SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 15 

Leaf Anatomy and Systematics 

of New World Velloziaceae 

Edward S. Ayensu 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1974 
ABSTRACT 

Edward S. Ayensu. Leaf Anatomy and Systematics of New World Velloziaceae. 

Smithsonian Contributions to Botany, number 15, 125 pages, frontispiece, 24 

figures, 51 plates, 1974.-The leaf anatomy of 106 species of New World Velloziaceae has been studied with the purpose of providing important characterstates in assessing the systematics of the family. Transverse section of the leaves 

have shown the type of sclerenchyma and mesophyll patterns that are assignable 

either to the genus Vellozia or Barbacenia (sensu lato). In addition to the light 

microscope, the scanning electron microscope was used to examine epidermal 

surfaces of the leaves as well as their internal structures. The SEM has served 

as a remarkable tool in allowing us to examine the topography of the leaf surface in three-dimension. Details of the structure of the stomata, the furrows in 

the leaf, and the types of hairs, including coalescent hairs, have been observed 

for the first time. The application of leaf anatomy in the taxonomy of the family 

has been stressed. Light and scanning electron micrographs are presented as an 

aid in the identification of each species. 

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

in the Institution’s annual report, Smzthsonian Year. SI PRESS &UMBER 4998. SERIES COVER DESIGN: Leaf clearing from the katsura tree Cercrdiphyllurn japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Ayensu, Edward S. 

Leaf anatomy and systematics of New World Velloziaceae. 

(Smithsonian contributions to botany, no. 15) 

1. Velloziaceae. 2. Leaves-Anatomy. 3. Botany-America. I. Title. 11. Series: Smithsonian 

QKl.SZ747 no. 15 [QK495.V4] 584’29 73-15606 

Institution. Smithsonian contributions to botany, no. 15. 

For sale by the Superintendent of Documents, US. Government Printing Office 

Washington. D.C. 20402 ~ Price $2.20 (paper cover) 
Contents 

Introduction . , . , , . , . . , , , . . , , . , , , . . . . , . . . , . . , . . , . . . . . . , . . . . . . . . . . . . Material Examined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . Methods ......................... ................. 

Characters Used in the Descriptions ... .................... 

Classification of the Species . . . . . . . . . . . . . . . . . . . . . . . , , . . . . . . . . . 

Acknowledgments . , , , , , , . . , , , , , , . . . , , . . . . . . . . ........... Vellozia , , , , , , . , , . , , . , , . , , . . , , , . , , , , . , . , , . , .................. Section I: Vellozia 

Vellozia abietina Martius , , 

Vellozia alata L. B. Smith . . . . . . . , . . . . , . . , , . . . 

Vellozia aloifolia Martius . . . , . . . . , . . . . . . . . . . . . 

Vellozia angustifolia Goethart & Henrard . . . . . . 

Vellozia asperula Martius , , . . . . . . , . . . . , , . . , . . . . . . . . . . . . . . . . . . . . 

Vellozia bicolor L. B. Smith 

Vellozia brachypoda L. B. Smith & Ayensu . . . , , . . . . . . . . . . . . . . . . . 

Vellozia burle-marxii L. B. Smith 

Vellozia caruncularis Martius & Seubert . . . . . . , , . . . . . . . . . . . . . . . . . 

Vellozia ciliata L. B. Smith 

Vellozia cinerascens Martius ex Schultes f. 

. . . , . . . . . . . , , . . . . . . . . . . . . 

. , . . , . . . ................ 

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

, , . . . . . . . . . . . . . . 

Vellozia compacta Martius ex Schultes f. . . . . . . . 

Vellozia costata L. B. Smith & Ayensu . . . 

Vellozia dasypus Seubert . . . . . . , , , , , , . . . . . . . . . 

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

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

Vellozia crassicaulis Martius ex Schultes f. . . . . . . . . . . . . . . . 

Vellozia crinita Goethart & Henrard . , , , , 

Vellozia declinans Goethart & Henrard . . . , , . . . . . . . . . . . . . . . . . 

Vellozia echinata Goethart & Henrard . . . . . . . . . . . . . . . . . Vellozia epidendroides Martius ex Schultes f. . . . . . . . 

Vellozia exilis Goethart & Henrard , . . , , , . 

