362 AMERICAN ANTIQUITY [Vol. 49, No.

2,1984]

A Comparison and Differentiation of Phytoliths from Maize and Wild Grasses: Use of Morphological Criteria
Author(s): Dolores R. Piperno Cross-shaped phytoliths, which occur only in the Panicoid sub-family of grasses and bamboos, were intensively evaluated because work by Pearsall (1978, 1979) and myself (1979, 1980) had indicated that they were most useful for distinguishing maize. (p.362) In a study of phytolith size in Ecuadorian grasses, Pearsall (1978, 1979) showed that cross-shaped phytoliths from maize were significantly larger than those of the native Panicoid grass cover. Cross-shaped phytoliths were defined as those with at least three indentations which have a dimension that falls between N by N and N by (N plus 2), with each micrometer unit (N) representing 4.58 microns. The expression N by (N plus 2) signifies that phytoliths more than 9.16 microns longer than wide are not classified as cross-shapes. This distinction is useful, because it eliminates phytoliths that appear to be transitional between dumbbells and crossshapes, and thus tightly defines the cross-shaped type. (p.362) The combination of cross-shaped phytolith size and cross-shaped phytolith threedimensional structure securely separated all wild grasses, including teosinte, from many of the maize races that were studied. Short-cell three-dimensional structure is also a highly promising area for the identification of some genera and species of wild Panicoid grasses and bamboos. (p.380)

Journal of Archaeological Science (1984) 11, 345-368

Prospects and Limits of a Phytolith Key for Grasses in the Central United States
Dwight A. Brown Ratios of certain trapezoid shapes to saddles or bilobates can provide information on the relative dominance of grasses possessing C4 and C3 carbon pathways. Fluctuations in these ratios found in stratified deposits may reflect shifts in midsummer temperature through time, but there are several sources of error that must be examined separately. Phytoliths that fit in the festucoid class of Twiss et al. (1969) (now Pooicoideae) have been seen in the tribes Aristideae, Eragrosteae, and Andropogoneae. Saddle shapes of Eragrosteae and Chlorideae are also common in Phragmites australis and bilobates (dumbbells) of the Panicoideae subfamily occur in species of Danthonia. (p.345) In summary, there are sufficient differences among the most important taxonomic groups in the central North American grasslands to allow us to identify the time of taxa arrival

during the Holocene, if stratified and datable deposits are available. Thus, it is possible for phytolith analysis to provide a better understanding of migration rates and sources of major species. (p.345) A summary of the general phytolith shape characteristics for subfamilies and tribes of grasses is shown in Table 6. The lack of both bilobates and saddles in the subfamily Pooideae is important. Paucity of these two phytolith shapes in earth materials, coupled with the abundance of trapezoids, probably indicates a dominance of C3 grasses. ()Variations within subfamilies and tribes in the degree of importance of these shapes may also be significant, but differences are not easily analyzed at this taxonomic level. (p.366)

Quaternary International 193 (2009) 8089 Phytolith analysis of Chusquea ramosissima Lindm. (Poaceae: Bambusoideae) and associated soils La Montti, Mariana Fernandez Honaine, Margarita Osterrieth, Dalva Graciano Ribeiro.

Leaves - The panicoid dumb-bells(bilobate flat/concave ends shortcell phytoliths) were the
second most abundant morphotype(Table3). They represent thesilica cells located in the costal zone of the epidermis. Chusquea bulliform phytoliths(cubic/parallepipedal sinuate bulliform cells) were abundant, both isolated or in articulated form. In a smaller proportion, the phytolith assemblage was characterized by the presence of saddles and tabular crenate phytoliths. These phytolith morphotypes correspond to the vertical silica bodies and long rectan- gular epidermic cells of winding walls of the costal and intercostal zone, respectively. Silicification of hairs was also observed. Within the articulated phytoliths, fragments formed by several of the morphotypes pre- viously mentioned, were observed. (p.83). In leaves, the main phytolith morphotypes described (concave dumb-bells, panicoid dumb-bell sand Chusquea bulliform) are the same as the ones observed by Piperno (1988) and Piperno and Pearsall(1998) for other species of the same genus or for other tropical bamboos. These authors argue that these phytolith can be diagnostic of Bambusoideae subfamily. (p.87)

