THE FINE SCALE ETHNOTAXA CLASSIFICATION OF MILLETS IN ...bidt.ca/maloles-et-al-2011.pdf · Millets...

THE FINE SCALE ETHNOTAXA CLASSIFICATION OFMILLETS IN SOUTHERN INDIA

Jose Rey Maloles, Kevan Berg, Subramanyam Ragupathy,Balasubramaniam C. Nirmala, Kabeer A. Althaf, Vadaman C. Palanisamy and

Steven G. Newmaster

This research explores variation in minor millets in the context of traditional knowledge (TK) and scientific knowledge(SK), including ethnobotany genomics, in southern India. In order to perceive biodiversity, we need to take a closer look at

the natural variation among species within the context of existing classifications using both TK and SK. Malayali

informants of the Kolli Hills in India were surveyed using 174 millet samples. We also collected seeds and grew millets in

greenhouse environments from which we recorded 96 morphological characters and extracted DNA for barcoding.Quantitative multivariate classification analysis of these plants revealed that the Malayali millet classification is

hierarchical and recognizes considerable fine scale variation with high consensus. In the field, the Malayali classified and

consistently identified 19 millet ethnotaxa (landraces). Variation in these same samples was analyzed using morphometricand molecular characters (DNA barcoding) but revealed fewer taxa. Some of the cryptic taxa identified by the Malayali,

including a potentially drought tolerant millet ethnotaxa, have considerable nutritional, medicinal, and ecological value.

Key words: ethnobotany genomics, ethnoclassification, millet, traditional knowledge, biodiversity

Jose Rey Maloles, Botany Division, Centre for Biodiversity Genomics, University of Guelph, Guelph,Ontario, Canada. N1G 2W1. Phone (519) 824-4120 ext 56002

Kevan Berg, Botany Division, Centre for Biodiversity Genomics, University of Guelph, Guelph,Ontario, Canada. N1G 2W1. Phone (519) 824-4120 ext 56002

Subramanyam Ragupathy, Botany Division, Centre for Biodiversity Genomics, University of Guelph,Guelph, Ontario, Canada. N1G 2W1. Phone (519) 824-4120 ext 56002 (e-mail: [email protected])

Balasubramaniam C. Nirmala, Department of Biotechnology, SRM Arts and Science College,University Central Facility, Tamil Nadu, India

Kabeer A. Althaf, Botanical Survey of India, Southern Circle, TNAU Campus Coimbatore 641 003,Tamil Nadu, India

Steven G. Newmaster, Botany Division, Centre for Biodiversity Genomics, University of Guelph, Guelph,Ontario, Canada. N1G 2W1. Phone (519) 824-4120 ext 56002 (e-mail: [email protected])

Vadaman C. Palanisamy, Centre for Biocultural Diversity, Kolli Hills. Chennai, India

Journal of Ethnobiology 31(2): 262–287 Fall/Winter 2011

Introduction

Millets are among the oldest cultivated grain crops and a source of food formillions of people living in the arid and semi-arid tropical areas of Asia andAfrica (Doggett 1989, 1993; Singh and Arora 1972). Millets have been cultivatedfor millennia (Fuller 2002, 2006a), and they are among the earliest archaeologicalplant remains (5500 BC) in areas such as India, Mexico, China and Africa (Weberand Fuller 2008). In southern India and Africa there is evidence that milletcultivation appears to be an independent process preceding the introduction ofother crops (Fuller 2005a, 2005b; Fuller et al. 2004; Neumann 2004). Theavailability of African varieties of millet during the last 400 hundred years withinIndia has marginalized the smaller, indigenous pearl millets that had beenutilized since 5500 BC (Fuller 2003a, 2005a; Weber 1998). Some ancient landracesor ‘‘lost millets’’ are no longer cultivated in some areas (Finnis 2007), butprevious research suggests that some were very important in local economiesand for the sustenance of many localized communities in southern India from5500 BC until at least 1600 AD (Austin 2006; De Wet 2000; Dove 1999).

Millet is a term that collectively refers to a number of small-seeded annualgrass species that are cultivated for both human and animal food. These grassesare adapted to dry infertile soils, have a short maturation period, perform well inmarginal environments and have considerable food value (Barbeau and Hilu1993; FAO and ICRISAT 1996). In India local cultures have long used millets andthese have been considered valuable by colonial and post-colonial governments(Fuller 2006a). Major millets like sorghum and non-indigenous pearl millet aretall growing and fairly drought tolerant. Minor millets are a group of grassyplants with short slender culms, small grains and they possess a remarkableability to survive under severe drought. At least three different species of minormillet are cultivated by tribal farmers of the Eastern Ghats: little millet (Panicumsumatrense Roth ex Roem & Schult.), Italian millet (Setaria italica (L.) P. Beauv.),and kodo millet (Paspalum scrobiculatum L.). Fuller (2006a) provides a full list ofsouth Indian millet cultivars, including those cultivated historically, andRenglakshmi (2005) lists the various landraces that are still used today in localcultures such as the Malayali.

Our research was conducted with the Malayali in the Kolli Hills, which lie inTamil Nadu’s Talaghat Plains (Bohle 1992), one of a series of hills of the EasternGhats (Figure 1). Covering approximately 282 square kilometers (Kumaran et al.1998), the Hills rise between 1000 and 1400 meters above sea level (Kumar-Range2001). Although surrounded by forest on the exterior, the interior has beenextensively cleared for agricultural purposes. Presently, the area is approximate-ly 51% agricultural land and 44% forest land (Kumar-Range 2001). The Malayaliare a scheduled tribe, a legally recognized tribal minority. The Malayali languageis a dialect that is not normally distinguished from mainstream Tamil by eitherlinguists or officials (Minatchisuntharan 1965). Renglakshmi (2005), Singh (1994)and Thurston and Rangachari (1975) provide detailed historical and linguisticaccounts of the Malayali. Until the last 20 to 30 years, the Kolli Hills Malayalicultivated most of their own foods, a tradition that has been sustained for

Fall/Winter 2011 JOURNAL OF ETHNOBIOLOGY 263

thousands of years. Several ethnotaxa of minor millets, including samai, thina,varagu, kambu, and keshvaragu (ragi), once played a key role in household foodsecurity and dietary diversity of the people in this region.

In this paper we investigate the agro-biodiversity knowledge of minor milletsin India by integrating the Malayali traditional knowledge (TK) with that ofscientific knowledge (SK). Several contributions to our understanding of theclassification of south Indian millets exist in the scientific literature (Kimata et al.1997, 2000; Kobayashi and Kimata 1989; Renglakshmi 2005). However,quantitative analysis of the TK classification, morphometric analysis or measureof genomic diversity among the minor millet taxa is lacking. Here, we addressthis deficiency and explore how tribal farmers classify minor millets usingvarious attributes (e.g., morphological, micro and macro habitat, agronomic andgastronomic characters), revealing considerable variation among landraces(Rengalakshmi 2005).

