Analysis of Starch Grains Produced in Select Taxa Encountered in Southwest Asia Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 135 Research Communica on Asian taxa except for Triticum durum Desf. (durum wheat), Triticum compactum L. (club wheat), Hordeum distichon L. (two-rowed barley), and Vicia ervilia (L.) Willd. (bitter vetch) and numerous wild grasses (Henry et al. 2011; Henry and Piperno 2008; Piperno et al. 2004). Unfortunately, not all of these publica- tions provide detailed descriptions of the taxa they discuss despite their pioneering endeavors. Having thorough descriptions of starch producing taxa included in publications provides information about which taxa do and do not produce starches thereby helping researchers in identifying their own archaeo- logical starch materials. Southwest Asian taxa are also described in archaeological publications from other parts of the world either because these taxa were introduced to the region or because their natural distribution overlaps with that of Southwest Asia. For example, Yang and Perry (2013) analyze starch grains from the tribe Triticeae that grow in north China. These taxa include introduced Southwest Asian domesticates, such as Triticum aestivum L. (bread wheat), and wild taxa that are native to both China and Southwest Asia, such as Aegilops tauschii Coss. (Tauschs goatgrass). A list of publications detailing starch grains from Poaceae taxa that grow in Southwest Asia can be found in Table 1. Introduction Recent starch grain analysis in Southwest Asia has provided insight into new areas of research such as beer brewing in ancient Egypt (Samuel 1996) and the diets of middle Holocene farmers (Henry and Piperno 2008), Upper Paleolithic hunters and gatherers (Piperno et al. 2004), and Neanderthals (Henry et al. 2011). Despite these promising strides in archaeological research, much remains to be done in regards to discovering which plants produce starches in Southwest Asia and whether or not these starch grains can be used to aid archaeological and paleoeco- logical endeavors. In this paper I seek to understand the research potential of archaeological starch grain research in Southwest Asia by: 1) centralizing where starch grain information about Southwest Asian taxa can be found; 2) examining 64 previously unstudied taxa from 22 families to assess their production patterns; and 3) examining the diagnostic potential of starches found in these new taxa if present. Organization of Comparative Southwest Asian Publications The most comprehensive and detailed information about Southwest Asian taxa are embedded within archaeological site reports from this region. These publications provide the best source of data because they cover almost all of the domesticated Southwest Analysis of Starch Grains Produced in Select Taxa Encountered in Southwest Asia Thomas C. Hart Author address: Department of Anthropology, University of Texas, 2201 Speedway C3200, University of Texas at Aus n, Aus n, Texas, 78712, U.S.A. Email: thomas.hart@utexas.edu Received: September 13, 2014 Volume: 5:135‐145 Published: December 15, 2014 © 2014 Society of Ethnobiology Abstract: Starch grain analysis is a rapidly growing field of archaeological research in Southwest Asia. However, much work s ll remains regarding which taxa produce starch grains that can be iden fied in the archaeological record. In this paper, I centralize what is known about starch produc on pa erns within regional flora and analyze 64 previously unstudied taxa from 22 families. The results of this study demonstrate that descrip ons of starch grains from Southwest Asian taxa are sca ered between archaeological and plant and food science publica ons. Ten of the species examined in this study, most of whom are grasses, produced starch grains that can be iden fied at varying taxonomic levels. Keywords: Paleoethnobotany, Starch Grains, Southwest Asia Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 136 Research Communica on Table 1. Poaceae (Gramineae) taxa that have been published. Note, not all of these publica ons include descrip ons of op cal proper es. Genus and Species Source Aegilops bicomis (Forsk.) Jaub. & Spach. Henry et al., 2011 Aegilops caudate auct. Reichert, 1913 Aegilops geniculata Roth Piperno et al., 2004 Aegilops peregrina (Hackel) Maire et Weiler Piperno et al., 2004 Aegilops speltoides Tausch Henry et al., 2011 Aegilops truincialis L. Reichert, 1913 