Soil phytolith assemblage - The dominant phytolith morphotype sinall the samples were the
concave dumb-bell phytoliths, except in the second sample of the Iguazu National Park close canopy (NPc2 sample),where the greater proportion of phytoliths were rectangular smooth(o30 m long),probably broken phytoliths, and degraded irregular phytoliths, possibly originated in trees or other dicotyledons species. The second dominant morphotype was the panicoid dumb-bells, while the Chusquea bulliform phytoliths predominate in some samples of gaps. (p.85)

Pl Syst Evol 270: 123 (2008) DOI 10.1007/s00606-007-0597-z Printed in The Netherlands

Diversity of lobate phytoliths in grass leaves from the Sahel region, West Tropical Africa: Tribe Paniceae
A. G. Fahmy

The subfamily Panicolideae is characterized by the presence of bilobate, polylobate and quadra-lobate phytoliths and absence of saddle phytoliths. (p.18) The subfamily Panicolideae is characterized by the presence of bilobate, polylobate and quadra-lobate phytoliths and absence of saddle phytoliths. Festucoideae produces orbicular and sinuous types while Chloridoideae yields saddles (Twiss et al. 1969, Mulholland 1989, Fearn 1998, Lu and Liu 2003, Gallego and Distel 2004). Bilobate and quadra-lobate shaped phytoliths signify subfamily Panicoideae, which includes tribes: Andropogoneae, Paniceae, Maydeae, Isachneae and Oryzeae (Twiss et al. 1969, Pearsall 2000). Piperno and Pearsall (1998) concluded that the bilobate-shape silicabodies are recorded from certain genera in other subfamilies, such as Aristida (Arundinoideae), Eragrostis (Chloridoideae) and Stipa (Pooideae). (p.18)

Modern Phytolith Assemblages from the North American Great Plains Author(s): Glen G. Fredlund and Larry T. Tieszen Source: Journal of Biogeography, Vol. 21, No. 3 (May, 1994), pp. 321-335 Published by: Blackwell Publishing Stable URL: http://www.jstor.org/stable/2845533 .Accessed: 30/07/2011 09:47 characteristic of the Stipa-type is that the smaller 'top' (typically the adaxial) surface is flat, lacking any ridges or other secondary facets (Mulholland, 1989:497). Panicoid grasses do produce small numbers of the Stipa-type. Con- versely, Stipa grasses commonly produce Lobate (or Dumb-bell) forms. While the distinction is not taxonomi- cally perfect, this use of the Stipa-type in classification should provide a means for distinguishing assemblages for Stipadominated grasslands from Panicoid-dominated com- munities. Four morphotypes typical of Panicoideae grasses are recognized in this study: Simple Lobate, Panicoid-type, Cross and Other Lobate form. (p.325 326) Saddles are abundantly produced by most genera within the Chloridoid tribe, especially Bouteloua and Bouchloe which dominate the short-grass prairies of the Great Plains (Metcalfe, 1960; Twiss et al., 1969; Mulholland, 1989). (327)

Conicals are roughly circular or oval in abaxialladaxial outline and exhibit a distinctly asymmetrical cross-section. Conicals include all conical and truncated conical short-cell forms, except those defined as Keeled below. Conicals are essentially the same as the 'Rondel' type (Mulholland, 1989:495). Conicals are commonly produced by Pooideae grasses including Poa, Festuca, Agropyron; all of which are common native C3 grasses (Twiss et al., 1969; Brown, 1984; Twiss, 1992). Although this form is produced pre- dominantly by the common C3 grasses, essentially all grass subfamilies produce trace amounts of this general morpho- type (Mulholland, 1989). (p.324)

Diversity and Distributions (2003) 9, 7387

Blackwell Science, Ltd

Morphological variations of lobate phytoliths from grasses in China and the south-eastern United States
HOUYUAN LU and KAM-BIU LIU

Early phytolith researchers noted that the different subfamilies of grasses produce different phytolith shapes; for example, grasses of the subfamily Panicoideae produce dumbbell and cross-shaped phytoliths, whereas Pooideae (Festucoideae) grasses produce rondels and sinuous types, and Chloridoideae grasses produce saddleshaped phytoliths (Twiss et al., 1969). All members of the grass subfamilies Panicoideae and Oryzoideae produce the bilobate (dumbbell) type of phytoliths. However, bilobate (dumbbell) phytoliths, thought previously to be the most diagnostic marker of the Panicoideae subfamily (Twiss et al., 1969), are also present in the Chloridoideae and Arundinoideae subfamilies (Mulholland, 1989; Lu, 1998; Piperno & Pearsall, 1998; Lu & Liu, in prep.). (p.74)

