Gene 545 (2014) 1–4

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Genetic, historical and linguistic perspectives on the origin of theKelantanese Malays

Keat Wei Loo a,b, Siew Hua Gan a,⁎a Human Genome Centre, School of Medical Sciences, Universiti Sains Malaysia, Kubang Kerian, Kelantan, Malaysiab Genomics Research Centre, Griffith Health Institute, Griffith University, Parklands Drive, Southport QLD 4222, Queensland, Australia

Abbreviations: BC, Before Christ; mtDNA, mitochondri⁎ Corresponding author at: Human Genome Centre

Universiti Sains Malaysia, 16150 Kubang Kerian, KelantanE-mail address: shgan@usm.my (S.H. Gan).

http://dx.doi.org/10.1016/j.gene.2014.05.0160378-1119/© 2014 Elsevier B.V. All rights reserved.

a b s t r a c t

a r t i c l e i n f o

Article history:Received 30 October 2013Received in revised form 4 May 2014Accepted 5 May 2014Available online 6 May 2014

Keywords:KelantanGeneticsHistoricalLinguisticsMalays

The Kelantanese Malays who resided in the remote northeastern regions of the Malay Peninsula in the Kelantanstate are believed to have a unique genetic signature. The objective of this review is to analyze the populationalsub-structure of the Kelantanese Malays from historical, genetic and linguistic perspectives. Historical data sug-gest that the Semang were composed of the Jahai, Bateq and Kensiu sub-tribes, whereas the Senoi were com-posed of only the Temiar sub-tribe. The Mendriq sub-tribe is believed to be the first group of aborigines toland in Kelantan. Subsequently, genetic analysis showed that the Kelantanese Malays are an independent cladeat the base of the phylogenetic tree and contain genetic material similar to that of the Semang, specifically theJahai and the Kensiu sub-tribes. The genetic data are supported by the fact that the Aslian language, a branchof the Austroasiatic languages that is widely spoken by the Semang, was potentially transmitted through agricul-tural activities. However, the potential limitation of this mini-review is the lack of primary reliable sources cov-ering the historical, linguistic and genetic features of the Kelantanese Malays.

© 2014 Elsevier B.V. All rights reserved.

1. Introduction

The population in Peninsular Malaysia consists of Deutero-Malays(modern Malays), Chinese, Indians and three main aboriginal tribes,namely, the Proto Malay, the Negrito (Semang) and the Senoi (Ismailet al., 2009). The Negrito (Semang) and the Senoi are known as ‘OrangAsal’ or ‘Orang Asli’ in Malaysia (Ismail et al., 2009). Based on themigra-tion and trading patterns from centuries ago, the Deutero-Malays wereconsidered to be descendants of the Proto Malays, with four main sub-ethnic groups consisting of the Melayu Minang, Melayu Jawa, MelayuBugis and Melayu-Kelantan or Kelantanese Malays (Hatin et al., 2011;Ismail et al., 2009), all of which are genetically heterogeneous. Accord-ing to Hatin et al. (2011), theMelayu Minang, Melayu Jawa andMelayuBugis are believed to have similar historical and cultural links withpopulations from the Indonesian archipelago when compared to theKelantanese Malays.

From 100 Before Christ (BC) to 1300 Anno Domini, PeninsularMalaysia, which was strategically located and connected the Indonesianarchipelagowith the kingdomof Indochina, was a famous port for tradersfrom China, India, Thailand, Arab and Sumatra. Consequently, thehigh frequency of intermarriage and integration with the localMalays attracted migrants from the surrounding areas (Marshall

al DNA., School of Medical Sciences,, Malaysia.

