Broomcorn and foxtail millet were cultivated in Taiwan about 5000 years ago
© The Author(s) 2017
Received: 19 May 2016
Accepted: 26 December 2016
Published: 2 January 2017
Archaeobotanical remains of millet were found at the Nan-kuan-li East site in Tainan Science Park, southern Taiwan. This site, dated around 5000–4300 BP, is characterized by remains of the Tapenkeng culture, the earliest Neolithic culture found so far in Taiwan. A large number of millet-like carbonized and charred seeds with varied sizes and shapes were unearthed from the site by the flotation method. Since no millet grain was ever found archaeologically in Taiwan previously, this discovery is of great importance and significance. This paper is in an attempt to further analyze these plant remains for a clearer understanding of the agricultural practice of the ancient inhabitants of the Nan-kuan-li East site.
We used light and scanning electron microscopy to examine the morphological features of some modern domesticated and unearthed seeds to compare and identify the archaeobotanical remains by three criteria: caryopsis shape, embryo notch, and morphology of lemma and palea. We also developed a new methodology for distinguishing the excavated foxtail and broomcorn millet seeds.
Two domesticated millet, including broomcorn millet (Panicum miliaceum) and foxtail millet (Setaria italica), as well as one wild millet species, yellow foxtail (Setaria glauca), were identified in the unearthed seeds. Together with the millet remains, rice was also cultivated in the area. Archaeological evidence shows that millet and rice farming may have been important food sources for people living about 5000 years ago in southern Taiwan.
The discovery of a large number of carbonized and charred millet grains from the NKLE site is especially noteworthy (Tsang and Li 2015). Because no earlier millet grains were ever found archaeologically in Taiwan, this discovery is apparently of great importance and significance, assuming that prehistoric people in Taiwan at 5000 BP probably already knew how to grow these plants, and the mode of their subsistence was no longer limited to hunting and gathering. This assumption seems to gain its support not only from direct evidence of millet grains and indirect evidence such as the types of tools used for agricultural activities at that time. The most distinguishing tools are shell knives. The knives were formed by cutting and polishing shells that are relatively flat and thin. On comparison with ethnographical records, these shell knives were probably used as a kind of grain-harvesting tool (Tsang and Li 2015).
The species of millet grains have not been genetically identified. However, they look similar to foxtail millet (Setaria italica L.), which is still cultivated by Austronesian-speaking people in Taiwan. To understand the agricultural practice of the ancient inhabitants of this site, this study aimed at a detailed analysis of the morphological characteristics of the millet grains and proper identification of their taxon.
In addition to millet, the plant remains collected from archaeological studies at unearthed sites in Taiwan revealed the existence of rice grains (Tsang et al. 2006; Hsieh et al. 2011; Tsang 2012; Li 2013; Tsang and Li 2015). Because these crops were domesticated in China (Nasu et al. 2007; Lu et al. 2009), we also aimed to reveal the early agriculture products grown in Taiwan. Two types of millet, including foxtail millet and broomcorn millet (Panicum miliaceum, also known as Proso millet), were widely cultivated in several northern Chinese cultures by 8000 BP (Liu and Kong 2004; Lu et al. 2009; Zhang et al. 2012; Weisskopf et al. 2015). Archaeobotany study of the Baligang site in north central China revealed mixed farming of rice and two kinds of millet (Weisskopf et al. 2015). We aimed to determine whether similar farming was present in Taiwan thousands of years ago.
More than 120,000 millet grains were collected from the NKLE (23º6′58″N, 120º16′35″E, altitude 0.5 m) site by using a floatation method. We selected about 3000 millet seeds as analytical samples from randomly chosen excavation units of the NKLE EB-Y series, including NKLE-E5T2P6L56F1, NKLE-E5T7P0L57F2, NKLE-F4T4P1L55F4, NKLE-E3T3P2L56F1-1, and NKLE-E4T1P3L56F1-1. The samples were used to estimate the ratio of large to small seeds. Some of these samples contained lemma and palea remains and were used to study the presence of different types of millet seeds. We then used 2000 complete millet seeds from NKLE-F4T3P7L57F1 for microscopy observation and morphological analysis of the angle of the embryo notch, width of the opening of the embryo notch, and depth of the embryo notch.
Characterization of millet seeds
A digimatic caliper (Model CD-6”ASX, Mitutoyo, Japan) was used to measure the width (w), length (l) and thickness (t) of unearthed seeds. The volume of seeds was estimated by w*l*t*π*1/6. We also used a scanner to record images of the millet seeds for ImageJ analysis. The output for each seed consisted of the longest width (w) and the longest length (l). The true seed area should be calculated as w*l*0.25*π, but we used the seed area index (w*l) instead.
