SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 88 

Opal Phytoliths 

in Southeast Asian Flora 

Lisa Kealho fer 

and Dolores R. Piperno 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1998 
ABSTRACT 

Kealhofer, Lisa, and Dolores R. Piperno. Opal Phytoliths in Southeast Asian Flora. 

Smithsonian Contributions to Botany, number 88,39 pages, 49 figures, 5 tables. 1998.-One of 

the major uses of phytolith analysis is the reconstruction of regional environmental histories. As 

a relatively new subset of paleoecology, reference collections and studies of phytolith 

distributions and morphology are still relatively few. This article summarizes a study of 

phytolith form and distribution across a broad spectrum of 77 families of both monocotyledons 

and dicotyledons. A total of 800 samples from different plant parts of 377 species were analyzed, 

with diagnostic phytoliths occurring in nine monocotyledon and 26 dicotyledon families. These 

diagnostic types are described and illustrated herein. Poaceae phytoliths were not included in 

this review because they warrant more detailed and systematic description. The wide 

distribution of diagnostic phytoliths across all basic habitats described for Thailand, 

demonstrated herein, indicates that phytolith analysis has great potential for paleoecological 

reconstruction. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution's annual report, Annals of the Smithsonian Institution. SERIES COVER 

DESIGN: Leaf clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging-in-Publication Data 

Kealhofer, Lisa. 

p. cm. - (Smithsonian contributions to botany ; no. 88) 

Opal phytoliths in Southeast Asian flora / Lisa Kealhofer and Dolores R. Piperno. 

Includes bibliographical references. 

1. Angiosperms-Cytotaxonomy-Thailand. 2. Angiosperms-Cytotaxonomy-Asia, Southeastern. 3. Phytoliths-Thailand. 4. Phytoliths-Asia, Southeastern. I. Piperno, Dolores 

R. 11. Title. 111. Series. 

QK364.K435 1998 

571.6'82'095Mc21 98-38793 

CIP 

@ The paper used in this publication meets the minimum requirements of the American 

National Standard for Permanence of Paper for Printed Library Materials 239.48-1984. 
Contents 

Page 

Introduction ....................................... 1 

Methods ...................................... 1 

Acknowledgments ................................. 1 

2 

Monocotyledons .................................... 2 

Order ALISMATALES ................................ 2 

Order ARECALES .................................. 2 

Order CYPERALES ................................. 5 

Order ZINGIBERALES ................................ 6 

Order LILIALES ................................... 6 

Order ORCHIDALES ................................. 6 

Monocotyledons Summary .............................. 6 

Dicotyledons ..................................... 9 

Subclass ASTERIDAE ................................ 9 

Subclass CARYOPHYLLIDAE ............................ 9 

Subclass DILLENIIDAE ............................... 9 

Subclass HAMAMELIDAE .............................. 13 

Subclass MAGNOLIIDAE .............................. 17 

Subclass ROSIDAE ................................. 17 

Dicotyledons Summary ................................ 20 

Conclusion ........................................ 22 

Literature Cited ..................................... 38 

Results of Phytolith Analysis .............................. 

... 

111 

Opal Phytoliths 

in Southeast Asian Flora 

Lisa Kea l h ofer 

and Dolores R. Piperno 

Introduction 

Phytolith analysis was first used in archaeology in the early 

1970s and came of age in the 1980s in New World archaeology 

(Rovner, 1971 ; Piperno, 1988; Pearsall, 1989). Phytolith 

analysts working in other parts of the world have contributed 

important but often isolated studies of specific families or 

issues (e.g., Fujiwara et al., 1985; Rosen, 1989). Recently, we 

have begun to build a comparative phytolith collection for 

Southeast Asian flora. Much of the phytolith work to date 

focuses on the reconstruction of prehistoric agricultural 

systems; however, phytolith studies are becoming increasingly 

important for issues in geology, paleoecology, and paleontology (Piperno, 1988; Piperno and Ciochon, 1990; Piperno and 

Pearsall, 1994; Kamanina, 1997; Madella, 1997; Pinilla et al., 

1997). 

The comparative collection reviewed herein was created to 

study environmental change and the development of land use in 

Southeast Asia (e.g., Kealhofer and Piperno, 1994; Kealhofer, 

1996a, 1996b; Kealhofer and Penny, in press). Phytolith 

analysis, only recently applied in Old World tropical studies, is 

one of the primary means of palaeoecological and human 

ecological analysis (e.g., Mercader et al., in press; Runge and 

Runge, 1997). In this paper, we present the initial results of an 

investigation of phytolith production and morphology in 

Southeast Asian flora, focusing in particular on Thai flora. We 

also assess the utility of phytolith analysis for the reconstruction of vegetation patterns and plant use in this critical region. 

Lisa Kealhofer, Department of Anthropology, The College of William 

and Mary, Williamsburg, Virginia, United States. Dolores R. Piperno, 

Smithsonian Tropical Research institute, Box 2072, Balboa, Republic 

of Panama (mailing address: Smithsonian Tropical Research Institute, 

Unit 0948, APO A A 34002-0948). 

Reviewers: Arlene Miller-Rosen, Department of Archaelogy, Ben 

Gurion University, P 0. Box 653, Beer Sheva, 84 105 Israel; Deborah 

Pearsall, Department of Anthropology-American Archaeology, University of Missouri, Columbia, 103 Swallow Hall, Columbia, Missouri 

6521 1, United States. 

METHODS.-Three hundred seventy-seven species from 77 

families were analyzed for opal phytoliths. The species 

analyzed were chosen on the basis of three criteria: (1) known 

phytolith production in related genera or families from other 

regions of the world (Piperno, 1988); (2) known economic uses 

(Yen, 1982; Jacquat, 1990; Harlan, 1992); and (3) the species’ 

specificity of habitat, and therefore its potential as an 

environmental indicator (Craib, 1912, 1913; Ogawa et al., 

196 1 ; Smitinand, 1968; Kiichler and Sawyer, 1967; Stott, 

1986; Maloney, 1992). 

The 377 species represent 17 monocotyledon families (1 1 

orders, four subclasses) and 59 dicotyledon families (27 orders, 

five subclasses). Samples were taken from specimens at the 

United States National Herbarium (USNH) or were fieldcollected in northeastern Thailand (J.C. White, private collection (JCW)). Herbarium samples were from Thai plants when 

possible, but when a Thai sample was unavailable, samples 

from other East and Southeast Asian locales were used. USNH 

vouchers are identified by herbarium sheet number. 

The species were subsampled by leaf, inflorescence, fruit, 

and root when possible. Piperno (1988) showed that size and 

form differences often distinguish phytoliths from different 

plant parts, and these can have taxonomic and economic 

significance. Standard phytolith extraction techniques (wet 

ashing), as described by Piperno (1 988) and Kealhofer (1 996b), 

were followed. Samples were mounted on slides, and the slides 

were scanned three times using a light microscope, 50 mm per 

scan, at x400, to count and identify assemblages of phytolith 

forms. Diagnostic phytolith types were photographed at 

x~OO-X~OO. (Figures 1-48 herein were reduced 1 l%.) 

The basic phytolith forms (i.e., silicified tissues) identified 

by Piperno (1985, 1988) also were found in this assemblage, 

including epidermis cells, hair-base cells, hair cells, stomata, 

mesophyll cells, sclerenchyma cells, and vascular tissue. The 

forms identified and discussed herein are based on these 

previous descriptions, with added distinctions as appropriate. 

ACKNOWLEDGMENTS.-This research was made possible by 

a grant from the Andrew W. Mellon Foundation to the 

Smithsonian Tropical Research Institute, Paleoecology Sec - 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

tion. The National Science Foundation also partially supported 

the collection and analysis of material for this study. We would 

particularly like to thank Vince Pigott for his generous 

assistance. Guiselle Mora and Roque Viera provided valuable 

technical support for this project. The Thai Fine Arts 

Department, the people of Huai Pong, Thailand, Joyce White, 

and the staff at both the United States National Herbarium and 

the Smithsonian Tropical Research Institute (STRI) also 

contributed significantly to this undertaking. Particular thanks 

are due to the Photographic Services Department at STRI. 

Results of Phytolith Analysis 

Eight hundred samples were analyzed from 377 species. Of 

these, 154 samples (47 monocotyledons, 107 dicotyledons) 

contained diagnostic phytolith forms, representing nine monocotyledon and 26 dicotyledon families. This distribution 

reinforces the utility of phytoliths as significant indicators of 

many types of vegetation, not only as indicators of grasses. 

The diversity of phytolith types present in grass species 

warrants a separate presentation; therefore, the grass species 

included in this project are not discussed herein but will be 

discussed in a subsequent publication on Southeast Asian 

Poaceae. When the Gramineae (Poaceae) are excluded, 29 

monocotyledon species with diagnostic phytoliths remain. 

Only those species with diagnostic phytolith forms are 

discussed below; species studied that had no phytoliths or had 

nondiagnostic phytoliths are summarized in Table 5. Results 

are presented by taxon. Table 1 summarizes the families in 

which diagnostic phytoliths occur. 

Economic and ecological information is discussed, where 

pertinent and available, based on general botanical reviews 

(Mabberly, 1987; Heywood, 1993) and specific discussions 

of Thai flora (Craib, 1912, 1913; Ogawa et al., 1961; h

and Sawyer, 1967; Smitinand, 1968; Smitinand and Larsen, 

1972, 1975; Maxwell, 1975; Ogino, 1976; Bunyavejchewin, 

1983; Stott, 1984, 1986; Maloney, 1992). Table 2 presents the 

primary habitats of the genera and families with diagnostic 

phytoliths. 

MONOCOTYLEDONS 

Eleven of 19 orders in four of five subclasses of monocotyledons were studied: Alismatales, Arales, Arecales, Cyperales, 

Eriocaulales, Liliales, Najadales, Orchidales, Poales, Typhales, 

and Zingiberales. Nine families in these orders have diagnostic 

phytoliths (Table 3). As noted above, phytoliths in Poaceae 

will be described elsewhere. No diagnostic phytoliths were 

found in Arales (Araceae), Eriocaulales (Eriocaulaceae), 

Typhales (Typha), or Najas sp. of Najadales (Table 5). The 

diagnostic forms are described below. 

Order ALISMATALES 

The species of family Alismataceae are of particular interest 

for environmental reconstruction because they are common in 

TABLE 1.-Families with diagnostic phytoliths. ~~ 

Monocotyledons Dicotyledons 

ALISMATACEAE 

CYPERACEAE 

DIOSCOREACEAE 

GRAMINEAE ( POACEAE) 

MARANTACEAE 

MU s A c E A E 

ORCHIDACEAE 

PALMAE 

ZINGIBERACEAE 

~~ 

ANACARDIACEAE 

ANNONACEAE 

BORAGINACEAE 

BURSERACEAE 

CHRYSOBALANACEAE 

CLUSIACEAE (GUTTIFERAE) 

CONNARACEAE 

CUCURBITACEAE 

DILLENIACEAE 

DIPTEROCARPACEAE 

EBENACEAE 

EUPHORBIACEAE 

FABACEAE ( LEGUMINOSAE) 

FAGACEAE 

FLACOURTIACEAE 

LORANTHACEAE 

MAGNOLI ACEAE 

MELIACEAE 

MORACEAE 

MORINGACEAE 

MYRTACEAE 

OLEACEAE 

RHIZOPHORACEAE 

SCROPHULARI ACEAE 

STERCULIACEAE 

ULMACEAE 

freshwater-swamp habitats. The two species analyzed each 

contain two types of spheres. Both species have folded spheres; 

additionally, Caldesia has a sphere 8 ym in diameter, with 

pentagonal facets, and Ranalisma has dimpled spheres (Figure 

l), ranging from 10 pm to 20 pm in diameter. 

Order ARECALES 

Family Palmae was studied in this order, with species chosen 

from seven of the eight Palmae subfamilies found in the Old 

World. Phytoliths are common to abundant in Palmae species. 

