SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 53 

A Scanning Electron Microscope 

Survey of the Epidermis 

of East African Grasses, I1 

Patricia G. Palmer 

and Alice E. Tucker 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1983 
ABSTRACT 

Palmer, Patricia G., and Alice E. Tucker. A Scanning Electron Microscope 

Survey of the Epidermis of East African Grasses, 11. Smithsonian Contributions 

to Botany, number 53, 72 pages, 52 plates, 1983.-In developing a reference 

collection as a standard for using fossil grass cuticles as a paleoecological tool, 

we have examined leaf epidermal features of grasses with SEM. Descriptions 

are presented of the abaxial and adaxial surfaces of the leaves of 26 East 

African species, comprising nine tribes of the subfamily Pooideae-Poeae. 

Bromeae. Brachypodieae, Triticeae, Meliceae, Aveneae, Phalarideae, Agrostideae, Stipeae. 

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

in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf clearing from the 

katsura tree Cercidiphjllurn japonicum Siebold & Zuccarini. 

Library of Congress Cataloging in Publication Data 

Palmer, Patricia G 

A scanning electron microscope survey of the epidermis of East African grasses. 

(Smithsonian contributions to botany ; no. 49, 53) 

Includes bibliographies and indexes. 

Supt. of Doc. no.: QK 1.S2747 

1. Grasses-Africa, East-Anatomy-Atlases. 2. Grasses-Anatomy-Atlases. 3. LeavesAnatomy-Atlases. 4. Epidermis-Atlases. 5. Ultrastructure (Biology)-Atlases. I. 

Tucker, Alice E., joint author. 11. Title. 111. Series: Smithsonian contributions to botany ; no. 49. 

QKl.S2747 no. 49 [QK495.G74] 581s 80-19201 [584’.909676] AACR2 
Contents 

Page 

1 

Materials and Methods 1 

Acknowledgments ............................................... 2 

Format for Species Descriptions ..................................... 2 

Tribe POEAE ......................................................... 2 

Briza maxima Linnaeus .................. 2 

Colpodium chionogeiton (Pilger) Tzvelev ...... ................ 3 

Dactylis glomerata Linnaeus ........................................ 3 

Festuca abyssinica A. Richard ........................................ 4 

Lolium temulentum Linnaeus ....................................... 4 

Poa schimperana A. Richard ......................................... 5 

Pseudobromus sylvaticus Schumann .................................. 5 

Vulpia bromoides (Linnaeus) S.F. Gray .............................. 6 

Tribe BROMEAE .............................................. 6 

Bromus leptoclados Nees ............................................ 6 

Tribe BRACHYPODIEAE ............................................... 7 

Brachypodium flexum V ees ........................................ 7 

Tribe TRITICEAE .......................................... 8 

Hordeum marinum Hudson .......................................... 8 

Secale africanum Stapf ............................................... 8 

Triticum aestivum Linnaeus . , , . ................................ 9 

Tribe MELICEAE ............................................. 9 

Streblochaete longiarista (A. Richard) Pilger .......................... 9 

Tribe AVENEAE ...................................................... 10 

Aira caryophyllea Linnaeus ......................................... 10 

Anthoxanthum nivale K. Schumann .............................. 11 

Auena abyssinica Hochstetter ..................................... 11 

Deschampsia caespitosa (Linnaeus) Beauvois var. oliveri Hubbard . . , , 12 

Helictotrichon elongatum (Hochstetter) Hubbard ..................... 12 

Koeleria capensis (Steudel) Nees . . , . , . , , . , , . . , . , . , . , , . , . , . . , , , , . , , , 13 

Tribe PHALARIDEAE ................................................. 13 

Phalaris arundinacea Linnaeus ...................................... 13 

Tribe ACROSTIDEAE .................................................. 14 

14 

Introduction ...................................................... 

.............................. 

......... 

Agrostis schimperiana Hochstetter .................................. 14 

Pohpogon monspeliensis (Linnaeus) Desfontaines ..................... 16 

Stipa dregeana Steudel .............................................. 16 

Calamagrostis epigeios (Linnaeus) Roth var. capensis Stapf ............ 

Gastridium phleoides (Nees and Meyen) Hubbard ............ 

Tribe STIPEAE ....................................................... 16 

... 

111 
iv SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Appendix: Material Studied .................................... 18 

Literature Cited .................................................... 19 

Plates .............................................................. 20 

Index to Grass Species .............................................. 72 
A Scanning Electron Microscope 

Survey of the Epidermis 

of East African Grasses, I1 

Patricia G. Palmer and Alice E. Tucker 

Introduction 

By comparing the anatomical features of fossil 

grass leaves from East African lake sediments 

with those of modern-day grasses, it is often possible to identify fossil leaf fragments to the genus 

(Palmer, 1976). Because of the importance of 

fossil grasses as paleoecological indicators in tropical Africa, it has been necessary to develop a 

reference collection of modern grasses as a standard for comparison. For our purposes we have 

found that examination of the grass leaves with 

scanning electron microscopy (SEM) is preferable 

to light microscopy. In an earlier paper, we discussed our system of describing the diagnostic 

features of grass leaves as viewed with SEM and 

gave descriptions of 24 East African species representing ten tribes (Palmer and Tucker, 1981). 

The reader is referred to this earlier paper for 

discussion of terminology and illustrations of diagnostic features. 

This paper represents the second part of a 

survey of East African grasses and includes 26 

genera representing nine tribes of the Pooideae: 

Poeae, Bromeae, Brachypodieae, Triticeae, Meliceae, Aveneae, Phalarideae, Agrostideae, Stipeae. We delimit East Africa to include Uganda, 

Patricia G. Palmer, Department of Biolou, Louisburg College, 

Louisburg, North Carolina 27549. Alice E. Tucker, Departmen1 of 

Zoolog, Duke UniLersity, Durham, North Carolina 27706. 

Kenya and Tanzania, the region dealt with in 

the Flora of Tropical East Africa, Gramineae by Clayton (1970, 1974, in prep.). We also follow the 

nomenclature of the Flora of Tropical East Africa. 

Hartley (1973) has shown that the pooid grasses 

are found in temperate regions of the world and 

in the tropics are limited to high altitudes. A 

recent study by Livingstone and Clayton (1980) 

indicates that temperature is the most likely environmental factor controlling the importance of 

pooid genera in the highlands of tropical Africa. 

Because the occurrence of pooid genera in the 

fossil record of East Africa will permit paleotemperature estimates (Livingstone and Clayton, 

1980), the early inclusion of the tribes treated in 

this paper is extremely important in our overall 

survey . 

MATERIALS AND METHoDs.-Leaf material for 

scanning electron microscopy was taken from 

dried herbarium material. ,4 section from the 

middle portion of a blade from an upper, mature, 

undamaged leaf was chosen for study. In a few 

cases, where the epidermis proved to have a heavy 

coat of epicuticular wax, the wax was removed 

by soaking in xylene for 12 to 14 hours. Two 

pieces of the leaf (one exposing the abaxial or 

lower surface, the other exposing the adaxial or 

upper surface) were mounted on stubs with double-coated Scotch tape. The specimens were sputter-coated with gold-palladium, observed with a 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

JEOL T20 or a Philips 501 scanning electron 

microscope and photographed, using Polaroid 

P/N 665 or P/N 55 film. Occasionally, specimens 

were examined by energy dispersive X-ray analysis using the EDAX 9100/60 to clarify certain 

anatomical features. The photomicrographs are 

deposited in the laboratory of D.A. Livingstone, 

Department of Zoology, Duke University. 

Each specimen was analyzed at the microscope 

using a standard check sheet of diagnostic features. By comparing the photomicrographs and 

the check sheets, a complete description of each 

taxon was compiled. At least one species of each 

genus was examined, and for some large genera 

several species were examined to determine intrageneric variability. In some cases more than one 

specimen of a species was examined to determine 

in t ras peci fic vari ab i 1 it y . 

Livingstone of the Department of Zoology, Duke 

University, for research support and encouragement in this study. We wish to thank W.D. 

Clayton and S. Renvoize of the Kew Herbarium, 

T. Soderstrom of the National Herbarium, and 

G. Davidse of the Herbarium of the Missouri 

Botanical Garden for providing material for 

study. We thank T. Soderstrom and S. Renvoize 

for their useful comments on the manuscript. We 

are grateful for technical assistance by Susan G. 

Jones, Tony Perdue, Susan Hutchison, Anne 

Luck, and Terri Williams. This research was 

supported by NSF grants DEB 76-80426, ATM80- 

035 16, DEB78- 17669, and GB-22858 to D.A. Livingstone and NSF Science Faculty Fellowship 

Grant to Patricia G. Palmer. 

ACKNOWLEDGMENTS.-We are grateful to D.A. 

Format for Species Descriptions 

STOMATA.-FreqUenCy; subsidiary cell shape; 

distribution; special features. 

INTERSTOMATAL CELs.-shape; end walls; outline of walls. Papillae: Number/cell, distribution 

on each cell, shape, size. 

LONG CELLs.-shape; outline of walls. Papillae: 

Number/cell, distribution on each cell, shape, 

size. 

PRICKLES-Frequency; location and type. 

SHORT CELLs.-Location and distribution. Szlica Cells: Location and frequency. Cork Cells: Location and frequency. 

SILICA BODIES.-Location and shape. 

MICROHAIRS.-Frequency; number of cells; 

MACROHAIRS.-Frequencv; location: nature of 

SPECIAL FEATURES.-Description of unique feashape; shape of apex. 

base. 

tures not covered in other categories. 

Tribe POEAE 

Briza maxima Linnaeus 

PLATES 1-2 

ABAXIAL 

STOMATA.-cOmmOn; parallel; 2 stomatal 

bands/intercostal zone, 2 rows/band. Shape of 

subsidiary cell often unclear because of collapsed 

cells; veins not obvious (Plate 16). 

INTERSTOMATAL CELs.-Rectangular to long, 

narrow, width uniform to nonuniform: ends 

straight to convex; nonsinuous. Papillae: None 

seen (Plate 16). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen 

(Plate la,d,e). 

PRIcKLEs.-None seen. 

SHORT CELLs.--Costal solitary; intercostal not 

seen. Silica Cells: Costal infrequent; intercostal 

not seen. Cork Cells: Costal infrequent; intercostal 

not seen (Plate If). 

SILICA BoDIEs.-costal elongated-smooth, 

elongated-sinuous; intercostal not seen (Plate lf). 

MIcRoHA1Rs.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

SToMATA.-Common; parallel; 2 stomatal 

bands/intercostal zone, 2 rows/band (Plate 241). 

