosome DNA haplogroup

• Molecular phylogenetics

• Paragroup

• Subclade

• Y-chromosome haplogroups in populations of the world

• Y-DNA haplogroups by ethnic group

• Y-DNA haplogroups in populations of East and Southeast Asia

Y-DNA O subclades
• O-47z

• O-M101

• O-M113

• O-M117

• O-M119

• O-M121

• O-M122

• O-M134

• O-M159

• O-M162

• O-M164

• O-M175

• O-M176

• O-M50

• O-M7

• O-M88

• O-M95

• O-MSY2.2

• O-P31