Agropyron cristatum (L.) Gaertn. Reichert, 1913 Agropyron rigidum (Schrad.) P. Beauv. Reichert, 1913 Agros s spica‐ven L. Reichert, 1913 Aira caespitosa L. Reichert, 1913 Alopecurus arundinaceus Poir Piperno et al., 2004 Alopecurus geniculatus L. Reichert, 1913 Alopecurus utriculatus Banks & Sol. Piperno et al., 2004; Reichert, 1913 Alopecurus pratensis L. Reichert, 1913 Avena barbata Po ex Link Piperno et al., 2004 Avena sterilis L. Henry et al., 2011 Brachypodium distachyon (L.) P.Beauv. Piperno et al., 2004 Bromus brachystachys Hornung Reichert, 1913 Bromus pseudobrachystachys H. Scholz Piperno et al., 2004 Bromus squarrosus L. Reichert, 1913 Gastridium ventricosum (G. australe) (Gouan) Schinz & Thell. Piperno et al., 2004; Reichert, 1913 Hordeum bulbosum L. Piperno et al., 2004 Hordeum glaucum Steudel Henry et al., 2011; Piperno et al., 2004 Hordeum hexas chon L. Henry et al., 2011 Hordeum marinum Huds. Piperno et al., 2004 Hordeum sa vum var. (Champion) Jess. Reichert, 1913 Hordeum spontaneum L. Henry et al., 2011; Piperno et al., 2004 Hordeum vulgare L. Henry et al., 2011; Reichert, 1913 Koeleria macrantha (Ledeb.) Schult. Messner, 2011 Lolium mul florum Lam. Piperno et al., 2004 Lolium rigidum Gaudin Piperno et al., 2004 Lolium temulentum var. speciosum L. Reichert, 1913 Phalaris minor Retz. Piperno et al., 2004 Phalaris paradoxa L. Piperno et al., 2004 Piptatherum holciforme (M.Bieb.) Roem. & Schult. Piperno et al., 2004 Poa pratensis L. Messner, 2011 Poa nemoralis L. Messner, 2011 Puccinellia distans (Jacq.) Parl. Piperno et al., 2004 Puccinellia gigantea (Grossh.) Grossh. Piperno et al., 2004 Secale cereale L. Reichert, 1913 Secale cereale var. MammothWinter L. Reichert, 1913 Secale cereale var. Spring L. Reichert, 1913 Secale cereale ssp. ancestrale L. Henry et al., 2011 Secale vavilovii Grossh. Henry et al., 2011 (con nued on next page) Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 137 Research Communica on Messner (2011) analyzes starch grains in seeds and underground storage organs (USOs) produced by taxa that grow in the Delaware River valley, USA. Two of these taxa, Typha latifolia L. (cattail) and Cyperus esculentus L. (yellow nutsedge), are also found in Southwest Asia (Davis 1965; Migahid 1988). Finally, a few Southwest Asian taxa are discussed in experimental archaeological publications where researchers examine how food-processing activities affect starch grain morphology and how these changes can be detected archaeologically (Ge et al. 2010; Henry et al. 2009). Food and plant science research on Southwest Asian taxa is extensive, focusing on understanding the chemical and physical attributes of main Southwest Asian domesticates: Triticum spp. (wheat), Hordeum spp. (barley), Secale spp. (rye), Vicia faba L. (faba bean), Lens culinaris Medikus (lentil), Pisum sativum L. (pea), and Cicer arietinum L. (chickpea). Other domesticates, such as Vicia sativa (common vetch) and Vicia ervilia (bitter vetch), have received little attention. Reichert (1913) provides the most comprehensive analysis of starch grains produced by taxa and remains one of the seminal publications used by many paleoethnobotanists. In this publication, he reviews the state of starch grain research at the beginning of the 20th century, discusses the chemical and physical properties of specific taxa, and provides an assessment on how these taxa can be identified based on their chemical and physical characteristics. Many of the taxa that he describes are found in Southwest Asia and can be referenced by comparing the list of species he covers with the species listed in one of the regional floras such as the Flora of Turkey and East Aegean Islands (Davis 1965). Materials and Methods Selecting species for analysis Sixty-four species representing 22 families that currently grow in Syria were collected from Professor Joy McCorriston’s extensive Southwest Asian herbari- um collection at Ohio State University. The 64 species were subdivided into their constituent parts resulting in eighty-two samples (Tables 2 and 3). These samples included seeds, pericarps, synconia, legumes, and legume capsules. In this study, the generic term “seed” is used for simplicity. No leaves, stems, or small roots (con nued from previous page) Tri cum aegilopoides (T. monococcum subsp aegilopoides) (Link) Balansa ex Körn. Henry et al., 2011 Tri cum aes vum (T. aes vum ssp aes vum) L. Henry et al., 2011; 2009 Tri cum dicoccum (T. turgidum ssp. dicoccum) Schrank ex Schübl Reichert, 1913 Tri cum dicoccoides Schrank ex Schübl Piperno et al., 2004 Tri cum monococcum L. Reichert, 1913 Tri cum monococcum subsp. aegilopoides Henry et al., 2011 Tri cum sa vum var.dicoccum (Schrank) Reichert, 1913 Tri cum sa vum var.vulgare Reichert, 1913 Tri cum turgidum Desf. Henry et al., 2011; Reichert, 1913 Tri cum urartu Tumanian ex Gandilyan Henry et al., 2011 Vulpia persica (Boiss. & Buhse) Krecz. & Bobrov Piperno et al., 2004 Bibliography for Poaceae of Southwest Asia: Henry, A. G., A. S. Brooks, D. R. Piperno. 