INTERSTOMATAL CELLs.-Rectangular to long, 
NUMBER 53 3 

narrow, width nonuniform; ends straight to convex; nonsinuous. Papillae: None seen (Plate 26). 

LONG CELLs.-Long, narrow, width nonuniform; nonsinuous. Papillae: None seen (Plate 2d,e). 

PR1cKLEs.-Infrequent; small costal prickles 

(Plate 2f). 

SHORT CELLs.-None seen. 

SILICA BoDIEs.-None seen. 

MICROHAIRS.-None seen. 

MACROHAIRS.-~One seen. 

Colpodiurn chionogeiton (Pilger) Tzvelev 

PLATES 3-4 

ABAXIAL 

SToMATA.-Abundant; parallel to sometimes 

low-dome; 4-6 stomatal bands/intercostal zone, 

1 or 2 rows/band (Plate 3a,b). 

IKTERSTOMATAL CELLs.-Rectangular to 

square; ends slightly concave; nonsinuous. Papillae: l/cell, globose, uniform. Appear to be thinwalled because usually deflated (Plate 3a,b,c). 

LONG CELLs.-Rectangular to long, narrow, 

width nonuniform; nonsinuous. Papillae: l/cell, 

globose, uniform (Plate 3a,c). 

PRIcKLEs.-Abundant; costal short, papillatetype (Plate 34. 

SHORT CELLs.-Costal solitary, paired; intercostal not seen. Silica Cells: Costal common; intercostal not seen. Cork Cells: Costal common; 

intercostal not seen (Plate 3ef). 

SILICA BoDIEs.-costal oblong, square; intercostal not seen (Plate 3e,f). 

MICROHAIRS.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

SToMATA.-Abundant; parallel; 4 stomatal 

bands/intercostal zone, 1 row/band. Bands difficult to detect because of collapsed long cells. 

Costal regions often indistinct (Plate 4a,b). 

INTERSTOMATAL CELLs.-Rectangular; ends 

slightly concave or sometimes straight to convex; 

nonsinuous. Papillae: l/cell, globose, uniform. Often deflated, probably thin-walled (Plate 4a,c). 

LONG CELLs.-Long, narrow, width nonuniform; nonsinuous. Papillae: l/cell, globose, uniform. Often deflated, probably thin-walled (Plate 

PRIcKLEs.-Abundant on margin, absent elsewhere; short unpointed with swollen bases. Several rows of cells near margin have large papillatetype prickles (Plate 4f). 

SHORT CELLS.-cOStal solitary; intercostal not 

seen. Silica Cells: Costal common; intercostal not 

seen. Cork Cells: Costal and intercostal not seen 

(Plate 44. 

SILICA BoDIEs.-costal oblong, elongatedsmooth, square; intercostal not seen (Plate 4d,e). 

MIcRoHArRs.-None seen. 

MAcRoHArRs.-None seen. 

44. 

Dactylis glornera ta Linnaeus 

PLATES 5-6 

ABAXIAL 

SToMATA.-Common: parallel; 2 or 3 stomatal 

bands/intercostal zone, 2 rows/band (Plate 5a,b). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform: ends straight to convex; nonsinuous. Papillae: None seen (Plate 56). 

LONG CELLs.-Long, narrow, width nonuniform; nonsinuous. Papillae: None seen (Plate 

5 a,d) . 

PRIcKLEs.-Common to abundant; costal prickles. Size variable (Plate 5d,e). 

SHORT CELLs.-Costal 3-5/row; intercostal not 

seen. Silica Cells: Costal common; intercostal not 

seen. Cork Cells: Costal common; intercostal not 

seen. Short cells indistinct because of wax (Plate 

SILICA BoDIEs.-costal oblong, elongatedsmooth, elongated-sinuous; intercostal not seen 

(Plate 5 e,f). 

5 e,f 1. 

MICROHAIRS.-None seen. 

MACROHAIRS.-None seen. 
4 

ADAXIAL 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ADAXIAL 

SToMATA.--Infrequent to common; parallel; 2 

stomatal bands/intercostal zone, 1-3 rows/band. 

Cells collapsed and shape difficult to ascertain 

(Plate 66). 

INTERSTOMATAL CELs.-Long, narrow, width 

nonuniform; ends straight to convex; nonsinuous. 

Papillae: None seen (Plate 6c). 

LONG CELLS.-LOng, narrow, width nonuniform; nonsinuous. Papillae: None seen (Plate 6a,c). 

PRrcKLEs.-Common: costal prickles. Size variable (Plate 664. 

SHORT CELLs.-Costal 3-5/row; intercostal not 

seen. Silica Cells: Costal infrequent to common; 

intercostal not seen. Cork Cells: Costal infrequent 

to common: intercostal not seen. Cells often obscured by wax (Plate 6e,f). 

SILICA BoDIEs.-costa~ elongated-sinuous, nodular; intercostal not seen (Plate 6e,f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

Festuca abyssinica A. Richard 

SToMATA.-Common; parallel; 1-2 stomatal 

banddintercostal zone, 2 or 3 rows/band. Stomata sunken among inflated long cells and obscured by long cells (Plate 86). 

IKTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform; ends straight to convex; nonsinuous. Papillae: None seen. Cells raised, 

appear inflated (Plate 86). 

LONG CELLs.-Long, narrow, width nonuniform; nonsinuous. Papillae: None seen. Cells 

raised, appear inflated. Difficult to distinguish 

between costal and intercostal region (Plate 

8a,c,e). 

PRIcKLEs.-Common; intercostal prickles. 

Prickles long; approaching short, macrohair type 

in length (Plate 8d,e). 

SHORT CELLs.-Costal not seen; intercostal solitary. Szlica Cells: Costal not seen; intercostal infrequent. Cork Cells: Costal and intercostal not 

seen (Plate 8f). 

SILICA BoDIm-Costal not seen; intercostal 

square to oblong (Plate 8f). 

h'fICROHAIRS.--NOne Seen. 

L~ACROHAIRS.--NOnC Seen. 

PLATES 7-8 

Loliurn ternulentum Linnaeus 

ABAXIAL 

STOMATA.-NOne seen. 

LOKG CELLs.-Long, narrow, width uniform: 

markedly sinuous, interlocking raised. Papillae: 

None seen (Plate 7a,c). 

PRIcKLEs.-Infrequent to common; costal and 

intercostal prickles (Plate 76,c). 

SHORT CELLS.-Costal and intercostal paired. 

Silica Cells: Costal and intercostal abundant. Cork 

Cells: Costal and intercostal abundant. Covered 

by reticulated wax (Plate 7c,e,f). 

SILICA BoDIEs.-Costal and intercostal elliptical or sometimes round, crescent-shaped (Plate 

7e,f 1. 

MicRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

PLATES 9- 10 

ABAXIAL 

STOMATA.-None seen. 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Occasionally, marginal 

cells have indistinctly sinuous outlines. Papillae: 

None seen (Plate 9a). 

PRIcKLEs.-Infrequent; costal prickles. Primarily in marginal areas (Plate 96,c). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal infrequent; intercostal 

not seen. Cork Cells: Costal and intercostal not 

seen. Silica cells found only in marginal areas and 

on mid-vein (Plate 9d-A. 

SILICA BoDIEs.-Costal elongated-sinuous, 

round; intercostal not seen (Plate 9e,f). 
NUMBER 53 5 

MIcRoHAIRs.-None seen. ADAXIAL 

MACROHAIRS.-None seen. 

ADAXIAL 

SToMATA.-Abundant: parallel: 2 stomatal 

bands/intercostal zone, 4 rows/band (Plate 

1 O6,c) . 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width uniform; ends straight to convex; 

nonsinuous. Papillae: None seen (Plate 1Oc). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform: nonsinuous. Papillae: None seen 

(Plate 10a,c). 

PRIcKLEs.-Abundant: costal and occasionally 

intercostal prickles (Plate 10d,e). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal common; intercostal not 

seen. Cork Cells: Costal and intercostal not seen 

(Plate lOf). 

SILICA BoDIEs.-Costal elongated-smooth, 

elongated-sinuous: intercostal not seen (Plate 

1Of). 

MICROHAIRS.-None seen. 

MACROHAIRS.-None seen. 

Poa schimperana A. Richard 

PLATES 11-12 

ABAXIAL 

STOMATA.-NOne Seen. 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen 

(Plate 1 la-c). 

PR1cKLEs.-Infrequent; costal prickles (Plate 

1 Id). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal common; intercostal not 

seen. Cork Cells: Costal and intercostal not seen 

(Plate 1 lef). 

SILICA BoDIEs.-costal elongated-sinuous; intercostal not seen. Occasionally very long (Plate 

11ef). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

STOMATA.-Abundant; parallel; 2 stomatal 

bands/intercostal zone, 2 rows/band (Plate 

12a-c) . 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform; ends straight to convex: nonsinuous. Papillae: None seen (Plate 12c). 

LONG CELLs.-Long, narrow. width nonuniform: nonsinuous. Papillae: None seen (Plate 

12a,c). 

PRIcKLEs.-Common: cost a1 and occasionally 

intercostal prickles (Plate 12d,e). 

SHORT CELLs.-Costal solitary or sometimes 

paired: intercostal not seen. Silzca Cells: Costal 

infrequent; intercostal not seen. Cork Cells: Costal 

and intercostal not seen (Plate 12f). 

SILICA BoDIEs.-costal elongated-sinuous; intercostal not seen (Plate 12f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

Pseudobromus sylvaticus Schumann 

PLATES 13-14 

ABAXIAL 

SToMATA.--Infrequent; parallel to occasionally 

low-dome: 2 stomatal bands/intercostal zone, 1 

row/band. Distribution difficult to determine because stomata not very numerous (Plate 136). 

INTERSTOMATAL CELLs.-Long, narrow, width 

uniform to nonuniform; ends straight to convex: 

nonsinuous. Papillae: None seen (Plate 136). 

LONG CELLs.-Long, narrow, width nonuniform or occasionally uniform: nonsinuous. Papillae: None seen (Plate 13a,c). 

PRICKLES ,-Abundant to common ; cost a1 

prickles. Abundant on margin and on two veins 

nearest margin but common on other veins (Plate 

13a,d). 

SHORT CELLS.-cOStal solitary, paired, 3-51 

row; intercostal not seen. Silica Cells: Costal common; intercostal not seen. Cork Cells: Costal infrequent; intercostal not seen (Plate 13eJ). 

SILICA BoDIEs.-costal oblong, elongated- 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

smooth or occasionally elongated-sinuous; intercostal not seen (Plate 13ef). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

MICROHAIRS.-None seen. SPECIAL FEATURES.-\’einS very prominent 

MACROHAIRS.-None seen. (Plate 15a). 