Previous research using morphological and genomic data only recognized fivedifferent types of millets: samai (Panicum sumatrense), thina (Setaria sp.), varagu(Paspalum scrobiculatum), kezhvaragu (Eleusine coracana (L.) Gaertn.) and solam(Sorghum bicolor (L.) Moench) (Singh 1994). The etymology of these terms providessome historical evidence of a long history of these crops’ use in southern India.Samai and varagu are featured in the oldest Tamil literature –the Sangam poemswhich date from 300 BC to 300 AD– as crops of pastoralists of the Deccan interior(the ‘mullai-tinai’ zone). These poems were composed by Dravidian Tamil poets,both men and women, from various professions and classes of society(Minatchisuntharan 1965). In Sangam literature, thina and varagu have beenmentioned as the staple food for the people. Archaeological evidence suggests that

Figure 1. Location of the study area on the Eastern Ghats of South India.

264 MALOLES et al. Vol. 31, No. 2

the millet landraces such as samai have been cultivated in Neolithic south India,such as at Hallur after 2000 BC (Fuller 2006a). It is found even earlier in the Gujaratand the northwest (Fuller 2006b). Keshvaragu (ragi) is not mentioned in Sangamliterature (2500-1800 BC) suggesting it was introduced from China and Korea, afterthe Sangam age. Many of the small-grained millets have been mistakenlyattributed to ragi, the finger millet of African origin (Eleusine coracana) (Fuller1999, 2002, 2003b, 2003c). The term ‘‘kezh’’, means red and was used todifferentiate the color of the grains. In the Sangam literature, people, land andplants, were classified based on habitat or land classification. Thus, the thina andvaragu are said to belong to kurunji, or fifth landscape of the mountain flower(Minatchisuntharan 1965). Kodo millet is a newer crop in south India from the IronAge (about 800 BC) onwards, and its name may originate from other grains, orperhaps from browntop millet (Brachiaria ramosa (L.) Stapf.; Fuller 2006a). Thenomenclature elements for Kodo millet (Paspalum scrobiculatum), little millet(Panicum sumatrense), ragi (Eleusine coracana), foxtail millets (Setaria italica), and(Setaria viridis (L.) P. Beauv.) are more ancient and widely shared across South,South Central and some Central Dravidian languages suggesting pre-Tamil (1000-300 BC, Megalithic age) ancestors of the Malayali have had names for these cropsfor millennia (Fuller 2006b), and have carried these names and these crops withthem. The Malayali community only arrived in the Kolli Hills 500 to 600 years ago(Kumaran 1993). It is widely claimed by the tribe members themselves that theybelonged to the Kongu Vellala caste of Dravidian Tamils when they settled in thehills. Moreover, the Malayali claim that they are the historical retainers andkeepers of millet practice in Kolli Hills. Therefore these five previously recordedterms for minor millets provide an important starting point for our study ofMalayali TK.

Because it is prudent to consider traditional knowledge of millet within acultural and historical-linguistic context alongside the natural variation in thesetaxa from the perspective of scientific knowledge, we worked with the Malayalitribal farmers to build a detailed TK classification of minor millets and surveytheir various uses within Malayali culture, but we also used a quantitativeapproach that incorporates SK including ethnobotany genomics (Newmaster andRagupathy 2009a, 2009b, 2010). Our research explores variation in minor milletsthrough several specific questions: 1) Is there high consensus of TK concerningclassification and use of minor millets? 2) Is the TK classification hierarchical andsufficiently fine scaled to recognize intraspecific variation as claimed byRengalakshmi (2005)? 3) Can millet species be classified using morphometricand molecular characters and if so, how fine scaled is this classification? 4) Is theclassification based on morphological and molecular characters consistent withthe TK classification of minor millets?

Methods

The methodology in this study supports the assemblage of SK and TK usinga quantitative approach. Consensus analysis provides quantitative evidence forthe classification of ethnotaxa and the associated TK. This forms a basis forexploring the complexity of the TK classification system by implementing


multivariate statistics that compared the TK classification with that of bothmorphological and molecular variation among these ethnotaxa. This approachallows us to assess the inherent variation among the taxa and ultimately addresshow the TK and SK classifications compare.

Study Site and People

Our study surveyed the Malayali people living in the Kolli Hills (Figure 1), anarea of about 473.3 km2, lying in Namakkal and Rasipuram taluks (counties) of theNamakkal district. We worked within 6 villages located in Kolli Hills: Aripalapatti,Kattankadupatti, Nakkamannu, Pallankuzhipatti, Soolavanthipatti, and Vilaram.The altitude varies from 300 m to 1,500 m, with an annual rainfall varying between800 mm and 1,300 mm. Rema and Indra (2003) have estimated that over 20,000 haof forests in Kolli Hills are under the Reserved Forest (RF) system.

According to the 1991 census, there were 37,000 people, 17,207 males and16,681 females, living in the Hills (Finnis 2007; Kumaran 1993; Surveyor’s Report1994). The Kolli Hills are predominantly populated by the Malayalis (meaning ‘‘hillhumans’’) living in the 14 nadus (a typical village). Malayali elders, who rememberthe move in historical times, recount that the people of the Kolli Hills were oncepeasants who inhabited areas near Kanchipuram. According to oral history, theMalayali people belonged to the lineage of three brothers who, when driven out oftheir lands in the 16th century, fled to the Kolli Hills, the Pachamalai and theKalrayan Hills (Kumaran 1993). Recent work by Finnis (2009) has raised a numberof points regarding changes to staple diets in the Kolli Hills, adding to the previouswork in the area (Finnis 2007), including Bohle’s (1992) discussion of the ways inwhich external influences and integration into a market economy has damagedfood security and dietary diversity in the Hills. These external influences include:water issues, household economic goals, house hold labor requirements, poverty,geographical isolation and the introduction of cash crops by policy makers.