Palm phytoliths commonly are distinctive spinulose to tabular 

spheres (Figure 2), but some subfamilies have conical to 

hat-shaped phytoliths (e.g., Caryota and Nypa; Table 3, Figure 

3). Except for these more diagnostic genera, the spinulose 

spheres vary little in shape from genus to genus, although 

sphere diameter varies between leaves and inflorescences in a 

few cases (e.g., Borassus and Calamus). Subfamilies also seem 

to produce spheres in overlapping but different size ranges 

(Table 3). For example, Borassus spheres were the largest 

phytoliths encountered, with inflorescence spheres ranging 

from 20 pm to 27 pm in diameter. Phoenicoideae spheres were 

the smallest phytoliths seen, ranging from 4 pm to 6 ym in 

diameter. Without study of more Palmae species no definite 

subfamily characterizations can be made; however, these data 

suggest a further study could reveal more diagnostic Palmae 

characteristics. 
NUMBER 88 

FIGURES 1-4 (x356).-1, Ranalisrna rostratum (Alismataceae): dimpled sphere from leaf sample, USNH 

1425829. 2, Areca catechu (Palmae): spinulose spheres from fruit sample, USNH 2455123. 3, Curyota mitis 

(Palmae): conical phytoliths from leaf and inflorescence samples, USNH 210777 I. 4, Cyperus corymbosus 

(Cyperaceae): regular, sharp peaks on leaf epidermis sample, JCW 285. 

3 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 2.-Environment-specifc genera and families with diagnostic phytoliths in Southeast Asia. Presence at 

the family level is indicated only when the family is associated in general with the given habitat. 

Taxon 

~~ ~ 

MONOCOTYLEDONS 

ALISMATACEAE 

PALMAE 

Areca 

Calamus 

COCOS 

Corypha 

Livistona 

Phoenix 

Rhapis 

[POACEAE] 

NYPa 

CYPERACEAE 

MARANTACEAE 

Phiynium 

MUSACEAE 

ZINGIBERACEAE 

Costus 

Curcuma 

Zingiber 

DIOSCOREACEAE 

ORCHIDACEAE 

Aerides 

Coelogyne 

Dendrobium 

BORAGINACEAE 

OLEACEAE 

Ligustrum 

SCROPHULARIACEAE 

Centranthera 

MORINGACEAE 

Moringa 

DILLENIACEAE 

Dillenia 

EBENACEAE 

Diospyros 

STERCULIACEAE 

CLUSIACEAE 

Calophyllum 

DIPTEROCAWACEAE 

Hopea 

Shorea 

CUCURBITACEAE 

Momordica 

Trichosanthes 

FLACOURTIACEAE 

Casearia 

Flacourtia 

Hydnocarpus 

FAGACEAE 

Lithocarpus 

Quercus 

Artocarpus 

Broussonetia 

Ficus 

Streblus 

DICOTYLEDONS 

MORACEAE 

Forest type Other habitats 

Tropical Dry Mixed Dry Hill/ Grassy/ Swamp/ 

evergreen evergreen Scrub deciduous deciduous Montane Disturbed Wet Coast 
NUMBER 88 

TABLE 2.4ontinued. 

5 

Taxon 

ULMACEAE 

Celris 

Trema 

ANNONACEAE 

Polyalihia 

Uvaria 

Manglieta 

Michelia 

Talauma 

Phyllanihus 

Sapium 

FABACEAE 

Acacia 

Albizia 

Mimosa 

Syzygium 

Podostemum 

MAGNOLIACEAE 

EUPHORBIACEAE 

MYRTACEAE 

PODOSTEMACEAE 

RHIZOPHORACEAE 

CHRYSOBALANACEAE 

Parinari 

LORANTHACEAE 

Elytranihe 

ANACARDIACEAE 

Mangifera 

BURSERACEAE 

Canarium 

Garuga 

Scutinanthe 

MELIACEAE 

Melia 

Forest type Other habitats 

Tropical Dry Mixed Dry Hill/ Grassy/ Swampi 

evergreen evergreen Scrub deciduous deciduous Montane Disturbed Wet Coast 

The typical Palmae habitat is the understory of tropical rain 

forests; however, individual species often have more specific 

habitats. For example, Nypa is found in mangrove habitats. 

Phoenix species occur in dry seasonal forests, swamp habitats, 

and mixed forests. Calamus, a climbing palm, is characteristically found in tropical rain forests but also is found in tall 

evergreen forests in northern Thailand. In the dry evergreen 

forest, Areca, Calamus, Corypha, Livistona, and Rhapis live 

near stream banks. 

Order CYPERALES 

Leaf and inflorescence samples of five species of Cyperaceae 

were studied (Table 3). Cyperaceae achenes (seeds) were not 

sampled; however, they often produce the most diagnostic 

phytolith forms. Except for Scirpus petelotii, phytoliths were 

common to abundant in both leaf and inflorescence samples. 

Although various forms are present in these species (see Table 

3), the diagnostic Cyperaceae phytolith is known as a “hat” or 

cone. Ollendorf (1992) discussed the typology of hat forms 

found in Cyperaceae but did not distinguish achene forms. The 

criteria described shape (top view), number of apices, sculpturing, presence of satellites, and cone occurrence either individually or on a platelet (Ollendorf, 1992:102). The leaf and 

inflorescence forms of the five species studied are distinguishable based on hat diameter (at base) or maximum width and on 

the shape of the hat (circular or square). Both Scirpus and 

Eleocharis species have square and round hat shapes, whereas 

Cyperus hats are round. Cyperus commonly has numerous 

satellites, either around the perimeter or distributed evenly 

across the cone. The epidermal tissue of Cyperus also has some 

diagnostic characteristics, with regular, sharp peaks on flat 

tissue (Figure 4). 

Cyperaceae species are herbaceous perennials, mainly found 

in marshes and swamps and disturbed, damp to wet habitats 

(Table 2). Cyperus in northern Thailand also occurs on 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

damp, marshy ground in evergreen forests and is occasionally 

associated with rice fields. 

Order ZINGIBERALES 

Ten species from three families of Zingiberales, Marantaceae, Musaceae, and Zingiberaceae, were analyzed (Table 3). 

In Marantaceae, distinctive folded and nodular spheres were 

found in leaf samples. Phytoliths are abundant in this family. 

Spheres are commonly 10-12 pm in diameter, but one species 

also has spheres of 5 pm (Figure 5). Inflorescence samples, on 

the other hand, had both hollow and infilled conical shapes. 

Diameters of these varied from 14 pm to 22 pm. Anticlinalepidermis forms also are characteristic of the genus Phrynium. 

The Marantaceae are most common in the New World, but 

two genera, Phrynium and Cucurligo, are common in Thai dry 

evergreen forests. Ph ynium is well represented in the 

herbaceous layer of evergreen forests. Phrynium pamiyorum 

also is a forb in higher-elevation tall evergreen forests in 

northern Thailand (Table 2). 

Musaceae phytoliths are notable for their highly diagnostic 

trough shapes (Figure 9). These shapes are abundant in leaves 

of Musa sp., but no diagnostic forms were found in the 

inflorescence samples. One sample also had multifacetedpolyhedral shapes. 

The Musaceae are known as “jungle weeds” and are 

characteristic of disturbed habitats in forests from India to 

Malesia. Musa is found in lowland dry evergreen forests. Musa 

acuminata also often grows in ravines in montane evergreen 

forests (Table 2). 

In Zingiberaceae, the most common phytolith form is a 

folded, decorated sphere. Size ranges vary by species (Figures 

6, 7), but without further analysis it is unclear if size is species 

or genus diagnostic. Sphere diameters are often quite uniform 

within species, varying by only 1-2 pm. The intensity of 

folding makes these spheres quite distinctive. Other nondiagnostic phytolith forms, particularly tracheids, are common. 

Large multifaceted polyhedrals occur in Zingiber sp. and 

Curcuma sp. Inflorescences of Zingiber sp. also have unusual, 

decorated ovoids (Figure 8). 

Zingiberales species in general are found in lowland tropical 

habitats. Zingiberaceae are perennial aromatic plants with 

multiple economic uses, including spices, medicines, and dyes. 

Curcuma is found from evergreen to deciduous forests in 

northern Thailand, whereas Zingiber primarily grows in damp 

evergreen forests. Costus is often a forb in tall evergreen 

forests. 

Order LILIALES 

Although phytoliths are rare in Dioscoreaceae, a family 

known for its economic species of yams (Dioscorea spp.), 

leaves in one species, Dioscorea membranacea, has small (5-6 

pm in diameter), tabular spheres (Figure lo). These could be 

confused with the small, ovoid, rough palm or orchid phytoliths 

because their diminutive size makes their decoration difficult to 

discern. Palm species, however, also have more diagnostic 

spinulose forms that allow them to be definitely identified. Two 

other Liliales families, Pontederiaceae and Taccaceae, either do 

not have phytoliths or have nondiagnostic phytoliths (Table 5). 

Dioscoreaceae species are herbaceous climbers in tropical 

forests, although temperate species do occur. Many Dioscorea 

species are climbers in hillside seasonal forests. Dioscorea 

membranacea is found in lowland mixed forests in Thailand 

(Table 2). 

Order ORCHIDALES 

Orchidaceae species produce small, rugulose, spherical and 

conical phytoliths (Table 3). Three species studied herein 

produce spheres in unique size ranges (4-6 pm, 8-10 pm, 

10-14 pm in diameter; Figure 11). There also are size 

differences between leaf and inflorescence spheres in Aerides. 

One species, CoelogyneJleuryi, has distinctive truncated cones 

(8-10 pm in diameter at base; Figure 12). Other than a 

silicified epidermis, few additional phytolith forms were found 

in these species. Two species, in Eria and Vanilla, do not have 

phytoliths. 

As with the Palmae, this family warrants more detailed 

investigation of phytolith variability among genera and species. 

Although Orchidaceae is a cosmopolitan family, the distribution of individual species is often ecologically constrained (see 

Table 2). 

MONOCOTYLEDONS SUMMARY 

The monocotyledon families studied herein can be identified 

using one or two of their diagnostic phytolith forms. Most 

commonly these forms are spherical or conical and show 

considerable potential to be diagnostic at least to genus. All 

monocotyledon families need detailed definition of phytolith 

variability within and between subfamilies, tribes, and species. 

This is particularly important for environmental reconstruction 

because a phytolith identified to genus in a sediment sequence 

provides more detailed environmental information than does 

family attribution. 

Although the diversity and abundance of silica present in a 

plant varies in relation to soil characteristics and plant 

senescence (Jones and Handreck, 1967; Iler, 1979; McNaughton et al., 1985), this rarely affects the presence of 

diagnostic phytolith forms. Even without considering the 

Poaceae, phytoliths from diagnostic monocotyledon genera can 

be used to distinguish a diverse set of specific habitats, as 

shown in Table 2. Orchidaceae and Zingiberaceae phytoliths 

have already aided in the identification of lowland forests, and 

phytoliths from Cyperaceae and Palmae genera have been used 

to identify coastal and swamp habitats in central Thailand 

(Kealhofer and Piperno, 1994; Kealhofer, in press). 
NUMBER 88 7 

FIGURES 5-8.-5, Phrynium pawiflorum (Marantaceae) (x 178): folded, rugulose, conical phytoliths from 

inflorescence sample, USNH 241 1368. 6, Costus speciosus (Zingiberaceae) (~356): folded, decorated sphere 

from leaf sample, USNH 2395216. 7, Elettaria cardamomum (Zingiberaceae) (x356): folded, decorated sphere 

from bract sample, USNH 206300. 8, Zingiber sp. (Zingiberaceae) (x356): decorated ovoids from inflorescence 

sample, USNH 2395209. 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 9-12 (x356).-9, Musa sp. (Musaceae): trough shapes from leaf sample, USNH 1512179. 10, 

Dioscorea membranacea (Dioscoreaceae): tabular spheres, 5-6 pm in diameter, from leaf sample, USNH 

1701266. 11, Dendrobium crumenaturn (Orchidaceae): rugulose spheres from leaf sample, USNH 221 1836. 12, 

Coelogvnefleuryi (Orchidaceae): rugulose, conical phytoliths from leaf sample, USNH 2532095. 
NUMBER 88 9 

DICOTYLEDONS 

Five of the six subclasses of dicotyledons studied herein 

have diagnostic phytoliths: Asteridae, Dilleniidae, Hamamelidae, Magnoliidae, and Rosidae (Table 4). Dicotyledon phytolith forms most commonly include hair cells or associated 

cells (e.g., hair-base cells and cystoliths); tissues related to 

respiration, such as stomata, tracheids, sclereids, and multifac - 

eted polyhedrals; and spheres, often from epidermal and 

subepidermal tissue. The sclereids, hair forms, multifaceted 

polyhedrals, and spheres have thus far proved to be the most 

diagnostic of the phytolith forms identified. 