2011. Microfossils in Calculus Demonstrate Consump on of Plants and Cooked Foods in Neanderthal Diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings of the Na onal Academy of Sciences 108:486‐491. Henry, A. G., H. F. Hudson, and D. R. Piperno. 2009. Changes in Starch Grain Morphologies from Cooking. Journal of Ar‐ chaeological Science 36:915–922. Messner, T. C. 2011. Acorns and Bi er Roots: Starch Grain Research in the Prehistoric Eastern Woodlands. University of Alabama Press, Tuscaloosa, AL. Piperno, D. R., E. Weiss, I. Holst, and D. Nadel. 2004. Processing of Wild Cereal Grains in the Upper Palaeolithic Revealed by Starch Grain Analysis. Nature 430:670‐673. Genus and Species Source Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 138 Research Communica on were analyzed because they rarely produce large storage starch grains (Haslam 2004). Underground storage organs of important wetland taxa from the Cyperaceae family (Ryan 2011) were not available for analysis because they are difficult to store and are rarely found in herbarium collections. Processing the samples Samples were cleaned according to the protocol outlined by Pearsall (2000: 436–437), cut into small pieces using a sterile scalpel, or gently crushed using a sterile mortar and pestle. Very little pressure was applied when using the mortar and pestle to minimize potential damage to the starch grains. Two drops of a one to one glycerol/distilled water mix were placed on a 25 × 75 × 1mm microscope slide for each compara- tive sample. This medium was chosen, as opposed to a more permanent medium such as Permount or Entellen, in order to allow potential starch grains to be rotated when examined. The sample was gently covered with a microscope cover slip and the edges were sealed using finger nail polish and allowed to dry before being examined. Recording methods Samples were examined at 500× magnification using a Zeiss AxioStar Plus microscope. Each starch grain was given an identification number, described according to terms defined in the International Code for Starch Grain Nomenclature (ICSN 2014) and measured using NIS Elements software. Photos of individual starch grains were taken at the Environ- mental Archaeology Lab at University of Texas. In order to minimize researcher bias, starch grains were chosen at random for description by using the random number generator function within Excel to provide x and y coordinates on the microscope stage. Fifty simple or half compound starch grains were described and photographed when present for each sample. Compound and aggregate starch grains were noted although excluded from the total count because clustering would often obscure their optical attributes making the individual starch grains difficult to describe and quantify. Starches less than five microns were typically omitted because their optical attributes were often hard to distinguish. Starch grains less than five microns in length were only counted in instances where they constituted the bulk of the starch grains produced. Results Ten of 64 species produced starch grains. All of the starches were produced in the seeds with the excep- tion of Moringa peregrina (Forssk.) Fiori (Yusor tree) that concentrated its starch in the pericarp (Table 2). The 54 species that did not produce starch grains were from wild taxa that were related to the domesticated grains and legumes or from other types of domesticat- Figure 1. Transmi ed and polarized views of starch at 400 × magnifica on from: a, b) Cyperus esculentus; c, d) Vicia ervil‐ ia; and e, f) Moringa peregrina. Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 139 Research Communica on ed taxa (Table 3). These taxa produced seeds that were very small and contained almost no starches. Cyperus esculentus The starches formed within Cyperus esculentus (yellow nutsedge) seeds have a mean length of four microns, range in size from one to eight microns and are mostly ovoid in shape (Figure 1a, b). They differ markedly in size and shape from the starches pro- duced in the tuber or root-stock, which have an average length of 12 to 14mm and are conical to oval in shape (Reichert 1913). The seed starches are diagnostic to Cyperaceae because of their size and rounded, oval, compressed lenticular, angular, or polyhedral shapes that are commonly associated with other Cypereraceae seeds discussed in Reichert (1913). Vicia ervilia Vicia ervilia (bitter vetch) starches have an average length of 16mm and range in size from five to 27mm (Reichert 1913) (Figure 1c, d). Vicia ervilia starches from seeds can be identified to the family level because they exhibit what Reichert (1913) refers to as “bean type” features (spherical to ovoid in shape, half to as broad as long, slightly compressed with a distinct longitudinal cleft) that are characteristic of the Fabaceae (Leguminosae) family. These starches are mostly ovoid to elliptical and reniform shape and have deep longitudinal clefts. Moringa peregrina Moringa peregrina starches are mostly angular rounded, range in size from four to 27mm, and have an average length of ten microns (Figure 1. e, f). It is hard to determine if these starches are diagnostic because there are no close relatives discussed in Reichert (1913) or any of the other publications mentioned in this study. More studies should be conducted on Moringa and closely related taxa to determine the diagnostic status of these starches. It is important to note that starch grains were extracted from the pericarp of the M. peregrina sample, and not the seed. This species suggests that tissues surrounding the seed, and not just the seed itself, need to be studied when conducting comparative starch grains research. Aegilops crassa, A. triaristata, Hordeum distichon, Triticum durum, and T. compactum The seeds from the species Aegilops crassa (Persian goatgrass) (Figure 2a, b), A. triaristata (three awn-goat Family Genus/Species Plant Part Cyperaceae Cyperus esculentus L. Seed Fabaceae (Leguminosae) Vicia ervilia Legume Moringaceae Moringa peregrina Pericarp Poaceae (Gramineae) Aegilops crassa Boiss Seed Aegilops triaristata Willd. Seed Aegilops vavilovii (Zhuk.) Chennav. Seed Hordeum dis chon L. Seed Pennisetum americanum (L.) Leeke Seed Tri cum compactum Host. Seed Tri cum durum Desf. Seed Table 2. Taxa that produced starch grains in abundance in this study. Figure 2. Transmi ed and polarized views of starch at 400× magnifica on from: a, b) Aegilops crassa; c, d) Ae‐ gilops triaristata; e, f) Aegilops vavilovii; g, h) Hordeum dis chon; i, j) Pennisetum americanum; k, l) Tri cum durum; and m, n) Tri cum compactum. Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 140 Research Communica on Table 3. Taxa that did not produce abundant starch grains. Family Genus/Species Plant Part Anacardiaceae Pistacia atlan ca Desf. Seed Pistacia khinjuk Stocks Seed Pistacia palaes na Boiss. Seed Pistacia terebinthus L. Seed Rhus coriaria L. Seed Apiaceae (Umbelliferae) Bupleurum lancifolium Hornem. Seeds Cuminum cyminum L. Seeds Arecaceae (Palmae) Phoenix dactylifera L. Seed Asteraceae (Compositae) Carthamus nctorius L. Seed Guizo a abyssinica (L.) Cass. Seed Helianthus annus L. Seed Notobasis syriaca (L.) Cass. Seed Onopordum illyricum L. Seed Onopordum palaes num Eig. Seed Silybum marianum (L.) Gaertn Seed Euphorbiaceae Chorozophora nctoria (L.) A. Juss. Seed Fabaceae (Leguminosae) Acacia farnesiana (L.) Willd. legume Acacia nilo ca (L.) Delile Seed Hymenocarpos circinnatus (L.) Savi Legume Prosopis farcta Banks & Sol.) J. F. Macbr. Legume capsule Trigonella foenum‐graecum L. Legume Trigonella monantha C. A. Mey. Legume Trigonella stellata Forssk. Legume Geraniaceae Erodium ciconium (L.) L'Hér. ex Aiton Seed Erodium gruinum (L.) L'Hér. ex Aiton Seed Malvaceae Malva parviflora L. Seed Moraceae Ficus carica L. Synconium, seed Moringaceae Moringa peregrina (Forssk.) Fiori Seed Oleaceae Olea europaea L. Pericarp, seed Pedaliaceae Sesamum indicum L. Seed Poaceae Bromus scoparius Scop. Seed Polygonaceae Polygonum patulum M. Bieb Seed Polygonum venan anum Clemen Seed Ranunculaceae Adonis dentata Delile Seed Rhamnaceae Rhamnus palaes nus Boiss. Pericarp, seed Zizyphus spina‐chris (L.) Desf. Exocarp, pericarp, seed Rosaceae Amygdalus arabica (Oliv.) Pericarp, seed Amygdalus communis