Cavendish Corporation, 2008). Recently, a study investigated thegenetic divergence among the Southeast East Asian population be-cause family ancestries are strongly influenced by geographicaland linguistic affiliations (Abdulla et al., 2009). For example, Hatin et al.(2011) hypothesized that the western region of the Melayu Minang andthe southern Malays (Melayu Jawa and Melayu Bugis) experienceddiverse cultural and historical links with the Indonesian archipelago, un-like the Kelantanese Malays who resided in the remote northeastern re-gions of the Malay Peninsula in the Kelantan state. In this review, wewill discuss the populational substructure of the Kelantanese Malaysfrom historical, genetic and linguistic perspectives to describe the originof the Kelantanese Malays. Overall, the objective of this review is to ana-lyze the populational sub-structure of the Kelantanese Malays from his-torical, genetic and linguistic perspectives (See Fig. 1).

2. Various populations in the state of Kelantan

Kelantan is located at the geographical coordinates of 6°75.12″ northlatitude and 102°14′ 23.25′ east longitude and consists of 10 districts,namely, Tumpat, Kota Bahru, Tanah Merah (Chi Tu), Pasir Mas, Bachok,Machang, Pasir Puteh, Kuala Krai, Jeli and Gua Musang. Kelantan isinhabited by Kelantanese Malays, aborigines, Chinese and Indians(http://www.kelantan.gov.my/v6/; Rentse, 1934).

Although the Semang and the Senoi are believed to have descendedfrom a single ancestry in Kelantan, they can be differentiated by their ad-aptations to the distinct ecological niches (Hill et al., 2006). The Semangare composed of the Jahai, Bateq, Kensiu andMendriq sub-tribes,whereas

Fig. 1. Genetic, Historical and Linguistic Perspectives on the Origin of the Kelantanese Malays and the Gaps that can be filled with future studies Abbreviations: SNP: single nucleotidepolymorphism, mtDNA: mirochrondria DNA.

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the Senoi are composed of only the Temiar sub-tribe. The Semang andSenoi are scattered in various districts of Kelantan (Hamid et al., 2013).The traditional Semang, who are of patrilineal descent, are rainforest for-agers and practice egalitarianism. The cognatic descent of the Senoi re-sulted in an appearance that is between the Negrito and Mongoloid. TheSenoi live in longhouses and practice slash-and-burn agriculture at higheraltitudes (Hill et al., 2006). The Senoi also practice egalitarianism (Hillet al., 2006).

Historically, theMendriq are believed to be the first group of aborig-ines who landed in the Gua Musang district of theMalay Peninsula. TheMendriq have physical features resembling the East African population.The Mendriq were foragers who harvested seasonal fruit and gatheredforest products for consumption and medical purposes (Mohamad,2010; Musharbash and Barber, 2012; Subramaniam, 2012). In 1972, re-settlement projects were launched by theMalaysian government to im-prove the lifestyles of the Mendriq. Economic transformation projectswere designed to help the Mendriq by focusing on commodity produc-tion (Musharbash and Barber, 2012). Unfortunately, this move indirect-ly demolished their traditional homeland, or saka, and their groupidentity. The Mendriq eventually became dispirited and suffered fromfrequent malnutrition, which finally led to their extinction (http://asiaharvest.org/people-group-profiles/malaysia/).

The Jahai or Jah Jehai initially settled along the Pos Rual River in theJeli district. The Jahai practiced hunting using blowpipes and poisondarts and scoured for fish (Mohamad, 2010). This indicated their

advancement from a shifting cultivation to a semi-nomadic way oflife. However, the Bateq inhabit the Kuala Krai and Gua Musang dis-tricts, mainly in the areas surrounding the national park. The Bateqlive as small nomadic foraging groups and are hunter-gathererswho trade rattan and resinous wood for tobacco and other staple foods(http://asiaharvest.org/people-group-profiles/malaysia/; Mohamad,2010).

Rather than settling in the Kelantan districts like the other groups, theKensiu Kelantan tend to practice cross-border migrations into Thailandand live in scattered areas, such as Kampung Baru Siong in Baling,Kedah and the Narathiwat and Yala provinces in Thailand (the southern-most town in Thailand at theMalaysia–Thailand border). This grouphas aclose resemblance to the Kensiw Thai and is reclusive, avoiding any socialinteraction with the outside world; however, their unique singing styleswere widely adopted by the locals. Because intermarriages (includingwith the Malays) are strongly prohibited, the genetic makeup of theKensiu has likely been highly conserved for generations, and thisconservation can be useful for the analysis of the Malay substructure(http://asiaharvest.org/people-group-profiles/malaysia/; Mohamad,2010).