Ethnobotanical Surveys

Ethnobotanical explorations were made in the study area in 2008 (Figure 1).Our research team included 118 tribal informants (traditional botanical experts)from local villages and 6 botanists from the Botany Division of the Centre forBiodiversity Genomics (CBG) at the University of Guelph and the BotanicalSurvey of India. Informed consent was obtained from all of the informants. Thesurvey used Bernard’s (1994) methodology, which identifies local experts intraditional botanical knowledge within the community. The field data werecollected in two stages. First, seeds and vouchers of whole plants were collectedwith tribal informants and surveys were administered based on the use of eachvoucher sample. Vouchers were collected and labeled for all taxa identified, andthen the voucher samples were the subject of interviews within the villages inorder to gather additional TK and validate the samples’ identifications. Datawere gathered in a series of structured, semi-structured and non-structuredinterviews, and a participatory approach was used regarding plant uses,identification, and nomenclature. Plants named by the Malayalis, ethnotaxa,are defined as variable plant populations that are adapted to local agro-climaticconditions. Traditional farmers name, select and maintain ethnotaxa to suit their


social, economic, cultural and ecological needs (Teshome et al. 1997). Interviewprotocols and data confirmation procedures were based on reports by Pelto andPelto (1990), Etkin (1993), Bernard (1994), Vogl et al. (2004), and Stepp andThomas (2005). To elucidate cultural domains and determine differences inknowledge about uses or classification of the different ethnotaxa amongindividuals, survey answers were cross-checked with those of other tribalrespondents by using various research protocols such as free recall lists, pilesorts, and consensus analysis (Weller and Romney 1988; Werner and Fenton1973). Plant samples collected from the tribal community were preserved for bothherbaria and DNA barcode analysis. Leaf, stem and flower parts collected in situwere fixed in silica gel, FAA solution (50% ethanol, 5% acetic acid, 10% formalin,35% water) and stored in 70% ethanol for morphological study ex situ. Thesesamples were used for measuring the variation in morphological characters andmolecular markers. Herbarium voucher specimens were deposited at the Centerfor Bio-Cultural Diversity (CBD) in Chennai, India. Scanned herbarium imageswere deposited in the Herbarium of the Biodiversity Institute of Ontario at theUniversity of Guelph, Ontario.

Identification and TK Consensus Analysis

Voucher samples including plants and seeds were collected from the sixcollection sites and were systematically identified by the taxonomists at theUniversity of Guelph Herbarium, Botanical Survey of India and by tribalinformants. Calculation of relative frequency (RF) and a consensus factor (Fic)was used to test homogeneity between SK and TK in identifying specimens pilesorting, revealing cryptic taxa (ethnotaxa) or limitations of the classificationwithout the use of molecular data. The relative frequency of each specimenmentioned during the interviews was calculated to determine a quantitativevalue for choosing a plant name (either the Latin binomial or ethnotaxa) from thepool of collected vouchers placed in a species concept (Newmaster et al. 2007;Ragupathy et al. 2007). RF is the simple calculation of the percentage ofspecimens associated with a taxon when taxonomists or tribal informants arepresented with a pool of vouchers and asked to perform pile sorting (Weller andRomney, 1988). Trotter and Logan (1986) provide the calculation of a ConsensusFactor [Fic 5Nur2Nt/(Nur21)], where Nur is the number of reported usagesclaimed by informants for a particular category (TK plant use) factor, where ause-report is a single record for use of a plant mentioned by an individual, andNt refers to the number of species used for that particular category for allinformants (Ali-Shtayeh et al. 2000; Camejo-Rodrigues et al. 2003). We adoptedthis measure to evaluate the degree of partitioning into categories (Heinrich2000). We have adapted this method to include ‘‘tribal use’’ by the tribalinformants (Heinrich 2000; Ragupathy and Newmaster 2009; Ragupathy et al.2008).

TK Classification Analyses

A classification analysis of traditional knowledge characteristics for all milletethnotaxa provided a quantitative assessment of how variation among milletethnotaxa is classified by the Malayali. We used Bray-Curtis average linkage to


classify minor millet ethnotaxa using the primary characteristics identified by theMalayali during the interviews. These characteristics include morphology,ecology, experience, gestalt, and use (Newmaster, Raguapthy et al. 2006;Newmaster et al. 2007). Morphological characters include visual features thatare either reproductive (e.g., seed size or color) or vegetative (e.g., leaf size).Ecological characters include those associated with sites where the Malayalicollect seeds and grow the various ethnotaxa, such as terrestrial, wetland, plains,hill slopes, upland, lowland, elevation, proximity to streams, and edaphicconditions. Personal experience refers to skills comprising smell, touch, and tastethat the Malayali use to identify ethnotaxa. Previous research (Newmaster et al.2007) has demonstrated that some cultures can recognize plants without the useof any specific characters. Often dubbed the ‘‘Gestalt’’ identification (Ellis 1999),we identify some things by their whole and not by their finer physical properties.We recorded the ethnotaxa that the Malayali distinguish without reference to anyother recognition characters. Plant use is defined as the use (e.g., nutritional,medicinal, technical or ritual) within the Malayali culture. The similarity matrixof species based on characters was used to build a cluster diagram (Figure 2),which is useful in assessing the capacity of each character to permit theclassification of all ethnotaxa.

Seed Collection and Greenhouse Protocols

Seed and voucher samples of landraces for little millet, Italian millet, andkodo millet were collected from the villages. Seeds of Indian finger millet(Eleusine coracana) and sorghum (Sorghum bicolor) were also collected and used asthe out group. Outgroup species are necessary to ‘‘root’’ a classification tree—that is, to indicate the last common ancestor (i.e., the deepest internal node)shared by the entire group of species. The seeds collected from Kolli Hills (fourpopulations of each ethnotaxa, including closely related taxa) were grown in 8’’plastic pots using standard Sunshine mix LA4 supplemented with Nutricote 13-13-13 Type 100 (with micronutrients). Fifty to one hundred seeds of eachpopulation were planted in each pot to ensure sufficient plant material. Theplants were grown in greenhouses at the Center for Biocultural Diversity fieldoffice (Kolli Hills) and at the Department of Biotechnology SRM (Sri RamasamyMemorial) Arts and Science College, SRM University Central Facility) in TamilNadu, for a period of five to six months. After a growth period of one month(September to October 2008), the plants were thinned to approximately 10 to 15plants per pot. The plants were grown at a temperature of approximately 24–26uC for 16 hours during the day, and 16–18u C for 8 hours at night. The plantsreceived 16 hours of photo-period through natural and supplementary lightingusing 600W high pressure sodium lamps. The irradiance of the greenhouse wasmeasured using a standard photometer. The bench was subdivided into a 237grid consisting of 14 sections, and irradiance levels were measured approxi-mately above pot level, 10’’ and 20’’ above pot level. Once a week, the pots wererepositioned randomly to different locations on the greenhouse bench tominimize the influence of micro-environments on the growth of the plants.Gibberellic acid (GA) was used to induce germination of the sengalivaragu,karumthina, and perumvaragu ethnotaxa. A GA solution with a concentration of


300 mmol/L was made using a 30 mmol/L GA + 30% ethanol stock solution andde-ionized water. The seeds of the three ethnotaxa were deposited on papertowels soaked in the GA solution and placed in an incubator at 25–27u C. Thesame protocols were repeated for another set of seeds from the same threeethnotaxa, but were incubated at 37u C.

Figure 2. Classification tree from tribal classification characters (i.e., morphology, ecology, experience,gestalt and utility, including 4 secondary classification mechanisms) of 19 ethnotaxa recognized as 5species (Latin binomials). Shaded area represents the finer scale of hierarchical TKclassification system.