L. Pericarp, seed Amygdalus orientalis Mill. Exocarp, pericarp, seed (con nued on next page) Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 141 Research Communica on grass) (Figure 2c, d), Hordeum distichon (two-rowed barley) (Figure 2g, h), Triticum durum (durum wheat) (Figure 2k, l), and T. compactum (club-wheat) (Figure 2m, n) in this study all exhibit features that are diagnostic of the tribe Triticeae within the Poaceae (Gramineae) family. In general, starch grains from this tribe have simple, lenticular, oval, kidney (reniform) or dicoid in shapes with small reticulate surface depressions (Piperno et al. 2004; Yang and Perry 2013). The five Triticeae taxa that yielded abundant starch grains within this study exhibited all of these features Seed starch grains from Aegilops, Hordeum, and Triticum taxa (AHT) and the Triticeae tribe are also much larger in general than the seed starch grains from non-Triticeae taxa. This feature can be used to identify individual starches at least to the tribe level when shape and size attributes are analyzed together. The mean length for the Poaceae starch grains observed in this study follow the pattern observed by Piperno et al. (2004) where AHT taxa can be distin- guished from other grass taxa, such as the Pennisetum americanum, based on their overall large size (Table 4). The average length of the 18 AHT seed starch grain taxa in Table 4 with a sample size of 50 is 17.7mm with a standard deviation of 5.7mm. This length is well above the average length of the 15 non-Triticeae with an average of 5.1mm and a standard deviation 2.6mm. Recent work by Yang and Perry (2013) on 38 grass species from China supports this hypothesis and goes one step further, suggesting that all members of the tribe Triticeae produce larger starches relative to other Poaceae. The one non-Triticeae grass in this study that yielded abundant seed starch, Pennesitum americanum yielded semi-compound to compound, flat, angular, or irregular shaped starch grains (Figure 2. i, j). This compares well with other studies of non-Triticeae grasses such as Bromus sp. and Pipatherum sp. where similar features were observed (Piperno et al. 2004). Discussion and Conclusions Chemical and physical properties of starch grains from over 100 species from Southwest Asia have been published in archaeological reports and food and plant science literature. An additional 64 species were examined here, ten of which produced abundant starch grains in their seeds and pericarps that are diagnostic at the tribe, family, and potentially genus and species level. This project adds to the growing body of knowledge regarding archaeological starch grain analysis in Southwest Asia by centralizing the published comparative literature for this region and describing the starches produced in domesticated and wild taxa. The starches from Cyperus esculentus seeds are Crataegus aronia (L.) DC Pericarp, seeds Prunus domes ca L. Seeds Prunus mahaleb L. Seeds Prunus persica (L.) Stokes Pericarp, seed Rosa canina L. Pericarp/seed, seeds Rosa phoenicea Boiss. Pericarp, seeds Sarcopoterium sinposum (L.) Spach. Seeds Rubiaceae Asperula arvensis L. Seeds Coffea arabica L. Beans Galium tricornutum Dandy Seeds Solanaceae Hyscamus mu cus L. Seed Physalis alkekengi L. Seed Physalis angulata L. Pericarp Solanum sepicula Dunal Seed, fruit Ur caceae Ur ca pilulifera L. Seed Zygophllaceae Balanites aegyp aca (L.) Delile Exocarp, pericarp, seed Family Genus/Species Plant Part (con nued from previous page) Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 142 Research Communica on Table 4. Mean (±s.d.) length (mm) and range for Poaceae starch grains divided by subfamily and tribe. Subfamily Tribe Genus/species Mean Range n Source Panicodae Paniceae Pennisetum americanum (L.) Leeke 5.7 (1.4) 3–10 50 This study Pooideae Aveneae Alopecurus arundinaceus Poir. 4 (0.9) 2–8 50 Piperno et al 2004 Alopecurus utriculatus Banks & Sol. 5 (1.5) 2–8 50 Piperno et al 2004 Avena barbata Po ex Link 12 (2.9) 6–18 50 Piperno et al 2004 Gastridium ventricosum (Gouan) Schinz & Thell. 4 (1.0) 2–6 50 Piperno et al 2004 Phalaris minor Retz. <2.0 ‐ 50 Piperno et al 2004 Phalaris paradoxa L. <4.0 ‐ 50 Piperno et al 2004 Brachypodieae Brachypodium distachyon (L.) P.Beauv. 