ADAXIAL ADAXIAL 

STOMATA.-common to abundant: parallel; 2 

stornatal bands/intercostal zone, 3 rows/band. 

Bands adjacent to veins (Plate 14a-c). 

INTERSTOMATAL cELLS.-LOng, narrow, width 

uniform; ends straight to convex; nonsinuous. 

Papillae: None seen (Plate 14c). 

LONG CELLs.-Rectangular to long, narrow, 

width uniform; nonsinuous. Papillae: None seen 

(Plate 14d). 

PRIcKLEs.-Common: costal prickles. Basal 

portion of prickles very long (Plate 14c,e,f). 

SHORT CELLs.-Costal solitary: intercostal not 

seen. Silica Cells: Costal infrequent; intercostal 

not seen. Cork Cells: Costal and intercostal not 

seen (Plate 14f). 

SILICA BoDIEs.-costal elongated-smooth; intercostal not seen (Plate l4f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

Vulpia bromoides (Linnaeus) S.F. Gray 

PLATES 15- 16 

STOMATA.-cOmmOn; parallel; 2 stornatal 

banddintercostal zone, 4 rows/band. Shape of 

subsidiary cell difficult to ascertain because of 

collapsed cells (Plate 166). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform: ends straight to convex; nonsinuous. Papillae: None seen. Cells collapsed (Plate 166). 

LONG CELLS.-LOng, narrow, width nonuniform; nonsinuous. Papillae: None seen. Cells collapsed (Plate 1 16a,c), 

PRIcKLEs.-Abundant; costal and intercostal 

prickles, occasionally short macrohair type. Often 

partially collapsed as if not heavily silicified. Prickles primarily costal (Plate 16d,e). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal infrequent; intercostal 

not seen. Cork Cells: Costal and intercostal not 

seen (Plate 16d,f). 

SILICA BoDrEs.-costal oblong to elongatedsmooth; intercostal not seen. Silica bodies have 

central horizontal ridge (Plate 16 d,f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

Tribe BROMEAE 

STOMATA.-NOnC seen. 

LONG CELLs.-Long, narrow, width uniform; 

markedly sinuous, interlocking, raised. Papillae: 

None seen (Plate 156). 

PRIcKLEs.-Infrequent; costal prickles (Plate 

156). 

Bromus leptoclados Nees 

PLATES 17-18 

ABAXIAL 

SHORT CELLs.-Costal paired; intercostal STOMATA.-NOne seen. 

paired or rarely 3-5/row. Silica Cells: Costal com- LONG CELLs.-Long, narrow, width nonunimon; intercostal abundant. Cork Cells: Costal form; nonsinuous. Papillae: None seen (Plate 

common; intercostal abundant (Plate 15d-f). 

SILICA BoDIEs.-Costal and intercostal cres- PRIcKLEs.-Abundant; costal prickles (Plate 

cent-shaped or sometimes tall and narrow, round 

(Plate 15d-f). SHORT CELLs.-Costal solitary, paired, 3-5,’ 

17a,b). 

1 76). 
NUMBER 53 7 

row; intercostal solitary. Silica Cells: Costal abundant; intercostal infrequent (only one seen). Cork 

Cells: Costal abundant; intercostal not seen (Plate 

17a,d,e). 

SILICA BoDIEs.-Costal and intercostal oblong, 

midhorizontal ridge sometimes present (Plate 

17a,d,e). 

MICROHAIRS.-None seen. 

MAcRoHAIRs.-Abundant; intercostal; basal 

cells absent (Plate 17f). 

ADAXIAL 

STOMATA.-Abundant; parallel to low-dome; 4 

stornatal bands/intercostal zone, 1 row/band. Intercostal zones wide and veins indistinct (Plate 

1 8a, b) . 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform; ends straight to convex; nonsinuous. Papillae: None seen (Plate 186). 

LONG CELLs.-Long, narrow, width nonuniform: nonsinuous. Papillae: None seen (Plate 

18a,c,e). 

PRIcKLEs.-Infrequent; costal prickles primarily in marginal area (Plate 184. 

SHORT CELLs.--Costal solitary; intercostal not 

seen. Silica Cells: Costal infrequent; intercostal 

not seen. Cork Cells: Costal and intercostal not 

seen (Plate 18e). 

SILICA BoDIEs.-costa~ elongated-smooth, often with a midhorizontal ridge: intercostal not 

seen (Plate 18e). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-cOmmOn; costal; basal cells absent (Plate 18a,f). 

Tribe BRACHYPODIEAE 

Brachypodium flexurn Nees 

PLATES 19-20 

ABAXIAL 

STOMATA.-NOne seen. 

LONG CELLs-Long, narrow, width uniform; 

markedly sinuous, interlocking, raised. Papillae: 

None to l/cell. Many long cells appear to have a 

raised area (papilla?) near the middle of cell 

opposite a silica body in an adjacent row of cells 

(Plate 19b,c). 

PRIcKLEs.-Infrequent to common: costal and 

occasionally intercostal on the sides of the veins. 

Veins indistinct (Plate 19a,d). 

SHORT CELLs.-Costal and intercostal solitary 

and paired. Silica Cells: Costal and intercostal 

common. Cork Cells: Costal and intercostal common (Plate 19e,f). 

SILICA BoDIEs.-costal and intercostal round 

or sometimes elliptical (Plate 19e,f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

Stomata.-Common; parallel; 2 stornatal 

bands/intercostal zone, 3 rows/band. Very collapsed and difficult to ascertain (Plate 20b). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform; ends straight to convex: nonsinuous. Papillae: None seen. Cells collapsed and shape not always obvious (Plate 206). 

LONG CELLs.-Pentagonal, hexagonal or occasionally rectangular or long, narrow, width nonuniform; nonsinuous. Papillae: None seen. Cells 

collapsed and shape not always obvious (Plate 

20a,c). 

PRIcKLEs.-Common; costal prickles, intercostal short macrohair type or occasionally hooks 

(Plate 206). 

SHORT CErLs.-Costal and intercostal solitary. 

Silica Cells: Costal and intercostal infrequent to 

common. Cork Cells: None seen (Plate 20d,e). 

SILICA BoDIEs.-costal tall and narrow or 

sometimes oblong, acutely angled, square; intercostal tall and narrow or sometimes acutely angled (Plate 20d,e). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-cOmmOn; costal and sometimes intercostal; raised basal cells absent (Plate 

20f 1 ’ 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Tribe TRITICEAE 

Hordeum marinum Hudson 

PLATES 2 1-22 

ABAXIAL 

SToMATA.-Infrequent to common; parallel; 3 

stomatal bands/intercostal zone, 1 row/band 

(Plate 2 la,b). 

INTERSTOMATAL CELs.-Rectangular; straight 

to convex; nonsinuous. Papillae: None seen. Shape 

and outline of walls difficult to determine because 

of collapsed cells and numerous prickles (Plate 

LONG CErLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen. 

Cells usually collapsed and shape difficult to 

determine (Plate 21c). 

PRIcKLEs.-Abundant; costal and intercostal 

short macrohair-type prickles. Prickles quite variable in length from regular macrohair length to 

mainly short macrohair type (Plate 2 ld,e). 

SHORT CELLs.-Costal and intercostal solitary. 

Silica Cells: Costal infrequent to common; intercostal infrequent. Cork Cells: Costal and intercostal absent (Plate 2 le,f). 

SILICA BoDIEs.-costal elongated-smooth; intercostal round (Plate 21ef). 

214. 

MICROHAIRS.-None seen. 

MACROHAIRS.-cOmmOn; intercostal and costal: raised basal cells absent. Most macrohairs are 

the short macrohair type (Plate 21d,e). 

ADAXIAL 

SToMATA.-Common; parallel to low-dome; 2 

stornatal bands/intercostal zone; 1-2 rows/band. 

Pattern difficult to ascertain; interstomatal areas 

narrow; subsidiary cells collapsed (Plate 22a-c) . 

INTERSTOMATAL CELLS.-Rectangular; ends 

straight to convex; nonsinuous. Papillae: None 

seen. Shape difficult to determine because of 

folding (Plate 22c). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen. 

Shape and outline difficult to determine because 

of wax and folding (Plate 22a,c). 

hcKLEs.-Abundant; costal and intercostal 

straight, short macrohair type. Variable in length 

and many long enough to be classified as macrohairs (See Macrohairs) (Plate 22a,d,e). 

SHORT CELLs.-Costal solitary, paired (?); intercostal not seen. Silica Cells: Costal common; 

intercostal not seen. Cork Cells: Costal infrequent; 

intercostal not seen (Plate 22d-f). 

SILICA BoDIEs.-costal oblong, elongatedsmooth, square; intercostal not seen. Some are 

very elongated (Plate 22d-f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-COmmOn; costal and intercostal; basal cells absent (See Prickles) (Plate 22a). 

Secale africanum Stapf 

PLATES 23-24 

ABAXIAL 

SToMATA.-Infrequent to common; parallel; 2 

stornatal bands/intercostal zone, 1 row/band. 

Stomata slightly sunken adjacent to veins and 

somewhat obscured by veins and macrohairs 

(Plate 23a-c). 

INTERSTOMATAL CELs.-Long, narrow, width 

uniform: end straight to convex; walls nonsinuous. Papillae: None seen (Plate 23c). 

LONG CELLS.-LOIIg, narrow, width uniform: 

nonsinuous. Papillae: None seen (Plate 23c). 

PRICKLES.-None seen. 

SHORT CELLs.-Costal solitary, paired, 3-5/ 

row: intercostal not seen. Silica Cells: Costal infrequent to common; intercostal not seen. Cork Cells: 

Costal infrequent to common; intercostal not seen 

(Plate 23d-f). 

SILICA BoDIEs.-costal round, oblong, elongated-sinuous; intercostal not seen (Plate 23d-f). 

MICRO HAIR^. -None seen, 

MAcRoHAms.-Abundant; intercostal and costal; raised basal cells absent. Major veins have 

macrohairs only on sides; minor veins have macrohairs on top (Plate 23a-c). 
NUMBER 53 9 

ADAXIAL 

SToMATA.-Infrequent to common; parallel to 

low-dome; 2 stomatal bands/intercostal zone, 1 

or 2 rows/band. Stomata slightly sunken adjacent 

to veins and somewhat obscured by veins and 

macrohairs (Plate 246,~). 

INTERSTOMATAL CELLS.-Square, rectangular, 

long, narrow, width uniform; ends straight to 

convex; nonsinuous. Papillae: None seen (Plate 

24c). 