Morphometric Analysis

A morphometric analysis was used to partition variation in morphologicalcharacteristics among the millet samples, which provided a classification of theethnotaxa. The 96 phenotypic characters for the morphometric analysis are listedin Table 1. These characters were obtained using digital imaging of ethnotaxapopulations collected from the field and those grown in the green house at twodifferent time intervals: three weeks and three months after the initial planting.We applied these morphometric criteria to the 174 specimens we collected in India.A matrix of 174 specimens and 96 morphological characters were used in amultivariate analysis (Pimentel 1979). Canonical ordination was used to detectgroups of specimens and to estimate the contribution of each variable to theordination. A principal component analysis (PCA; ter Braak 1998) was used toidentify the length of the ordination axis and the need for either a linear orunimodal ordination technique. The results of the PCA indicated the need for aunimodal, indirect ordination detrended correspondence analysis (DCA) toexplore variation in species scores in this study. A cluster analysis was used toclassify the specimens, as it is better in representing distances among similarspecimens, whereas DCA is better in representing distances among groups ofspecimens (Sneath and Sokal 1973). Cluster analysis was performed with NTSYS(Rohlf 2000). A distance matrix was generated using an arithmetic average(UPGMA) clustering algorithm and standardized data based on averagetaxonomic distance subjected to the un-weighted pair-group method (Morrison1967). A discriminant function analysis (DFA; SPSS 1999) was used to rigorouslytest the classification of specimens provided in the cluster analysis. The object ofDFA is to predict multivariate responses that best discriminate subjects among

Table 1. List of the 96 characters and criteria for morphometric analysis.

PhenotypicCharacters Criteria

Culm Stature, straightness, heightCulm node Number of nodes, rooting, constriction, color, indumentumCulm internode Diameter of lower internode, length, hollowness, length of base internode,

color, variegation, surface relief, indumentumCulm sheath Persistence, length at base, length at top, as a fraction of internode, color when

fresh, variegation, surface indumentum, hair location, hair inclination, haircolor, presence of marginal hairs, apex shape

Culm sheath blade Outline, basal constriction, persistence, width, indumentum, shape of apexLigule Height, color, apical ornament, structure, consistency, apex incision, shape of apexCollar Color, length of hairsLeaves Basal or cauline, heterophyllyLeaf sheaths Involute margins, length, presence of keel, venation, roughness, surface

indumentum, hair type, hair color, presence of marginal hairs,Leaf blades Shape of base, base symmetry, carriage, straightness, direction of taper,

vernation, length, width, consistency, stiffness, color, midrib distinctness,midrib lower emergence, clarity of venation, surface roughness, surfaceindumentum, surface hair density, surface location of hairs, surface hairtype, marginal thickening, marginal roughness, marginal hairiness, locationof marginal hairs, apex shape, apex pungency

Oral hairs Presence, shape, carriage, length, colorCaryopsis Outline, compression, transverse section, length, color, roughness, shape of

apex, presence of apical appendageEmbryo As a fraction of caryopsis


different groups (Ramsey and Schafer 1997). The 174 specimens were used as inputfor the DFA and were coded as belonging to one of the designated a priori groupswhich 1) determined if the classification was accurate, 2) provided discriminantfunctions for the classification of the taxa and, 3) indicated if there are importantmorphological characters for each of the canonical discriminate functions.

Ethnobotany Genomics

DNA barcoding was used to classify genetic variation among the milletethnotaxa, providing a complimentary classification to the morphometric analysisand the TK/SK classification of millets. For the DNA analysis of the milletethnotaxa, fresh samples of specimens were collected from the populationsgrowing in the greenhouse. After approximately 2.5 months of growth, 5individual plants were selected from each pot containing one ethnotaxapopulation. The youngest leaves were removed from the selected plants and wereplaced into silica gel bags to ensure rapid drying and minimal DNA degradation.Several DNA regions were selected for exploring variation among the milletsamples. Previous analyses of molecular data suggest that rbcL, matK, trnH-psbA(CBOL Plant Working Group 2009; Fazekas et al. 2008; Kress and Erikson 2007;Kress et al. 2005; Lahaye et al. 2008; Newmaster, Fazekas et al. 2006; Newmaster etal. 2008; Newmaster, Murugesan et al 2009; Newmaster, Rugapathy et al 2009;Ragupathy et al. 2009) and the ribosomal DNA intergenic spacer region (IGS; Kenjiet al. 2006) are the most variable regions for land plants, which is why they werechosen for this study. Total genomic DNA was isolated from approximately 10 mgof dried leaf material from each sample using the NucleoSpinH 96 Plant II(MACHEREY-NAGEL). Extracted DNA was stored in sterile micro-centrifugetubes at 220u C. The selected loci were amplified by PCR on a PTC–100thermocycler (Bio-Rad). DNA was amplified in 20 ml reaction mixtures containing1 U AmpliTaq Gold Polymerase with GeneAmp 106 PCR Buffer II (100 mM Tris-HCl pH 8.3, 500 mM KCl) and 2.5 mM MgCl2 (Applied Biosystems, Foster City,CA), 0.2 mM dNTPs, 0.1 mM of each primer (0.5 mM for matK), and 20 ng/ mltemplate DNA. Amplification products were sequenced directly in both directionswith the primers used for amplification, following the protocols specified inFazekas et al. (2008). Sequence products were cleaned from each specimen onSephadex columns and the samples were run on an ABI 3730 sequencer (AppliedBiosystems, Foster City, CA). Bidirectional sequence reads were obtained for allPCR products. Sequencher 4.7 (Gene Codes Corp, Ann Arbor, MI) was used toassemble and base-call sequences, and alignment was completed manually usingBioedit version 7.0.9. In order to obtain an estimate of variation in the regionsexamined, we calculated pairwise uncorrected p-distance for each region usingmega 3.1 (Kumar et al. 2004). The sequences were submitted to BOLD (Barcode ofLife Database - http://www.boldsystems.org/views/login.php) and GenBank.

Results

Malayali Millet Classification and Consensus of TK

The Malayali millet classification recognizes considerable fine scale variationamong all of the millet samples. A hierarchical cluster analysis resulted in a


diagram (Figure 2) with well-developed branches and clusters representing 19ethnotaxa using the TK classification characters. Their classification is hierarchi-cal, with a primary tier congruent with the five species in SK classification;Malayali recognize these five primary taxa as: sama (little millet; Panicumsumatrense), varagu (Kodo millet; Paspalum scrobiculatum), keshvaragu (Indianfinger millet; Eleusine coracana), thina (Italian millet; Setaria italica), and solam(sorghum; Sorghum bicolor). The Malayali classification is much finer than thescientific one in that a secondary tier of the hierarchical classification recognizes afiner classification comprised of 19 ethnotaxa (shaded section in Figure 2). Oneethnotaxon (senthina) has variants that are clearly defined by the Malayaliclassification and they have distinctive uses and habitat requirements (shown assenthina (1) and (2) in Figure 2).

