Archaeological SCIENCE

Journal of Archaeological Science 30 (2003) 10851094 http://www.elsevier.com/locate/jas

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Assessing the feasibility of identifying maize through the analysis of cross-shaped size and three-dimensional morphology of phytoliths in the grasslands of southeastern South America
Jose Iriarte *
Department of Anthropology, University of Kentucky, 211 Laerty Hall, Lexington, KY 40506-0024, USA Received 22 February 2002; received in revised form 18 April 2002

Abstract This paper tests the feasibility of applying a technique developed by Piperno and Pearsall (Am. Antiquity 49 (1984) 361; Phytolith Analysis: An Archaelolgical and Geological Perspective, 1988, Academic Press: San Diego; Paleoethnobotany, 2000, Academic Press: San Diego) based on size and three-dimensional morphology criteria of cross-shaped phytoliths to identify maize in a previously unexplored region outside of the Neotropics; the grasslands of southeastern Uruguay. Because the area is dominated by subtropical Panicoid grasses that produce abundant cross-shaped phytoliths, intensive studies of the regional Panicoid grasses are needed to ensure that no wild taxa have phytoliths that are potentially confusable with maize. With this in mind, I carried out analysis of cross-shaped phytoliths in 35 Panicoid, 5 Oryzoid and 1 Bambusoid grasses, as well as on nine modern soil samples that belong to the most representative vegetation formations of the area. This study demonstrates that an application of multivariate (linear discriminant function) analysis together with qualitative and other assessments of cross-shaped phytolith assemblages as originally described by Piperno and Pearsall can be successfully used to distinguish the presence of maize in the grasslands of southeastern Uruguay. The technique provides a useful tool to trace the dispersal of maize into the southern cone of South America.  2003 Elsevier Ltd. All rights reserved.
Keywords: Uruguay; Pampas; South America; Phytoliths; Palaeobotany; Maize

1. Introduction Microfossil botanical analyses are increasingly being applied to studies of the origin and early dispersals of plant domesticates [24]. Renements in the recognition of micro-morphological features, together with the application of multivariate statistical analyses to phytolith and starch grain assemblages, are allowing palaeoethnobotanists to distinguish certain phytolith and starch morphotypes to specic taxonomic levels, and to dierentiate wild from domesticated species. Examples include maize [19,2125,27], squashes and gourds [4,29], rice [43], wheat and barley [3,16,39], banana [17,42], manioc, and yams [30]. Central to these advances has been the construction of extensive regional plant

* Tel.: +1-859-323-9957; fax: +1-859-323-1959 E-mail address: jiria0@uky.edu (F. Iriarte). 0305-4403/03/$ - see front matter  2003 Elsevier Ltd. All rights reserved. doi:10.1016/S0305-4403(02)00164-4

reference collections, which enable analysts to more securely characterize and classify palaeobotanical remains. This paper tests the feasibility of applying a crossshaped phytolith size and three-dimensional (3-D) technique developed by Piperno and Pearsall [20,22,23] to identify maize in a previously unexplored region outside of the Neotropics; the grasslands of southeastern Uruguay. Because the area is dominated by Panicoid grasses that produce abundant cross-shaped phytoliths [2,10,33], intensive studies of the regional ora are needed to ensure that no wild grasses potentially confusable with maize occur. At the same time, the presence of several of the same species of cross-shaped producing grasses in the Uruguayan ora previously analyzed by Piperno and Pearsall permits an independent, replicative analysis to verify that the size and 3-D morphology of cross-shaped phytoliths remain fairly constant across regional and environmental boundaries.