Lobate phytoliths are originated from the short cells of grasses with identifiable shape characteristics. Metcalfe (1960) identified three types of dumbbell or bilobate phytoliths. The Panicoid division in Twisss classification is composed of 11 types of dumbbells and crosses (Twiss et al., 1969). Brown (1984) recognized bilobates, polylobates and crosses. Mulholland & Rapp (1992) proposed the lobate class to denote phytoliths with definite lobes, including the cross, sinuate and dumbbell types. Based on our observation and statistics of the lobate phytoliths from 85 species of modern grass plants, we found that two important parameters can be used to characterize the morphological variations in lobate phytoliths: (1) the shape of the outer margins of the two lobes and (2) the length of the shank in the lobate structure. (p.78)

Panicoideae produces dumbbell and cross-shaped phytoliths; Festucoid forms rondels and sinuous types, and Chloridoideae yields saddles (Brown, 1984; Mulholland, 1989; Fearn, 1998). The shape of individual phytoliths

from grasses can be used as an indication of C3 or C4 photosynthetic pathways. (p.84)

Journal of Archaeological Science 1989,16,489-5 11

Phytolith Shape Frequencies in North Dakota Grasses: A Comparison to General Patterns

Susan C. Mulholland

Dumb-bells and crosses were identified as characteristic of the Panicoideae; they are described as halter-shaped. (p. 491) Saddles were found in four species of the Chlorideae (Chloridoideae), but a bamboo (Bambusoideae) also contained saddles. Dumb-bells and crosses were observed in all Panicoid species examined but also occurred in Aristida, a non-Panicoid. Hilaria, which produced dumbbells, was then classified as a Panicoid and did fit the pattern; it has since been reclassified as a Chloridoid (Pohl, 1978), becoming an anomaly. Although the correlation of the three subfamilies to three phytolith shapes may hold in general, exceptions do occur. Twiss et al. (1969: 112) concluded that phytolith morphology and grass taxa must be correlated cautiously. (p. 492)

Trapezoids occur in many Chloridoid species; dumb-bells were found in some species. (p.492) The three major classifications of grass silica-cells identify similar shape types. Twiss et al. (1969) proposed three major divisions: dumb-bell/cross (Panicoid), saddle (Chloridoid), and rectangular/oblong/circular (Festucoid). Most of Metcalfes (1960) 20 shapes can fit with th ese divisions. He recognizes dumb-bells, crosses, and saddles, as well as other shapes that appear similar to Festucoid types. Brown (1984) also distinguishes crosses, dumb -bells (bilobates and polylobates), saddles, and other forms (trapezoids). Based on illustrations and descriptions in this literature, the following correlations can be made between classifications. (p. 495) The Panicoideae are dominated by dumb-bells and crosses, whereas the Chloridoideae have saddles and saddle-like dumb-bells and crosses. Most deviations from these patterns are less than 5% of the assemblage. Saddles occur regularly as a minor component of Pooids. Sinuates are not unique to the Pooideae; minor amounts are present in three species of other subfamilies. (p. 497) ()rondels occur frequently in subfamilies other than the Pooideae. Rondels, sinuates, and rectangles had been thought indicative of this subfamily. ()enough occur elsewhere so that rondels in sediments cannot be automatically attributed to the Pooideae. However, sinuates and rectangles are largely confined to the Pooideae. (p. 497-498) Chloridoid inflorescences are dominated by saddles to an even greater extent than seen in the leaf. ()Panicoid inflorescences generally continue a trend of dominant dumb -bells and crosses; however, dumb-bells decrease with respect to other forms. (p. 498)