Surveys of TK revealed a classification for minor millets includingintraspecific variation and various uses of millet among 19 ethnotaxa (Figure 2).High consensus factors (Fic . 0.95) among the 118 informants provide confidencein the TK classification system. The informants could classify the samples veryeasily as they regularly deal with these ethnotaxa as part of their livelihood.Traditional knowledge surveys supported by high consensus factors (Fic 0.85–0.92) revealed that many of the ethnotaxa are used for a variety of purposes:nutritional, forage and structural (e.g., roofs, huts; Table 2). The informantsimpressed upon us the cultural importance of these ethnotaxa while sharingdetailed accounts of their uses (Table 3).

Morphometric Analysis

Based on morphometric analysis of collected ethnotaxa specimens, ordina-tion analyses (using DCA) distinguished 15 of the 19 ethnotaxa. Four ethnotaxashowed no morphological differences even though the Malayali coulddistinguish these species using other characteristics (i.e., palatability, digestion,growing requirements, etc.). Principal components analysis (PCA) provided acharacter gradient that was unimodal (3.5 Standard Deviations SD), violating theassumption of a linear model (ter Braak 1998). Consequently, a DCA was used toclassify the 174 specimens resulting in distinct groups representing 19 ethnotaxa.High eigen values for the X-axis (0.847) and the Y-axis (0.672) indicated that thegradient axes were of considerable length and justified the use of DCA. The X-axis (axis 1) was strongly correlated with 61 characters; these include theduration of life cycle, number of nodes, ligule color, and direction of taper of leafblades (Figure 3). The Y-axis (axis 2) was strongly correlated with 52 characters;some of these include internode color, culm sheath color, collar color, ligulestructure, and caryopsis compression (Figure 3).

A discriminant function analysis (DFA) used 96 quantitative characters toclassify heterogeneity in 174 specimens into what is currently considered 5known taxa of millet (Panicum sumatrense, Paspalum scrobiculatum, Setaria italica,Eleusine coracana and Sorghum bicolor). The canonical correlation from thediscriminant functions is the ratio of the between group sums of squares to thetotal sums of squares. Thus, the first discriminant function is responsible for 60%of the between group differences (variability in the discriminant scores). Thesecond function is responsible for an additional 14% of the between group


variance and the third function is responsible for an additional 11% of thevariance. Wilk’s Lambda was used to test the hypothesis that there is nodifference in variance (p , 0.001) between the groups of taxa, which representdifferent species (SPSS 1999). There were significant differences (p , 0.001) forthe first two canonical functions; 100% of the specimens were correctly classifiedusing DFA into 18 distinct groups of taxa. This result indicates that TKclassification recognizes fine-scale morphological variation among millet taxa atthe infra-specific level in the SK classification.

Ethnobotany Genomic Analysis

The molecular data was used to test whether the ethnotaxa were consistentwith SK species, and to find out whether there was genetic variation among themillet species that the Malayali identified that SK classification did not at thelevel of species. The molecular data was used to discriminate all five species(Panicum sumatrense, Paspalum scrobiculatum, Setaria italica, Eleusine coracana andSorghum bicolor). Amplifications were highly specific with a clear background inthe agarose gel. A classification tree of the rbcL, matK and trnH-psbA plastidintergenic spacer was congruent with the primary tier of the Malayaliclassification resolving the five species in SK classification (Figure 4). Theinterspecific variation (p-distance 0.0234) is considerable, but there is nointraspecific variation and therefore no differentiation among any ethnotaxa asin the Malayali classification (Figure 3). The ribosomal DNA intergenic spacerregion sequence was only obtained for the samples in the genus Setaria. Therewas no intraspecific variation in the sequence data from this region and thereforeno differentiation of the ethnotaxa within the genus Setaria that were classifiedusing the morphometric analyses.

Identification Analysis

Although the ability of field taxonomists and informants to identify milletswas considerable, the respective classifications of SK and TK are nothomogeneous. Taxonomists identified 5 species from the 174 specimens with96% (RF) accuracy among individuals. A DNA barcode (rbcL, matK and trnH-psbA) was used to make accurate species assignments for all 174 samples. Weused the ‘‘best match’’ and ‘‘best close match’’ functions of the program

Table 2. Ethnobotanical consensus index for TK ethnotaxa use categories.

Millets Use category Number of Taxa (Nt) Number of use-reports (Nur) Fica

Nutritional 14 160 0.92Nutraceutical 9 29 0.71Forage 14 151 0.91Ecological (Soil binder) 4 10 0.67Ceremony 4 7 0.50Spiritual 5 18 0.76Mechanical 7 41 0.85

Total b 57 416

aFic 5 Nur 2 Nt / (Nur 2 1), providing a value between 0 and 1, where high value indicates high rate of informant

consensus.bA taxon may be listed in several of the categories of indigenous use.


Table 3. Malayali traditional knowledge (TK) of 19 minor millet (small millet) TK ethnotaxa includingTK identification characters (ID), habitat, nutrition and ethnomedicine. Herbarium accession vouchernumbers are for the Center for Biocultural Diversity, Chennai, India and ARK collection, BiodiversityInstitute of Ontario University of Guelph.

Ethnotaxa, Species, & CBD(Herbarium AccessionNumber) Habitats Root Name Etymology and Main Uses

MalliasamaPanicum sumatrense(CBD07, CBD08, CBD09,

CBD10)

Terrestrial hill slopes. ID —Small size; copper colored seeds; goodpalatability.

Etymology —Malli (coriander), looks similar tothe coriander plant.

Nutrition —Used as a main dish and as a drink(see kottapattisama).

Ethnomedicine —Used to treat the shivers ormild hypothermia (see kottapattisama).

Kottapattisama/kottakattisama

Panicum sumatrense(CBD19, CBD20, CBD21,

CBD23)

Terrestrial hill slopes(800 MSL) andnear streams(moist soils).

ID —Castle-like inflorescence; moderateheight; late maturity; poor palatability.

Etymology – Kottampatti or kottakatti (castle-like), refers to the castle-like inflorescence.

Nutrition —Seeds are cooked in water and saltand water is then decanted and consumedas a healthy drink in the evening duringsocial gatherings. Seeds are ground intoflour to make cake or pudding.

Ethnomedicine —Used to treat the shivers ormild hypothermia. Flour is mixed withhoney and consumed directly withoutcooking to raise the body temperature. Acommon remedy in winter and rainyseasons.

VellaperumsamaPanicum sumatrense(CBD24, CBD25, CBD26,

CBD27)

Terrestrial plains andhill slopes.

ID —Large size; pale colored (pallid) seeds;excellent yield and palatability.

Etymology —Vellaperum (white big), refers tothe white color of seed grains and tallgrowth.