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Through a study of size and 3-D morphology of cross-shaped phytoliths occurring in 23 races of maize, 6 varieties of teosinte, and more than 350 species of wild grasses, Piperno and Pearsall demonstrated that it is possible to diagnose the presence of maize in soil phytolith assemblages in the Neotropics (see Ref. [20] for a detailed history of the technique). Cross-shaped phytoliths are separated into eight or more morphological categories based on their 3-D characteristics (from Variant 1 (= mirror image cross-shape) to Variant 8), and the mean width and percentage occurrence of each crossshaped Variant is determined. Maize phytolith assemblages are characterized by high proportions of large, Variant 1 cross-shapes, distinguishing them from wild taxa. Piperno [23, pp. 7983] reported a multivariate, Fishers linear discriminant function analysis that formalized these criteria and divided maize and wild grasses into two separate groups. The function is based on three variables: mean size of Variant 1 cross-shaped phytoliths, mean size of Variant 6 cross-shaped phytoliths and percentage of Variant 1 cross-shaped phytoliths H size Var 1)+0.1025 (X H size Var 6)+0.0215 (df0.8082 (X (% Var 1)). When used together with a qualitative trait assessment to rule out the presence of a few wild grasses that also possess some large-sized (greater than 16 m), Variant 1 cross-shapes, this technique appears to successfully identify maize presence in Preceramic and Formative-period archaeological sites of lower Central and South America [20,24]. The presence of extra-largesized Variant 1 cross-shaped phytoliths (greater than 20.60 wide) in archaeological assemblages was often considered sucient to identify maize [19,23]. Another important fact about the cross-shaped technique is that it is a conservative method of identifying maize. In other words, it probably will not identify all maize races [23, pp. 173174], but neither should it result in inaccurate identications of wild grasses as maize. Piperno and Pearsalls (P&P) maize phytolith technique has not been free of critics (e.g. Refs. [8,34]). For reasons I will briey discuss here and demonstrate below, the critiques do not appear to be supported by adequate data and/or are based on misconceptions about how the technique is applied (see also a detailed discussion by Piperno [24, pp. 427434]). For example, Doolittle and Frederick [8] claimed that they could not duplicate the P&P technique, but they were unable to identify cross-shaped phytoliths in the maize leaves they studied. The phytolith illustrations in Ref. [8 Fig. 1] clearly show that cross-shaped phytoliths were commonly present in their specimens, as they are in all other examples of modern maize and wild Zea reported on to date (e.g. Refs. [15,1820,23]). Perhaps, as discussed by Piperno [24pp 428429], Doolittle and Fredericks inexperience with phytolith studies led to their inability to identify cross-shaped phytoliths.

Rovner [34] stated that cross-shaped 3-D variants could be articial products created by using conventional microscopy. For reasons that will be made clear below, I do not agree, and other phytolith researchers in addition to Piperno and Pearsall have documented that variability in 3-D attributes is a valuable taxonomic tool in cross-shaped (e.g. Refs. [18,40,41]) and other [5,18,37] grass phytolith identication. Most recently, Staller and Thompson [37] questioned the P&P cross-shaped technique, but they did so by citing the previous criticisms just discussed without attempting a replication by themselves. For example, in voicing doubts about the utility of cross-shaped 3-D forms, these authors cite Doolittle and Frederick [8], but in light of that studys failure to identify and study cross-shaped phytoliths at all, it is dicult to understand what merit they found in it. It also seems odd that they appear to agree with Rovners [34] comments on crossshaped 3-D attributes when their criteria for developing a taxonomy for phytoliths from maize cha relies heavily on the same kinds of 3-D features exhibited by cross-shaped phytoliths (e.g. dierences in the shapes of the two faces of the phytoliths) (see also Ref. [38]). Staller and Thompson [37, p. 34] also state that the P&P typology . is based on the identication of forms only produced by maize, adding that P&Ps eorts . have focused upon extra-large crosses, usually produced in the leaves of grasses. These statements are erroneous and misleading. Cross-shaped phytoliths and the critical cross-body 3-D variants are not exclusive to maize, as P&P have consistently stated and demonstrated [22, pp. 36467, Table 1; 23, pp. 7277, Tables 3.13.6; 25, pp. 34043, Tables 25; 26, pp. 1415, Tables 25]. This realization combined with the fact that low percentages of large Variant 1 cross-bodies occur in a few wild grasses, led Piperno [23, pp. 8083] to apply a Fisher linear discriminant function analysis, which eectively separated cross-shaped phytoliths from maize and wild grasses into two discrete clusters. The discriminant function analysis has been upgraded by Piperno and Pearsall (in Ref. [20]) with the inclusion of 100 additional wild grasses, yielding the same statistically robust results. Furthermore, no wild Neotropical grass has been found to produce extra-large (wider that 21 ) Variant 1 cross-shapes. Nevertheless, as has been pointed out on many occasions (e.g. Ref. [20]), large reference collections of the regional ora are necessary before archaeobotanical methods seeking to identify major domesticated species can be condently applied to any study area. 2. The study area and its vegetation formations Mound complexes located in the seasonally humid wetlands of southeastern Uruguay investigated by the