Subclass ASTERIDAE 

Boraginaceae species (Lamiales) have short, armed hair cells 

and spheroids (Figure 13). The hair cells found in Cordia are of 

medium length, heavily armed, and side attached, lying flush 

with the epidermis. The spheres in Ehretia are very smooth and 

are associated with epidermal cells. Their size range is highly 

variable, from 6 pm to 22 pm in diameter, although most are 

10-12 ym. 

This family is most common in temperate regions; however, 

Southeast Asian genera are predominantly tropical trees. 

Lamiaceae and Verbenaceae species (Lamiales) either did 

not produce phytoliths or contained a few rare, nondiagnostic 

forms (see Table 5). 

Ligustrum (Scrophulariales: Oleaceae) contained an unusual 

hemispherical clump of subepidermal cells that may be 

diagnostic (Figure 14). 

This shrubby genus has a disjunct distribution in Europe and 

AsiaAndomalesia. In Thailand, Ligustrum is found in higher 

elevation evergreen forests. 

The leaves of one genus of Scrophulariaceae, Centranthera, 

also have diagnostic armed-hair phytoliths. This species was 

collected in deciduous tropical forests. Another Scrophulariales 

family, Lentibulariaceae, does not produce phytoliths. 

The species in other orders of Asteridae sampled, Gentianales (Apocynaceae and Asclepiadaceae), Rubiales (Rubiaceae), and Solanales (Convolvulaceae and Solanaceae), did 

not yield diagnostic phytoliths (Table 5). 

Subclass CARYOPHYLLIDAE 

In the two families sampled, Amaranthaceae and Basellaceae, phytoliths were not found in leaves or in fruits 

(Table 5). 

Subclass DILLENIIDAE 

In Dilleniidae, diagnostic phytoliths were identified in six 

orders: Capparidales, Dilleniales, Ebenales, Malvales, Theales, 

and Violales (Table 4). Thus far, families in Violales, 

specifically Cucurbitaceae and Flacourtiaceae, have yielded the 

most species with diagnostic phytolith forms. Several families 

within the Malvales, Theales, and Violales do not produce 

diagnostic phytoliths. 

Phytolith production in Capparidales is not consistent, 

CIeome (Capparidaceae) does not produce phytoliths, but 

Moringa (Moringaceae) inflorescences have very distinctive 

armed hair cells, similar to those of Streblus (Moraceae). 

Among the Dilleniales, Dillenia has unusual hair cells and 

hair bases. Both D. ovata and D. robusta have hair cells with 

either square distal or square proximal ends (see Figure 16). 

Dillenia robusta has an additional hair-cell type with a 

proximal end shaped like an arrow point (Figure 17). Both 

species have angular, fused hair-base cells that may be 

diagnostic as well (Figure 18). 

Dillenia species are tropical trees and are most common 

from China to Australia. In Southeast Asia they are found in 

evergreen to mixed deciduous forests, and certain species are 

persistent in degraded dry dipterocarp (seasonal) forests 

(Blasco, 1983). 

Both Diospyros (Ebenaceae) and Madhuca (Sapotaceae) in 

Ebenales produce phytoliths. Diospyros ebenum has large 

(19-20 pm in diameter) and small (7-9 pm in diameter) 

spheres both in leaf and seed samples. Madhuca phytoliths are 

predominantly found in vascular tissue, including sclereids and 

tracheids. Some epidermal tissue also is silicified. 

Madhuca and Diospyros species are both used for timber and 

fruit. Diospyros is often found in mixed deciduous and 

semievergreen forests. Diospyros siamensis was identified in a 

disturbed, open seasonal forest in northern Thailand. 

In Malvales few families produce phytoliths. A few species 

of Sterculiaceae have rare, diagnostic types. Eriolaena has a 

rugulose sphere 15 pm in diameter, Pterospermum and 

Sterculia species both produce sclereids, and Sterculia has 

other vascular- and hair-related forms. Melochia has a rare, 

unusual form of linked spheres (Figure 19). 

The species sampled from other Malvales families, Bombacaceae, Malvaceae, and Tiliaceae, produce phytoliths only 

rarely, and none are diagnostic (Table 5). 

In Theales, the genus Calophyllum in Clusiaceae (Guttiferae) 

has abundant phytoliths. A spherical form, shaped somewhat 

like a seed, is the most distinctive shape (Figure 20). Another 

diagnostic, slightly spinulose, elongate epidermal form also is 

common and has an unusual spike on one of its longer sides 

(Figure 21). In six other genera of Clusiaceae, phytoliths are 

absent or are not diagnostic (Table 5); however, Mammea 

produces tiny (4 pm in diameter) spheres, and Ochrocarpus 

has tiny sclereids. 

Species of Calophyllum, like those of Dillenia, are predominantly tropical trees, the timber of which is useful for its 

durability and ease of working. The species analyzed herein are 

most common in hill evergreen forests. 

The family Dipterocarpaceae (Theales), known for habitatspecific tropical trees, yielded few diagnostic phytoliths. 

Leaves from two genera, Shorea and Hopea, however, did have 

rare, decorated, spherical phytolith forms that are at least 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 13-16.--13, Cordia grandis (Boraginaceae) (x 178): short, armed hair cells from leaf sample, USNH 

1213307. 14, Ligustrum robustus (Oleaceae) (x356): subepidermal hemispherical clump From leaf sample, 

USNH 1214625. 15, Centranthem hispida (Scrophulariaceae) (x 178): armed hair from leaf sample, USNH 

294 1665. 16, Dillenia ovata (Dilleniaceae) (x 178): square proximal hair cells from leaf sample, USNH 1700567. 
NUMBER 88 11 

FIGURES 17-20.-17, Dillenia robustu (Dilleniaceae) (x356): proximal end of arrow-point hair cell from leaf 

sample, USNH 1700660. 18, Dillenia ovafa (Dilleniaceae) (x 178): angular, fused hair-base cells from leaf 

sample, USNH 1700567. 19, Melochia umbellafa (Sterculiaceae) (x 178): stalk-linked spheres from leaf sample, 

USNH 3083385. 20, Culophyllum burmanii (Clusiaceae) (~356): seed-like sphere from leaf sample, USNH 

1668055. 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 2 1-24.-21, Calophyllum burmanii (Clusiaceae) (x 178): spinulose phytolith (spike end) from leaf 

sample, USNH 1668055. 22, Hopea odorata (Dipterocarpaceae) (x356): decorated sphere from leaf sample, 

USNH I701 199. 23, Shorea obtusa (Dipterocarpaceae) (x356): decorated sphere from fruit sample, USNH 

22 1 181 9. 24, Solena heterophylla (Cucurbitaceae) (x 178): armed hair cells from leaf sample, USNH 2553066. 
NUMBER 88 13 

genus-specific (Figures 22, 23) and have been found in 

sediments (Kealhofer, 1996b). The Vatica and Dipterocarpus 

species investigated, however, did not have diagnostic phytolith forms. 

Dipterocarp species, most abundant in Malesia, also are 

widespread in Thailand (dipterocarp species comprise 45% of 

Thai forest species). The culturally modified dry dipterocarp 

forest type is defined by a few dominant Dipterocarpaceae 

species (Pentacme siamensis, Shorea obtusa, Dipterocarpus 

tuberculatus, D. obtusifolius). Ogawa and coworkers (1 961) 

identified two dipterocarp forest subhabitats: Pentacme-Shorea 

(drier) and Dipterocarpus tuberculatus-Dipterocarpus obtusifolius (wetter). In general, the Pentacme-Shorea habitat is 

species poor, including only a few grass species (such as 

Arundinaria spp.), Cycas siamensis, Phoenix humilis, P: 

acaulis, and relatively infrequent arboreal species. Composition 

varies regionally. Different Dipterocarpaceae (particularly 

Dipterocarpus spp.) are found in mixed and evergreen forests, 

but their relative abundance is much lower as species diversity 

increases (Ogawa et al., 1961:67). Hopea odorata is common 

in dry evergreen forests. 

Elatinaceae species (Theales) do not produce diagnostic 

phytoliths. 

By far the most abundant and distinctive phytoliths of the 

Dilleniidae are found in the Violales, particularly in Cucurbitaceae and Flacourtiaceae (Table 4). The Cucurbitaceae are 

known for their segmented hair cells, some of which are armed. 

Eight genera of Cucurbitaceae from two subfamilies were 

analyzed. Only Momordica did not produce armed or segmented hair cells in their leaves. None of the inflorescences 

produced diagnostic phytoliths. No fruits were analyzed, but 

distinctive, faceted spheres have been identified with certain 

parts of the fruit (Bozarth, 1986). 

Citrullus, Cucumis, Gymnopetalum, Lufla, and Trichosanthes all have distinctive segmented hair cells of various sizes, 

whereas Solena and Mukia have armed hair cells (Figures 24, 

25). Other diagnostic features include curved hair tips 

(Gymnopetalum) and unique hair bases (Citrullus, Cucumis). 

Momordica contains cystoliths, often seen linked at the top to 

form a tripodal arrangement (Figure 26). In only one genus, 

Solena, were two species examined, and these appeared to have 

species-specific forms. Further analyses within these genera 

may yield more phytolith forms diagnostic at the species level. 

These Cucurbitaceae genera are all from the Asian tropics 

and are most commonly lianas, or climbing vines. They are 

often weeds in disturbed habitats. Members of this family were 

some of the earliest species domesticated. 

Samples from members of Flacourtiaceae (Violales) only 

sporadically produced phytoliths. Nondiagnostic phytoliths are 

common in Flacourtiaceae leaves, and diagnostic forms are 

relatively rare. The fruits of several species contain spherical 

shapes, possibly associated with epidermal cells. These spheres 

are 10-14 pm in diameter, with a dimple (Casearia) or with 

flat tissue attached (Hydnocarpus). Scolopia flowers have a 

rare, spheroid, kidney-bean-shaped phytolith. In leaves, epidermal forms also are diagnostic in a few cases, particularly for 

Hydnocarpus (Figure 27). Zdesia has an unusual, multifacetedpolyhedral form (Figure 28). 

The Flacourtiaceae are fairly cosmopolitan trees and shrubs 

with wide distribution in the tropics and subtropics. Casearia 

and Flacourtia are both found in dry mixed forests and 

occasionally in dry deciduous forests. Hydnocarpus is a 

common shrub in tropical evergreen to dry evergreen forests. 

Only a few species, namely Scolopia and Casearia (usefil 

timber) and Flacourtia (edible fruit), have economic uses. 

Samples from species in two other families in the Violales, 

Caricaceae and Passifloraceae, did not yield diagnostic phytoliths (Table 5). 

Subclass HAMAMELIDAE 

Three of the four Hamamelidae families analyzed revealed 

characteristic phytoliths: Fagaceae (Fagales) and Moraceae and 

Ulmaceae (Urticales) (Table 4). The one Juglandaceae species 

tested did not produce diagnostic phytoliths (Juglandales; 

Table 5). 

In Fagaceae (Fagales), the leaves of Lithocarpus have 

distinctive, faceted, spherical polyhedrals (Figure 29) and 

spheres. Quercus samples produced spheres from the nut-shell. 

The Fagaceae are, in general, a temperate family (oaks), but 

in the tropics they appear in montane evergreen forests. Three 

Quercus species and one Lithocarpus species have been 

identified as particularly common on the higher slopes of 

montane forests in northern Thailand (Kuchler and Sawyer, 

1967). Other Quercus species, such as Q. mespilifoliodes and 

Q. kerrii, however, are found in dry dipterocarp forest 

understory. 

The order Urticales was more intensively investigated 

because it is more extensively represented in the tropics. Two 

families, Moraceae and Ulmaceae, demonstrated a wide variety 

of phytoliths. In both families, hair cells and cystoliths are the 

most diagnostic forms. 