Sadansama or samaiPanicum sumatrense(CBD11, CBD12, CBD13,

CBD14)

Terrestrial plains andhill slopes (optimalgrowth in drysoils).

ID —Small size; spike-like inflorescence; darkbrown colored seeds; poor palatability.

Etymology —Sada (plait-like), refers to thespikelet cobs that are twisted or plait-like inappearance.



ThirikulasamaPanicum sumatrense(CBD15, CBD16, CBD17,

CBD18)

Hill slopes in very drysoils.

ID —Twisted plait-like spikelets; tolerant ofdrought; long ancestral use; poorpalatability.

Etymology —Thirikula (twisted plait likespikelets).

Nutrition —Good nutrition in ancientgenerations; used the same askottapattisama.

Ethnomedicine —Ancient medicine, same askottapattisama.


Table 3. Continued.


PerumathinaSetaria italica(CBD58, CBD59, CBD60,

CBD61, CBD62)

Terrestrial plains andwetlands.

ID —Tall plants with large grains; good yield;excellent palatability.

Etymology —Perum (large), refers to the robustgrowth of the plant and large grain size.

Nutrition —Eaten as a paste (see palanthinaabove).

Ethnomedicine —Flour is mixed with honeyand consumed to treat the shivers or mildhypothermia (see kottapattisama).

PalanthinaSetaria italica(CBD43, CBD44, CBD45,

CBD46, CBD47)


ID —Needle-like seeds; moderate height; earlymaturity; poor palatability.

Etymology —Palan (short), refers to the awnedglumes.

Nutrition —Seeds of all ‘thina’ are ground intoflour and mixed with ghee and sugar.Offerings and prayers are made to LordMuruga before eating this pasty dough-likemixture.

Ethnomedicine —The flour is mixed with honeyand consumed to treat the shivers or mildhypothermia (see kottapattisama).

KoranthinaSetaria italica(CBD38, CBD39, CBD40,

CBD41, CBD42)


ID —Inflorescence cob-like; very soft and finetextured; yellow seeds; susceptible tolodging.

Etymology —Koran (cob-like), refers to theshape of the inflorescence.

Nutrition —Seeds are ground into flour andused to make cake or pudding.

Ethnomedicine —Flour is mixed with honeyand consumed to treat the shivers or mildhypothermia.

KillanthinaSetaria italica(CBD53, CBD54, CBD55,

CBD56, CBD57)


ID —Base of inflorescence with beak-likestructure that is convenient for plucking/picking during harvesting; late maturity;good yield and palatability.

Etymology —Killan (easy picking), refers tohow the inflorescence is harvested usingbeak-like structure.



SenthinaSetaria italica(CBD68, CBD69, CBD70,

CBD71, CBD72)

Terrestrial plains (drysoils).

ID —Heavy, orange-brown seeds; susceptibleto lodging; good yield and palatability.

Etymology —Sen (red), refers to the reddishcolor of the grain.




Table 3. Continued.


KarumthinaSetaria viridis(CBD78, CBD79, CBD80,

CBD81, CBD82)

Terrestrial plains andhill slopes.

ID —Small, dark-brown seeds; moderatelodging; good yield; poor palatability.

Etymology —Karum (dark-brownish), refers tothe brown color of the grain.



ThirivaraguPaspalum scrobiculatum(CBD33, CBD34, CBD35,

CBD36, CBD37)

Terrestrial plains andhill slopes (good indry soils).

ID —Twisted, cob-like inflorescence;susceptible to lodging; early maturity; goodyield.

Etymology —Thiri (twisted), refers to thetwisted inflorescence; varagu refers to lightweighted grains.

Nutrition —Cooked grains are served as mainmeal.

Ethnomedicine— Skinned and polished grainsare cooked in water, which is decanted andmixed with salt; the warm water isconsumed as a diuretic, bowel cleanser andto reduce inflammation. Fine powders onseed husks are used as vitaminsupplements to treat weak children.Following harvest, seed husks are removedand ground into a fine powder using astone hand mill; thin layer of very fine, siltyor somewhat slimy powder on edges ofhand mill is gently rolled into small balls (131 cm) and stored in cool places to beconsumed as ‘‘vitamin balls’’ before theydry out and crack.

SengalivaraguPaspalum scrobiculatum(CBD92, CBD95, CBD96,

CBD99)

Terrestrial plains andhill slopes with drysoils.

ID —Reddish awned seeds; good yield andpalatability.

Etymology —Seng (reddish), gali (awn), varagurefers to light weighted grains.


Ethnomedicine —Diuretic qualities, anti-inflammatory and the fine powders on theseed husks are used as vitaminsupplements for treating weak children (seethirivaragu).

PeruvaraguPaspalum scrobiculatum(CBD72, CBD75, CBD78,

CBD98)

Terrestrial plains andhill slopes with drysoils.

ID —Tall plants with large, dark-brown, heavyseeds; large, thick cob-like inflorescences;late maturity; excellent yield; poorpalatability.

Etymology —Peru (large size), refers to plantheight and grain size; varagu refers to lightweighted grains.

Nutrition — Excellent nutritional quality andtaste.

Ethnomedicine —see thirivaragu.


Table 3. Continued.


SensolumSorghum bicolor(CBD63, CBD64, CBD65,

CBD66, CBD67)


ID —Very tall plants with red, heavy seeds;susceptible to lodging; late maturity; goodyield; excellent palatability.

Etymology —Sen (red), refers to the reddishcolor of the grain.

Nutrition —Seeds can be cooked whole inwater and eaten or ground into flour, whichis often used to make fermented food(fermented foods derived from the‘sensolam’ are hard to digest, butconsidered to be more nutritious than the‘vellaisolum’). ‘Sensolam’ is difficult to findin local markets as it does not grow in largeproductions; small patches are quicklyconsumed by the local tribal people.

Ethnomedicine —Mild fevers brought on bymeasles, pleurisy and smallpox are treatedby consuming the fermented foods.

VellaisolamSorghum bicolor(CBD73, CBD74, CBD75,

CBD76, CBD77)


ID —Very tall plants with white, light seeds;large compact cob-like inflorescence;susceptible to lodging; late maturity;excellent yield/palatability; producesattractive flour.

Etymology —Vella (white), refers to white colorof the grain.

Nutrition — Generally used to make fancyfoods; breads and cakes made from thisflour are soft, have a good taste and areeasily digested. Seeds can be cooked wholein water and eaten or ground into flour,which is often used to make fermentedfoods. Velsolum production capacity is highand therefore readably available in the localmarkets.

Ethnomedicine —Mild fevers brought on bymeasles, pleurisy and small pox are treatedby consuming the fermented foods.

KeshvaraguEleusine coracana(CBD28, CBD29,CBD30,

CBD31, CBD32)

Wetlands, uplands, hillslopes and alkalinesoils.