Most sinuates, rectangles, and rondels occur in the Pooideae; saddles dominate the Chloridoideae; and the Panicoideae contain abundant dumb-bells and crosses. (p.499) The Panicoideae have long been characterized by dumb-bells and crosses [Table 1 l(a)]. Twiss et al. (1969) indicated occurrence in all Panicoid species examined. Brown (1984) found dumb-bells in all but one of his Panicoids (Panicum obtusum). Metcalfe (1960) recorded dumb-bells and crosses consistently in the Panicoideae, although Capillipedium (Andropogon venustus) lacks any of these forms. Gould & Shaw (1983) also indicated that the Panicoideae produce dumb-bells and crosses. (p.506) However, dumb-bells are not restricted to the Panicoideae [Table 10(b)]. Twiss et al. (1969) found them in a Chloridoid (Hilaria) and an Arundinoid (Aristida). Brown (1984) also recorded them in Chloridoids (Eragrosteae) and Arundinoids (Danthonia and Aristida). Metcalfe (1960) found dumb-bell and cross forms in the Arundinoids (Danthonia, Arundo, Aristida), Chloridoids (Diarrhena, Uniola, Eragrostis, Muhlenbergia, Enneapogonf, and four Pooids (Phalaris, Calamagrostis, and two Stipoids). Gould & Shaw (1983) recorded dumb-bells in four tribes of the Gramineae (Stipeae, Brachyelytreae, Nardeae, and Monermeae) as well as Arundinoids and Chloridoids. ()This study indicates that Chloridoid dumb -bells may possess a distinctive three-dimensional morphology, resembling saddles. Stipoid dumb-bells may also be distinguishable from typical Panicoid dumb-bells, although the morphology is not as distinct. In addition, Aristida may contain dumb-bells with longer shafts than in Panicoids. (p. 506)

Annals of Botany 98: 11551165, 2006 doi:10.1093/aob/mcl207, available online at www.aob.oxfordjournals.org

Phytolith Assemblages and Systematic Associations in Grassland Species of the South-Eastern Pampean Plains, Argentina
MARIANA FERNNDEZ HONAINE, ALEJANDRO F. ZUCOL and MARGARITA L. OSTERRIETH

The phytolith assemblages of the subfamily Panicoideae show two tendencies: the first characterizes the panicoid species (Paspalum quadrifarium) and is dominated by the presence of short-shank, convex-ended dumb-bells; the other is present in andropogonoid species (Sorghastrum pellitum and Bothriochloa laguroides) and is characterized by short-shank, concave/straight-ended dumb-bells. (p. 1160) Short-shank, convex-ended dumb-bells not only characterize species of the

Panicoideae subfamilies, and therefore mostly of C4 species, but are also present in species of the Pooideae (Melica brasiliana). (p. 1161)

Estuarine, Coastal and Shelf Science 58 (2003) 587 600

Phytoliths of common grasses in the coastal environments of southeastern USA

Houyuan Lu, Kam-biu Liu

The dumbbell (lobate) phytolith originates from the epidermal cells (shortcells) of Panicoideae and Oryzoideae, some Arundinoideae, and Chloridoideae subfamilies. (p. 588) The Chloridoid class (Twiss et al., 1969) consists of only two types of saddle-shaped bodies; one is the Chloridoid type (which originates from epidermal cells (short cells) of Chloridoideae, and some Bambusoideae and Arundinoideae subfamilies. (p. 588) The Festucoid class (Twiss et al., 1969) contains several geometrical types that are circular, rectangular, elliptical, or oblong. Kondo et al. (1994) and Pearsall (1989) used the same term of Festucoid class to define the short-cell phytolith types as originating from the epidermal cells of the Pooideae subfamily. (p. 588) All specimens of plants from Panicoideae (Fig. 1 and Plate 1A, D) and Oryzoideae (Plate 1B) grasses in this study have typical dumbbell type. The crosses are mostly associated with the Panicoideae subfamily. Despite being the most characteristic markers of the Panicoideae (Twiss et al., 1969), dumbbell and cross phytoliths are also found to be present in the Chloridoideae (Aristida desmantha and Ctenium aromaticum). (p. 593) The short saddle morpho-type is produced in a high proportion by the Chloridoideae. The descriptive term, _battle-axes with double edges_ (Prat, 1936), refers to the outline of the base when the body is oriented in top view. Saddle-shaped phytoliths are the dominant class of the Chloridoideae subfamily. (p. 594) A great quantity of _rondel/saddle ellipsoid_ morphotypes are isolated from Spartina alterniflora (Chloridoideae), with both round and saddle tendencies. Spartina alterniflora (cordgrass) is a tall, stiff-stemmed grass typically growing in salt marshes. (p.595) Rondels (circular to oval phytoliths) and wavy trapezoids (Mulholland, 1989) are most closely associated with Pooideae. Wavy trapezoid (Plate 3B), in particular, appears to be unique to Pooideae. (p. 597)