9 (2.2) 4–16 50 Piperno et al 2004 Bromeae Bromus pseudobrachystachys H. Scholz 5 (1.4) 4–8 50 Piperno et al 2004 Poeae Lolium mul florum Lam. <6.0 ‐ 50 Piperno et al 2004 Lolium rigidum Gaudin <6.0 ‐ 50 Piperno et al 2004 Puccinellia distans (Jacq.) Parl. <4.0 ‐ 50 Piperno et al 2004 Puccinellia gigantea (Grossh.) Grossh. <4.0 ‐ 50 Piperno et al 2004 Vulpia persica (Boiss. & Buhse) Krecz. & Bobrov <2.0 ‐ 50 Piperno et al 2004 S peae Piptatherum holciforme (M.Bieb.) Roem. & Schult. 3 (1.0) 2–4 50 Piperno et al 2004 Tri ceae Aegilops crassa Boiss 16 (7.6) 5–31 50 This study Aegilops geniculata Roth 21 (6.4 ) 10–36 50 Piperno et al 2004 Aegilops peregrina Hack. 25 (8.0) 12–52 50 Piperno et al 2004 Aegilops speltoides Tausch 22 (4.5) 10–32 50 Henry et al 2011 Aegilops triaristata Willd. 10 (3.4) 5–20 50 This study Aegilops vavilovii (Zhuk.) Chennav. 13 (6.2) 5–35 50 This study Hordeum bulbosum L. 17 (3.7) 10–24 50 Piperno et al 2004 Hordeum bulbosum (with lamellae only) 21 (1.6) 18–24 50 Piperno et al 2004 Hordeum dis chon L. 11 (2.7) 5–18 50 This study Hordeum glaucum Steudel 18 (3.5) 10–30 39 Henry et al 2011 Hordeum glaucum Steudel 18 (3.9) 8–24 50 Piperno et al 2004 Hordeum glaucum (with lamellae only) 22 (1.4) 18–26 50 Piperno et al 2004 (con nued on next page) Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 143 Research Communica on distinct from the starches produced in its tubers but are similar to the seeds of other related taxa within the Cyperaceae family making them diagnostic to this family. Vicia ervilia starches exhibit “bean type” features and can be identified to the genus and species level due to their small size and Fabaceae (Leguminosae)-like properties. Although the diagnos- tic ability of Moringa peregrina starches remains unclear, their production in the pericarp, and not the seed challenges assumptions originally made in this project, and in the general literature, about perceived starch production in particular plant parts and illustrates the importance of testing every part of a plant when possible. Finally, the Poaceae taxa in this study can be distinguished from each other at the tribe level by size and overall shape. Centralization of information about taxa that produce starch grains will help specialists narrow down identification of unknown starch grains encountered in the archaeological record. The discovery of starch grains within important domesti- cated taxa such as Hordeum distichon, Triticum durum, and wild taxa such as Cyperus esculentus provides a clearer understanding of what can be identified within Southwest Asia and within these families and genera. There are many avenues of comparative starch grain research that can be pursued to better aid archaeologists in their reconstruction of plant use in Southwest Asia. With a few exceptions, very little research has been conducted on starch grains pro- duced by underground storage organs such as bulbs, corms, rhizomes, and tubers (Henry et al. 2009, 2011; Messner 2011; Piperno et al. 2004; Reichert 1913; Yang and Perry 2013). Macrobotanical and phytolith evidence suggests that wetland taxa played an important role as a source of food in Southwest Asia during the Epipaleolithic (Wollstonecroft et al. 2008), Pre-Pottery Neolithic (Balbo et al. 2012), Pottery Neolithic (Rosen 2005), and Ubaid (Kennett and Kennett 2006) periods. Aside from the research by Hather (1991, 1993), very little work has been conducted to establishd criteria for identifying underground storage organs at archaeological sites. Recovering and identifying starch grains associated with USO’s would open a whole new avenue of research into wild resource exploitation, complement existing datasets, and allow for archaeologists to explore new topics through the analysis of starches contained in artifact residues and dental calculus. The research on Triticeae taxa from China (Yang and Perry, 2013) and taxa from the Delaware River Valley, USA (Messner 2008, 2011) are excellent examples of how a regional synthesis can lead to the construction of standardized dichotomous keys for a region. In both of these papers, the researchers develop an easy to use dichotomous key that allows for quick identification of archaeological starch grains. Further research into starch grain production patterns of other taxa found in Southwest Asia and the identification of Southwest Asian taxa discussed in Reichert (1913) would eventually lead to the develop- Pooideae Tri ceae Hordeum hexas chon L. 20 (3.5) 10–30 52 Henry et al 2011 Hordeum marinum Huds. 10 (1.8) 6–14 50 Piperno et al 2004 Hordeum spontaneum L. 18 (3.8) 12–30 27 Henry et al 2011 Hordeum spontaneum L. 20 (4.7) 10–26 50 Piperno et al 2004 Hordeum spontaneum (with lamellae only) 28 (2.9) 18–26 50 Piperno et al 2004 Secale vavilovii Grossh. 