The Temiar or Sakai is the fourth largest aboriginal group. Unlikeother aborigines, the lives of the Temiar are highly dependent on bam-boo for making weapons and receptacles and for building houses (Ben-jamin, unpublished data, Nanyang Technological University Singapore;Mohamad, 2010). The Temiar live in longhouses in extended family

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structures, mainly in the regions of Kuala Yai, Gamala, Kuala Betis andPos Blau in Gua Musang.

3. Genetic signature of the Kelantanese Malays

Structural genetic analysis of the Malay population was used to pre-dict the relationships among the aboriginal populations in the MalayPeninsula. Single nucleotide polymorphism (SNP) data analysis stronglyindicated that the Kelantanese Malays are an independent clade at thebase of the phylogenetic tree and are composed of a mixture of Semangand Indian populations, unlike other Malay subgroups (Hatin et al.,2011). As reported by Hatin et al. (2011), the genetic materials ofother Malay subgroups, for example, theMelayu Minang may resemblethat of the Indonesian Malays and Torajawhile theMelayu Jawamay re-semble that of the Indonesian Jawa and Chinese.

The above hypothesis is plausible because Kelantan was once ruledby the Patani Kingdom, which had a strong Indian influence for centu-ries in the early millennia before the introduction of Islam. In addition,an epic titled Hikayat Patani suggests that after Patani's invasion of Ke-lantan, part of the Patani's throne was given to the first Kelantan kingRaja Sakti (Maier, 2001a,b; Rahman, 2013; Syukri, 2002). However, aSNP study (Hatin et al., 2011) indicated that the Kelantanese Malaysare not in the same clade as the Indians. This is plausible because theKelantanese Malays were an agrarian society who resided in areaswith lush paddy fields devoid of rubber and oil palm plantation indus-tries after British colonization. It has been reported that the absence ofthese industries failed to attract Indian laborers, eventually leading to amore favorable natural selection for the Kelantanese Malays (MarshallCavendish Corporation, 2008).

Apart from the Indian admixture, the genes of the KelantaneseMalays were reported to be similar to the Semang (specifically theJahai and the Kensiu) (Hatin et al., 2011). However, no similar ge-netic study was conducted for the Bateq and Mendriq tribes. In ad-dition, according to an ancient Malay text called the HikayatMerong Mahawangsa, the Semang appeared to be the oldest aborig-inal group in the Malay Peninsula (Nagata, 2006). These nomadichunter-gatherers have frizzy hair, dark skin (ranging from copperto black color), short stature (≤1.5 m), rounded eyes, broad noses andlow cheekbones (Ghani unpublished data, Universiti Sains Malaysia),leading to a debate among geneticists onwhether theirmigrationwas as-sociatedwith thefirstwave of humanmigration fromAfrica or the secondwave of Asian human migration (Ghani, unpublished data, UniversitiSains Malaysia; Hatin et al., 2011).

During the first wave of migration, the Semangmigrated from Africato Southeast Asia before colonizing Polynesia, coastal Melanesia andMadagascar (Hill et al., 2006; Stoneking andDelfin, 2010). Furthermore,based on linguistic and archeological evidence for the period between4000 BC and 3000 BC, the Proto-Malays inhabited Taiwan and were in-volved in the first wave of human migration towards Sulawesi, Borneo,Central Jawa and Indonesia via the Philippines (Andaya, 2001; Husseinet al., 2007). This migration was followed by a second wave of humanmigration between 1500 BC and 500 BC involving the Senoi who origi-nated from South Asian regions and moved from central Jawa to theMalay Peninsula via the Straits of Malacca.