Among the Moraceae, three Artocarpus species were 

studied. Leaves of all species of Artocarpus revealed armed hair 

cells (Figure 30). One species of Ficus and one species of 

Streblus also have diagnostic armed hair cells, and these are the 

only two genera with diagnostic phytoliths in either fruit or 

inflorescence parts. Streblus has various armed and unarmed 

hair cells (Figures 3 1,32). Broussonetia and Malaisia hair cells 

also are distinctive but are not armed (Figure 33). Some species 

contain up to five different types of hair-cell phytoliths, with 

characteristic attachment locations, curvature, striations, and 

other features. Other genera, such as Taotrophis, have simpler 

but still diagnostic hair cells. Ficus annulata, Broussonetia 

kasinoki, Artocarpus elasticus, and Morus alba leaves all 

contain cystoliths as well (e.g., Figure 34). Ficus hispida 

epidermis has unusual hexagonal epidermal(?) cells, and F. 

annulata fruit epidermal cells have a wrinkled surface 

(Figure 35). 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 25-28.-25, Mukia maderaspatana (Cucurbitaceae) (x 178): armed hair cells from leaf sample, USNH 

2039897. 26, Momordica charantia (Cucurbitaceae) (x 178): top-linked cystoliths from leaf sample, USNH 

2039869. 27, Hydnocarpus anthelmintheca (Flacourtiaceae) (x356): epidermal phytoliths from leaf sample, 

USNH 1427948.28, Idesiapolycarpa (Flacourtiaceae) (~356): multifaceted polyhedral from leaf sample, USNH 

2986601. 
NUMBER 88 

FIGURES 29-32.-29, Lithocurpus acuminatissima (Fagaceae) (x356): multifaceted polyhedral from leaf sample, 

USNH 1701025.30, Artocarpus elusticus (Moraceae) (x 178): armed hair cells from leafsample, USNH 29391 50. 

31, Streblus asper (Moraceae) (x356): armed hair cells from leaf sample, USNH 2064818. 32, Streblus asper 

(Moraceae) (x356): armed hair cell from leaf sample, USNH 20648 18. 

15 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 33-36.-33, Broussonetia kasinoki (Moraceae) (x 178): unarmed hair cells from leaf sample, USNH 

1597387. 34, Arrocarpus elasticus (Moraceae) (x 178): cystolith from leaf sample, USNH 2939 150. 35, Ficus 

annulara (Moraceae) (x 178): wrinkled epidermal cells from leaf sample, USNH 2602693.36, Celtis cinnamomea 

(Ulmaceae) (x356): unusual cystoliths from leaf sample, USNH 1212982. 
NUMBER 88 17 

Silicified epidermal cells, hair bases, stomata, tracheids, 

mesophyll, and nondiagnostic hair cells are common in this 

fami 1 y . Members of the Moraceae, and particularly Ficus species, 

are widespread but are often associated with river or stream 

habitats in gallery forests. Moraceae species include a variety 

of economically important tropical plants. Both Artocarpus and 

Ficus species grow in evergreen to semievergreen forests as 

well as often being found in middle and higher elevations. 

Ficus is a huge genus, with a broad variety of trees, shrubs, and 

lianas. Broussonetia is often in the shrub understory of 

evergreen forests, and Streblus has been identified in open 

disturbed seasonal forests in northern Thailand. 

Many characteristics of the Moraceae also were found in the 

Ulmaceae. Hair-cell, hair-base, and cystolith forms are common in both leaves and fruit (Table 4). Gironniera has armed 

hair cells, and the two species of Celtis have unusual cystoliths 

(Figure 36). A spiny, irregular phytolith form was found in 

Celtis. Two Trema species have distinctive, small, anticlinal 

epidermal cells (Figure 37). Trema orientalis also revealed an 

unusual, pitted, striated phytolith form (Figure 38). Gironniera 

leaf epidermis is distinctive. No phytoliths were found in 

Holoptelea leaves. 

The tropical and subtropical tribe Celtideae of Ulmaceae is 

composed predominantly of trees, with some shrubs. Many of 

these genera produce decay-resistant timber. Celtis is common 

in scrub forests, whereas species of Trema are often found in 

evergreen forests. 

Subclass MAGNOLIIDAE 

Six of the eight orders of Magnoliidae were sampled, but 

only one, Magnoliales (3 1 species), produced diagnostic 

phytoliths (Table 4). No phytoliths were found in Aristolochiales. Some samples from the families Illiciales, Laurales, 

Piperales, and Ranunculales did produce phytoliths, but they 

were rare to uncommon. 

Among the Magnoliales, only three families are present in 

Thailand. Two of these families, Magnoliaceae and Annonaceae, contain characteristic phytoliths. Species sampled from 

the third family, Myristicaceae, did not produce silica bodies. 

In Annonaceae, the characteristic phytoliths are relatively 

large, multifaceted polyhedrals (Figure 39). Artabotlys has 

multifaceted spheres, and other multifaceted polyhedrals were 

found in Polyalthia species and in Sageraea elliptica. Sageraea 

phytolith forms are unique in having tiny spinulose protrusions 

as decoration. Sclereid forms are present in Annonaceae, and 

diagnostic phytoliths occur in leaves of Goniothalamus, 

Sageraea, and Uvaria (Figure 40). Uvaria fruit has large, 

irregular spheroids (1 8 pm in diameter). Vascular-tissue and 

stomata forms also are well represented in these species. 

Annonaceae species include both trees and shrubs, several of 

which produce important edible fruit. All species studied herein 

are most common in tropical, lowland evergreen forests and 

often occur in mixed deciduous forests. Polyalthia grows in 

tropical evergreen forests. Uvaria was identified in scrub 

forests, but it also is found in evergreen forests. 

Magnoliaceae species also have multifaceted bodies, particularly Michelia. Distinctive decorated and faceted epidermal 

phytolith forms were found in Talauma and Manglieta (Figure 

41). Phytolith production, however, seems to vary by species 

because no diagnostic phytoliths were found in Michelia 

Jloribunda. 

The Magnoliaceae are most abundant in southeast Asia, and 

certain species are used for timber. Genera analyzed include 

trees of evergreen forests; Michelia is often found at the edge of 

lowland forests as well as in (mossy) montane evergreen 

forests, whereas Talauma is associated with streams or hills, 

and Manglieta is typical in the diverse, higher-slope forests of 

northern Thailand. 

Subclass ROSIDAE 

The last subclass studied, Rosidae, comprises 18 orders, 11 

of which were sampled. Species in four of these orders do not 

produce diagnostic phytoliths: Apiales, Celastrales, Podostemales (see below), and Rhamnales (Table 5). Phytolith 

production is not consistent in those orders that do produce 

phytoliths. For example, in Santalales, some families have 

phytoliths, and others do not. 

In the Euphorbiaceae family (Euphorbiales), phytoliths were 

common in three of the four genera sampled, Sapium, 

Phyllanthus, and Manihot. In Manihot esculenta leaves, 

slightly irregular spheres, 6-10 pm in diameter, were common 

(Figure 42). Fruit samples of Phyllanthus also produced 

spheres, but these are nodular and 6-15 pm in diameter. The 

epidermal cells of Sapium also are diagnostic, with an irregular, 

rough surface decoration (Figure 43). 

Euphorbiaceae is a large and cosmopolitan family, but its 

richest concentration of genera is in Indomalaysia. Two of the 

genera with diagnostic phytoliths are economically useful, 

particularly Manihot esculenta (cassava, a New World species), 

an important edible root crop, and Phyllanthus, used medicinally and for dyes. Sapium is most common in tall broadleaf 

evergreen forests. 

In the Fabales, many species of Leguminosae (Fabaceae) 

produce abundant phytoliths, but few forms are diagnostic 

(Tables 4, 5). Three subfamilies of Fabaceae were analyzed: 

Caesalpinoideae, Mimosoideae, and Papilionoideae. None of 

the species tested in Caesalpinoideae produced diagnostic 

phytoliths. 

In the Mimosoideae, the stomata may need further investigation because they seem to take some unusual shapes (Albizia, 

Pithecellobium; Figure 44). Acacia also may have diagnostic 

hair cells and hair-base cells. The epidermis in many species, 

particularly in subfamily Papilionoideae, also is distinctive. 

Tracheid or hair-base-cell forms are common to all subfamilies 

in which silica is present (-50% of those sampled). 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 37-40.-37, Trema cannabina (Ulmaceae) (x356): anticlinal epidermal cells from fruit sample, USNH 

2880555. 38, Trema orientalis (Ulmaceae) (~89): pitted, striated epidermis from leaf sample, USNH 2769297. 

39, Polyalthia suberosa (Annonaceae) (~356): large, multifaceted polyhedrals from leaf sample, USNH 1 171 29 1. 

40, Goniothalamus marcani (Annonaceae) (x 178): sclereid from leaf sample, USNH 1700786. 
NUMBER 88 

FIGURES 4 1-44 (x 356).4 1, Tuluuma longifolia (Magnoliaceae): decorated, faceted phytolith from leaf sample, 

USNH 2407533. 42, Manihot esculenra (Euphorbiaceae): nodular spheres, 6-10 pm in diameter, from leaf 

sample, USNH 2395207. 43, Supium indicum (Euphorbiaceae): rough surface decoration on fruit epidermis 

sample, USNH 1701 697. 44, Pithecellobium duke (Fabaceae: Mimosoideae): stomata from leaf sample, USNH 

2423 177. 

19 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Fabaceae is a very large family and is distributed in a wide 

range of habitats. The subfamilies of Fabaceae have more 

restricted distributions, however, and Mimosoideae and Caesalpinoideae include tropical to subtropical trees and shrubs. 

Particular species are of enormous economic importance, 

especially in the Papilionoideae. The species analyzed are 

common in scrub and evergreen forests. Acacia catechu and 

Albizzia are frequent in mixed deciduous forests. Vines of 

Acacia and Mimosa inhabit the scrub understory of disturbed, 

open, seasonal forests. 

In Myrtales, two of the nine families present in Southeast 

Asia were investigated: Combretaceae and Myrtaceae. No 

diagnostic phytoliths were found in the three Combretaceae 

species studied, although phytoliths were present (Table 5). Of 

the two species of Syzygium (Myrtaceae) tested, one, S. 

ieptantha, has an unusual, very tall stomata phytolith form. 

This family is predominantly tropical and is composed 

mostly of large shrubs and trees. Syzygium is commonly found 

in dry evergreen forests on hillsides in Thailand and has a 

broader distribution elsewhere. 

Podostemales are known to produce phytoliths; however, the 

species studied herein, Podostemum subulatus, does not 

(Piperno, 1988). Species in this order are found in flowing, 

fresh-water environments. 

Three species in the family Rhizophoraceae were sampled. 

Phytoliths were not common in any of these. Sphere forms are 

present in all three species, although sphere size and surface 

decoration vary by species (Table 4). 

Rhizophoraceae species are most common in coastal 

mangrove forests in Thailand. Many taxa are used economically for timber or charcoal. 

Two of the six families of Rosales present in Thailand, 

Rosaceae and Chrysobalanaceae, were tested; Rosaceae did not 

have diagnostic phytoliths (Table 5). In Chrysobalanaceae one 

species, Parinari annamense, incorporates irregular but smooth 

spheres of 6-12 pm diameter in the leaves and 6-10 pm 

diameter in the inflorescence. The fruit has very large spheres 

(20-85 pm in diameter). 

Chrysobalanaceae species, trees and shrubs, are found in the 

lowland tropics (mostly in the New World), and Parinari 

includes important fruit and timber trees. This genus ranges in 

habitat from deciduous to evergreen forests, but in Thailand it 

seems to be most common in mixed deciduous forests. 

One sample tested seems to be from the Connaraceae 

(labeled “Connarc?’ in the United States National Herbarium), 

and it also has diagnostic epidermal forms, suggesting that this 

family warrants further investigation. 

In Santalales, two of the four families present in Thailand 

were investigated. No phytoliths were found in the Olacaceae 

species tested. In Loranthaceae, only leaves produced phytoliths. Two distinctive stomata forms were found in Elyrranthe, a set of elongated/palisaded cells and a set of subepidermal 

spherical cells in a clump (Figure 45). Unusual epidermal-cell 

forms also were found in this genus. 

Loranthaceae includes shrubby tropical parasites. Elytranthe 

is found in evergreen forests and often impedes the regrowth of 

hardwood trees. 

Five of the 10 families of Sapindales present in Thailand 

were sampled, and species in two of the families, Anacardiaceae and Burseraceae, have diagnostic phytoliths. All of the 

other families (Meliaceae, Rutaceae, and Sapindaceae) are 

phytolith producers, but the forms are rare or nondiagnostic 

(Table 5). Vascular tissue in both Rutaceae and Sapindaceae, 

particularly tracheids and adjoining tissue, occasionally includes unusual spiny forms. 