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Fig. 1. (ac) Oryzoid cross-shaped phytoliths: (a) Luziola peruviana (20 27 ); (b) Oryza glandiglumis (17.3 19.8 ); and (c), Zizaniopsis bonaerensis (16 22 ). (d) Variant 1 cross-shaped from Paspalum plicatulum (14.8 20.9 ). (e) Variant 2 cross-shaped from Andropogon condensatus (12.4 14.8 ). (fi) Variant 5&6 cross-shaped phytoliths: (f) Panicum prionitis (13.6 14.8 ); (g) Paspalum dilatatum (9.9 11.2 ); Rottboellia selloana (12.4 14.8 ); Leptocoryphium lanatum (12.4 12.4 ). (jl) Variant 1 cross-shaped phytoliths: (j) Panicum decipiens (9.9 12.4 ); (k) Setaria geniculata (11.1 12.4 ); and (l) Axonopus compressus (9.9 11.1 ).

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author and others [11] that date to ca. 4000 BP are characterized by the same, poor preservation of macrobotanical remains common to sites located in the lowland Neotropics. Despite the implementation of extensive otation programmes, the only macrobotanical remains recovered from these contexts are carbonized palm nut shells, which typically survive because they are extremely hard. No macrofossil remains of domesticated plants have thus far been recovered. Phytoliths, being resistant to decay over time, potentially provide a means to document some aspects of crop history in this region. Previous classications of grass phytoliths in the Cerrados of Central Brazil (e.g. Ref. [35]) and the pampas of east-central Argentina (e.g. Ref. [44]) did not take into account the size and 3-D morphology of cross-shaped phytoliths. In addition, previous studies conducted in Uruguay (e.g. Ref. [7]) have not suciently addressed this problem due to the lack of an extensive reference collection. In this paper, I present the results of an analysis of cross-shaped phytoliths from the 35 most abundant Panicoid species of southeastern Uruguay, as well as 5 Oryzoid and 1 Bambusoid species that occur in the region. Nine modern soil samples that belong to the most representative vegetation formations of southeastern Uruguay were also analyzed, in order to examine patterns of cross-shaped phytolith deposition from dierent grasses in sedimentary contexts (Table 1 ). The study region corresponds to the subtropical and humid (total annual rainfall of 1200 mm) of the Rio de la Plata Grasslands extending along the eastern Atlantic coast of South America. The grassland vegetation of southeastern Uruguay is well studied and arguably one of the best known of the worlds regions. Agronomists have carried out exhaustive inventories of the native grasses in order to assess their grazing capacity [2,33]. More recently, the area has also been subjected to intensive studies for conservation purposes and the development of sustainable development programmes [32]. The vegetation formations of the area are abundant producers of cross-shaped phytoliths, being characterized by the dominance of such subtropical grass genera as Andropogon, Axonopus, Panicum, Paspalum, Schizachyrium and Bothriochloa [14,33]. Our own study of modern vegetation in the area and its phytolith record conrms these reports (Iriarte and Alonso, in preparation). Based on previous studies [1,32], soil samples from the most representative modern vegetation formations were collected. Nine modern soil samples were analyzed from wetlands, wet prairies, hilly prairies and riparian forest vegetation formations. Wetlands occur in the low oodplains, which remain covered with shallow water most of the year, and are characterized by deep, poorly drained, clayey or peat supercial horizons. These areas are characterized by the presence of emergent