ID —Large, heavy red-brown seeds; latematurity; good yield and palatability.

Etymology —Kesh (red-color), refers to the redseeds; varagu refers to the light weight ofthe grains.

Nutrition —Seeds can be cooked whole,ground into flour, or soaked and fermentedfor use in various foods.

Ethnomedicine —Mild fevers brought on bymeasles, pleurisy and smallpox are treatedby consuming a fermented, oatmeal typemixture of keshvaragu with buttermilk.Diabetic patients are encouraged toconsume any form of food derived fromkeshvaragu. Ulcers are treated using thegreen seeds before they are fully mature,which are ground into a paste using a smallquantity of water; a milky white liquid isdecanted or pressed from the paste andconsumed on an empty stomach. The freshgreen grains from the field are directlyconsumed with or without sesame seeds toreduce stomach ulcer pain.


TAXONDNA (Meier et al. 2006) to differentiate the samples into 5 distinct groupsof barcodes representing the 5 species (Panicum sumatrense, Setaria italica,Paspalum scrobiculatum, Eleusine coracana, Sorghum bicolor). In our study, no twoindividuals of different species share identical sequences and the percentage ofcorrect identifications of all pairwise comparisons was 100% for five distinctgroups of taxa. The Malayali classification was hierarchical and fine scaled withthe primary tier congruent with the barcode analysis representing five taxa. Thesecond tier was a fine scale classification recognizing considerable intraspecificvariation, which was only partially resolved by the morphometric analysis.Although the morphometric analysis resolved 15 taxa, the Malayali informantsidentified 19 ethnotaxa from the same 174 specimens with 98% RF among theinformants.

Discussion

Recent research has revealed that tribal classifications are complex and canlead to the discovery of new species and cryptic diversity. The exploration ofbiological diversity defined by morphological and genomic variation has beenassessed alongside TK classifications within several tribal cultures. A recentproject with Irulas and Malasars of India used morphometric (Ragupathy et al.2008) and genomic analyses (Newmaster and Ragupathy 2009a, 2009b) toidentify a new species of Tripogon. Similarly, a study of Irulas and Malasarsrevealed three new cryptic species of Byophyum (Newmaster, Murugeshan, et al.2009). In our study, the Malayali classification recognized considerable variation

Figure 3. An ordination from a Detrended Correspondence Analysis (DCA) for 96 quantitativevariables (morphometric characters) of 174 specimens (classification of 12 minor millet (small millet)ethnotaxa with 3 closely related taxa).


in minor millets, 19 ethnotaxa, that were not apparent in previous research usingmorphological and genomic data, which only identified 5 species. Ourmorphometric analysis provides support for 15 of the 19 ethnotaxa.

Rengalakshmi (2005) suggested that the Malayali farmer’s classification ishierarchical and that morphology is important in the primary tier of thehierarchy as reflected in the Malayali nomenclature. Our research providesquantitative evidence to support these claims. Using DNA barcoding weprovided genomic evidence that the primary tier of the Malayali classificationsuggests a deep understanding of phylogenetic relationships among the minormillets. Our classification analysis provides quantitative support for Renga-lakshmi’s (2005) claim that the secondary tier of the Malayali’s hierarchicalclassification is based on practical use of the ethnotaxa including several sub-classifications based on cultural value.

The variation recognized and described by the Malayali includes a suite ofcharacters not recorded in previous literature. The basis of the Malayaliclassification includes at least a primary and secondary level of characters thatcan be used to distinguish all of the TK ethnotaxa (Table 3). This ethno-classification of minor millets is based on a complex interaction of phenotypes,agricultural use and food quality. Phenotypic characters include color, shape,and size of grains (figure 5), height of plants, and size and shape of the cob.Agricultural use includes the number of days to maturity, yield, resistance tolodging (plants that are not prone to tip over in wind or heavy rain), and need forwater (Table 3). Food quality includes not only taste, but also the ability to usethe plant for a variety of dishes and the digestibility of the grain. Although thebroad criteria of classification and distinctiveness of major groups based on a SK

Figure 4. A classification tree of the rbcL, matK and trnH-psbA plastid intergenic spacer for the 18ethnotaxa in the Malayali classification representing 5 species in the SK classification. Pairwise p-distance provided on scale.


classification has been defined (Renglakshmi 2005), our research provides somenew knowledge on how finer scale variation in TK ethnotaxa is differentiated.Boster (1985, 1996) argued that any cultivar that fills a gap in the morphologicalcontinuum of existing cultivars will be incorporated in the repertoire of existingvarieties. The theoretical underpinning is that the tribal farmer’s perception is avisual process and takes place prior to and independently of any assessmentregarding the use of particular landraces as a crop or food (Boster 1985).However, cultures have selected for desirable characters and this selectionprocess has been recorded in the historical nomenclature that categorizesvariation. Some clues to the ancestral history of these crops are provided in adeeper understanding of the etymology of millet terms in Tamil.

In this study, the Malayali classification reveals ethnotaxa that relate toconsiderable TK, including the nutritional and medicinal values of minormillets. In this study, the consensus and relative frequency (RF) of TK for eachindividual landrace from the interviews was high (mean RF 5 0.98 ±0.04). Theinformant consensus on nutraceutical plant usage resulted in Fic ranging from0.50 to 0.92 (Table 2). We found that use of ethnotaxa with the category‘‘ceremony’’ (e.g., millets are gifted at ceremonies such as weddings) had thelowest consensus with an Fic of 0.50, which reflects the variation found amongthe different communities although within a community, the Fic was notvariable (Fic 5 1.00). The average Fic value for both nutraceutical and nutritionalcategories was 0.82, indicating a high level of informant consensus compared tosimilar studies from other countries (Amiguet et al. 2005; Heinrich 2000). In theliterature, high informant consensus was also recorded among studies with

Figure 5. Morphological diversity of seeds among traditional millet landraces. Photograph bySubramanyam Ragupathy.


many nutraceutical categories, for instance a study with the Malasars (Fic 5

0.71) in India (Ragupathy et al. 2008) and several others focused on treatments,healing and disease: the snakebite healers of Kamba in Africa (Fic 5 0.88)(Owuor et al. 2006); the treatment of ‘mich’ or febrile diseases (Fic 5 0.80)(Teklehaymanot and Giday 2007) and respiratory disorders (Fic 5 0.88) amongInuit in Nunavut (Bernard, 1994). In a recent study, Ragupathy et al. (2008)utilized a consensus analysis of the ethnomedicine of the Malasar peopleoccupying the forests of the Velliangiri holy hills in the Western Ghats. Theresults of this research revealed the Malasars used 95 ethnotaxa for medicinaland general health purposes, with high consensus (Fic . 0.80) amongrespondents regarding many ailments (e.g., respiratory problems and jaundice).