LONG CELLS.-Pentagonal to long, narrow, 

width uniform; nonsinuous. Papillae: None seen 

(Plate 24a,c,e). 

PRIcKLEs.-None seen. 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal infrequent; intercostal 

not seen. Cork Cells: Costal and intercostal not 

seen (Plate 24d). 

SILICA BoDIEs.-costal oblong; intercostal not 

seen (Plate 24d). 

MIcRoHAIRs.-None seen. 

MAcRoHAIRs.-Abundant; intercostal and costal; raised epidermal cells absent. Macrohairs 

vary in length (Plate 24a,e,f). 

Triticum aestivum Linnaeus 

PLATES 25-26 

ABAXIAL 

STOMATA.- Common; low-dome; 2(?) stomatal 

bands/intercostal zone, 1 row/band. Veins broad 

and indistinct, distribution not clear (Plate 25a,b). 

INTERSTOMATAL CELLS.-Square, rectangular, 

long, narrow, width uniform; ends straight to 

convex; nonsinuous. Papillae: None seen (Plate 

25c). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen 

(Plate 25a,d). 

PRICKLES. -None Seen. 

SHORT CELLs.-Costal solitary, paired(?). Silica 

Cells: Costal infrequent to common; intercostal 

not seen. Cork Cells: Costal infrequent to common(?); intercostal not seen. Cork cells indistinct; 

some suggest a silica cell paired with a cork cell 

(Plate 23e,f). 

SILICA BoDIEs.-oblong, or sometimes elongated-sinuous (Plate 25e,f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAI 

SToMATA.-Abundant: parallel to low-dome; 2 

stomatal bands/intercostal zone, 1 or sometimes 

2 rows/band (Plate 26a,b). 

INTERSTOMATAL CELLS .- Rec t angu lar ; ends 

straight to convex; nonsinuous. Papillae: None 

seen (Plate 26c). 

LONG CELLs.-Long, narrow, width uniform; 

nonsinuous. Papzllae: None seen. Mid-intercostal 

band of long cells with indistinct walls (Plate 

26a,d). 

PRICKLES.-None seen. 

SHORT CELLs.-Solitary, sometimes paired, 3- 

5/row(?). Silica Cells: Costal common; intercostal 

not seen. Cork Cells: Costal infrequent (?): intercostal not seen. Occasional paired silica cells 

(Plate 26ef). 

SILICA BoDIEs.-costal elongated-sinuous; intercostal not seen (Plate 26e,f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

Tribe MELICEAE 

Streblochaete longiarista (A. Richard) Pilger 

PLATES 27-28 

ABAXIAL 

SToMATA.-Infrequent; parallel to low-dome: 

distribution indeterminable because stomata too 

infrequent (Plate 276). 

LONG CELLs.-Long, narrow, width uniform; 

markedly sinuous, interlocking, raised. Papillae: 

None seen (Plate 27a,c). 

PRIcKLEs.-Common; costal and occasionally 

intercostal prickles (Plate 27a,d). 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

SHORT CELLs.-Costal and intercostal solitary. 

Silica Cells: Costal infrequent to common; intercostal not seen. Cork Cells: Costal and intercostal 

not seen (Plate 27e, f). 

SILICA BoDIEs.-costal oblong, elongated-sinuous; intercostal square (Plate 27e,f). 

MICROHAIRS.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

SToMATA.-Common; parallel to low-dome; 2 

stomatal bands/intercostal zone, 2 rows/band 

(Plate 28a,b). 

INTERSTOMATAL CELLS.-Rectangular or sometimes square or long, narrow, width uniform; 

ends straight to convex; nonsinuous. Papillae: 

None seen (Plate 286,~). 

LONG CELLs.-Pentagonal to long, narrow, 

width nonuniform; nonsinuous. Papillae: None 

seen (Plate 28a,c,e). 

PRrcKLEs.-Abundant; costal prickles and 

short, straight, macrohair type; intercostal hooks 

and short, straight, macrohair type. Some of the 

short macrohair type have sunken bases (Plate 

28d-f). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal infrequent to common; 

intercostal not seen. Cork Cells: Costal and intercostal not seen (Plate 28d). 

SILICA BoDIEs.-~osta~ oblong, elongatedsmooth, square; intercostal not seen (Plate 28d). 

MICROHAIRS.-None seen. 

MACROHAIRS.-cOmmOn; costal; basal cells absent (Plate 28e). 

Tribe AVENEAE 

Aira caryophyllea Linnaeus 

PLATES 29-30 

ABAXIAL 

SToMATA.-Abundant; parallel; 1 stomatal 

band/intercostal zone, 3 or 4 rows/band. Intercostal areas very narrow. Veins prominent and 

numerous. Shape of subsidiary cell difficult to 

ascertain because of collapsed cells (Plate 29a,b). 

INTERSTOMATAL CELLs.-Rectangular to occasionally long, narrow, width uniform; ends 

straight to convex; nonsinuous. Papillae: None 

seen (Plate 29c). 

LONG CELLS.-LOng, narrow, width uniform; 

nonsinuous. Papillae: None seen (Plate 29c,d). 

PRIcKLEs.-Abundant; costal prickles and 

short macrohair-type prickles (Plate 29d). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal common; intercostal not 

seen. Cork Cells: Costal and intercostal not seen 

(Plate 29d-f ). 

SILICA BoDIEs.-costal elongated-sinuous to 

occasionally elongated-smooth, square; intercostal not seen (Plate 29e, f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-seC section on Prickles. 

ADAXIAL 

STOMATA.-cOmmOn; parallel: 1 stomatal 

band/intercostal zone, 3 or sometimes 1 row/ 

band. Costal areas very prominent, intercostal 

very narrow (Plate 30a,b). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width uniform; ends straight to convex; 

nonsinuous. Papillae: None seen (Plate 30c). 

LONG CELLs.-Long, narrow, width uniform to 

occasionally nonuniform; nonsinuous. Papillae: 

None seen (Plate 30a,c). 

PRIcKLEs.-Abundant; costal prickles and occasional hooks (Plate 30d). 

SHORT CELLs.-Costal solitary, paired, 3-5/ 

row; intercostal not seen. Silica Cells: Costal abundant; intercostal not seen. Cork Cells: Costal common; intercostal not seen (Plate 30e, f). 

SILICA BoDIEs.-costal nodular or sometimes 

elongated-sinuous, square; intercostal not seen 

(Plate 30e, f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 
NUMBER 33 11 

Anthoxanthum nivale K. Schumann 

PLATES 3 1-32 

ABAXIAL 

SToMATA.--Infrequent to common; parallel; 2 

stomatal bands/intercostal zone, 2 rows/band 

(Plate 31a). 

INTERSTOMATAL CELLs.-Long, narrow, width 

uniform (often appearing nonuniform because of 

collapsed cells) : ends straight to convex; nonsinuous. Papillae: None seen (Plate 3 lb). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen 

(Plate 31c,d). 

PR1cKLEs.-Infrequent: costal prickles (Plate 

31e). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Szlica Cells: Costal infrequent; intercostal not 

seen. Cork Cells: Costal and intercostal not seen 

(Plate 3 lf). 

SILICA BoDIEs.-Costal elongated-smooth; intercostal not seen (Plate 3lf). 

MrcRoriAIRs.-None seen. 

MACROHAIRS.-NO~~ Seen. 

ADAXIAL 

STOMATA.-NOne seen. 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous (sometimes collapsed). 

Zones (approximately five cells wide) of uniform 

width cells alternate with nonuniform width cells. 

Papillae: None seen (Plate 32a-c). 

PRIcKLEs.-Infrequent; costal prickles (Plate 

32d-f). 

SHORT CELLs.-None seen. 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-NOne Seen. 

Avena abyssinica Hochstetter 

PLATES 33-34 

ABAXIAL 

SToMATA.-Common; parallel to low-dome; 2 

stomatal bands/intercostal zone, 1-3 rows/band. 

Stomata1 bands adjacent to veins (Plate 33a-c). 

INTERSTOMATAL CELs.-Long, narrow, width 

uniform to nonuniform (perhaps an artifact of 

folding) ; ends straight to convex; nonsinuous. 

Papillae: None seen (Plate 33a). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform (perhaps an artifact of folding); nonsinuous to slightly sinuous (marginal area). 1’0- 

pillae: None seen (Plate 33a). 

PRIcKLEs.-Common; costal prickles (Plate 

SHORT CELLs.-Costal paired. 3-5/row; intercostal not seen. Silica CellJ: Costal abundant; 

intercostal not seen. Cork Cell,: Costal abundant; 

intercostal not seen (Plate 334. 

SILICA BoDIEs.-costal elongated-sinuous; intercostal not seen (Plate 334. 

MICROHAIRS.-NOne seen. 

MACROHAIRS.-Infrequent to common: intercostal: basal cells absent. Primarily ad-jacent to 

midvein (Plate 33e,f), 

33c). 

ADAXIAL 

SToMATA.-Common; parallel: 2 stomatal 

bands/intercostal zone, 1-3 rows/band (Plate 

34b). 

INTERSTOMATAL CELs.-Long, narroic, width 

uniform to nonuniform; ends straight to convex; 

nonsinuous. Papillae: None seen (Plate 34a,d). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. I’upillae: None seen 

(Plate 34~4). 

PRIcKLEs.-Common; costal and intercostal 

prickles. Intercostal infrequent (Plate 34c). 

SHORT CELLs.-Costal solitary, paired, 3-5/ 

row; intercostal not seen. Silica Cells: Costal abundant; intercostal not seen. Coik CellJ: Costal abundant: intercostal not seen (Plate 34a,d). 

SILICA BoDIEs.-costa~ elongated-sinuous; intercostal not seen (Plate 34a,d). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-COmmOn; costal; basal cells absent (Plate 34eJ). 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Deschampsia caespitosa (Linnaeus) Beauvois 

var. oliveri Hubbard 

PLATES 35-36 

ABAXIAL 

STOMATA.-NOne seen. 

LONG CELLs.-Long, narrow, width uniform; 

markedly sinuous, interlocking, raised. Papillae: 

None seen (Plate 356). 

PRIcKLEs.-Abundant; costal and intercostal 

prickles. Extremely abundant and uniformly distributed (Plate 35a,c,d). 

SHORT CELLs.-Costal and intercostal solitary 

and paired. Silica Cells: Costal and intercostal 

abundant. Cork Cells: Costal and intercostal abundant (Plate 35c,ed ). 

SILICA BoDIEs.-Costal and intercostal elliptical to round (Plate 35c,ef). 

MIcROHAIRs.-None seen. 