Vellozia fibrosa Goethart & Henrard . , . , , , . . . . . . . . . . . . . . . . . . . . . . 

........... 

Vellozia flavicans Martius ex Schultes f. 

Vellozia glabra Mikan , , . . . . . . . . , . . . . . . . . . . . . . . . 

Vellozia glandulifera Goethart & Henrard . , . . . . . . . . . . . . . . . . . . . . . . 

Vellozia glauca Pohl . . . . . . . . 

Vellozia glochidea Pohl-Heringer 

Vellozia granulata Goethart & Henrard . . . . . . 

Vellozia grisea Goethart & Henrard . . . . . . . . . . . . . . . . . . . . 

Vellozia hatschbachii L. B. Smith & Ayensu . . . . . 

Vellozia hemisphaerica Seubert in Martius . . , . . . . . . . . . . . . . . . . . . . . 

Vellozia hypoxoides L, B. Smith . . . . . . , , . . . . 

Vellozia incuruata Martius ex Schultes f. 

Vellozia intermedia Seubert in Martius , , . . . . . . . . . . . . . . . . . . . . . . . . 

Vellozia irwinii L. B. Smith , , 

. , . . . . . . . . . , . . . 

..... ..... 

,.. ............ ...... 

, , . . . . . . . . . . . ............ 

. . . . . . . . . . 

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

... ,. ....... ... ...... 

Vellozia lappa L. B. Smith & Ayensu . . . . . . . . . . . .......... 

posr 

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Vellozia leptopetala Goethart & Henrard ........................ 30 

31 

Vellozia metzgerae L. B. Smith ..... 31 

Vellozia minima Pohl 33 

Vellozia modesta L. B. Smith & Ayensu ................. 33 

Vellozia nuda L. B. Smith & Ayensu , , . . , . 33 

Vellozia ornata Martius ex Schultes f. .......................... 34 

Vellozia phalocarpa Pohl ...................................... 34 

Vellozia pilosa Goethart & Henrard ............... ' 35 

Vellozia piresiana L. B. Smith ' 35 

Vellozia plicata Martius ...................................... 35 

Vellozia pumila Goethart & Henrard ............................ 37 

Vellozia punctulata Seubert in Martius .......... ... .. 37 

Vellozia resinosa Martius ex Schultes f. 37 

Vellozia sellovii Seubert in Martius 38 

Vellozia squalida Martius ex Seubert f. .................... 38 

Vellozia swallenii L. B. Smith ........... ............ 39 

Vellozia taxifolia Martius ex Schultes f. .................. 39 

Vellozia tenella Martius ex Schultes f. ........................... 39 

Venozia teres L. B. Smith & Ayensu .................... 

Vellozia tragacantha Martius ex Seubert . . 

Vellozia variabilis Martiu ..................... 44 

Vellozia verruculosa Martius ex Schultes f. .... ........ 44 

Vellozia virgata Goethart & Henrard . . ........................ 45 

Vellozia wasshausenii L. B. Sm 

Vellozia cachimbensis L. B. Smith .......... ............ 45 

Vellozia cana Goethart & Henrard ......................... 46 

Vellozia candida Mikan ...... 

Vellozia dawsonii L. B. Smith ............... 48 

Vellozia dumitiana Schultes ... 

Vellozia hirsuta Goethart & He 

Vellozia lanata Pohl ........................... 

Vellozia machrisiana L. B. Smith 

Vellozia maculata Goethart & Henrard .......................... 50 

Vellozia riedeliana Goethart & Henrard ......... 

Vellozia tubifEora A. Richard ............ 

Vellozia magdalenae L. B. Smith & Ayensu ........ 

....... 

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

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

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

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

Vellozia triquetra Pohl . . ......................... 42 

Section 11: Radia ................................... 

Vellozia caput-ardeae L. B. Smith & Ayensu ...... 

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

Vellozia lithophila Schultes .............. ............ 49 

Barbacenia . , ............................................... 53 

............................................... 53 

............. 53 

Barbacenia gaveensis Goethart & Henrard ........................ 53 

Barbacenia gounelleana Beauverd ............... 

Barbacenia hatschbachii L. B. Smith & Ayensu ................... 54 

Barbacenia irwiniana L. B. Smith .............................. 55 

55 

Barbacenia delicatula L. B. Smith & Ayensu . , 

Barbacenia longiscapa Goethart & Henrard ............... 