hydrophytic vegetation, which give rise to diverse vegetation formations locally known by the popular name of the species that dominate the habitat. The wetland vegetation formations examined in this study correspond to Juncal of Schoenoplectus californicus (junco), Totoral of Thypa spp. (totora), and Espadan al of Rynchospora corymbosa (espadan a). The latter name is usually given also to communities dominated by Zizaniopsis bonaerensis, which also occur in the area. The most common grass species in the herbaceous strata of these formations are the Panicoid grasses Ehinochloa helodes, Paspalum spp. and Paspalidium paludivagum and the Oryzoid grasses Leersia hexandra, Luziola peruviana and Z. bonaerensis [1]. Wet prairies occur in areas slightly higher than wetlands. These areas are characterized by at and poorly drained soils that remain waterlogged during the winter months and are exceptionally inundated. The wet prairie vegetation formations sampled for this study were Pradera Uliginosa, Pajonal, Pradera Uliginosa con Palmas, and Caraguatal. Praderas uliginosa is characterized by the dominance of Panicoid grasses such as Axonopus compressus, Axonopus anis, Stenotapharum secundatum, Paspalum notatum. Pajonal correspond to associations of tall perennial grasses like Panicum prionitis, Paspalum quadrifarium, and Cortadeira selloana. Caraguatal is a vegetation formation dominated by Eryngium spp. (Apicaceae) whose herbaceous strata is very similar to the one characteristic of Praderas Uliginosas. Pradera Uliginosa con Palmas constitute the herbaceous strata of palm forest of Butia capitata mainly characterized by associations of A. anis and Panicum gounii, among other non-Panicoid species. In the area the riparian forest comprises a narrow strip bordering the main streams, and is mainly composed of Celtis spp., Blepharocalyx salicifolius, Sebastiania brasiliensis, Schinus longifolius, among others. Upland prairies, called Sierra in this study, are located in hills and knolls, and are characterized by well-drained supercial soils, which suer from a hydric decit during the summer months. The abundant Panicoid species in these habitats are P. notatum, Setaria geniculata, A. compressus, and Paspalum dilatatum [2,32,33]. 3. Field and laboratory procedures The grass species selected for study were chosen on the basis of three main criteria: aliation with the cross-shaped producing Panicoid and Oryzoid subfamilies, known production of some large Variant 1 cross-shapes (i.e. Andropogon spp. and Paspalum spp.), and high percentages of Variant 1 cross-shapes (i.e. Axonopus spp. and Panicum spp.) [22,23]. In addition, one of the two bamboos occurring in the area that produce cross-shapes, Chusquea ramosissima [12,28,32]

Table 1 Cross-shaped phytolith size, 3-D morphology and discriminant function value in wild Panicoid, Oryzoid and Bambusoid grasses and modern vegetation formations in southeastern Uruguay Three-dimensional morphology Cross-shaped phytolith size Small Medium 10.62 9.92 9.74 47 9.68 51 Large L. Var 1 1 2 3 4 5&6 7 8 H1 Oryz. Chusq. X %Var 1 H6 X 3-D morphology Variant Discriminant function d.f. Value 10.67 10.03 11.66 11.06 11.07 11.27 11.24 42 12.19 11.32 80 11.65 12.35 77 11.8 10.26 71 11.9 10.01 37 12.51 12.8 60 12.7 12.23 56 11.68 10.02 39 13.4 11.47 10.64 11.6 7 10.5 68 9.74 70 12.35 10.08 22 10.13 9.71 50 11.16 12.73 12.33 12.11 11.43 12.71 12.72 11.31 12.15 11.80 11.10 11.48 10.26 N 51 53 33 53 52 50 30 30 79 30 50 50 28 30 31 30 51 50 (continued on next page) 1089

Grasses

Panicoid Agenium nessi Nees

Agenium villosum (Nees) Pilg.

Andropogon condensatus Kunth

11.21 10.73 70 11.04 10.17 51 11.65 11.68 21

Andropogon consanguineus Kunth

Andropogon lateralis Nees

Andropogon selloanus (Hack) Hack.