SPECIAL FEATURES.-Regular raised areas that 

appear to be veins, but epidermis in these costal 

zones and in the intercostal regions uniform 

(Plate 35a). 

MACROHAIRS.-None seen. 

ADAXIAL 

STOMATA.-COmmOn; low-dome; 2 Stomata1 

bands/intercostal zone, ca. 15 row/band. Intercostal zones very wide (Plate 36a,b). 

INTERSTOMATAL CELLs.-Long, narrow, width 

uniform; ends straight to convex: markedly sinuous, interlocking, raised. Papillae: None seen 

(Plate 36a). 

LONG CELLs.-Long, narrow, width uniform; 

markedly sinuous, interlocking raised. Papillae: 

None seen. In the center of each intercostal zone 

a narrow belt (2 or 3 cells wide) of waxy, nonsinuous long cells. On each side of the narrow belt 

a region containing markedly sinuous long cells 

(Plate 366). 

PRIcKLEs.-Common; costal prickles. Only 

along top of veins; usually a double row of mainly 

large but a few small ones on the side (Plate 

36d,e). 

SHORT CELLS.-cOStal solitary or sometimes 

paired; intercostal solitary. Silica Cells: Costal and 

intercostal common. Cork Cells: Costal infrequent: 

intercostal not seen (Plate 36d, f). 

SILICA BoDIEs.-costal and 

MICROHAIRS.-None seen. 

MACROHAIRs.-None seen. 

cal, round (Plate 36f). 

Helictotrichon elongatum 

Hubbard 

intercostal ellipti- 

(Hochstetter) 

PLATES 37-38 

ABAXIAL 

SToMATA.-Common: parallel to low-dome; 2 

stomatal bands/intercostal zone, 1 or 2 rows/ 

band (Plate 37a,b). 

INTERSTOMATAL CELLS.-LOng, narrow, width 

uniform to nonuniform; ends straight to convex; 

nonsinuous. Papillae: None seen (Plate 37a). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen 

(Plate 37a,c). 

PRrcKLEs.-Common; costal prickles (on sides 

of veins) (Plate 374. 

SHORT CELLs.-Costal solitary, paired, 3-5/ 

row; intercostal not seen. Silica Cells: Costal common; intercostal not seen. Cork Cells: Costal infrequent to common; intercostal not seen (Plate 

SILICA BoDIEs.-Costa~ elongated-sinuous; intercostal not seen (Plate 37e,f). 

MIcRoHAIRs.-None seen. 

37e,ff). 

MACROHAIRS.-None seen. 

ADAXIAL 

SroMATA.-Common; parallel to low-dome; 2 

stomatal bands/intercostal zone, 1-3 rows/band. 

Bands adjacent to veins (Plate 386,~). 

INTERSTOMATAL CELs.-Long, narrow, width 

uniform to nonuniform; ends straight to convex; 

nonsinuous. Papillae: None seen (Plate 38c). 

LONG CELLs.-Long, narrow, width uniform to 
NUMBER 53 13 

nonuniform; nonsinuous. Papillae: None seen 

(Plate 38a,d). 

PRIcKLEs.-Common; costal prickles (Plate 

38e). 

SHORT CELLs.-Costal solitary, paired, 3-5/ 

row; intercostal not seen. Silica Cells: Costal common; intercostal not seen. Silica cells frequently 

solitary, alternating with prickles. Cork Cells: Costal infrequent to common; intercostal not seen 

(Plate 38c, f) . 

SILICA BoDIEs.-costal elongated-sinuous 

(many very long); intercostal not seen (Plate 

38c,f?. 

MrcRoHAIRs.-~one seen. 

MACROHAIRS .-None Seen. 

ADAXIAL 

STOMATA.-NOne Seen. 

LONG CELLs.-Long, narrow, width uniform: 

nonsinuous. Papillae: None seen (Plate 406,~). 

PRIcKLEs.-Abundant; costal prickles and 

short macrohairs (Plate 40a-c). 

SHORT CELLs.-Costal paired, rarely solitary or 

3-5/row; intercostal not seen. Silica Cells: Costal 

abundant; intercostal not seen. Cork Cells: Costal 

abundant; intercostal not seen (Plate 406-e). 

SILICA BoDIEs.-costal oblong, elongatedsmooth, and square; intercostal not seen (Plate 

40c-e) . 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-cOmmOn; intercostal; basal 

cells absent. Collapsed macrohairs are twisted 

(Plate 40a,cf). 

SPECIAL FEATuREs.-surface ridged because of 

wide, raised costal zones and narrow, sunken 

intercostal zones (Plate 40a,c!. 

Koeleria capensis (Steudel) Nees 

PLATES 39-40 

Tribe PHALARIDEAE ABAXIAL 

STOMATA.-cOmmOn; low-dome; 2 Stomata1 

bands/intercostal zone, 3 rows/band. Stomata 

are small, often collapsed, and difficult to observe 

in narrow intercostal zones sunken between large 

veins (Plate 39a,6). 

INTERSTOMATAL CELs.-Probably long, narrow. Shape and nature of walls not discernible 

because of collapsed cells. Papillae: None seen. 

LONG CELLs.-Long, narrow, width uniform; 

nonsinuous. Papillae: None seen (Plate 396). 

PRIcKLEs.-Abundant; costal prickles and 

short macrohairs (Plate 39c,d). 

SHORT CELLs.-Costal solitary, rarely paired; 

intercostal not seen. Silica Cells: Costal abundant: 

intercostal not seen. Cork Cells: Costal and intercostal not seen (Plate 39c,e). 

SILICA BoDIEs.-costal oblong, elongatedsmooth, and square; intercostal cot seen (Plate 

39e). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-common; costal; basal cells absent (Plate 396,dJ). 

Phalaris arundinacea Linnaeus 

PLATES 4 1 ~ 42 

ABAXIAL 

SToMATA.-Abundant; parallel or sometimes 

low-dome; 2 stomata1 bands/intercostal zone, 4 

rows/band (Plate 41a-c). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width uniform; ends straight to convex; 

nonsinuous. Papillae: None seen. Cell slightly 

raised (Plate 416). 

LONG CELLs.-Long, narrow, width uniform; 

nonsinuous. Papillae: None seen (Plate 4 la,b,e). 

PRIcKLEs.-Common; costal prickles and intercostal hooks (Plate 4la,d). 

SHORT CELLS.-cOStal 3--5/row, !)/row, or 

sometimes paired; intercostal solitary, paired. Silica Cells: Costal common to abundant; intercostal 

common. Cork Cells: Costal common to abundant; 

intercostal infrequent (Plate 4 Id-f). 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

SILICA BoDIEs.-Costal square; intercostal 

square, round, crescent-shaped (Plate 4 Id-f). 

MIcRoHAIRs.-~one seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

STOMATA.-Abundant; parallel to low-dome; 1 

stornatal band/intercostal zone, 1 or sometimes 2 

rows/band. Wax somewhat obscures the shape 

(Plate 42a,b). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width uniform: ends straight to convex; 

nonsinuous. Papzllae: None seen. Cell slightly 

raised (Plate 42c). 

LONG CELrs.-Rectangular to long. narrow, 

width uniform; nonsinuous. Papzllae: None seen. 

Long cells adjacent to stomata slightly raised. 

Wax obscures cell outline (Plate 42a,cf). 

PRIcKLEs.-Infrequent; costal prickles and occasionally intercostal hooks (Plate 42d). 

SHORT CELLs.-Costal solitary, paired (often 

two silica cells), 3-3/row; intercostal solitary. Szlzca Cells: Costal abundant; intercostal infrequent. 

Cork Cells: Costal abundant; intercostal not seen 

(Plate 42 d-f). 

SILICA BoDIEs.--Costal oblong, square; intercostal square. crescent-shaped (Plate 42d-f). 

MICROHAIRS.-None seen. 

MACROHAIRS.-~One seen. 

SPECIAL FEATURES.-SpeCimen Very waxy. 

Tribe AGROSTIDEAE 

Agrostis schimperiana Hochstetter 

PLATES 43-44 

A B AXI A L 

SToMATA.-Infrequent; parallel: 2 stornatal 

bands/intercostal zone, 1 row/band. Cells collapsed; intercostal zones very narrow; distribution 

not obvious (Plate 436). 

INTERSTOMATAL CErs.-Long, narrow, width 

nonuniform; ends straight to convex: nonsinuous. 

Papillae: None seen (Plate 43b). 

LONG CELLS.-LOng, narrow, width nonuniform; ends straight to convex; nonsinuous. Papillae: None seen. Cells often collapsed; width 

variable, with wider cells often over veins (Plate 

43a,c). 

PRIcKLEs.-Abundant; costal prickles (Plate 

43a,bf). 

SHORT CELLS.-Costal solitary, paired, 3-5/ 

row; intercostal not seen. Silica Cells: Costal common: intercostal not seen. Cork Cells: Costal and 

intercostal not seen (Plate 43d-f). 

SILICA BoDIEs.-Costai nodular; intercostal not 

seen. Some very long (Plate 43d-f). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

SToMATA.-Common; parallel; 2 stornatal 

bands/intercostal zone, 2 rows/band (Plate 

446,~). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform; ends straight to convex; nonsinuous. Papillae: None seen (Plate 44c). 

LONG CELLs.-Long, narrow, width nonuniform; nonsinuous. Papillae: None seen. Veins numerous; long cells more collapsed in narrow intercostal regions (Plate 4464. 

PRIcKLEs.-Abundant; costal prickles. Many 

prickles rather long and thin (Plate 44e). 

SHORT CELLs.-Costal solitary; intercostal not 

seen. Silica Cells: Costal common; intercostal not 

seen. Cork Cells: Costal and intercostal not seen 

(Plate 44df). 

SILICA BoDIEs.-Costal nodular; intercostal not 

seen. Silica bodies very long, often indistinct 

(Plate 44df). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-NO~~ seen.

Calamagrostis epigeios (Linnaeus) Roth 

var. capensis Stapf 

PLAT~S 45-46 

ABAXIAL 

STOMATA.-COmmOn; parallel 

stomatal band/intercostal zone, 

(Plate 45a,b,c). 

to low-dome; 1 

1-3 rows/band 
NUMBER 53 15 

INTERSTOMATAL CELs.-Rectangular to long, 

narrow, width uniform; ends straight to convex 

or slightly concave: nonsinuous. Papillae: None 

seen (Plate 45c). 

LONG CELLs.-Long, narrow, width uniform; 

nonsinuous. Papillae: Cells adjacent to stoma have 

1 papilla/cell; all other cells have none (Plate 

456). 