Barbacenia purpurea Hooker ....... .................... 55 

iv 
Barbacenia seubertiana Goethart & Henrard . . . . . , , , . . . . . . . , . . . . . Barbacenia stenophylla Goethart & Henrard . . . . . , . . ..... ... Subkey 11. . . . . . . . . , . . . . . . , . . , . . . . . . .................. Barbacenia celiae Maguire ..................................... 

Barbacenia flava Martius ex Schultes f. . . . . . . , , , . , . . . . . . . . . . . . . . . 

Barbacenia involucrata L. B. Smith . . . , , . . . . . . . . . . . . . . . . . . . . . . . . 

Barbacenia schwackei Goethart & Henrard . . . . . . . . . . 

Barbacenia sellovii Goethart & Henrard , . . . . . . . . . . Barbacenia vandellii Pohl ex Seubert in 

Barbacenia williamsii L. B. Smith . . . . . . . . . . . . . . . ... . . . . . . . . . . . . . . 

Subkey 111. . . . , . . . . . . . . , , . , , . . . . . . . . . . . , . .. . . . . . . . . . . .. . .. . . . . . . Barbacenia blackii L. B. Smith . . . . . . . . . . 

Barbacenia conicostigma Goethart & Henrard , , . . . . . . . . . . . . . . . . . . . Barbacenia exscapa Martius .................................... 

Barbacenia ignea Martius ex Schultes f. . ............ 

........... *. 

Barbacenia fragrans Goethart & Henrard . . . . . 

Barbacenia gardneri Seubert in Martius . . 

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

.............. Barbacenia gentianoides Goethart & Henra 

Barbacenia longiflora Martius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

Barbacenia lurulifolia Martius ex Schultes f. . . . . 

Barbacenia macrantha Lem ................................... .. Barbacenia nana L. B. Smith & Ayensu . . . . . . . . . , , . . . . . . . . . 

Barbacenia paranuensis L. B. Smith . . . . . . . . 

Barbacenia riedeliana Goethart & Henrard 

Barbacenia rubro-virens Martius . , 

Barbacenia tomentosa Martius . . . , ..... ... 

Barbacenia viscosissima Goethart & 

,....*........ 

Observations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .............. 

Light Microscopic Observations . . 

Scanning Electron Microscopic 0 

Literature Cited ........................ 

Plates .............................. 

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V 

Leaf Anatomy and Systematics 

of New World Velloziaceae 

Edward S. Ayensu 

Introduction 

This study is a sequel to an earlier anatomical 

treatment of the Old World Velloziaceae (Ayensu 

1969a) prepared primarily to help solve taxonomic 

problems in this family, Unlike the Old World 

species, the New World members have been largely 

brought up to date taxonomically by Smith (1962), 

including his subsequent publications. Except for 

the publication by Warming (1893) on the biology 

and anatomy of the leaf of Velloziaceae, the work 

of Menezes (1971a) and my studies of Barbaceniopsis (Ayensu, 1968), and Velloria uaipanensis 

(Ayensu, 1969b), no systematic attempt has been 

made to bring together the anatomical data of the 

New World species. 

As I have indicated in other publications 

(Ayensu 1969a, 1973), features of the vegetative 

anatomy of the leaves in this family provide very 

important character-states in separating the genera. 

The most important anatomical characters are the 

sclerenchyma patterns in the leaves. In species of 

Vellozia the vascular bundles are partially surrounded by sclerenchyma on the adaxial side in 

the form of an inverted crescentiform or V-shaped 

cap. The abaxial sclerenchyma is either U- or Yshaped. In Barbacenia the vascular bundles are 

partially surrounded on both the adaxial and 

abaxial sides by Y-shaped sclerenchyma, the adaxial 

one being inverted. 

Apart from these broad sclerenchyma patterns 

that separate the genera, each species exhibits disEdward S. Ayensu, Department of Botany, Smithsonian Institution, Washington, D.C. 20560. 

tinct mesophyll and epidermal features that permit 

their individual identification. The descriptions of 

the leaf anatomy of the New World Velloziaceae, 

as well as the accompanying line drawings and 

photomicrographs of the anatomical features, will 

hopefully provide additional clarification for the 

interpretation of the relationships of taxa in this 

family. This study will also serve as an atlas of 

the leaf anatomy of the New World species. 