Axonopus anis Chase

Axonopus aranjoi Nees

Axonopus compressus (Sw.) P. Beauv.

Axonopus suultus (Mikan ex Trin.) Parodi

Bothriochloa laguroides (D.C.) Herter

Bothriochloa saccharoides (Sw.) Rydb.

J. Iriarte / Journal of Archaeological Science 30 (2003) 10851094

Eriochloa punctata (L.) Desv. ex Ham.

Leptocoryphium lanatum (Kunth) Nees

Panicum bergii Arechav.

Panicum decipiens Nees ex Trin.

Panicum dilatatum Steud.

Panicum gouinii E. Fourn.

38 75% 47 89% 11 33% 19 36% 24 46% 25 50% 4 13% 9 30% 33 42% 19 63% 10 20% 14 28% 19 68% 11 37% 16 52% 20 67% 37 73% 45 90%

13 25% 6 11% 22 67% 33 62% 27 52% 25 50% 26 87% 21 70% 45 57% 11 37% 37 74% 33 66% 9 32% 19 63% 15 48% 10 33% 14 27% 5 10%

1 2% 1 2% 1 1% 3 6% 3 6%

2 7% 2 7%

26 51% 25 47% 23 4 70% 12% 27 3 51% 6% 11 2 2 21% 4% 4% 21 1 42% 2% 24 80% 23 77% 56 71% 11 37% 30 60% 28 56% 11 39% 2 7% 21 2 68% 6% 21 1 70% 3% 11 3 22% 6% 25 3 50% 6%

25 49% 28 53% 6 18% 23 43% 36 69% 28 56% 6 20% 7 23% 22 28% 19 63% 20 40% 22 44% 17 61% 28 93% 8 26% 8 27% 37 73% 22 44%

1 2% 1 3%

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Table 1 (continued) Three-dimensional morphology Cross-shaped phytolith size Small Medium 12.8 12.61 41 11.99 10.56 66 10.24 10.3 9.47 33 10.7 46 12.51 12.18 9.96 10.41 12.01 10.21 10.29 11.87 10.57 35 10.78 10.39 21 12.84 11.39 15 10.7 9.53 18 12.82 10.85 62 12.39 11.83 62 12.48 11.22 45 12.96 11.82 40 11.31 9.81 63 9.5 11.99 60 11.44 10.22 11.87 10.00 12.81 12.56 12.20 12.55 11.49 10.20 Large L. Var 1 1 2 3 4 5&6 7 8 H1 Oryz. Chusq. X %Var 1 H6 X d.f. Value 3-D morphology Variant Discriminant function N 51 32 51 50 82 30 31 51 53 80 34 50 50 40 47 48 30

Grasses

Panicum grumosum Nees

Panicum prionitis Nees

Panicum .sabulorum Lam.

Paspalidium gemiculata Croat

Paspalum dilatatum Poir.

12.47 10.69 39

Paspalum distichum L.

Paspalum exalum Presl.

10.3 10.18 40 11.12 10 13

Paspalum lividum Trin. ex Schltdl

Paspalum notatum Flugge

Paspalum plicatulum Pers.

Paspalum quadrifarium Lam.

Rottboellia selloana Hack.

Schizachyrium intermedium Nees

J. Iriarte / Journal of Archaeological Science 30 (2003) 10851094

Schizachyrium tenerum Nees

Setaria geniculata P. Beauv.

Setaria vaginata Spreng.