PRIcKLEs.-Common: costal prickles (Plate 

43d). 

SHORT CELLs.-costal solitary, 3-3/row, >j/ 

row: intercostal solitary. Silica Cells: Costal infrequent to common: intercostal not seen. Cork Cells: 

Costal infrequent to common; intercostal common (Plate 45ef). 

SILICA BoDIEs.-Costal dumbbell with ends 

straight, middle wide and short, occasionally oblong or nodular: intercostal not seen (Plate 45,). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

SToMATA.-Abundant; parallel to low-dome; 2 

stomatal bands/intercostal zone, 3-4 rows/band 

(Plate 466). 

INTERSTOMATAL CELLs.-Rectangular to 

square; ends straight to convex; nonsinuous. Papillae: None seen (Plate 46c). 

LOKG CELLs.-Long, narrow, width uniform; 

nonsinuous. Papillae: Occasionally cells adjacent 

to stoma have raised area: all other cells have 

none. Midintercostal area of raised long cells with 

no distinct boundaries, i.e., appears to be one 

enormous cell (Plate 46a,d). 

PR1cKLEs.-Infrequent to common; costal and 

intercostal prickles, intercostal hooks. Intercostal 

larger and more frequent than costal prickles 

(Plate 46e). 

SHORT CELLS.-costal >5/row; intercostal solitary. Silica Cells: Costal abundant; intercostal not 

seen. Cork Cells: Costal abundant; intercostal common. Short cells are waxy (Plate 46f). 

SILICA BoDIEs.-costal dumbbell with ends 

straight, middle wide and short; intercostal not 

seen (Plate 46f). 

MICROHAIRS.-None seen. 

MACROHAIRS.-None Seen. 

Gastridium phleoides (Nees and Meyen) 

Hubbard 

PLAT~S 47 18 

ABAXIAL 

SToMATA.-Common; parallel; 2 stomatal 

bands/intercostal zone. 1 row/band. Intercostal 

zones very narrow and sunken (Plate 47a,b). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width uniform; ends straight to convex: 

nonsinuous. Papillae: None seen (Plate 17c). 

LONG CELLs.-Long, narrou.. width uniform: 

nonsinuous. Papillae: None seen (Plate 47a,c). 

hcKLEs.-Abundant: costal prickles, intercostal prickles and hooks (Plate 15d,c). 

SHORT CELLS.-cOStal solitary or sometimes 

paired, 3-5/row; intercostal paired. Silica Cells: 

Costal common; intercostal infrequent. Cork Cells: 

Costal infrequent to common: intercostal infrequent (Plate 47c-f). 

SILICA BoDrEs.-Costal nodular, sometimes 

very short: intercostal not seen (Plate 47c;f’). 

MICROHAIRS.--NOne seen. 

MACROHAIRS.-~One seen. 

ADAXIAL 

STOMATA.-cOmmOIl; parallel to low-dome: 2 

stomatal bands/intercostal zone, 1 row/band 

(Plate 48a,b). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width uniform or sometimes nonuniform; 

ends straight to convex; nonsinuous. Papillae: 

None seen (Plate 48c). 

LONG CELLs.-Long, narrow, width uniform to 

nonuniform; nonsinuous. Papillae: None seen 

(Plate 48c). 

hicKLEs.-Abundant; costal prickles; intercostal prickles and hooks (Plate 48c,d). 

SHORT CELLs.-Costal solitary and occasionally paired; intercostal not seen. Silica Cells: Costal 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

common; intercostal not seen. Cork Cells: Costal 

and intercostal not seen. The occasional pairs 

appear to be two silica cells (Plate 48ef). 

SILICA BoDIEs.-Costal nodular; intercostal not 

seen (Plate 48ef). 

MICROHAIRS.-None seen. 

MACROHAIRS.-None seen. 

Polypogon rnonspeliensis (Linnaeus) 

Desfontaines 

PLATES 49-50 

ABAXIAL 

STOMATA.-Abundant ; parallel to occasionally 

low-dome; 1 stomatal band/intercostal zone, 1 or 

2 rows/band. Difficult to distinguish costal and 

intercostal zones (Plate 496,~). 

INTERSTOMATAL CELLs.-Long, narrow, width 

nonuniform to occasionally rectangular or 

square; ends straight to convex; nonsinuous. Papillae: None seen (Plate 49c). 

LOKG CELrs.-Long, narrow, width nonuniform; nonsinuous. Papillae: None seen (Plate 

49a,c). 

PRIcKLEs.-Common; costal and occasional 

intercostal prickles (Plate 49d,e). 

SHORT CErLs.-Costal solitary, paired; intercostal not seen. Silica Cells: Costal infrequent to 

common; intercostal not seen. Cork Cells: Costal 

infrequent to common; intercostal not seen. Costal zones difficult to distinguish from intercostal 

regions. Cork cells difficult to detect (Plate 49e,f). 

SILICA BoDIEs.-Costal elongated-sinuous to 

nodular; intercostal not seen (Plate 49ef). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-None seen. 

ADAXIAL 

SToMATA.-Common; parallel; 1 stomatal 

band/intercostal zone, 1-3 rows/band (Plate 

50a,b). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width nonuniform; ends straight to convex; nonsinuous. Papillae: None seen (Plate 504. 

LONG CELLs.-Long, narrow, width nonuniform; nonsinuous. Papillae: None seen (Plate 

PRIcKLEs.-Abundant; costal prickles and 

short, straight, macrohair-like prickles (Plate 

5Od, f) . 

SHORT CELLs.-Costal solitary, 3-5/row; intercostal not seen. Silica Cells: Costal common; intercostal not seen. Cork Cells: Costal common; intercostal not seen (Plate 50ef). 

SILICA BoDIEs.-costal elongated-sinuous; intercostal not seen. Some very long (Plate 5Oef). 

MIcRoHAIRs.-None seen. 

MACROHAIRS.-see section on Prickles. 

5Oa,c). 

Tribe STIPEAE 

Stipa dregeana Steudel 

PLATES .3 1-52 

ABAXIAL 

SToMATA.-Common; parallel to low-dome; 2 

stomatal bands/intercostal zone, 1 row/band 

(Plate 5 la,b). 

INTERSTOMATAL CELs.-Long, narrow, width 

uniform; ends straight to convex or slightly concave; markedly sinuous, interlocking, raised. Papillae: None seen (Plate 5 lc). 

LONG CELLs.-Long, narrow, width uniform; 

markedly sinuous, interlocking, raised. Papillae: 

None seen (Plate 5la,e). 

PRIcKLEs.-Infrequent; costal prickles (Plate 

5ld). 

SHORT CELLs.-Costal 3-5 row, >5/row; intercostal solitary. Silica Cells: Costal abundant; intercostal common. Cork Cells: Costal abundant; 

intercostal not seen (Plate 5 1 c,e,f). 

SILICA BoDIEs.-costal dumbbell, ends 

rounded (with small terminal lobe), middle wide, 

long or short; intercostal elliptical (Plate 5 lc,e,f). 

MIcRoHAIRs.-None seen. 

MAcRoHAIRs.-None seen. 
NUMBER 53 

ADAXIAL 

STOMATA.-Abundant; parallel to low-dome: 

1-3 stomata1 bands/intercostal zone, 1-4(?) rows/ 

band. Costal regions hard to distinguish (Plate 

52a-c). 

INTERSTOMATAL CELLs.-Rectangular to long, 

narrow, width uniform, ends straight to convex 

to slightly concave; markedly sinuous, interlocking, raised. Papillae: None seen (Plate 526). 

LONG CELLs.-Long, narrow, width uniform; 

markedly sinuous, interlocking, raised. Papillae: 

None seen (Plate 52a,c). 

PRIcKLEs.-Infrequent; 

SHORT CELLs.-Costal 

52a). 

17 

costal prickles (Plate 

paired, 3-5/row: >5/ 

row; intercostal paired, 3-5/row. Silica Cells: Costal abundant; intercostal common. Cork Cells: 

Costal abundant; intercostal common (Plate 

52d-f). 

SILICA BoDIEs.-costal dumbbell, ends 

rounded, middle wide, long or short, intercostal 

square, round, dumbbell, ends rounded, middle 

wide, long or short (Plate 52d-f). 

MIcRoHAms.-None seen. 

MACROHAIRS.-common; intercostal and costal; basal cells absent (Plate 52f). 
Species 

Agrostis schimperiana 

Aira caryophyllea 

Anthoxanthum nivale 

Avena abyssinica 

Brachypodium flexum 

Briza maxima 

Bromus leptoclados 

Calamagrostis epigeios 

Colpodium chionogeiton 

Dactylis glomerata 

Deschampsia caespitosa 

var. olioeri 

Fes t uca a by ss in ica 

Gastridium phleoides 

Helictotrichon elongatum 

Hordeum marinum 

Koeleria capensis 

Lolium temulentum 

Phalaris arundinacea 

Poa schimperana 

Polypogon monspelienszs 

Pseudobromus sylvaticus 

Secale africanum 

Stipa dregeana 

Streblochaete longiarista 

Triticum aestivum 

Vulpia bromoides 

Appendix 

Material Studied 

Collection 

Schlieben 4645 

Morton 7069 

Amshoff 1970 

Pappi 4901 

Davidse 5825 

Simon 2330 

Maitland 126 1 

Greenway and 

Wood 923 

Gillett 15753 

Osmaston 3779 

Kanuri 14923 

Kerfoot 1732 

Loubser 3224 

Liebenberg 12 

Bornmutter 11 102 

Taylor 3 7 7 7 

Amshoff 1972 

Toosseus 1042 

Davidse 7070 

Adam 18808 

Robinson 5278 

Schweickerdt 7482 

Schweickerdt 1818 

Brenan 95 12 

Harshberger 1139 

Mooney 6348 

Origin 

Tanganyika 

Cameroon 

Ethiopia 

Ethiopia 

South Africa 

Rhodesia 

Cameroon 

Kenya 

Tanzania 

Jordan 

Uganda 

Tanzania 

South Africa 

Uganda 

Kenya 

Ethiopia 

South Africa 

Kenya 

Mauritania 

Nyasaland 

South Africa 

South Africa 

British 

Cameroons 

Algeria 

Ethiopia 

Egypt 

Herbarium 

MO 

K 

MO 

US 

MO 

MO 

K 

MO 

K 

us 

US 

K 

MO 

u S 

US 

MO 

MO 

US 

MO 

K 

MO 

US 

US 

us 

US 

K 

18 
Literature Cited 

Clayton. W.D. 

1970. Gramineae, Part 1. In E. Milne-Redhead and 

R.M. Polhill, editors, Flora of Tropical East Africa, 

pages 1-176. London: Crown Agents for OLVersea 

Governments and .4dministrations. 