Three Anacardiaceae species were tested, and the leaves of 

all three produced phytoliths; however, those in Spondias 

pinnata are not diagnostic (Table 5). Mangifera indica (mango) 

and Rhus sp. both have unique folded spheres (Figure 46). 

Mangifera species are common in upland and lowland 

evergreen forests. 

Burseraceae species show a wide range of unique phytoliths. 

Epidermal phytoliths from both fruit and leaves are the most 

distinctive. Silicified anticlinal and polyhedral epidermal cell 

shapes are both represented. Canarium, Commiphora, Daciyodes, Garuga, and Scutinathe species all have thick, decorated 

epidermal cells (Figure 47). Commonly, one surface is smooth 

and the opposite surface is spikey. Canarium fruits often have 

unusual, very smooth, large spheroid shapes as well (Figure 

48). One Canarium species has a distinctive acorn-shaped hair 

cell (Figure 49); a related form was found in Garuga. Silicified 

stomata, tracheids, hair cells, and hair-base cells are common to 

abundant in Burseraceae. 

Burseraceae is a tropical family of trees and shrubs. It forms 

a common constituent of lowland deciduous and dipterocarp 

forests in southeast Asia. Canarium, Daciyodes, and Garuga 

are the three Burseraceae genera most common in this lowland 

habitat, and all have tree species with usehl timber. Garuga 

pinnata has been identified in mixed deciduous forests. 

Canarium kerrii and C. subulatum are common constituents of 

the dry dipterocarp forest. Protium has been identified in 

deciduous, closed-canopy forests in northern Thailand. 

Meliaceae species only rarely produce phytoliths, but Melia 

inflorescences have rare, small spheres (5-7 pm in diameter). 

Species in this family are known for producing good timber, 

such as mahogany, in lowland tropical forests. 

DICOTYLEDONS SUMMARY 

The most common of the distinctive dicotyledon phytolith 

types are spheres (including spheroids), armed and segmented 

hair cells, faceted polyhedrals, and decorated epidermal cells. 

The variability seen in dicotyledons differs from that seen in 

monocotyledon families. Phytolith presence is not as predictable, and orders commonly have families and species both with 

and without phytoliths. A more limited range of phytolith 

forms also is represented, with greater frequency of vascular 
NUMBER 88 

FIGURES 45-48 (x356).45, Elytranthe umpullaceae (Loranthaceae): stomata from leaf sample, USNH 

1700671, 46, Mangifera indica (Anacardiaceae): folded spheres from leaf sample, USNH 595090. 47, 

Commiphoru cuuduta (Burseraceae): thick, decorated epidermis from hit sample, USNH 2805849. 48, 

Canarium kerrii (Burseraceae): large, smooth spheroids from fruit sample, USNH 2436020. 

21 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cones of the monocotyledons. Dicotyledon sphere types, 

however, appear to be good indicators of trees in various 

forested habitats. The genera and families with diagnostic 

phytoliths include a wide range of habitats but dominate the 

lowland evergreen, mixed deciduous, and dry deciduous 

forests. Specific habitats are discernable based on the unique 

complement of genera identified in soil samples. 

Conclusion 

The results presented herein demonstrate the abundance and 

diversity of diagnostic phytoliths in Old World tropical flora. 

These phytoliths are consistent with those described by Piperno 

(1 985, 1989) in New World taxa, confirming the uniformity of 

phytolith forms among taxa with Old and New World 

distributions. 

Based on the habitat distribution of their parent plants, 

phytoliths can be correlated with a wide range of specific 

environments. Plant age, soil chemistry, and other environmental factors may influence phytolith abundance and the 

diversity of forms present in a given plant; however, diagnostic 

forms are produced despite variable conditions. This is clearly 

seen when the patterns of production and morphology 

presented herein are compared with New World taxa (Piperno, 

1988, 1989). These results hrther demonstrate that different 

kinds of habitats and economic patterns are identifiable in soil 

phytolith sequences. The increasing definition of diagnostic 

phytolith types enhances the value of phytolith sediment 

sequences for complementing and expanding the reconstruc - 

tion of paleo-environments, and the human interactions with 

these environments, currently available from pollen and 

geomorphological analyses. 

FIGURE 49.--Cunarium kerrii (Burseraceae) (x400): acom-shaped hair cell 

from leaf sample, USNH 2436020. 

forms (e.g., sclereids) than in monocotyledons. Short cells are 

not represented, nor are the distinctive tabular spheres and 
TABLE 3.-Phytolith forms identified in monocotyledon species. Some forms (elongates, mesophyll, stoma, tracheids) are not 

diagnostic. Phytoliths listed under “Other forms” provide information on additional characteristic forms. Measurements are of 

the diameter. (an = angular; cu = cupped; dc = decorated; fc = faceted; fd = folded; ge = geometric; gr = grainy; Ig = large; 

MFP = multifaceted polyhedrals; nd = nodular; nr = narrow; ov = ovoid; rd = round; rg = rugulose; tr = truncate; sc = scalloped; 

sm = smooth; sp = spinulose; sq = square; tb = tabular; strip = long, thin polyhedrals; us = unsegmented.) 

Taxon Phytolith Intra-/Intercellular forms Hair Epidermal forms 

abundance sphere conical elongate bag-shaped Hair base Stoma Tracheid anticlinal polyhedral 

ALISMATIDAE: ALISMATALES 

ALISMATACEAE 

Caldesia oligococca (F. Muell.) Buche 

Ranalisma rostratum Stapf in Hook. 

leaf 

leaf 

ARECIDAE: ARECALES 

PALMAE (ARACEAE) 

Areca catechu L. 

ARECOIDEAE 

fruit 

leaf 

BORASSOIDEAE 

Borassus Jlabelrifer L. 

leaf 

inflorescence 

CARYOTOIDEAE 

Catyota miris Lour. 

leaf 

inflorescence 

fruit 

COCOSOIDEAE 

Cocos nucifea L. 

leaf 

inflorescence 

fruit 

leaf 

fruit 

Elaeis guineensis Jacq. 

LEPIWCARYOIDEAE 

Calamus walkeri Hance 

leaf 

inflorescence 

NYPOIDEAE 

Nypa fruticans Wurmb 

leaf 

fruit 

PHOENICOIDEAE 

Phoenix paludosa Roxb. 

leaf 

CYPERACEAE 

COMMELINIDAE: CYPERALES 

Cyperus compactus Retz. 

inflorescence 

rare 

rare 

fd, 8 pm 

fd, 10-20 pm 

- 

- rd 

- 

rd 

- 

sp, 6-10 pm 

sp, 5-1 1 pm 

common 

common 

sp, 10-15 pm 

sp, 20-27 pm 

common 

common 

common 

common 

common 

13-18 pm 

nd, 10-18 pm 

hats 

- 

- 

sp, 6-1 1 pm 

common 

common 

common 

sp, 5-8 pm 

sp, 5-8 pm 

sp, 5-8 pm 

sp, 6-10 pm 

sp, 4- 10 pm 

common 

common 

abundant 

abundant 

sp, 8- 12 pm 

sp, 6-10 pm X 

7-10 pm 

7-8 pm 

common 

abundant 

sp, 4-6 pm common 

common 
TABLE 3.4ontinued. 

'hytolith Intra-/lntercellular forms Hair Epidermal forms 

ibundance sphere conical elongate bag-shaped Hair base Stoma Tracheid anticlinal polyhedral 

Taxon 

Cyperus corymbosus Rottb. 

leaf 

inflorescence 

leaf 

inflorescence 

leaf 

inflorescence 

Scirpus peieloiii 

inflorescence 

Cyperus pulcherrimus Willd. ex Kunth 

Eleochuris dulcis Trinus ex Hensch. 

ZINGIBERIDAE: ZINGIBERALES 

Phrynium purviflorum Roxb. 

MARANTACEAE 

leaf 

inflorescence 

leaf 

inflorescence 

MUSACEAE 

Musa sp. 

leaf 

Musu sp. 

leaf 

Phrynium placenturium (Lour.) Merr. 

ZINGIBERACEAE 

Alpiniu speciosa (J.C. Wendl.) K. Schun 

Amomum cf. pavieunum Pierre 

Costus speciosus (Koen.) Sm. 

leaf 

leaf 

leaf 

fruitmract 

Curcumu sp. 

leaf 

Elelfurin curdumomum (L.) Maton 

bract 

Lanpus gulunga (L.) Merr. 

leaf 

Zingiber sp. 

leaf 

Zingiber sp. 

inflorescence 

LILIIDAE: LILIALES 

DIOSCOREACEAE 

Dioscoreu membranacea Pierre 

leaf 

LILIIDAE: ORCHIDALES 

ibundant 

ibundant 

:ommon 

ibundant 

:ommon 

:ommon 

rare 

abundant 

abundant 

common 

abundant 

abundant 

abundant 

abundant 

20 pm 

20 pm 

15-18 pm 

15 pm 

sq, 6-8 pm 

sq, 8, 12 pm 

s9 

X X 

X - 

ov, 4-1 lpm 

X 

X 

fd/rg, 5 pm 

17-28 pm 

fd, 10- 12 pm 

- 

8-22 pm 

- 

dc, 4-6 pm 

uncommon 

abundant dc, 15-16 pm 

abundant fd, 10 pm 

common 

common rg, 8-15 pm 

abundant tb, 9-15 pm 

uncommon 

uncommon 25-35 

- 

- 

- 

pm 

x- dc X - 

- sm 

common tb, 5-6 fim 
TABLE 3 .--Continued. 

Taxon 

ORCHIDACEAE 

Aerides fulcatum Lindl. & Paxt. 

leaf 

inflorescence 

leaf 

leaf 

Coelogynefleuryi Gagnep. 

Dendrobium crumenatum Sw. 

Phytolith Intra-/Intercellular forms Hair Epidermal forms 

abundance sphere conical elongate bag-shaped Hair base Stoma Tracheid anticlinal polyhedral 

- - - - -- - - bumpy 

uncommon rg/tb, 9 pm 

rare rg, 10-14 pm 

common - tr, 8-10 pm 

common rg, 4-6 pm 

- - - -- - - - - 

- - - - X 

strip 

-- 

- - - -- - - - - 

TABLE 4,Phytolith forms identified in dicotyledon species. Phytoliths listed under “Other forms” provide information on 

unusual characteristic forms. Some forms (elongates, mesophyll, stoma, tracheids) are not diagnostic. Measurements are of 

diameter. (2 = two forms present; cr = circular; dc = decorated; dm = dimpled; fd = folded; ge = geometric; gr = grainy; 

ir = irregular; MFP = multifaceted polyhedrals; nd = nodular; rg = rugulose; sc = scalloped; sf = short, fat; sg = segmented; 

sk = spikey; sm = smooth; sm sk = smooth top, spikey bottom; sp = spiney; sq = square; st = striated; strip = long, thin 

pol yhedrals.) 

Taxon Phytolith Intra-/Intercellular forms Hair Cysto- Epidermal forms M

abundance sphere elongate Hair base lith Stoma Tracheid Sclereid anticlinal polyhedral 

ASTERIDAE 

LAMIALES 

BORAGINACEAE 

Cordia grandis Cham. 

Ehretia sp. 

leaf 

leaf 

SCROPHULAIUALES 

OLEACEAE 

leaf 

Ligustrum robustus (Roxb.) Blume 

SCROPHULARIACEAE 

Centrunthera hispida Benth. 

leaf 

inflorescence 

DILLENIIDAE 

CAPPARIDALES 

MORINGACEAE 

inflorescence 

DILLENIACEAE 

Moringa oleifera Lamarck 

DILLENIALES 

Dillenia ovata H0ok.f. & Thomson 

leaf 

abundant - - annedx - X X 

common sm,6-22pm - X 2 - X - 

- - - x? - uncommon X X 

uncommon - - armed x - X X 

rare - - med x - - - 

common - 

rare 

- - - 

- X X 

- - X 
TABLE 4.4ontinued. 

hytolith Intra-/Intercellular forms Hair Cysto- Epidermal forms M

bundance sphere elongate Hair base lith Stoma Tracheid Sclereid anticlinal polyhedral p

Taxon 

Dilleniu robustu 

leaf 

EBENALES 

EBENACEAE 

Diospyros ebenum Koenig 

leaf 

Seed 

leaf 

MALVALES 

Madhuca sp. 

STERCULIACEAE 

Erioluena cuudollei Wall. 

Melochiu umbellatu (Houtt.) Stapf 

Pterospermum semisugittuJblium Ham. 