Trachypogon montufari (Kunth) Nees

9 18% 22 69% 45 88% 31 62% 41 50% 24 80% 22 71% 31 61% 39 74% 36 45% 28 82% 14 28% 14 28% 14 35% 12 26% 30 63% 23 77% 18 36% 9 18% 23 4% 16 32% 30 60% 4 8% 11.25 10.25 36

40 78% 10 31% 6 12% 19 38% 41 50% 6 20% 9 29% 20 39% 14 26% 42 53% 6 18% 35 70% 34 68% 25 63% 33 70% 18 38% 7 23%

2 4% 2 3% 1 2% 2 4% 1 3% 2 4%

2 9% 1 8% 1 3% 2 6%

21 41% 21 66% 17 33% 23 46% 32 39% 12 40% 4 13% 18 35% 11 21% 12 15% 6 18% 31 62% 31 62% 18 45% 12 26% 0 63% 18 60%

1 2% 2 6% 1 2% 2 2% 1 3% 1 2% 3 6% 7 9% 2 6%

29 57% 9 28% 33 65% 27 54% 48 59% 18 60% 26 84% 32 63% 39 74% 61 76% 26 76% 19 38% 19 38% 22 55% 18 38% 18 38% 12 40%

Bambusoid Chusquea ramosissima Lindm.

10.91

50

Table 1 (continued) Three-dimensional morphology Cross-shaped phytolith size Small Medium Large L. Var 1 1 2 3 4 5&6 7 8 H1 Oryz. Chusq. X %Var 1 H6 X d.f. Value 3-D morphology Variant Discriminant function N 50 11.44 11.73 16 11.73 11.73 20 11.54 10.48 27 12.53 10.23 10 10.79 10.72 10.98 11.39 60 30 30 50

Grasses

Oryzoid Leersia hexandra Sw.

Luziola peruviana Juss. ex J.F.Gmel.

Oryza grandiglumis (Doll) Prod.

1 2% 2 6%

Oryza latifolia Desv.

Zizaniopsis bonaerensis (Balansa & Poitr.) Speg.

20 40% 12 20% 4 13% 7 23% 14 23%

27 54% 45 75% 24 80% 22 73% 37 62%

3 6% 3 5% 2 7% 1 4% 9 15%

10 16% 6 20% 8 27% 6 10%

6 10% 5 17% 3 5%

50 100% 43 72% 22 78% 17 56% 51 85%

Vegetation formation Wetlands Juncal

11.25 10.36

9.05 46 9.6 42 11.4 10.53 36

11.01 10.27 11.07

52 52 36

Totoral

Espadan al

39 75% 44 85% 24 67%

11 21% 6 12% 11 31%

2 4% 1 3%

1 5%

24 46% 22 42% 13 36%

1 3%

24 46% 27 52% 18 50%

1 2% 1 2%

Wet prairies Pradera uliginosa

11.41 10.18 44 12.78 10.04 40 10.79 10.43 40

11.21 12.22 10.64

50 50 48

Pajonal

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Pradera Uliginosa con Palmas 35 70% 19 37% 28 55% 4 8% 2 10% 13 26% 2 4% 1 5% 20 2 40% 4% 21 2 41% 4%