Results and discussion
Both carbonized and charred seeds were present in the unearthed NKLE sites
Characterization of the millet seeds at NKLE
Some archaeologists and archaeobotanists have investigated the seed remains of broomcorn and foxtail millet discovered from archaeological sites in eastern Asia or the Middle East and established criteria to distinguish the caryopses of these two millets (e.g., Nesbitt and Summers 1988; Liu and Kong 2004; Fuller and Zhang 2007). Three features were identified. The first is caryopsis shape. Grains of broomcorn millet have a pointed distal end and relatively blunt proximal end, whereas grains of foxtail millet are gently rounded at both ends (Nesbitt and Summers 1988). The second criterion is the embryo notch. The embryo notch is short and wide for broomcorn millet but longer and more narrow for foxtail millet (Knorzer 1971; Nesbitt and Summers 1988). The third criterion is the morphology of the lemma and palea. Carbonized husk fragments of broomcorn millet are smooth and glossy, and foxtail millet fragments vary from finely rugose to punctuate (Knorzer 1971; Nesbitt and Summers 1988). Several features, such as the length-to-width ratio of the grain and surface sculpture of the lemma (especially the upper lemma), have been used to distinguish foxtail millet from its wild relatives (Musil 1963; Renfrew 1973; Nasu et al. 2007).
Millet seed features
111.6 ± 11.3
37 ± 6
53 ± 7
96.5 ± 25.8
43 ± 8
48 ± 5
88.8 ± 25.6
66 ± 7
31 ± 6
72.6 ± 22.9
55 ± 11
33 ± 7
Broomcorn millet (P. miliaceum) seeds at NKLE
S. italica and S. glauca seeds at NKLE
Two kinds of the present Setaria seeds are shown in Fig. 3 (S. italica in e to h and S. glauca in i to l). The ratio of grain length to breadth for the two kinds of seed differs, with yellow foxtail having a slender shape and foxtail millet a round shape. Figure 7e and f are the same seeds, as are g and h, i and j, and k and l, one from the ventral side and the other the dorsal side. Yellow foxtail shows distinct horizontal ridges on the lemma at the dorsal sides (Fig. 3j, l). Thus, these characteristics provide reliable ways of separating the two species. Most of the smaller unearthed grains were not intact and had punctated husk fragments (Fig. 7g–l), especially on SEM (Fig. 8e–i). Because the sample chamber has to be in vacuum status for SEM observation, many of the Setaria seeds were crushed.
The unearthed millet seeds show two types: three (Fig. 7g–i) with a round shape and another three (Fig. 7j–l) with a slender shape. In addition, the three slender ones have the typical horizontal ridges on the upper lemma, the hallmark of yellow foxtail. Thus, the three round ones are foxtail millet and the other three are yellow foxtail. The SEM images of Fig. 8g–i illustrate that they should be seeds of yellow foxtail because of the horizontal ridges on the upper lemma (all dorsal sides). Figure 8d to f illustrates that these carbonized seeds are foxtail millet because of (1) the round caryopsis shape and (2) the punctate rather than smooth palea surface.
Both foxtail and yellow foxtail are still commonly consumed in India and Southern Asia (Sakamoto 1987; Kimata et al. 2000), the NKLE residents might have also consumed yellow foxtail seeds thousand years ago. More than 10,000 yellow foxtail seeds were unearthed, indicating these plants, a wild species, were abundant near the NKLE village. According to the records for aboriginal agriculture in the Ching Dynasty, rice, broomcorn and foxtail millet but not yellow foxtail were cultivated in the village (Chiang 1694). Yellow foxtail grows well in the countryside these days, according to a herbarium record in Taiwan (http://hast.sinica.edu.tw/). Thus, yellow foxtail may have been common near the NKLE village thousands of years ago. Even though the seeds were from wild species, the residents may have harvested and consumed them. Thus, their presence has implications for human subsistence practices.