Evidence for both waves of human migration consists of observa-tions made by Andaya (2001) and Hussein et al. (2007) that the mito-chondrial DNA (mtDNA) of the inhabitants originating from SoutheastAsia [Malaysia, Indonesia, Singapore, Borneo, Philippines and Oceania(Micronesia and Melanesia)] is linked to those of Eastern Austronesia(i.e., Taiwan, Philippines and Eastern Indonesia). The subsequent colo-nization of the Indo-Malay archipelago brought the light-skinned andstraight-haired Proto Malays into the Malay Peninsula and representedthe first influx of the Mongoloids (Hill et al., 2006).

Genealogy studied usingmtDNA transmitted viamatrilineal lineagesprovides vast information for phylogeography because the detectedmutation rate of mtDNA is higher than the mutation rate of nuclear

DNA in the absence of a recombination event (Rokas et al., 2003). Thevariations in mtDNA detected among the tribe of Semang strongly sug-gest that the tribe of Semang was the descendants of the Hoabinhian,who migrated from South India via the southern route, and providestrong evidence for the aboriginal origin of the tribe of Semang in theMalay Peninsula between the late Pleistocene and mid-Holocene, ap-proximately 60,000 years ago. This observation again indicates thatthe tribe of Semang was among the earliest aboriginal settlers in thecoastal areas of the Malay Peninsula. However, the tribe of Semangmay have fled inland in the fear of newcomers, and this movementeventually led to their isolation in remote forested hilly regions suchas the national park that spans a large region in the northern MalayPeninsula (Hatin et al., 2011).

A recentmtDNA analysis of some inhabitants in East Asia indicates ahigh frequency ofM21mtDNA lineages among the tribe of Semang. Thisobservation suggests an early dispersal of this haplotype group in di-verse parts of the Malay Peninsula and reflects the early modernhuman's migration from Africa (Stoneking and Delfin, 2010). The flowof genes from the islands in Southeast Asia indicated the arrival of theN9a lineages during the Holocene; the tribes of Semang and Senoi ab-originals predominantly had M21 and R21 lineages, whereas the N21and N22 lineages were limited to the Proto Malays (Hill et al., 2006).The tribe of Senoi also contained a considerable Holocene componentfrom Indochina in the F1a1a lineage. The Bateq sub-tribe most likelyoriginated from Southeast Asia Island, with B5b, N21 and N22 lineages,and the ProtoMalays were represented by theM7c1c lineage. The pres-ence of theM7c1c, N21, andN22 lineages represents the recent offshorearrival of the Proto Malays and is related to the influx of Austronesian-speaking people since the late Holocene. The R21 haplogroup observedamong the aborigines may have spread from Indochina to the MalayPeninsula during the early to mid-Holocene (Hill et al., 2006). In addi-tion, the mtDNA diversity suggests that genetic drift occurred amongthe aborigines and the Proto-Malays, which may have been due to aninitial ethnogenetic process or a subsequent geographical effect (Hillet al., 2006). Analysis of the mtDNA not only supports the theory ofgene flow, but also indicates the settlement of Semang and Senoi tribesand their respective subtribes in theMalay Peninsula. Since mtDNA lin-eages can act as disease indicators, it is hypothesized that the precisecorrelation between genetic materials (in this case, the mtDNA line-ages)with tribes and/or subtribes of the KelantaneseMalays has the po-tential to provide a new insight for personalized medicine based onethnicity.Mohseni et al. (2014) in their research reported an interestingfinding on the complex re-arrangement in the arginase I gene amongfive unrelated Malay families residing in the state of Kelantan. Eventhough they have not stratified the Malay subgroup into their re-spective tribes and sub-tribes, five novel mutations detected withinthe arginase I gene (designated as g.insIVS1 + 1899GTTTTATCAT;g.invIVS1 + 1933_ + 1953; g.delIVS1 + 1954_IVS2 + 914; c.del116_188; p.Pro20SerfsX4) acted as a common indicator for hyperargininemia.In addition, the researchers also suggested that the five novel mutationsare derived from similar ancestral family irrespective of the tribes andsub-tribes indicating that the genes within the Kelantanese Malays areunique and well-conserved.