25 (4.2) 15–36 50 Henry et al 2011 Tri cum aes vum L. 24 (4.4) 15–35 52 Henry et al 2011 Tri cum compactum Host. 12 (4.9) 5–22 50 This study Tri cum dicoccoides Schrank ex Schübl. 17 (6.1) 8–30 50 Piperno et al 2004 Tri cum durum Desf. 11 (4.0) 5–23 50 This study Tri cum monococcum subsp. aegilo‐ poides (Link.) Thell. 15 (1.7) 10–20 46 Henry et al 2011 Subfamily Tribe Genus/species Mean Range n Source (con nued from previous page) Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 144 Research Communica on ment of a dichotomous key and the establishment of regional diagnostic starch grain types that archaeologists could use in this important area of the world. Acknowledgements I owe Professor Joy McCorriston a massive debt of gratitude for allowing me to study and use her Near Eastern macrobotanical comparative collection for this project. I would also like to thank Masoumeh Kimiaie and Matthew Senn for their assistance and hospitality while working in the McCorriston Laboratory at Ohio State University. This project would not have been possible without the aid of my undergraduate assistants Kathleen Hammel, Andrew Ritz, Joyce Fountain, Stephen McKay, and Jessica Lundquist. Without them, I never would have been able to create and study such a wonderful starch grain comparative collection. Finally, thank you to my committee members Dr. Alexia Smith, Dr. Natalie Munro, Professor Deborah Pearsall, Professor Gil Stein, and Professor Sally McBrearty, as well as the anonymous reviewers for the helpful comments on the manuscript. Declarations Permissions: None declared. Sources of funding: National Science Foundation Dissertation Improvement Grant. Conflicts of interest: None declared. References Cited Balbo, A. L., E. Iriarte, A. Arranz, L. Zapata, C. Lancelotti, M. Madella, L. Teira, M. Jiménez, F. Braemer, and J. J. Ibáñez. 2012. Squaring the Circle. Social and Environmental Implications of Pre-Pottery Neolithic Building Technology at Tell Qarassa (South Syria). PLoS One 7:e42109-e42109. Cummings, L. S., and A. Magennis. 1997. A phytolith and starch record of food and grit in Mayan human tooth tartar. In Estado Actual De Los Esudios De Fitolitos en Suelos Y Plantas, edited by A. Pinilla, J. Juan-Tresserras, and M. J. Machado, pp. 211-218. Centro de Ciencias Medioambientales, Madrid. Davis, P. H. ed. 1965. Flora of Turkey and the East Aegean Islands, Vol 1. Edinburgh University Press, Edinburgh. Duncan, N. A., D. M. Pearsall, and R. A. Benfer, Jr. 2009. Gourd and Squash Artifacts Yield Starch Grains of Feasting Foods from Preceramic Peru. Proceedings of the National Academy of Sciences 106:13202 -13206. Ge, W., L. Liu, X. Chen, J. Zhengyao. 2010. Can Noodles Be Made From Millet? An Experimental Investigation of Noodle Manufacture Together with Starch Grain Analyses. Archaeometry 53:194-204. Gott, B., H. Barton, D. Samuel, and R. Torrence. 2006. Biology of Starch, In Ancient Starch Research, edited by R. Torrence, and H. Barton, pp. 35-45. Left Coast Press, Walnut Creek, CA. Haslam, M. 2004. The Decomposition of Starch Grains in Soils: Implications for Archaeological Residue Analyses. Journal of Archaeological Science 31:1715-1734. Hather, J. G. 1991. The Identification of Charred Archaeological Remains of Vegetative Parenchy- mous Tissue. Journal of Archaeological Science 18:661- 675. Hather, J. G. 1993. An Archaeobotanical Guide to Root and Tuber Identification: Europe and South West Asia. Oxbow Books Limited, Oxford. Henry, A. G., A. S. Brooks, and D. R. Piperno. 2011. Microfossils in Calculus Demonstrate Consumption of Plants and Cooked Foods in Neanderthal Diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings of the National Academy of the Sciences 108:486-491. Henry, A. G., H. F. Hudson, and D. R. Piperno. 2009. Changes in Starch Grain Morphologies from Cooking. Journal of Archaeological Science 36:915-922. Henry, A. G., and D. R. Piperno. 2008. Using Plant Microfossils from Dental Calculus to Recover Human Diet: A Case Study from Tell al-Raqā'i, Syria. Journal of Archaeological Science 35:1943-1950. Horrocks, M., G. Irwin, M. Jones, and D. Sutton. 