Journal ofArchaeologicalScience37(2010)1953e1967

Poaceae phytolithsfromtheNiassaRift,Mozambique
Julio Mercader , FernandoAstudillo , Mary Barkworth , Tim Bennett , Chris Esselmont , Rahab Kinyanjui , Dyan Laskin Grossman , Steven Simpson , Dale Walde

We also document polylobates, which are frequent in the Panicoideae. (p. 1965)

SMITHSONIANCONTRIBUTIONSTOBOTANYNUMBER85

The Silica Bodies of Tropical American Grasses: Morphology, Taxonomy, and Implications for Grass Systematics and Fossil Phytolith Identification
Dolores R. Piperno and Deborah M. Pearsall Twiss et al. (1969) proposed three major divisions of short-cell phytoliths corresponding to three dominant subfamilies native to the Great Plains of the United States: bilobate/cross = Panicoideae, saddle = Chloridoideae, and circulariovall rectangular = Pooideae. (p. 2) Circular to oval phytoliths or rondels (Mulholland, 1989), which are most closely associated with the Pooideae , also are found in the Arundinoideae, Panicoideae, and, most prominently, in the inflorescences of the Bambusoideae. Bilobate phytoliths, the most characteristic markers of the Panicoideae, also are present in the Arundinoideae, Pooideae, Bambusoideae, and Chloridoideae . Indeed, bilobates are the most common kind of silica body in phytolith assemblages from certain genera in these subfamilies, such as Aristida (Arundinoideae), Eragrostis

(Chloridoideae), and Stipa (Pooideae). The crossshaped phytolith, another panicoid marker, is common in certain Bambusoideae (tribe Olyreae) (Table 1) and occurs in small numbers in the Chloridoideae, Arundinoideae, and Pooideae (Brachypodium, Polypogon). (p.2) Many bilobates isolated from panicoid grasses exceed 20 microns in length, whereas those from the Bambusoideae, Cloridoideae, and Pooideae (with the exception of Stipa) are almost without exception shorter. (p.4) Complex bilobates or trilobates and polylobates are common in phytolith assemblages from panicoid taxa. (p.4)

Some primitive (see below) panicoids, such as Zsachne, also have assemblages dominated by tall saddles (Figure 73), but they lack the other, irregular forms. A phytolith assemblage of very tall (longer than 15 microns) saddles also suggests bamboo presence. (p.5) Furthermore, a phytolith assemblage that contains bilobates with (1) semirounded or rounded lobes and long, thin shafts (Figures 5, S), (2) squared lobes and distinct, thin or moderately thin shafts (Figure 7), or (3) lengths consistently (with a proportion of greater than 20% of all bilobates) longer than 20 microns probably contains representation from the Panicoideae, Arundinoideae (Aristida), or a genus in the Pooideae, Stipa. Presence of complex bilobates in this same assemblage indicates contribution by the Panicoideae andor Stipa (actually, such a combination is precluded by the ecological habitats of these grasses, see below). (p.5) It can be deduced that the earliest grass phytoliths were often thick pieces of silica with only one saddle- or bilobate-type face. Leaf epidermes may have contained a general assortment

of saddle-, bilobate-, and rondel-like siliceous bodies, with forms assuming traits intermediate between these major classes also being common. Most regular saddles were tall. Leaf epidermal silicifications that result in a dominance of planar, mirror-image saddles and bilobates characteristic of mainstream genera in the Chloridoideae and Panicoideae appear to be later developments. It also can be argued that chloridoid ancestors had leaf epidermes with bilobate silica bodies and that panicoid ancestors had leaf epidennes with saddle-shaped silica bodies. (p.7)

Reprinted from the Soil Science Society of America Proceedings Vol. 33, no. 1, January-February 1969 677 South Segoe Road, Madison, Wisconsin 53711 USA Purchased by Agricultural Research Service, U. S. Department of Agriculture, for official use.

Morphological Classification of Grass Phytoliths

P. C. Twiss, ERWIN SUESS, AND R. M. SMITH

The Chloridoid class contains two types of saddle-shaped bodies. The Panicoid class contains 11 types that are variations of crosses and dumbbells. (p.109) The Chloridoid class (Class 2) consists of only two types of saddle-shaped bodies. Prat (21) described them as "battle axes with double edges" and designated them as a type in the subfamily Panicoideae. (p.111)