Sterculiu foetida L. 

Sterculiu pexu Pierre 

fruit 

leaf 

leaf 

leaf 

leaf 

THEALES 

CLUSIACEAE (GUTTIFERAE) 

Culophyllum burmunii Wight 

leaf 

DIPTEROCARPACEAE 

Hopea odorutu Roxb. 

Shorea obtusa Wall. 

leaf 

fruit 

VIOLALES 

CUCURBITACEAE 

Citrullus vulguris Eckl. & Zeyh. 

Cucumis trigonis Roxb. 

Gymnopetalum cochinchinensis (Lour.) 

leaf 

leaf 

Merr. 

leaf 

leaf 

leaf 

leaf 

leaf 

leaf 

Luffs cylindricu (L.) M. Roem. 

Momordicu charantiu L. 

Mukiu muderusputuna (L.) M. Roem. 

Solenu amplexicaulis (Lam.) Gandhi 

2 fused ncommon 

20 pm, 7 pm 

19pm. 9pm 

ncommon 

ncommon 

incommon X - 

are 

are 

:ommon 

incommon 

:ommon 

X - 

ibundant 20-24 pm 

dc, 20-24 

fd, 18-21 pm 

X 

- 

:ommon 

very rare 

- X 

- - 

Zommon 

uncommon - 

x nucleate 

- - 

- - 

- 

- 

common - 

- rare 

common 

rare 

- 

- 

- X 

common 

abundant 
abundance sphere elongate Hair base lith Stoma Tracheid Sclereid anticlinal polyhedral p

TABLE 4.4ontinued. 

Phytolith Intra-/Intercellular forms Hair Cysto- Epidermal forms M

Taxon 

Solena heterophylla Lour. 

Trichosanthes anaimalaiensis Bedd. 

leaf 

leaf 

FLACOURTIACEAE 

Casearia virescens Jacq. 

Flacourtia jangonias (Lour.) Steud. 

Hydnocarpus anthelmintheca Laness. 

fruit 

fruit 

leaf 

fruit 

leaf 

leaf 

fruit 

flower 

HAMAMELIDAE 

FAGACEAE 

Hydnocarpus ilicijolius SK 

Idesia polycarpa Max. 

Scolopia acuminata Clos. 

FAG ALES 

Lithocarpus acuminatissima (BI.) Rehdc 

leaf 

inflorescence 

exocarp 

URTICALES 

MORACEAE 

Quercus dumosa Nutt. 

Artocarpus dadah Miq. 

Artocarpus elasticus Reinw. ex Blume 

Artocarpus nitidus Trec. 

Broussonetia kasinoki Siebold & Zucc. 

Ficus annulata BI. 

leaf 

leaf 

leaf 

leaf 

leaf 

fruit 

leaf 

seed 

leaf 

Ficus hispida L.f. 

Maluisia scandens (Lour.) Planch. 

uncommon 

abundant - - 

- - 

very rare dm, 10-14 pm - 

very rare 

abundant 

uncommon 

abundant 

uncommon 

very rare 

very rare 

- X 

uncommon 

uncommon 

X 

6-16 pm 

4-10 pm uncommon 

rare 

rare 

rare 

common 

common 

rare 

abundant 

abundant 

abundant 

X 

wrinkled 

hexagon 

hexagon 

X 
TABLE 4.4ontinued. 

'hytolith ha-llntercellular forms Hair Cysto- Epidermal forms M

ibundance sphere elongate Hair base lith Stoma Tracheid Sclereid anticlinal polyhedral p

Taxon 

Morus ulbu L. 

Streblus usper Lour. 

leaf 

leaf 

inflorescence 

leaf 

Taxofrophis macrophylla (BI.) Merr. 

ULMACEAE 

Celtis cinnumomeu Lindl. 

leaf 

fruit 

leaf 

fruit 

leaf 

fruit 

fruit 

leaf 

inflorescence 

MAGNOLIIDAE 

Celtis collinsae Craib 

Gironnieru subuequalis Planch. 

Trema cunnubina Lour. 

Tremu orientulis (L.) Blume 

MAGNOLIALES 

ANNONACEAE 

Alphonsea philastreana Finet & Gagnep 

Artabofrys siamensis Miq. 

Goniothalumus murcuni (Blume) 

leaf 

leaf 

Ho0k.f. & Thomson 

leaf 

inflorescence 

leaf 

Miliusu bulunrae Lcschen. ex A. DC. 

Mitrephoru collinsue (Blwne) Ho0k.f. 

& Thomson 

leaf 

leaf 

leaf 

Polyulthiu jucunda Blume 

Polyulthiu suberosa (Roxb.) Thwaites 

Sugerueu ellipfica (A. DC.) Ho0k.f. 

& Thompson 

leaf 

(Ivuriu micrunthu (A. DC.) Ho0k.f. & 

Thomson 

leaf 

fruit 

:ommon X 

incommon 

are 

X 

- 

ibundant strip 

are 

ibundant 

- 

sk bottom 

:ommon 

uncommon 

- 

dc 

uncommon 

common 

- X 

- curved 

X 

- 

common 

common 

uncommon 

common 

common 

- - 

faceted - 

X 

X 

- 

sk - 

- 

uncommon 

rare - X 

uncommon 

uncommon 

rare 

sk X - 

uncommon 

rare 

rare 
TABLE 4.4ontinued. 

Phytolith Intra-/Intercellular forms Hair Cysto- Epidermal forms M

abundance sphere elongate Hair base lith Stoma Tracheid Sclereid anticlinal polyhedral p

Taxon 

MAGNOLIACEAE 

Munglietu gurrettii 

leaf 

Micheliu chnmpucu L. 

exocarp 

Micheliu sp. 

leaf 

fruit 

leaf 

Tuluumu longifoliu (Blume) Ridl. 

ROSIDAE 

EUPHORBIALES 

EUPHORBIACEAE 

Munihot esculentu Crantz 

Phyllunthus ucidus (L.) Skeels 

Supium indicum P. Browne 

leaf 

fruit 

leaf 

fruit 

FABALES 

FABACEAE: MIMOSOIDEAE 

Acacia cutechu (L.f.) Willd. 

Albizia gurretti I.C. Nelson 

Pithecellobium duke Roxb. 

FABACEAE: PAPILIONOIDEAE 

fruit 

leaf 

leaf 

Cunavuliu cuthurticu Thouars 

Millettia pendula Brown 

leaf 

leaf 

MYRTALES 

MYRTACEAE 

S’ygium ieptuntha Gaertner 

leaf 

RHIZOPHORALES 

RHIZOPHORACEAE 

inflorescence 

leaf 

leaf 

Bruguieru cylindricu (L.) Blume 

Ceriops tugal Perrottet 

Rhizophoru upiculutu Blume 

uncommon 

rare 

uncommon 

rare 

uncommon 

- 

- 

- 

- 

- 

common ir, 6- I0 pm 

uncommon nd, 6-1 5 pm 

- 

abundant 

uncommon 

SP 

SP 

- 

- 

- 

uncommon 

uncommon 

common 

gr 

X 

rare 

abundant 

X 

X 

- 

- angular 

rare 

uncommon 

uncommon 

uncommon 
TABLE 4.4ontinued. 

‘hytolith Intra-llntercellular forms Hair Cysto- Epidermal forms M

bundance sphere elongate Hair base lith Stoma Tracheid Sclereid anticlinal polyhedral p

Taxon 

ROSALES 

CHRYSOBALANCEAE 

Parinari annamense Aublet 

leaf 

inflorescence 

fruit 

CONNARACEAE 

“Connarc??” sp. 

fruit 

SANTALALES 

LORANTHACEAE 

Elytranthe ampullaceae (Roxb.) G. Don 

leaf 

SAPINDALES 

ANACARDIACEAE 

Mangifea indica L. 

leaf 

bundant 

ommon 

ommon 

:ommon 

tbundant 

are 

are 

are 

ibundant 

:ommon 

rare 

abundant 

Eommon 

common 

common 

rare 

ir,6-12pm - xx X - 

ir,6-lOpm - - - - - X 

- 

20-85pm - - - - - X 

- xx - X X X 

Rhus sp. 

leaf 

BURSERACEAE 

Canarium kerrii L. 

Canarium litforale BI. 

fruit 

leaf 

leaf 

leaf 

Canarium zeylaninrm L. 

Commiphora caudata (Wight & 

Am.) Engl. 

fruit 

leaf 

fruit 

leaf 

Dactyodes rostrata (Blume) H.J. Lam 

Garuga pinnata Roxb. 

Scutinathe brunnea Thw. 

- 

X 

- - X - X X X 

sk bottom 

- 

sm sk 

knobby 

MELIACEAE 

Melia azedarach L. 

inflorescence 
NUMBER 88 31 

TABLE 5.-Species without phytoliths or with nondiagnostic phytoliths. Measurements are of diameter. 

Taxon 

MONOCOTYLEDONS 

ALISTMATIDAE 

ALISMATALES: ALISMATACEAE 

Caldesia ologococea 

NAJADALES: NAJADACEAE 

Najas sp. 

NAJADALES: POTAMOGETONACEAE 

Potamogeton hens L. 

ARECIDAE 

ARALES: ARACEAE 

Alocasia macrorrhiza (L.) G. Don f. 

Amorphophallus corrugatus Decne 

Colocasia esculenta Schott 

Cyrtosperma johnstonii Gri ffith 

ERIOCAULALES: ERIOCAULACEAE 

Eriocaulon luzulaefolium L. 

Eriocaulon siamense L. 

Eriocaulon siamense 

TYPHALES: TYPHACEAE 

Dpha angustifolia L. 

COMMELINIDAE 

ZIGIBERIDAE 

Musa sp. 

Curcuma sp. 

Elettaria cardamomum (L.) Maton 

Hedychium ellipticum J. Koenig 

Kaempferia pulchra L. 

LILIALES: DIOSCOREACEAE 

Dioscorea alata L. 

Dioscorea collinsae L. 

Dioscorea membranacea Pierre 

Dioscorea sp. 

LILIALES: PONTEDERIACEAE 

Eichhornia crassipa Solms 

Monochoria vaginalis Presl. 

Monochoria vaginalis 

LILIALES: TACCACEAE 

Tacca sp. 

ORCHIDALES: ORCHIDACEAE 

Coelogvnejleuryi Torrey 

Eria dasyphylla Lindley 

Vanilla griflthii Miller 

ZINGIBERALES: MUSACEAE 

ZINGIBERALES: ZINGIBERACEAE 

LILIIDAE 

DICOTYLEDONS 

ASTERIDAE 

GENTIANALES: APOCYNACEAE 

Aganonieron polymorphum Spire 

Alstonia scholaris R. Br. 

Rauvo&a verticillata L. 

Atherolepis pierrei Cost. 

Telosma minor Craib 

LAMIALES: BORAGINACEAE 

Cordia grandis Cham. 

Ehretia sp. 

Perilla frutescens L. 

GENTIANALES: ASCLEPIADACEAE 

LAMIALES: LABIATEAE (LAMIACEAE) 

Present(P)/ 

Plant part absent(A) Phytolith forms present 

seed 

leaf 

leaf, seed 

leaf 

leaf 

leaf 

leaf 

leaf, infloresence 

infloresence 

seed 

reed, seed 

infloresence 

infloresence 

infloresence 

leaf, infloresence 

leaf 

leaf, seed 

leaf, infloresence, seed 

infloresence 

root 

leaf, root 

leaf 

infloresence 

leaf, infloresence 

root 

leaf, root 

leaf 

leaf 

leaf 

leaf 

leaf 

leaf 

infloresence 

fruit 

leaf, infloresence 

A 

P 

A 

P 

A 

A 

A 

A 

P 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

P 

P? 

A 

P 

A 

A 

irregular, rugulose spheroid 

curved, short unsegmented hair cells; tracheids 

long, thin epidermal polyhedrals with central spikes 

cupped cell (intrusives) 

8- 15 pm spheres, could be intrusive 

lots of intrusives 

rugulose, globular, irregular body 

P hair-base cells? segments, sphere (10 pm), hexagonal 

P epidermis, stomata 

P sphere (10 pm), tracheid 

bodies, anticlinal epidermis 

A 

P tracheids 

A 

A 

A 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 5.4ontinued. 

Taxon 

Pogostemon glaber Desf. 