Mixed farming of millet and rice at NKLE
Were the rice and millet grain in NKLE from domesticated crops or wild species? That is, we wondered whether a farming system existed. The development of the rice grain has a unique characteristic. The maturation degree of florets differs in one single panicle (i.e., the flowering time of each floret in one panicle differs from the earliest to the latest for about 10 days), and the use of shell knives in NKLE indicated that these seeds did not shatter on maturation. If the seed of the rice and two millets did not shatter, they were not wild species. In addition, the site contained 8387 and 110,860 rice and millet grain, respectively, which indicates relatively extensive cultivation of both rice and millet. Thus, we suggest the existence of rice and mixed millet agriculture about 5000 years ago in NKLE, with millet species including broomcorn and foxtail millet. There was no field ridge or clayed soil, the index of paddy field practice (Tsang and Li 2015; Li 2013). In addition, no barnyard grass seed was found in the plant remains, again indicating an upland practice (Yang et al. 2015). Thus, the field was an upland practice. Mixed farming of millet and rice was also evident in central–north China at several unearthed sites. For instance, early mixed farming of millet and rice 7800 years ago (Peiligang Culture) was discovered in the middle Yellow River Region (Zhang et al. 2012); rice, broomcorn millet and foxtail millet were present. Further analysis will be necessary to determine where the NKLE residents and crops came from.
We identified two domesticated millet, broomcorn millet (Panicum miliaceum) and foxtail millet (Setaria italica), as well as one wild millet species, yellow foxtail (Setaria glauca), at the NKLE archeological site, which, along with the previously identified rice remains, indicates that four kinds of cereals were consumed about 5000 BP during the Tapenkeng Culture. All four species were abundant in the unearthed site (i.e., 8717; 33,000; 77,000; and about 10,000 for rice, broomcorn millet, foxtail millet, and yellow foxtail seeds, respectively). Such high quantity indicates that these seeds were consumed thousands of years ago, in addition to food from fishing and hunting. The mixed millet–rice farming might have played an important role in the early civilization period. Our results indicate that NKLE was an important location for early hunter-gatherer-fisher-farmers about 5000 years ago in southern Taiwan.
CHT, KTL provided the analytical samples and participated in discussions, TZH, YCT, and YICH participated in discussions; TZH, PHF and YCT analyzed the seeds; YICH wrote the manuscript. All authors read and approved the final manuscript.
We thank the members who unearthed the Nan-kuan-Li East site, Dr. Wai-Chao Leong for seed identification, and Ms. Laura Smales (BioMedEditing, Toronto, Canada) for English editing. This work was supported by NSC 95-2420-H-001-010 to CHT, NSC99-2420-H-001-008 and NSC100-2420-H-001-014 to KTL and the Investigator Award to YICH.
The authors declare that they have no competing interests.
Open AccessThis article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.
- Chiang Y-Y (1694) Taiwan Prefecture Gazetteer. Version IGoogle Scholar
- Fukunaga K, Kawase M, Sakamoto S (1997) Variation of caryopsis length and width among landraces of foxtail millet, Setaria italica (L.) P. Beauv Jpn J Trop Agric 41:235–240Google Scholar
- Fuller DQ, Zhang H (2007) A preliminary report of the survey archaeobotany of the upper Ying Valley (Henan Province). In: Sampu S, Hpiacr H (eds) Dengfeng Wangchenggang yizhi de faxian yu yanjiu (2002–2005) [Archaeological discovery and research at the Wangchenggang site in Dengfeng (2002–2005), in Chinese]. Great Elephant, Zhengzhou, pp 916–958Google Scholar
- Hodson M, Parry DW (1982) The ultrastructure and analytical microscopy of silicon deposition in the aleurone layer of the caryopsis of Setaria italica (L.) Beauv. Ann Bot 50:221–228Google Scholar
- Hsieh JS, Hsing YIC, Hsu TF, Li JKP, Li KT, Tsang CH (2011) Studies on ancient rice—where botanists, agronomists, archeologists, linguists, and ethnologists meet. Rice 4:178–183View ArticleGoogle Scholar
- Kimata M, Ashok EG, Seetharam A (2000) Domestication, cultivation and utilization of two small millets, Brachiaria ramose and Setaria glauca (Poaceae), South India. Econ Bot 54:217–227View ArticleGoogle Scholar