2004. Starch Grains and Xylem Cells of Sweet Potato (Ipomoea batatas) and Bracken (Pteridium esculentum) in Archaeological Deposits from North- ern New Zealand. Journal of Archaeological Science 31:251-258. ICSN. 2014. The International Code for Starch Nomenclature [WWW Document]. Foundation for Archaeobotanical Research in Microfossils. Availa- ble at: http://fossilfarm.org/ICSN/Code.html. Accessed on February 2, 2014. Ethnobiology Le ers. 2014. 5: 135‐145. DOI: 10.14237/ebl.5.2014.251. 145 Research Communica on Kennett, D., and J. Kennett. 2006. Early State Formation in Southern Mesopotamia: Sea Levels, Shorelines, and Climate Change. The Journal of Island and Coastal Archaeology 1:67-99. Li, M., X. X. Yang, H. Wang, Q. Wang, X. Jia, and Q. Ge. 2010. Starch Grains from Dental Calculus Reveal Ancient Plant Foodstuffs at Chenqimogou Site, Gansu Province. Science China Earth Sciences 53:694-699. Messner, T. C. 2008. Woodland Period People and Plant Interactions: New Insights from Starch Grain Analysis. Unpublished Doctoral Dissertation, Department of Anthropology, Temple University, Philadelphia, PA. Messner, T. C. 2011. Acorns and Bitter Roots: Starch Grain Research in the Prehistoric Eastern Woodlands. University of Alabama Press, Tuscaloosa, AL. Migahid, Ahmed Mohammed. 1988. Flora of Saudi Arabia, Vol 1., King Saud University, Riyadh. Parr, J. F., and M. Carter. 2003. Phytolith and Starch Analysis of Sediment Samples from Two Archaeo- logical Sites on Dauar Island, Torres Strait, North- eastern Australia. Vegetation History and Archaeobotany 12:131-141. Pearsall, D. M. 2000. Paleoethnobotany: A Handbook of Procedures, 2nd ed. Academic Press, San Diego, CA. Perry, L. 2004. Starch Analyses Reveal the Relation- ship between Tool Type and Function: An Example from the Orinoco Valley of Venezuela. Journal of Archaeological Science 31:1069-1081. Piperno, D. R., and T. D. Dillehay. 2008. Starch Grains on Human Teeth Reveal Early Broad Crop Diet in Northern Peru. Proceedings of the National Academy of Sciences 105:19622-19627. Piperno, D. R., A. J. Ranere, I. Holst, and P. Hansell. 2000. Starch Grains Reveal Early Root Crop Horticulture in the Panamanian Tropical Forest. Nature 407:894-897. Piperno, D. R., E. Weiss, I. Holst, and D. Nadel. 2004. Processing of Wild Cereal Grains in the Upper Palaeolithic Revealed by Starch Grain Analysis. Nature 430:670-673. Riehl, S. 1999. Bronze Age Environment and Economy in the Troad: The Archaeobotany of Kumtepe and Troy. Unpublished Doctoral Disser- tation, Faculty of Geosciences, University of Tübingen, Tübingen, Germany. Reichert, E. T. 1913. The Differentiation and Specificity of Starches in Relation to Genera, Species, Etc., Stereochemis- try Applied to Protoplasmic Processes and Products and as a Strictly Scientific Basis for the Classification of Plants and Animals. The Carnegie Institute of Washington, Washington D. C. Rosen, A. M. 2005. Phytolith Indicators of Plant and Land Use at Çatalhöyük, In Inhabiting Çatalhöyük; Reports From the 1995-99 Seasons, edited by I. Hodder, pp. 203–212. McDonald Institute of Archaeology, Cambridge. Ryan, P. 2011. Plants as Material Culture in the Near Eastern Neolithic: Perspectives from the Silica Skeleton Artifactual Remains at Çatalhöyük. Journal of Anthropological Archaeology 30:292-305. Samuel, D. 1996. Investigation of Ancient Egyptian Baking and Brewing Methods by Correlative Microscopy. Science 273:488-490. Sivak, M. N. S., and J. Preiss, eds. 1998. Starch: Basic Science to Biotechnology. Academic Press, San Diego, CA. Tester, R. F., J. Karkalas, and X. Qi. 2004. Starch— Composition, Fine Structure and Architecture. Journal of Cereal Science 39:151-165. Torrence, R., and H. Barton, eds. 2006. Ancient Starch Research. Left Coast Press, Walnut Creek, CA. Wollstonecroft, M. M., P. R. Ellis, G. C. Hillman, and D. Q. Fuller. 2008. Advances in Plant Food Processing in the Near Eastern Epipalaeolithic and Implications for Improved Edibility and Nutrient Bioaccessibility: An Experimental Assessment of Bolboschoenus maritimus (L.) Palla (sea club-rush). Vegetetation History and Archaeobotany 17:19-27. Yang, X., and L. Perry. 2013. Identification of Ancient Starch Grains from the Tribe Triticeae in the North China Plain. Journal of Archaeological Science 40:3170-3177. Zohary, D., M. Hopf, M., and E. Weiss. 2012. Domestication of Plants in the Old World. Oxford University Press, Oxford. Biosketch Thomas C. Hart is the laboratory manager/research scien st for the Environmental Archaeology Laboratory at the University of Texas at Aus n.