MATERIAL EXAMINED.-The following 24 1 specimens (106 species of two genera) were examined 

for this study. The herbarium vouchers are deposited either in the United States National 

Herbarium, National Museum of Natural History, 

Smithsonian Institution (US) or the New York 

Botanical Garden (NY). 

Vellozia abietina Martius.-Maguire, Magalhaes & Maguire 

49115, (1960) Concepcao, Minas Gerais, Brazil, NY & US. 

Vellozia alata L. B. Smith.-Maguire, Magalhaes & Maguire 

49049, (1960) Serra do Cip6, 1200 m, Minas Gerais, Brazil, 

NY & us. 

Vellozia aloifolia Martius.-Y. Mexia 5870 (1931) Serra do 

Capao, 1240 m, Minas Gerais, Brazil, US. 

Vellozia angustifolia Goethart & Henrard.-Riedel (1824) 

Serra da Lapa, Brazil, US; W. A. Archer 4092 (1936) highway between Diamantina and Guinda, Minas Gerais, Brazil, 

US; Centro Pesq. (R) 1411, Brazil, NY. 

Vellozia asperula Martius.-G. Gardner 5228, Minas Gerais, 

Brazil, US; Occhioui s.n., Brazil, NY. 

Vellozia bicolor L. B. Smith.-B. Maguire et al. 49047 (1960) 

Serra Cipo, 1200 m, Minas Gerais, Brazil, NY & US. 

Vellozia brachypoda L. B. Smith & Ayemu.-Irwin 23534, 

Brazil, NY. 

Vellozia burle-marxii L. B. Smith.-R. Burle-marx m., 

Brazil, US; Irwin et al. 32520 (1971) Rio Ferra Doido, 18 

km E of Morro do Chapeu, ca. 1100 m, Bahia, Brazil, US. 

Vellozia cachimbensis L. B. Smith.-Pires, Black, Wurdack & 

Silva 6235 (1956) Serra do Chachimbo, Para Brazil, US: 

1 
2 

Pires 6111 (1956) Serra do Chachimbo, Para Brazil, NY & 

us. 

Vellozia cana Goethart & Henrard.-H. S. Irwin, R. Souza & 

R. Reis dos Santos 11734, Brazil, NY & US; H. S. Irwin 

et al. 12556, Brazil, NY & US. 

Vellozia candrda Mikan.-Kuhlmann s.n. (1922) Brazil, NY; 

j. P. P. Carauta 160 (1963) Pedra da Gavea, Guanabara, 

Brazil, US; Porto 91195, Brazil, NY; Jardim Bot., Rio de 

Janeiro 2485, Brazil, NY; J. N. Rose & P. G. Russell 20669 

(1915) Corcovado, Rio de Janeiro, Brazil, US. 

Vellozia caput-ardeae L. B. Smith.-L. B. Smith, Ayensu & 

Hatschbach 15989, (1972) Giunda, Diamantina, Minas 

Gerais, Brazil, US. 

Vellozia caruncularis Martius & Seubert.-B. Maguire, J. M. 

Magalhaes, C. K. Maguire 49013 (1960) Serro do Cipo, 

1200 m, Minas Gerais, Brazil, NY & US; E. P. Heringer 

5265 (1956) Serra do Cipo, Riacho da Serra, Minas Gerais, 

Brazil, US; J. M. Pires & G. A. Black 2721 (1951) Serra do 

Cipo, Minas Gerais, Brazil, US; L. B. Smith 6855 (1952) 

Serra do Cipo, 1200 m, Minas Gerais, Brazil, US; A. Macedo 

2959 (1951) Serra do Cipo, Minas Gerais, Brazil, US; L. B. 

Smith & Ayensu 15988 (1972) 9 km W of Serro, Datas, 

Minas Gerais, Brazil, US. 

Vellozia ciliata L. B. Smith.-L. B. Smith & Ayensu 15995 

(1972) 20 km from Diamantina, Minas Gerais, Brazil, US. 

Vetlozia cinerascens Martius ex Schultes f.-Lehnkner 323 

(1912) Serra de Tinga, Bahia, Brazil, NY. 