32 64% 33 66% 35 73%

18 36% 15 30% 16 33%

2 4% 1 2%

2 9% 2 10%

22 44% 20 40% 19 40%

1 2%

28 56% 28 56% 29 60%

1 2% 28 56% 27 1 53% 2%

Riparian forest

11.33

9.97 40

11.04

50

Hills Sierra

12.85 12.39 41

12.54

51

H size Var 1)0.1025(X H size Var 6)0.0215(%Var 1). df0.8082(X

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was analyzed. The idiosynchratic crosses produced by the other occurring bamboo, Olyra latifolia, are well described in Ref. [8]. Five Oryzoid grasses [12] were examined. Three of them are common in the area: L. hexandra, Lu. peruviana, and Z. bonaerensis. Two of them are not reported for the immediate area but are present in the regional vegetation: Oryza grandiglumis and Oryza latifolia [6]. Voucher specimens were collected at the National Museum of Natural History Herbarium-Smithsonian Institution in Washington DC, the Missouri Botanical Garden in Saint Louis, the Herbario de la Facultad de Agronom a, Universidad de la Repu blica, Uruguay, and the herbarium of the Smithsonian Tropical Research Institute. Additionally, some species where collected in the eld by project botanist Eduardo Alonso and identied with comparative specimens curated at the Hervario de la Facultad de Qu mica, Universidad de la Repu blica, Uruguay. Each of the samples for the modern soils analyzed is a composite of four A1-horizon (upper 13 cm) samples taken from the corners of a 100 m2 square plot in wetland areas and a 1 m2 plot in prairie areas. The extraction of phytoliths and the slide preparation of modern reference grasses and modern soils followed standard procedures described in Ref. [23]. Crossshaped phytoliths were identied and measured using Pearsalls [19] criteria to separate crosses from dumbells. Only leaf phytoliths were considered. Cross-shaped 3-D morphology was determined using Pipernos [22,23] descriptions. One additional type of cross-shaped phytolith Variant was described in this study. These comprise the idiosynchratic forms produced by Oryzoid grasses (see Fig. 1). Phytoliths were rotated, measured with an eyepiece micrometer, and counted at 400 , using an Olympus BH2 microscope. As with any identication technique, learning to rotate and identify the dierent 3-D Variants of cross-shaped phytoliths took practice and good supervision, but these various forms became readily identiable. At least 30 crosses were counted per species and at least two specimens per species from dierent regions were analyzed whenever possible. 4. Results and discussion The results of this analysis are in good agreement with Pearsall and Pipernos determinations of crossshaped phytolith size and morphological characteristics of Neotropical grasses, and indicate that cross-shaped phytoliths can be used to identify maize in archaeological contexts from southeastern Uruguay (Table 1). The most important results can be summarized as follows: (a) Unlike maize, neither the wild grasses nor the modern soils presented a df value (discriminant

function) higher than 13 (see Refs. [20, Fig. 5.1719; 23, Fig. 3.8]). The grasses that contain some large Variant 1 cross-shapes (e.g. greater than 16 in width), such as Leptocoryphium lanatum, have high percentages of Variant 5 and 6 crosses (Fig. 1fi) (e.g. 93% in Lep. lanatum), which distinguish them from maize. (b) Considering all the cross-shaped variants together, 24 (58%) of wild grasses do not produce any large crosses and the only large crosses produced in Oryzoid and Bambusoid grasses are idiosyncratic forms discussed below and illustrated in Fig. 1ac. In addition, unlike maize, which commonly produces between 12 and 39% of cross-shaped phytoliths with a width of 16 and larger [19,22,23], the wild Panicoid grasses and modern soil assemblages do not produce more than 8% cross-shapes with widths greater than 16 and larger. (c) While maize commonly produces high numbers of large, Variant 1 crosses, only 6 (17%) of the Panicoid grasses studied produced any large Variant 1 crosses, with an average of 7% of large Variant 1s (range 7 to 9%). This factor also leads to a higher average size for Variant 1 crosses in maize than in wild grasses. The highest percentage of large Variant 1 crosses is produced by the Pajonal (Pa. prionitis) vegetation formation, reaching 10% in this context. This vegetation formation, however, produced only 40% Variant 1 crosses, making it separable from maize on a morphological basis, and giving it a df value of 12.22, which results in a classication as a wild grass. (d) In contrast to maize, which produces an average of 3% of extra-large crosses (greater than about 20.60 wide), none of the wild grasses nor the modern soils tested produced extra-large crosses. It should be remembered that the presence of extra-large crosses is a condition that is sucient but not necessary to prove the presence of maize, since not all maize races produce cross-shaped phytoliths of this size. (e) Many of the wild grasses produced high percentages of Variant 1 crosses, as does maize, but unlike maize, these wild grasses (e.g. Axonopus selloanus, A. anis, Axonopus suultus, Agenium spp., Andropogon selloanus, Andropogon condensatus, all Panicum species with the exception of Panicum grumosum and Trachypogon montufari) do not produce any large crosses. Furthermore, unlike maize, 16 (39%) of the wild Uruguayan grasses were characterized by high proportions of Variants 5 and 6 cross-shapes. Importantly, in modern soils, the percentage of Variant 1 cross-shapes does not exceed 46%. (f) Grass species from Uruguay previously analyzed by Piperno and Pearsall (e.g. A. compressus, Paspalum plicatulum, among others) from the Neotropics demonstrated very similar size and 3-D characteristics to the tropical species, indicating that these features of grass phytoliths are essentially stable from region to region (compare Table 1 with Ref. [23, Table 3.1]).