4. Linguistics among the Kelantanese Malays

Language and genetics have a methodological history of cross-fertilization (Abdulla et al., 2009; Dunn et al., 2011). The Y-chromosome types and mtDNA are likely to represent the survival oflanguages and ancient settlements, respectively (Forster and Renfrew,2011). The gender-specific transmission of language indicates the dis-persal of farming terminology on domesticated animals and plants byfarmers speaking a protolanguage (Forster and Renfrew, 2011). For ex-ample, the Austroasiatic and Austronesian languages widely spoken bythe Kelantanese Malays and the aborigines in Kelantan were reported tooriginate from South China and spread to the Malay Peninsula via the

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Mongoloid farmers during the Neolithic and mid-Holocene periods (Hillet al., 2006). The Austroasiatic language was reported to spread south-wards from the mainland, whereas the Austronesian language was re-ported to spread from Taiwan, the Philippines and Indonesia to theMalay Peninsula via the island arc (Hill et al., 2006). However, the Austro-asiatic speakers in Southeast Asia are likely to have dispersed to SouthAsia during the Neolithic area because an analysis of the uniparentallyinherited markers of 610,000 common SNP loci in the nuclear genomeshowed that the Y chromosome haplogroup O2a has significantly higherdiversity and coalescent time (17–28 thousand years ago) in SoutheastAsia relative to South Asia (Chaubey et al., 2010).

In Kelantan, the Kelantanese Malays speak the Austronesianlanguage. It has been known that the Austronesian language withinthe Malayo-Polynesian subfamily is branched into the Malayan andMalayic languages as well as the local Malay dialect (Benjamin, 2009;Dunn et al., 2013; Pan-Asian SNP Consortium). The Aslian languagewidely spoken by the Semang is in the Mon Khmer subfamily underthe division of Austro-Asiatic languages (Dunn et al., 2011). Potentiallybecause of the strong agricultural background from 4000 to 5000 yearsago, the Aslian languagewas introduced to this agrarian state during thesecond wave of Southeast Asian migration. Ethnographically, the no-madic Semang (Jahai, Bateq, Kensiu and Mendriq) spoke northernAslian, the semi-sedentary horticulturalists Senoi (Temiar) spoke cen-tral Aslian, and the Proto-Malay spoke the southern Aslian language(Dunn et al., 2011).

The unique Aslian languages inherited by the Semang belong to abasic semantic category with a high degree of idiolectal variations as aresult of intermarriage (Ghani, unpublished data, Universiti SainsMalaysia). However, based on their marriage and place of living andthe continuous modernization among agricultural communities, thelanguage spoken by the Semang was influenced by the Malay and Thailanguages and the Mon-Khmer language from Vietnam. This observa-tion is consistent with the fact that the subsequent residency patternsand social practices could affect the patterns of genetic variation. Thehaplotypes that contain a group of genetic variations tend to cluster to-gether in populations that have a similar linguistic group and, possibly,similar historical backgrounds (Stoneking and Delfin, 2010). The poten-tial limitation of this mini-review is the severe lack of primary reliablesources covering the historical, linguistic and genetic features of theKelantanese Malays which clearly indicates the need for further re-search in this area.

5. Conclusion

We conclude that a populational substructure analysis from his-torical, genetic and linguistic perspectives may elucidate the originof the Kelantanese Malays and help to differentiate the mechanismsof cultural expansion by social selection and the expansion ofagriculture-related populations. The historical, genetic and linguis-tic perspectives on the origin of the Kelantanese Malays have cometo a common point where the tribes and sub-tribes of KelantaneseMalays could be explained from different perspectives thus highlightingthe uniqueness of the untainted genetic materials. In addition, the impor-tance of biolingohistorical perspectives in this mini-review may inspirefuture research on evolutionary and personalized genomics because it re-veals the untainted genetic signature of the Kelantanese Malays which isunique.

Conflicts of interest

None declared.

Acknowledgments

This researchwasfinancially supported by a ResearchUniversity grant(1001/PPSP/815073). First author was the recipient of an EndeavourResearch Fellowship and a MyBrain 15 (MyPhD) scholarship.

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