Although all ethnotaxa were used for food, use varies according to landraceavailability and season. Traditional knowledge revealed the ethnotaxa are usedto treat a variety of illnesses and promote general good health in theircommunities. Malayali women prepare several dishes with millets for dailyconsumption and for festivals. Millets’ use as a special food for women’s prenatalcare and the landrace quality are important factors in maintaining this traditionalresource. Pregnant and lactating women in many households prefer a millet-based diet because it provides energy and prevents weight gain. They believethat during pregnancy, the consumption of millet helps to induce lactation andmaintain body temperature and energy levels after delivery. Some ethnotaxa arealso used for medicinal purposes. For example, people attempt to treat diabetesby consuming more keshvaragu in the form of fermented bread (oatmeal type).Because modakathan (balloon vine) is used to soothe the pain of rheumatoidarthritis (Ragupathy et al. 2007) but the leaves are bitter, many people eat it withkeshvaragu bread. They collect freshly grown matured seeds of keshvaragu andcrush them into a green paste with the leaves of modakathan. The paste is eatenwith bread made from various millet ethnotaxa. Other ethnotaxa, such asthirivarugu, are used as vitamin supplements. They grind the husk into a finepowder using a hand stone mill. Malayali mothers make small balls (about 1 cmin size) from the fine powder, and store them in cool places. These vitamin ballsare usually consumed before they dry, mainly by weak children and sometimesmothers suffering from malnutrition.

The ethnotaxa were also used as forage, as sillage, hay or haylage, for cows,goats and buffalo. The animals like to eat thirivaragu while the plants are youngand green. Hay from thirivarugu is not palatable for animals, but is used asmulch around beds of Capsicum, tomatoes, and eggplant, or as roof thatching(Figure 6). The mulch is prepared by making piles of this millet, and then byluring many millipedes and centipedes from the forest. The millipedes andcentipedes then convert the tough stems and leaves into rich, humus-like mulch.For human consumption, sensolum and vellaisolum are more palatable thenother millets in this study. These ethnotaxa are grown with other crops (e.g.,coriander, garlic and onion) to maintain soil fertility and prevent soil erosion.These and other ethnotaxa are used on specific sites such as steep slopes, erosionand drought conditions, that challenge traditional agriculture.

Molecular variation in millet has been resolved at the rank of subfamily, butlittle research has been completed at either the specific or intraspecific scales. A


study by Bouchenak-Khelladi et al. (2008) looked at 3 plastid regions, rbcL, matKand trnF-L, in creating phylogenetic trees of the Poaceae family. The study was ableto resolve most of the subfamily relationships and groups including tribes andgenera. This is consistent with many classification studies of plants at the rank offamily. However, the study did not focus on resolving species relationships. In ourstudy, there was resolution of the genera, which has been shown before withinonly the foxtail millets (Setaria italica; Li et al. 1995). However, it was not surprisingto see unresolved groupings of the ethnotaxa because only a few regions whereutilized: rbcL, matK, trnH-psbA and the ribosomal DNA intergenic spacer. All of theseregions are known to provide limited intraspecific resolution, particularly withintaxonomically difficult groups (Fazekas et al. 2008). We are conducting furtherresearch to explore other regions in the millet genome including single copynuclear regions, which may have faster rates of evolution. Alternatively, there areother molecular techniques (e.g., Pyrosequencing, use of random fragments, andDNA finger printing) that could be used to detect differences among the ethnotaxa.

Conclusion

Studies of biodiversity benefit from considering traditional knowledgebecause it represents a more complete picture of how humans interpretbiological variation in the landscape. This approach includes a broaderperspective than that provided by the scientific lens, and seeks to embrace theancient body of knowledge still found in cultures throughout the world. Perhapsthe greatest benefit of this approach is the assemblage of new hypotheses

Figure 6. Traditional harvesting of millet in Kolli Hills, Eastern Ghats, India. Photograph bySubramanyam Ragupathy.


concerning variation in taxa in the landscape, which provides a new perspectiveon biodiversity. We suggest that biodiversity projects should consider theperspective of both traditional and scientific knowledge when assessing,managing and conserving biodiversity.

Weber and Fuller (2008) call for further research combining linguistic andethnobotanical research. Here we do just that to explore the subtleties of Malayaliclassification and use of minor millets. Their fine-scaled classification suggeststhat tribal classifications are complex and should be considered when exploringbiodiversity. Ethnotaxa are the outcome of the complex interaction between thenatural processes of evolution, as well as the conscious or unconscious selectionby the tribal farmer over time (Teshome et al. 1997). This supports themultimechanistic ethno-taxonomic hypothesis (Newmaster et al. 2007), whichsuggests that traditional knowledge has evolved to include multifariousmechanisms in order to classify fine-scaled phenotypic variation in plantsbecause it has use and is valued within a particular community.

Weber and Fuller (2008) consider whether or not millets should play a moreprominent role in modern agricultural practices. The Malayali recognize andutilize ethnotaxa that have considerable value in their communities andrepresent intraspecific variation in minor millets that occupy specific habitatsin the landscape. These ethnotaxa would not exist in the present ecologicaldynamics if not for the tribal farmers, demonstrating the complex relationshipsbetween people and plants (Teshome et al. 1997). Our research revealedtraditional ecological knowledge for some ethnotaxa classified as one scientificspecies, Panicum sumatrense; thirikulasama is drought tolerant, while kottapat-tisama (meaning ‘‘castle hills’’) is not, but has a desirable and distinctive tastethat is served as food to high ranking royals in the local villages. We are currentlytesting the drought tolerance of these ethnotaxa to validate these TEK claims.Perhaps these intraspecific traits of Panicum sumatrense could be linked tophysiological processes and genomic diversity. The application to society isconsiderable if a minor millet ethnotaxa could be grown in drought-ridden areassuch as sub-Saharan Africa. This TK could serve society at large for manynutritional and medicinal purposes.

Acknowledgements

This research was supported by the grants to Dr. Steven G. Newmaster from theSocial Sciences and Humanities Research Council of Canada, Genome Canada through theOntario Genomics Institute, the Canadian Foundation for Innovation, the CanadianInternational Food Security Research Fund (CIFSRF) from the International DevelopmentResearch Centre (IDRC) and the Canadian International Development Agency (CIDA). Weacknowledge and give special thanks to all the tribal informants who shared theirknowledge with us. We would like to thank Dr. Manish Raizada, Dr. Aron Fazekas, IanSmith, Michael Mucci and Tannis Slimmon at the University of Guelph. We thankDirectors of the Botanical Survey of India, Central and Southern Circle for their help andsupport. A special thanks to Dr. K. Sambandhan, Women’s Arts and Science College,Karaiakal, and A. Sivakumar, S.N. Suresh, V. Balasubarmaniam and N. Nagarajan at theKongunad Arts and Science College, Coimbatore, and SRM Arts and Science College,Chennai, India for help during several field trips.


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