- Knorzer KH (1971) Eisenzeitliche pflanzenfunde im rheinland. Bonn Jahrb 171:40–58Google Scholar
- Li KT (2013) First farmers and their coastal adaptation in prehistoric Taiwan. In: Underhill AP (ed) A Companion to Chinese archaeology. Wiley-Blackwell Press, New York, pp 612–633Google Scholar
- Liu C, Kong Z (2004) Morphological comparison of foxtail millet and broomcorn millet and its significance in archaeological identification. Kaogu (Archaeology) 443:76–83 (in Chinese)Google Scholar
- Lu H, Zhang J, Liu KB, Wu N, Li Y, Zhou K, Ye M, Zhang T, Zhang H, Yang X, Shen L, Xu D, Li Q (2009) Earliest domestication of common millet (Panicum miliaceum) in East Asia extended to 10,000 years ago. Proc Natl Acad Sci USA 106:7367–7372View ArticlePubMedPubMed CentralGoogle Scholar
- Matsuo T (1952) Genecological studies on cultivated rice. Bull Natl Inst Agr Sci D 3:1–111 (in Japanese) Google Scholar
- Matsuo T, Hoshikawa K (1993) Science of the rice plant. Morphology, vol 1. Food and Agriculture Policy Research Center, Tokyo, p 686Google Scholar
- Musil AF (1963) Identification of crop and weed seeds (Agriculture Handbook 219). Department of Agriculture, Agricultural Marketing Service, WashingtonGoogle Scholar
- Nasu H, Momohara A, Yasuda Y, He JJ (2007) The occurrence and identification of Setaria italica (L.) P. Beauv. (foxtail millet) grains from the Chengtoushan site (ca. 5800 cal B.P.) in central China, with reference to the domestication centre in Asia. Veget Hist Archaeobot 16:481–494View ArticleGoogle Scholar
- Nesbitt M, Summers GD (1988) Some recent discoveries of millet (Panicum miliaceum L. and Setaria italica (L.) P. Beauv.) at excavations in Turkey and Iran. Anatol Stud 38:85–97View ArticleGoogle Scholar
- Renfrew J (1973) Palaeoethnobotany. The prehistoric food plants of the near East and Europe. Butler & Tanner, LondonGoogle Scholar
- Sakamoto S (1987) Origin and dispersal of common millet and foxtail millet. Jpn Agr Res Q 21:84–89Google Scholar
- Schmuller J (2013) Statistical analysis with excel for dummies. Wiley, New YorkGoogle Scholar
- Schneider CA, Rasband WS, Eliceiri KW (2012) NIH Image to ImageJ: 25 years of image analysis. Nat Methods 9:671–675View ArticlePubMedGoogle Scholar
- Tsang CH (2005) Recent discoveries at a Tapenkeng Culture site in Taiwan: Implications for the problem of Austronesian origins. In: Blench R, Sanchez-Mazas A (eds) Sagart L. The peopling of east Asia Routledge Curzon, London, pp 63–73Google Scholar
- Tsang CH (2012) Issues relating to the ancient rice and millet grains from the archaeological sites in Tainan Science Park. Jour Chin Diet Cult 8:1–24 (in Chinese with English abstract) Google Scholar
- Tsang CH (2013) New archaeological discoveries at Baxiandong and the issues related to the chronology and affinities of the Paleolithic culture in Taiwan. In: Chen KT, Tsang CH (eds) The 4th international conference on sinology New lights on East Asian archaeology. Academia Sinica, Taipei, pp 379–416 (in Chinese with English abstract) Google Scholar
- Tsang CH, Li KT (2015) Archaeological heritage in Tainan Science Park of Taiwan. National Museum of Prehistory, TaitungGoogle Scholar
- Tsang CH, Li KT, Chu CI (2006) The Ancestors’ Footprint: The Discoveries of the Rescue Archaeology in Tainan Science Park. Tainan County Cultural Bureau, Taiwan (in Chinese with English abstract) Google Scholar
- Weisskopf A, Deng Z, Qin L, Fuller DQ (2015) The interplay of millets and rice in Neolithic central China: integrating phytoliths into the archaeobotany of Baligang. Archaeol Res Asia 4:36–45View ArticleGoogle Scholar
- Yang X, Fuller DQ, Huan X, Perry L, Li Q, Li Z, Zhang J, Ma Z, Zhuang Y, Jiang L, Ge Y (2015) Barnyard grasses were processed with rice around 10,000 years ago. Sci Rep 5:16251View ArticlePubMedPubMed CentralGoogle Scholar
- Zarnkow M, Mauch A, Back W, Arendt EK, Kreisz S (2007) Proso millet (Panicum miliaceum L.): evaluation of the microstructure changes in the endosperm during malting by using scanning-electro and confocal laser microscopy. J Inst Brew 113:355–364View ArticleGoogle Scholar
- Zhang J, Lu H, Gu W, Wu N, Zhou K, Hu Y, Xin Y, Wang C (2012) Early mixed farming of millet and rice 7800 years ago in the Middle Yellow River region, China. PLoS ONE 7:e52146View ArticlePubMedPubMed CentralGoogle Scholar