1974. Gramineae, Part 2. In E. Milne-Redhead and 

R.M. Polhill, editors, Flora of Tropical East Africa, 

pages 177-449. London: Crown .4gents for Oversea Governments and .4dministrations. 

Gramineae, Part 3. In E. Milne-Redhead and 

R.M. Polhill. editors. Flora qf Tropical East Africa. 

London: Crown Agents for Oversea Governments 

and Administrations. 

In prep. 

Hartley, W. 

1973. Studies on the Origin, Evolution, and Distribution of the Gramineae, V: The Subfamily 

Festucoideae. Australian Journal of Botany, 2 1 :20 1- 

234. 

An Altitudinal Cline in Tropical African Grass 

Floras and Its Paleoecological Significance. Quaternay Research, 13:392-402. 

Grass Cuticles: A New Paleoecological Tool for 

East PIfrican Lake Sediments. Cunadian Journal of 

Botany, 54(15): 1725-1734. 

.A Scanning Electron Microscope Survey of the 

Epidermis of East African Grasses, I. Stn~thso~irarr 

Contributions to Botanl, 49: 81 pages. 

Livingstone, D..4., and W.D. Clayton 

1980. 

Palmer, P.G. 

1976. 

Palmer, P.G.. and .4.E. Tucker 

1981. 

19 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 1.-Abaxial epidermis Brim maxima, Simon 2330: a, overview, X 160: b, stoma, X 640: 

c, interstomatal cell, X 320; d, uniform long cells, X 640; e, nonuniform long cells, X 640: J 

silica body (arrow), X 867. 
NUMBER 53 

PLATE 2.-Adaxial epidermis Brzta maxima, Simon 2330: a, overview, X 161; b, stoma, X 1285; 

c, interstornatal cell, X 416; d, long cells, X 295: e, long cells, X 625;J prickle, X 1696. 

21 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE S.-Abaxial epidermis Colpodium chionogeiton, Wood 923: a, overview, X 295; 6, stoma, X 

867; c, interstornatal cells (arrow), long cells, X 867; d, costal papillate prickles, X 416: e, costal 

silica body (arrow), X 1285;J costal silica body (sb) and cork cell (cc). X 1285. 
NUMBER 53 

PLATE 4.-Adaxial epidermis Colpodium chionogeiton, Wood 923: a, overview, X 416; b, stoma, X 

1696: c, long cells and interstomatal cell (arrow), X 625: d, short cells (arrow), X 416: e, silica 

body, X 1696; J marginal papillate-type prickles, X 29.5. 

23 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 5.-Abaxial epidermis DacQlis glornerata, Gillett 15753: a, overview, X 295; b, stoma, X 

1285; c, interstornatal cell (arrow), X 867; d, prickle, X 1285: e, marginal area, prickles, short 

cells, X 295;J silica bodies (arrow), X 625. 
NUMBER 53 

PLATE 6.-Adaxial epidermis DacQlzs glomerata, Gillett 15753: a, overview, X 295: b, stoma. X 

1285; c, interstornatal cell (arrow), X 416; d, prickle, X 867; e, silica body (arrow), X 867: 1; 

silica bodies (sb), cork cells (cc), X 416. 

25 
26 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

PLATE 7.-Abaxial epidermis Festuca abyssinzca, Kerfoot 1732: a, overview, X 161; b, long cells, 

X 867; c, long cells. intercostal prickles. X 295; d, costal prickle, X 867; e, costal silica body (sb) 

and cork cell (cc), X 1696;J costal silica body (sb) and cork cell (cc), X 1696. 
NUMBER 53 

PLATE 8.-Adaxial epidermis Fesluca abyssznzca, Kerfoot 1732: a, overview. X 161. b, stoma. X 

867; c, intercostal long cells, X 867; d, short macrohair-type prickle, X 1285: e, overview with 

prickles, X 416;J intercostal silica body, X 1696. 

27 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 9.-Abaxial epidermis Lolium &mdentum, Amshoff 1972: a, overview, long cells, X 295; b, 

marginal zone, prickle, X 295; c, marginal prickle, X 1285; d, midvein with short cells (arrow), 

X 295; e, marginal silica body (arrow), X 1696:J silica body (arrow) on midvein X 1285. 
NUMBER 53 

PLATE 10.-Adaxial epidermis Lolium temulentum, Amshoff 1972: a, overview, X 295: b, stoma, 

X 1285; c, stomata1 bands, X 416; d, costal prickles, silica body (arrow), X 625; e, intercostal 

prickle (arrow), X 295:J; costal silica bodies (arrow), X 867. 

29 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 1 1.-Abaxial epidermis I’ou schimperana, Davidse 7070: a, overview, X 161: b, overview, 

prickles, X 295; c, long cells, X 625; d, costal prickle. X 1285; e, costal silica body (arrow), X 

1285:f; very long costal silica body (arrow), X 867. 
NUMBER 53 31 

PLATE 12.-Adaxial epidermis f’oa schzmperana, Davidse 7070: a, overview, X 295; b, stoma, X 

1285; c, interstomatal cell (arrow), X 416; d, costal prickle, X 1285; e, intercostal prickle, X 

1285:f; costal silica bodies, X 867. 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 13.-Abaxial epidermis I’.\eudohromus Aylualicu.c, Robinson 5278: u, o\wview, X 161: h, 

stoma. X 867: c. long cells, X 416; d, costal prickles. X 121; e, silica body. X 1696:J silica bodies 

(sb). cork cell (cc). X 625. 
NUMBER 53 

PLATE 14.--Adaxial epidermis Pseudobromus sylvaticus, Robinson 5278: a, overview, X 16 1 : b, 

stoma. X 867; c, interstornatal and long cells, X 295: d, long cells, X 416; e, costal prickle, X 

416:J; costal silica body (arrow), X 416. 

33 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 15.-Abaxial epidermis Vulpia bromoides, Mooney 6348: a, overview, X 295: b, long cells, 

X 867; c, costal prickle, X 1696; d, costal silica bodies (sb) and cork cells (cc), X 1285; e, 

intercostal silica bodies (sb) and cork cells (cc). X 1285;J intercostal silica bodies (sb) and cork 

cell (cc). X 1696. 
NUMBER 53 35 

PLATE 16.-Adaxial epidermis Vulpia 67ornotdes, Mooney 6348: a, overview, X 161; 6, stoma 

(arrow). interstornatal cell (ic), X 625; c, long cells, X 625; d, costal and intercostal (arrow) 

prickles, silica body (sb). X 625: e, intercostal prickle, X 1285; J; costal silica body with 

horizontal ridge (arrow), X 1696. 
36 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

PLATE 17.-.4baxial epidermis Bromus leptoclados, Maitland 1261: a, overview, X 295: b. long 

cells. macrohair, X 416; c, costal prickle, X 1285; d, costal silica body (sb) and cork cells (cc), 

X 867: e, costal silica body, X 1285;J base of macrohair, X 1285. 
NUMBER 53 

PLATE 18.--Adaxial epidermis Brornus leptoclados, Maitland 1261: a, overview, X 161; 6: stoma, 

X 1285; c, interstomatal cell (arrow), X 625; d, marginal area, X 295; e, costal silica body, X 

867:S, base of macrohair. X 867. 

37 
38 SMITHSONIAN CONTRIBITIONS TO BOTANY 

PLATE 19.-Abaxial epidermis Brachypodzum flexurn, Davidse 5825: a, overview. X 295; b, long 

cells, X 1696; c, papillae on long cells, X 1696; d, costal prickle, X 1696; e, costal silica body (sb) 

and cork cell (cc), X 2970;J intercostal silica body (arrow), X 2970. 
NUMBER 53 

PmrE XI.--lidaxial epidermis Bmrh~P:pudiurn,~exurn, Davidse 5825: a, overview, X 161: b, )tollla. 

X 1696: c, intercostal short macrohairs. X 867; d. costal silica body. X 2970; P, intercostal silica 

body. X 2970:,L macrohairs, X 80. 

39 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 21.-Abaxial epidermis Hordeum marinum, Bornmutter 11 102: a, overview, X 161: b, 

stoma, X 1696; c, interstomatal cell (arrowj. long cells, X 867; d, short macrohair-type prickle. 

X 867; e, macrohair, short macrohairs, intercostal silica bodies (arrow), X 625; J; costal silica 

body (arrow), X 1696. 
NUMBER 33 

PLATE ??.--Adaxial epidermis Hordeum marinum, Bornmutter 11 102: a, overview, X 295: b, 

stoma, X 1696: c, stomata, interstomatal cell, long cells, X 867: d, costal silica body (arrow), X 

1285; e, costal silica body (sb) and cork cell (cc), X 1285:J costal silica body (arrow'], X 1696. 

41 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 23.-Abaxial epidermis Secale aJrzcanum, Schweickerdt 7482: a, overview, X 80: b, stoma, 

X 867; c, long cells, macrohairs, X 295; d, costal silica body, X 1696; e, costal silica body, X 

1696;J costal silica bodies (sb) and cork cells (cc) (?), X 867. 
PLATE 24.-Adaxial epidermis Secale africanum, Schweickerdt 7482: a, overview. X 80: b, stoma, 

X 128.5; c, interstomatal cells, X 625; d, costal silica body, X 867; e, macrohairs, X 295;f; base 

of macrohair, X 1696. 

43 
44 SMITHSOh-IAN CONTRIBUTIONS TO BOTANY 

PLATE 25.--i-\baxial epidermis Triticurn aesfiuurn, Harshberger 1139: a, overview. X 161: b. stoma. 

X 1285; c, interstomatal cell (arrow), X 867; d, long cells, X 867: e, costal silica body (arrow). 

X 1696;f; costal silica body (sb) and cork cell (cc) (T), X 1696. 
NUMBER 33 

PLATE 26.-Adaxial epidermis Trztzcum aestzuum, Harshberger 1139: a, overview, X 295; b, stoma, 

X 1285; c, interstomatal cell, (arrow), X 867; d, long cells, X 867; e, costal silica bodies, X 1285: 

J costal silica bodies (sb) and cork cells (cc) (?), X 625. 

45 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATL 27.-.4baxial epidermis Streblochaete longzaruta, Brenan 95 12: a, overview, X 295; b. stoma, 

X 1696: c, long cells, X 1285; d, intercostal prickle, X 1285; e, costal silica body (arrow), X 2970; 

J intercostal silica body, X 2970. 
NUMBER 53 

PLATE 28.-Adaxial epidermis Streblochaete longiarista, Brenan 95 12: a, overview, X 295: b, stoma. 