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(g) Last but not least, none of the modern soils, where phytolith assemblages from dierent species are mixed, presented maize cross-shaped phytolith characteristics. This study demonstrates that the application of a multivariate discriminant function analysis described by Piperno and Pearsall in conjunction with qualitative and other assessments of cross-shaped phytolith assemblages can distinguish the presence of maize from wild grasses in the grasslands of southeastern Uruguay. Doubts expressed by a few investigators about the reliability and replicability of 3-D structure and size variation in cross-shaped phytoliths [34,37] are not supported here. Cross-shaped phytolith assemblages having a df value higher than 13 and/or containing: (a) more than 15% of large-sized (greater than 16 wide) Variant 1 crosses; (b) high (greater than ca. 50%) percentages of Variant 1 and low percentages of Variants 2 and 6 crosses, and (c) extra-large Variant 1 cross-shaped phytoliths (a sucient but not necessary condition), appear to be securely identiable as maize-bearing assemblages. Application of a discriminant function analysis based on size and shape variables allows robust statistical input into the problem, contra unsupported statements (e.g. Refs. [36, p. 159; 37]) that the technique is statistically fragile. A new type of cross-shaped phytolith is also described in this study for the Oryzoid subfamily (Fig. 1ac). The distinctive characteristics of Oryzoid dumbells and crosses have been previously described (e.g. Refs. [13,15,43]). Oryzoid crosses are characterized by the presence of pointed triangular projections extending from the lobes of the nontype-tier face of the cross, lobes are characteristically angular but often rounded, indentations usually are arch-like between two pointed lobes, and usually there is no clear demarcation in the middle of the cross-shape phytolith. The three types displayed in Fig. 1ac correspond to the Oryzoid type. Within the study region, the Oryzoid cross-shaped phytolith illustrated in Fig. 1a is only produced in Lu. peruviana. The phytolith in Fig. 1b is predominantly produced in L. hexandra, and O. glandiglumis, whereas O. latifolia and Z. bonaerensis mainly produce the type illustrated in Fig. 1c, but also produce the Fig. 1b type. These new types not only are useful for dierentiating wild Oryzoid grasses from maize; they also have considerable potential as important indicator phytoliths in palaeoenvironmental reconstruction because these species are limited to seasonally inundated, wetland environments [6]. It is also important to note that the distribution of these idiosyncratic phytolith types closely mirrors phylogenetic relationships postulated for grasses [12], further supporting beliefs that there is a strong genetic inuence on phytolith morphology [24]. This study indicates that maize leaf phytoliths will be an important avenue in tracing the dispersal of maize

into southern South America. Importantly, recent studies also indicate that diagnostic phytolith morphotypes occur in the reproductive structures (cobs and cha) of maize and teosinte [5,18,25,37,38], and the production and assemblage trait characteristics of these phytoliths is known to be genetically controlled [9,24]. The common, decay-in-place depositional nature of phytoliths combined with the production of diagnostic phytoliths from multiple parts of the maize plant (e.g. leaves and cobs) provides researchers with the potential to identify dierent activity areas, like gardens and household consumption areas, in archaeological contexts. Furthermore, the examination of sediments from wetlands and wet prairies in southern South America and elsewhere (e.g. Ref. [31]) may allow us to assess where and when maize was planted using ood-recessional agricultural practices.

Acknowledgements This research was supported in part by a grant from the Wenner-Gren Foundation for Anthropological Research-Gr. 6614 and the Smithsonian Tropical Research Institute in Panama as part of my dissertation research on HumanEnvironment Interactions in the Wetlands of southeastern Uruguay during the MidHolocene. I am grateful to the National Museum of Natural History Herbarium-Smithsonian Institution, the Missouri Botanical Garden and the Herbario de la Facultad de Agronomia-Universidad de la Republica in Uruguay who facilitated access to their collections for the extraction of voucher specimens.

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