Pogostemon glaber 

LAMIALES: VERBENACEAE 

Callicarpa arborea L. 

Tectona grandis L.f. 

RUBIALES: RUBIACEAE 

Paedena foetida L. 

SCROPHULARIALES: LENTIBULARIACEAE 

Ultricularia aurea Lour. 

SCROPHULARIALES: OLEACEAE 

Jasminum nervosum L. 

Jasminum nervosum 

Jasminum sambac Aiton 

Ligustrum robustus L. 

SOLANALES: CONVOLVULACEAE 

Ipomoea batatas (L.) Lam. 

Ipomoeapes-caprae (L.) R. Br. 

SOLANALES:OLANACEAE 

Capsicum fnctescens L. 

Solanum album L. 

CARYOPHYLLIDAE 

CARYOPHYLLALES: AMARANTHACEAE 

CARYOPHYLLALES: BASELLACEAE 

Aerva scandens Forssk. 

Basella alba L. 

DILLENI IDAE 

CAPPARIDALES: CAPPARIDACEAE 

DILLENIALES: DILLENIACEAE 

Cleome gynandra L. 

Dillenia ovata L. 

Dillenia robusta L. 

EBENALES: EBENACEAE 

Diospyros kaki L.f. 

Madhuca sp. 

MALVALES: BOMBACACEAE 

Bombax ceiba L. 

Bombax valentonii L. 

Durio zibethinus Murray 

Durio zibethinus 

Ochroma pyramidale Sw. 

Gossypium arboreum L. 

Gossypium herbaceum L. 

Hibiscus sabdarifa L. 

MALVALES: STERCULIACEAE 

Ambroma augusta L.f. 

Byttneria pilosa Roxb. 

Byttneria pilosa Roxb. 

Commersonia bartrania (L.) Men. 

Eriolaena caudollei Wall. 

Helicteres elongata Wall. 

Helicteres isora L. 

Helicteres isora 

Heritiera littoralis Dryand 

Melochia umbellata 

Pterospermum diversifolium 

Pterospermum semisagittafolium 

Sterculia pexa Pierre 

MALVALES: MALVACEAE 

Ham. 

Present(P)/ 

Plant part absent(A) Phytolith forms Present 

leaf 

infloresence 

leaf, infloresence 

leaf, infloresence 

leaf, infloresence 

leaf 

leaf 

infloresence 

leaf 

fruit, branch 

leaf 

leaf 

leaf, infloresence, fruit 

leaf 

leaf, seed 

leaf, fruit 

leaf, pod 

in fl o r e s e n c e 

infloresence 

leaf 

fruit 

infloresence, seed, pod 

leaf 

leaf 

infloresence 

leaf 

leaf, bract 

leaf 

leaf. infloresence 

leaf, seed 

leaf 

infloresence 

leaf, infloresence 

leaf 

leaf, infloresence 

leaf 

infloresence 

leaf 

leaf, infloresence 

leaf, infloresence 

leaf 

infloresence 

P 

A 

A 

P 

A 

A 

P 

P 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

P 

P 

A 

A 

P 

P 

A 

A 

A 

A 

A 

P 

A 

A 

P 

A 

P 

A 

A 

A 

A 

P 

A 

cupped epidermal cells, anticlinal epidermis, hair cells 

epidermis with hemispherical bumps; short, unsegmented 

hairs; cell clumps 

long, thin epidermal polyhedrals; tracheids; spiny, sockshaped 

tracheids 

tracheids, long, thin epidermal polyhedrals 

tracheids, epidermal cells 

hair cell bases? segments 

epidermal cells, small spheres 

irregular epidermal cells, tracheids 

epidermal cells, mesophyll, tracheids, stomata 

tracheids 

tracheids, sclereids 
NUMBER 88 

TABLE 5.4ontinued. 

33 

Taxon 

MALVALES: TILIACEAE 

Berrya mollis Roxb. 

Bertya mollis 

Colona flagrocarpa Cav. 

Grewia acuminata L. 

Grewia acuminata 

Grewia conforta L. 

Grewia conforta 

Triumfetta rhomboidea L. 

THEALES: CLUSIACEAE (Guttiferae) 

Calophyllum burmanii Wight 

Calophyllum thorelli Pierre 

Calophyllum thorelli 

Clusia rosea L. 

Cratoxylon polyanthum Blume 

Garcinia gracilis Pierre 

Garcinia gracilis 

Garcinia mangostana L. 

Garcinia xanthochymus T. AnHypericum japonicum Thunb. 

Mammea siamensis L. 

Mammea siamensis 

Ochrocarpus siamensis L. 

Ochrocarpus siamensis 

THEALES: DIPTEROCARPACEAE 

Dipterocarpus obtusifolius Miq. 

Dipterocarpus obtusifolius 

Hopea odorata Roxb. 

Shorea henryi Roxb. ex Gaertner 

Shorea henryi 

Shorea siamensis Roxb. ex Gaertner 

Shorea obtusa Wall. 

Vatica cinerea L. 

Vatica cinerea 

Vatica sp. 

Bergia ammanioides L. 

Bergia ammanioides 

VIOLALES: CARICACEAE 

Carica papaya L. 

VIOLALES: CUCURBITACEAE 

Citrullus vulgaris Shard. 

Coccinia cordifolia L. Cogn. 

Gynostemma sp. 

Hodgsonia heteroclita Ho0k.f. & 

Lagenaria siceraria Ser. 

Luffa cylindrica (L.) M. Roem. 

Momordica charantin L. 

Trichosanthes anaimalaiensis Bedd. 

VIOLALES: FLACOURTIACEAE 

Casearia grewiaefolia Vent. 

Casearia grewiaefolia 

Casearia virescens Pierre 

Etythrospermum zeylanicum (GaertFlacourtia indica (Bum. f.) Merr. 

Flacourtia indica 

Flacourtia jangonias 

Flacourtia sepiaria Roxb. 

Homalium damrongianum Craib 

Homalium tomentosum Benth. 

derson 

THEALES: ELATINACEAE 

Thomson 

ner) Alston 

Presen t(P)/ 

Plant part absent(A) Phytolith forms present 

leaf 

infloresence 

leaf 

leaf 

seeds 

infloresence 

leaf 

leaf, infloresence 

fruit 

leaf 

infloresence 

leaf, infloresence 

leaf, fruit 

leaf 

infloresence 

leaf 

leaf, fruit 

leaf, infloresence 

leaf 

fruit 

leaf 

infloresence, fruit 

leaf 

fruit 

infloresence 

leaf 

infloresence 

leaf, infloresence 

leaf 

leaf 

infloresence 

leaf, infloresence 

infloresence, root 

leaf, stem 

leaf 

infloresence 

leaf, infloresence 

leaf 

leaf 

leaf 

seeds, pod 

infloresence 

infloresence 

fruit 

leaf 

leaf 

leaf, fruit 

leaf 

fruitbract 

leaf 

leaf, infloresence 

leaf, infloresence 

leaf, infloresence 

P 

A 

P 

P 

A 

P 

A 

A 

P 

P 

A 

A 

A 

P 

A 

A 

A 

A 

A 

P 

P 

A 

P 

A 

P 

P 

A 

A 

A 

P 

A 

P 

A 

P 

A 

P 

A 

P 

A 

A 

A 

A 

P 

A 

P 

A 

P 

P 

A 

P 

A 

P 

A 

stomata, tracheids, epidermal cells, cell clumps 

long hair cells, spherical epidermal cells 

spherical epidermal cells 

tracheids, barely silicified stomata 

tracheids, rugulose spheres 

tracheids 

tracheids 

tiny spheres (4 pm) 

tiny sclereids? same as Mammea? 

epidermal cells, stomata, hair-base cells 

tracheids (fat) 

tracheids (fat), hair-base cells 

stomata, epidermal cells, hair-base cells 

unsegmented V-shaped hair cells, stomata, hair-base cells 

epidermal cells, spheres 

hair-cell fragments 

hair-base cell clumps 

segmented hair cells 

tracheids, dendritic forms 

tracheids, stomata 

tracheids 

tracheids, grainy elongates 

tracheids. hair cells 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 5.--Continued. 

Taxon 

Hydnocarpus ilicifolius King 

Itoa orientalis Hemsl. 

Ryparosa javanica Men. 

Ryparosa javanica 

Scolopia acuminata Clos. 

Scolopia spinosa (Roxb.) Warb. 

Trichadenia zeylanica Thw. 

Xylosma sp. 

Passijlora bijlora L. 

Passijlora biflora 

FAGALES: FAGACEAE 

VIOLALES: PASSIFLORACEAE 

HAMAMELIDAE 

Castanopsis diversifolia (D. Don) 

Quercus lobaia Nee 

Quercus dumosa L. 

JUGLANDALES: JUGLANDACEAE 

Engelhardia serrata Blume 

Engelhardia serrata 

URTICALES: MORACEAE 

Allaeanthus hizii Standley 

Artocarpus dadah Miq. 

Artocaips nitidus Trec. 

Broussonetia kasinoki Seibold & 

Broussonetia papyrifera Vent. 

Broussonetia papyr ifera 

Cudrania poilanei Gagnep. 

Cudrania poilanei 

Cudrania cochinchininsis Lour. 

Spach. 

zucc. 

Cudrania cochinchininsis 

Ficus sp. 

Ficus sp. 

Malaisia scandens (Lour.) Planch. 

Morus alba L. 

Moms sp. 

Taxotrophis macrophylla (BI.) Men. 

Holoptelea intregifolia (Roxb.) PI. 

Trema cannabina Lour. 

URTICALES: ULMACEAE 

MAGNOLIIDAE 

ARISTOLOCHIALES: ARISTOLOCHIACEAE 

ILLICIALES: ILLICIACEAE 

Aristolochia pothieri L. 

Illicium sp. 

Illicium sp. 

Kadsura chinensis Juss. 

Schisandra perulata Michaux 

Schisandra pemlata 

LAURALES: LAURACEAE 

Beilschmiedia glauca Nees 

Beilschmiedia glomerata Nees 

Cinnamomum iners Schaeffer 

Endiandra macrophylla R. Br. 

Endiandra macrophylla 

Litsea panamanja Lam. 

Litsea panamanja 

ILLICIALES: SCHISANDRACEAE 

Present(P)/ 

Plant part absent(A) Phytolith forms present 

fruit 

leaf 

leaf 

fruit 

leaf 

leaf, infloresence 

leaf 

leaf. flower 

leaf 

infloresence 

leaf 

leaf 

leaf, nut 

leaf 

seed 

leaf 

fruit 

fruit 

infloresence 

leaf 

fruit 

leaf 

fruit 

leaf 

fruit 

leaf 

fruit 

infloresence 

infloresence 

leaf 

infloresence 

leaf 

leaf 

leaf, fruit 

leaf 

fruit 

leaf, fruit 

leaf 

infloresence 

leaf, fruit 

fruit 

leaf, infloresence 

leaf 

infloresence 

leaf 

infloresence 

A 

P 

P 

A 

A 

A 

A 

A 

P 

A 

P 

A 

A 

P 

A 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

A 

P 

P 

P 

A 

A 

A 

A 

P 

A 

A 

A 

P 

A 

A 

A 

P 

A 

P 

A 

unsegmented hair cells, grainy elongates 

grainy elongates, tracheids, stomata, polyhedral epidermal 

cells 

stomata, epidermal cells with central sphere 

stomata, hair-base cells, epidermal cells 

anticlinal epidermis 

tracheids, anticlinal epidermis 

V-shaped small hair cells 

epidermis 

unsegmented hair cells of variable size 

long, curved fiber-like hair cells and hair-base cells 

small hair cells with cystoliths 

epidermis, stomata 

tiny hair cells, tracheids, hair-base cells 

stomata, tracheids, polyhedral epidermal cells, fused hairsmall, unsegmented hair cells; hair-base cells 

pronged hair cells; armed, unsegmented hair cells; tracheids; 

base cells? 

unsegmented hair cells 

funnel-shaped, unsegmented hair cells of variable length 

tracheids, 2 types of unsegmented hairs with cystoliths 

long, curved, armed and short hair cells; tracheids; cystoliths 

stomata, MFPs, tracheids 

long, thin epidermal polyhedrals 

epidermal cells? 

tracheids 
NUMBER 88 

TABLE 5.4ontinued. 

35 

Taxon 

MAGNOLIALES: ANNONACEAE 

Anomianthus heterocarpus Zoll. 