X 1696; c, interstomatal cell, X 625; d, costal prickles and silica body (arrow), X 867: e, 

macrohair, prickles, and hooks (arrow), X 416; J; short macrohair-type prickle with sunken 

base, X 867. 

47 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 29.-Abaxial epidermis Aira caryophyllra, Morton 7069: a, overview, X 161; b, stoma, X 

1285; c, interstornatal cell, X 416; d, costal prickles and short cells, X 416; e, costal silica body 

(arrow), X 625:J; costal silica body (arrow), X 867. 
NUMBER .53 

PLATE 30.--Adaxial epidermis Azra caiyophyllea, Morton 7069: a, overview, X 295; b, stoma, X 

867; c, interstomatal cell (arrow), X 416; d, costal prickles, X 625; e, costal silica bodies (sb) and 

cork cells (cc), X 625:J costal silica body, X 1285. 

49 
50 SMITHSONIAN CONTRIBUTIOKS TO BOTANY 

PLATE 3 1.-Abaxial epidermis Anthoxanthum niuale, Amshoff 1970: a, overview, X 295; b, stoma. 

X 1696; c, interstornatal cell (arrow), X 29.5: d, long cells, X 416: P, prickle, X 1285:J silica body 

(arrou,), X 1285. 
NUMBER 53 51 

PLATE 32.-Adaxial epidermis Anthoxanthum ntuale, Amshoff 1970: a, overview. X 161: b, uniform 

and nonuniform width long cells. X 867; c, end walls (arrows) of long cells. X 867: d, prickles 

and long cells, X 295: e, prickles, X 62.i;J; costal prickle, X 1285. 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 33.--Abaxial epidermis Acena abyssznzca, Pappi 4901: Q, overview, X 295; b, stoma, X 

1285; c, prickles. X 625; d, costal short cells, X 625; e, macrohairs, X 63:J base of macrohair. 

x 625. 
NUMBER 53 

PLATE 34.-Adaxial epidermis Auena abyssinzca, Pappi 490 1 : u, overview, silica bodies (arrow), 

X '295: b. stoma, X 867; c, prickles, X 867; d, long cells. silica body (arrow), X 625; e, base of 

macrohair. X 625:J; macrohairs, X 161. 

53 
54 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

PLATE 35.-Abaxial epidermis Desrhampsia caespitosa var. diwri, Osmaston 3779: a, overvieLv, 

X 161; h, long cells, X 1285: c, long cells, short cells. prickles, X 625: d, prickles, X 1285: P, long 

cells, short cells, X 867;J; silica body (sb). cork cell (cc). X 2970. 
NUMBER 53 

PLATE %-.Adaxial epidermis DeJchumpJia raespilosa var. oliueri, Osmaston 3779: u, overview. 

X 416: b, stoma. X 1696: c. long cells, X 1285: d, prickles, long cells. short cells (arrow). X 416: 

e. prickles. X 62.i:J; silica body (sb). X 2970. 

55 
56 SILIITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 37.-Abaxial epidermis Helictotrichon elongaturn, Liebenberg 12: a, overview. X 295: b, 

stoma, X 867; c, long cells, X 867; d, prickle, X 1691: e, long cells, silica body (arrow), X 416: 

J silica body. X 867. 
NUMBER 53 

PLATE 38,--.4daxial epidermis Heiictotrichon eiongalum, Liebenberg 12: a, overview; X 161 : 6, 

htoma. X 1'285: c, interstornatal cells. silica body (arrow), X 416; d, long cells. X 867: e, prickles. 

X 1285;J silica body (sb) and cork cell (cc), X 867. 

57 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 39.-Abaxial epidermis Koeleriu cupensis, Taylor 3777: a, overview and macrohairs, X 80; 

b, stoma, X 1696: c, long cells, X 295; d, prickle and short macrohair, X 867: e, silica bodies 

(arrow). X 867:J base of macrohair, X 867. 
NUMBER 53 

PurE 4O.-Adaxial epidermis Koeleria capensis, Taylor 3777: a, overview, X 161; b, prickles, X 

128.5: c, macrohairs, short macrohairs, and short cells, X 295; d, silica bodies (sb), cork cells (cc). 

X 1285: e. silica body (sb) and cork cell (cc). X 1696;l; base of macrohair, X 625. 

59 
60 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 4 1 .-Abaxial epidermis Phalaris arundznacen. Toosseus 1042: a, o\-erview. X 295: b. stoma, 

X 867; c, interstomatal cell (arrow), X 867; d, costal prickles, silica bodies (arroiv). X 416: P, 

intercostal silica body (sb) and cork cell (cc), long cells, X 867;,1; intercostal silica body (sb) and 

cork cell (cc) (?), X 1696. 
NUMBER 53 

PLATE 42.-Adaxial epidermis Phalarts arundinacea, Toosseus 1042: a, overview,, X 295; 6, stoma, 

X 1285, c, long cells, interstomatal cells, X 416; d, costal prickle and silica bodies (arrow). X 

625: e, costal silica bodies, X 1285;J; intercostal silica body (arrow), X 1285. 

61 
62 SMITHSONIAN CONTRIBUTIONS TO BOTAKY 

PLATL 43.--.4baxial epidermis .4gruJ/z.\ schzmnperiana, Schlieben 4645: a, overviav, X 295: 6, stoma 

(arrow). prickle, X 867: c, long cells. X 62.5: d, long cells. silica bodies. X 29.5: e, silica body, X 

1696;J; silica bodies (arrow). X 416. 
NUMBER 53 

PLATE 44.-Adaxial epidermis Aprostis schimperzana, Schlieben 4645: a, overview, X 295; h, 

stoma, X 1696: c, long cells, stornatal bands, X 416; d, long cells, silica body (arrow). X 867: e, 

costal prickle, X 867:f; costal silica body (arrow), X 867. 

63 
64 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 45.-Abaxial epidermis Culumagrostis epigeios, Greenway and Kanuri 14923: a, ovenview. 

X 161; b, stoma, papillate adjacent long cells, X 867: c, interstomatal cell, X 867; d, prickles, 

X 625; e, costal silica bodies (sb) and cork cells (cc), intercostal cork cells (arrow), X 625: 1; 

costal cork cells (arrow), X 625. 
NUMBER 53 

PLATE 46.-Adaxial epidermis CaLamagroslis epigeios, Greenway and Kanuri 14923: a, overview. 

mid-intercostal zone (arrow). X 161; 6, stoma, X 1696: c, interstomatal cell. X 1285: d, midintercostal long cells, X 625: e, intercostal prickles. hook (arrobv). cork cell (cc)~ X 625:j; cohtal 

silica bodies (sb) and cork cells (cc), X 867. 

65 
66 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

PLATE 47.--Abaxial epidermis Gastrzdzum phleozdes, Loubser 3223: a, overview, X 295: b, stoma, 

X 1696; c, interstornatal cell, X 625: d, costal prickle, short cells, X 867; e, costal silica body. 

hook, X 867;J costal silica bodies, X 867. 
NUMBER 53 

PLATE 48.-Adaxial epidermis Gastridium phleoides, Loubser 3224: a, overview. X 295: b, stoma, 

x 1285; c, interstomatal cell, X 416; d, prickles and hook, X 625; e, costal silica body, X 867:,f; 

paired costal silica bodies, X 625. 

67 
68 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 49.--Abaxial epidermis Polypogon monspeliensis, Adam 18808: a, overview, X 295; b, stoma, 

X 1285: c, interstomatal and long cells, X 416; d, costal prickle, X 867; e. costal silica body 

(arrow). X 625;J costal silica body (arrow), X 867. 
NUMBER 53 

PLATE 50.--.4daxial epidermis Polypogon monspeliensis, Adam 18808: a, overview, X 295; b, stoma, 

X 1285: c, interstornatal cell, X 416: d, short, niacrohair-type prickle. X 416: e, costal silica body 

(sb) and cork cell (cc). X 625;J; costal silica body (arrow). X 416. 

69 
70 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 5l.-Abaxial epidermis Stzpa dregeana, Schweickerdt 1818: a, overview, X 416; h, stoma. 

X 1285; c, interstomatal cell (arrow), silica bodies. X 867; d, costal prickle, X 867, e, intercostal 

silica body, X l696;J costal silica body. X 1696. 
NUMBER 53 

PLATE 5'2.-Adaxial epidermis Shpa dregeana, Schweickerdt 1818: a, overview. X 295: h, stoma, 

X 1696: c, interstomatal and long cells, X 867; d, costal silica bodies (sb), cork cell (cc). X 1'285; 

e, intercostal silica bodies. X 1696;J base of macrohair, X 867. 

71 
Index to Grass Species 

Agrostis schimperiana. 14. 62(pl.), 63(pl.) 

Aira caryophyllea, 10, 48(pl.), 49(pl.) 

Anthoxanthum nivale, 11. 5O(pl,), 5l(pl.) 

Avena abyssinica, 11. 52(pl.). 53(pl.) 

Brachypodium flexum. 7, 38(pl.). 39(pl.) 

Briza maxima, 2. 2O(pl.). 21(pl.) 

Bromus leptoclados, 6. 36(pl.), 37(pl.) 

Calamagrostis epigeios, 14, 64(pl.), 65(pl.) 

Colpodium chionogeiton. 3. 22(pl.), 23(pl.) 

Dactylis glomerata. 3. 24(pl.). 25(pl.) 

Deschampsia caespitosa var. oliveri. 12, 54(pl.). 55(pl.) 

Pestuca abyssinica. 4. 26(pl.). 27(pl.) 

Gastridium phleoides. 15. 66(pl.). 67(pl.) 

Helictotrichon elongatum. 12. .56(pl.). 5i(pl.) 

Hordeum marinum. 8. 4O(pl.). 4l(pl.) 

Koeleria capensis, 13. 58(pl.). 59(pl.) 

Lolium temulentum. 4. 28(pl.). 29(pl.) 

Phalaris arundinacea. 13. GO(p1.). 61(pl.) 

Poa schimperana, 5, 30(pI.). 3l(pl.) 

Polypogon monspeliensis. 16. 68(pl.). 69(pl.) 

Pseudobromus sylvaticua. .3. 32(pl.l. 33(pl.) 

Secale africanum, 8. 42(pl.). 43(pl.) 

Stipa dregeana. 16, iO(p1.). il(p1.) 

Streblochaete longiarista. 9, 46(pl.). 4i(pl.) 

Triticum aestivum, 9, 44(pl.). 45(pl.) 

\'ulpia bromoides. 6. 34(pl,). 33(pl.) 

72