Artabotrys siamensis Miq. 

Cananga sp. 

Cananga sp. 

Mitrephora collinsae Craib 

Orophea polycarpa A. DC. 

Polyalthia suberosa (Roxb.) Thwaites 

Popowia fomentosa Endl. 

Unona? chinensis 

Unona chinensis 

Manglietia garrettii Craib 

Michelia champaca L. 

MAGNOLIALES: MAGNOLIACEAE 

Michelia champaca 

Michelia floribunda Finet & 

Gagnep. 

MAGNOLIALES: MYRISTICACEAE 

PIPERALES: PIPERACEAE 

Myristica fragrans Houtt. 

Piper longum L. 

Piper longum 

Piper sarmentosum Roxb. 

RANUNCULALES: MENISPERMACEAE 

Tiliacora triandra Diels 

ROSIDAE 

APIALES: APIACEAE (Umbelliferae) 

Coriandrum sativum L. 

Eryngium foetidum L. 

Eryngium foetidum 

Foeniculum vulgare Miller 

Foeniculum vulgare 

Pimpinella wallichians L. 

CELASTRALES: AQUIFOLIACEAE 

Ilex trijlora L. 

EUPHORBIALES: EUPHORBIACEAE 

Cladogynos orientalis Zipp. ex Span. 

Manihot esculenta Crank. 

Phyllanthus acidus Skeels 

Phyllanthus emblica L. 

Phyllanthus emblica 

Sapium baccatum P. Browne 

FABALES: FABACEAE (Leguminosae) 

CAESALPINOIDEAE 

Bauhinia bracteata L. 

Cassia occidentalis L. 

Cassia tora L. 

Crudia caudata Schreber 

Tamarindus indica L. 

Tamarindus indica 

Acacia catechu Willd. 

Acacia catechu 

Acacia farnesiana (L.) Willd. 

Acacia rugata Voight 

Acacia tomentosa Willd. 

MIMOSOIDEAE 

Present(P)/ 

Plant part absent(A) Phytolith forms present 

leaf, fruit 

infloresence 

leaf 

fruit 

infloresence 

leaf 

fruit 

leaf 

leaf 

fruit 

fruit 

leaf 

fruit 

leaf, fruit 

leaf, infloresence 

leaf 

infloresence 

leaf 

leaf 

leaf, infloresence 

leaf 

infloresence 

leaf 

infloresence 

leaf, infloresence 

leaf 

leaf, infloresence, seed 

seed 

leaf 

leaf 

fruit 

leaf, infloresence 

leaf, infloresence 

leaf, infloresence, fruit 

leaf 

leaf 

leaf 

infloresence 

leaf 

infloresence 

leaf 

infloresence 

leaf 

A 

P 

P 

A 

P 

A 

A 

A 

P 

A 

A 

P 

A 

P 

A 

P 

P 

P 

P 

A 

P 

A 

A 

P 

A 

A 

A 

P 

A 

P 

A 

P 

P 

A 

A 

P 

P 

A 

P 

A 

A 

A 

P 

epidermal cells 

tracheids, polyhedral epidermal cells 

tracheids 

tracheids, stomata 

tracheids; stomata; barely silicified, polyhedral epidermal 

cells; elongated, multifaceted bodies with tracheid impressions 

stomata 

small, irregular MFPs; epidermal cells 

geometric epidermal cells; curved, irregular MFPs; domeshort, unsegmented, fat hairs; geometric epidermal cells; 

shaped cells? 

cells with spheres 

tissue with embedded spheres 

tracheids 

anticlinal epidermis 

subrectangular bodies, epidermal cells 

spherical epidermal cells, grainy elongates, cupped cells 

stomata 

tracheids 

spikey epidermal cells, stomata, folded spheres, rugulose 

epidermal cells, stomata 

spheres 

tracheids, unsegmented hairs, cupped cells 

fat tracheids, stomata, knobby epidermal cells, hair-base 

cells, square-base hair cells 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

~ 

Taxon 

Adenanthera pavonina L. 

Albizia garretti F. Nielsen 

Albizia lucidior Durazz. 

Albizia lucidior 

Albizia vialeana Pierre 

Leucaena glauca Benth. 

Mimosa pudica L. 

Neptunia oleracea Lour. 

Neptunia oleracea 

Prosopis julrjlora (SW.) DC. 

Xylia dolabrifornus Benth. 

Xylia dolabrifornus 

Arachis hypogaea L. 

Cajanus cajan (L.) Huth. 

Cajanus cajan 

Cajanus cajan 

Canavalia ensiformis (L.) DC 

Dalbergia dongnaensis L. f. 

Glycine mar Merr. 

Indigofera galegoides L. 

Lablab purpureus (L.) Sweet 

Lablab purpureus 

PAPILIONOIDEAE 

Lablab purpureus 

Lathyrus sativa L. 

Millettia pendula Brown 

Millettia pendula 

Phaseolus sp. 

Phaseolus sp. 

Pterocarpus indicus Willd. 

Pterocarpus indicus 

Sesbania grandiyora Dew. 

Vicia sativa L. 

Vicia sativa 

MYRTALES: COMBRETACEAE 

Combretum grrffhii Loefl. 

Combretum griffithii 

Lumniizera racemosa Willd. 

Lumnitzera racemosa 

Rrminalia bellerica (Gaertner) Roxb. 

MYRTALES: MYRTACEAE 

Syzygium ieptantha Gaertner 

Syzygium aromaticum (L.) Men. & 

Perry 

PODOSTEMALES: PODOSTEMACEAE 

Podosiemum subulatus Michaux 

RHAMNALES: RHAMNACEAE 

Colubrina longipes Back. 

Ziziphus jujuba Miller 

Bruguiera cylindrica Lam. 

Ceriops tagal (Perrottet) C. Robinson 

Rhizophora apiculata L. 

Agrimonia eupatoria L. 

Eriobotva iengyuehensis Lindley 

Potentilla fulgens Wall. 

Potentilla kleiniana Wight 

RHIZOPHORALES: RHIZOPHORACEAE 

ROSALES: ROSACEAE 

TABLE 5.--Continued. 

Present( P)/ 

Plant part absent(A) Phytolith forms present 

leaf, pod 

fruit 

leaf 

seed 

leaf 

leaf, fruit 

leaf 

leaf 

leaf 

leaf 

Pod 

leaf 

shell, nut 

inflorescence 

pod with seeds 

leaf 

leaf 

leaf 

leaf 

leaf, infloresence 

leaf 

pods 

flower 

leaf 

seed 

Pod 

leaf 

Pod 

leaf 

infloresence 

leaf, infloresence 

leaf 

seeds 

leaf 

infloresence 

leaf 

infloresence 

leaf, infloresence 

infloresence 

leaf 

plant 

leaf 

leaf, seed 

leaf 

infloresence 

infloresence 

leaf 

leaf, infloresence 

leaf, infloresence 

leaf 

A 

P 

P 

A 

P 

A 

A 

P 

A 

P 

A 

P 

A 

P 

A 

P 

P 

A 

P 

A 

P 

P 

A 

A 

P 

P 

P 

A 

A 

P 

A 

A 

P 

P 

P 

A 

P 

A 

P 

A 

A 

A 

A 

P 

A 

A 

A 

A 

A 

P 

subepidermal cells 

tracheids, square-base hair cells, stomata, cupped cells 

grainy elongates 

anticlinal epidermis 

tracheids, anticlinal epidermis, stomata 

tracheids 

long hair cells 

hair cells, stomata, mesophyll, concave cells 

epidermal cells, tracheids 

circular striated cells, tracheids 

hair cells, mesophyll, stomata, hair-base cells 

unsegmented hair cells, epidermal cells with central protrusion 

irregular rugulose spheres 

tiny spikey spheres 

long hair cells with arrow tip, anticlinal epidermis, tracheids, 

hair-base cells 

amorphous silica bodies 

tracheids 

tracheids, angular epidermal cells, stomata, cupped cells 

long, unsegmented hairs; tracheids; long, thin epidermal 

polyhedrals 

epidermal cells with spheroids attached 

hair-base cells? mesophyll? 

tracheids, epidermal cells 

hair cells 
NUMBER 88 

TABLE 5.--Continued. 

37 

Taxon 

Prunus arborea L. 

Prunus hosscusii L. 

Pyracantha angustijolia (Franch.) 

Raphiolepis indica (L.) Lindl. 

Rubus ellipticus Sm. 

Rubus efferatus Craib 

Spiraea cantonensis L. 

SANTALALES: LORANTHACEAE 

Laranthus pentapetalus Jacq. 

Loranthus pentapetalus 

Elytranthe ampullaceae (Blume) 

Schneid. 

Blume 

SANTALALES: OLACACEAE 

Olax scandens L. 

Scorodocarpus borneensis (Baillon) 

Becc. 

SAPINDALES: ANACARDIACEAE 

Mangijera indica L. 

Rhus sp. 

Spondias pinnata Kurz. 

SAPINDALES: BURSERACEAE 

Canarium album Raeuschel 

Canarium commune L. 

Canarium kerrii L. 

Canarium littorale BI. 

Canarium nigrum L. 

Canarium pimela Koen 

Canarium pimela 

Canarium subulatum Guillaum 

Canarium sumatranum Benth. 

Canarium zeylanicum L. 

Commiphora caudata (Wight & Am.) 

Garuga pinnata Roxb. 

Garuga pinnata 

Engl. 

Santiria apiculata A.W. Benn 

Santiria laevigata Blume 

Scutinathe brunnea Thw. 

SAPINDALES: MELIACEAE 

Aglaia chandocensis Lour. 

Aglaia chandocensis 

Azadirachta indica Juss. 

Dysoxylem cochinchinensis Blume 

Dysoxylem cochinchinensis 

Melia azedarach L. 

SAPINDALES: RUTACEAE 

Clausena excavata Burm.f. 

Clausena excavata 

SAPINDALES: SAPINDACEAE 

Allophyllus cobbe (L.) Rausch. 

Nephelium hypoleucum L. 

Nephelium hypoleucum 

Present(P)/ 

Plant part absent(A) Phytolith forms present 

leaf, fruit 

infloresence 

leaf. infloresence 

leaf, fruit 

leaf, infloresence 

leaf, infloresence 

leaf, infloresence 

leaf 

infloresence 

fruit 

leaf, infloresence, fruit 

leaf 

infloresence 

fruit 

leaf 

leaf, infloresence 

leaf, infloresence 

leaf 

fruit 

leaf 

leaf 

infloresence 

leaf, infloresence 

leaf, infloresence 

infloresence 

leaf 

leaf 

leaf, infloresence 

leaf, infloresence 

leaf, infloresence 

fruit 

leaf 

infloresence 

leaf, infloresence 

leaf 

infloresence 

leaf 

leaf 

infloresence 

leaf 

leaf 

fruit 

A 

A 

A 

A 

A 

A 

A 

P 

A 

A 

A 

A 

A 

A 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

P 

A 

A 

P 

A 

P 

P 

P 

P 

P 

A 

hair-base cells, spheres? (rare) 

stomata, geometric epidermal cells, tracheids, cell clumps 

with spheres 

hair cells; hair-base cells; stomata; tracheids; long, thin 

hair cells, stomata, hair-base cells 

hair cells; hair-base cells; stomata; MFPs, 1 side spiculate; 

mesophyll; armed, unsegmented hair cells; conical cells (see 

stomata, tracheids 

stomata, hair cells, hair-base cells 

hair cells 

stomata, tracheids, hair cells, hair-base cells, epidermal cells 

stomata, tracheids, MFPs, hair cells, hair-base cells 

hair cells, tracheids, stomata, hair-base cells 

tracheids, stomata 

epidermal polyhedrals 

epidermal cells 

Table 4) 

unsegmented hair cells (size variable), stomata, tracheids 

curved, unsegmented hair cells; tracheids; cystoliths; hairstomata, tracheids, hats (round crown), knobby epidermal 

grainy, irregular epidermal cells; tracheids; hair cells; 

stomata, tracheids, hair-base cells 

base cells; stomata 

cells 

hair-base cells; stomata 

tracheids, cell clumps (stomata?) 

tracheids; long, thin epidermal polyhedrals; cupped cells 

epidermal cells 

stomata, tracheids, epidermal cells, cupped cells 

tracheids 

stomata, tracheids, epidermal cells, mesophyll, spheres 

tracheids, vascular forms?, stomata, cupped cells 
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