Vellozia compacta Martius ex Schultes f.-B. Maguire, G. M. 

Magalhaes, C. K. Maguire 49132 (1960) 3-5 km E of Serra 

on road to Diamantina, Minas Gerais, Brazil, NY & US; 

H. S. Irwin 2407 (1959) Serra da Piedade, 5500 ft, Minas 

Gerais, Brazil, NY & US; L. B. Smith 7005, Brazil, US; 

Segadas-Vianna & Loredo 1085, Brazil, US. 

Vellozia costata L. B. Smith & Ayensu.-H. S. Irwin et al. 

20999, Brazil, NY. 

Vellozia crassicaulis Martius ex Schultes f.-W. A. Archer 

4076 (1936) Mendanha, Minas Gerais, Brazil, CS; W. A. 

Archer 4079 (1936) Mendanha, Minas Gerais, Brazil, NY 

& US; H. S. Irwin, R. Souza, & R. Reis dos Santos 9975 

(1965) 5 km S of Cristalina, 1175 m, Goias, Brazil, NY & 

US; H. S. Irwin, R. Souza, & R. Reis dos Santos 9744 (1965) 

5 km S of Cristalina, Goias, Brazil, NY & US; H. S. Irwin, 

R. Souza, & R. Reis dos Santos 11699 (1966) Serra Dourada, 

20 km SE of Goias Velho, Goai s, Brazil, NY & US; H. S. 

Irwin, J. W. Grear, Jr., R. Souza, & R. Reis dos Santos 

13377 (1966) ca. 3 km S of Cristalina, 1200 m, Goias, 

Brazil, NY & US; L. B. Smith & Ayensu 15965 (1972) 1 km 

N of Palacios, Minas Gerais, Brazil, US. 

Vellozia crinita Goethart & Henrard.-Glaziou 16388, Brazil, 

NY. 

Vellozia dasypus Seubert.-Blanchet 3558, Brazil, NY; R. de 

Lemos Froes 20246 (1943) Serra de Sincora, Baiia Brazil, 

US; J. N. Rose & P. G. Russell 19697 (1915) Bahia, Brazil, 

us. 

Vellozia dawsonii L. B. Smith.-H. S. Irwin, J. W. Grear, R. 

Souza & R. Reis dos Santos 12648 (1966) Chapada dos 

Veadeiros, 1000 m, Goias, Brazil, NY & US; H. S. Irwin, 

J. W. Grear, Jr., R. Souza, R. Reis dos Santos 12622 (1966) 

Chapada dos Veadeiros, Goias, Brazil, US. 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Vdhia declinans Goethart & Henrard.-Hatschbach 27842, 

Brazil, NY. 

Vehzia durnitiana Schultes.Schultes & Cabrera 1505OA 

(1952) Cerro Isibukuri, Rio Kananari, Vaupes, Colombia, 

US; H. Garcia-Barriga 13804 (1951) Rio Kananari y Cerro 

Isibukuri, Vaupes, Colombia, US. 

Vellozia echinata Goethart & Henrard.-Riedel 1780 (1824) 

near Paranna, Minas Gerais, Brazil, US. 

Vellozin epidendroides Martius ex Schultes f.-E. .Pereira 

2229 (1956) Espirito Santo, Forno Grande, 1700 m, Brazil, 

US; A. P. Duarte 4519 (1958) Serra do Cipo, Minas Gerais, 

Brazil, NY; B. Maguire, G. M. Magalhaes, C. K. Maguire 

49060 (1960) Serra do Cipo, 1200-1300 m, Minas Gerais, 

Brazil, US; A. Macedo 2993 (1951) Cipo, Minas Gerais, 

Brazil, US; L. B. Smith 6762 (1952) Serra do Cipo, 1200 m, 

hlinas Gerais, Brazil, US; Duarte 6537, Brazil, NY; Brade 

19136, Brazil, NY; Schwacke 10321, Brazil, NY; Lima & 

Brade 13297A, Brazil, NY; Schwacke 8354, Brazil, NY. 

T'ellozia exilis Goethart & Henrard.-H. S. Irwin, J. W. 

Grear, Jr., R. Souza, & R. Reis dos Santos 12696 (1966) 

Chapada dos Veadeiros, 1000 m, Goiais, Brazil, US; H. S. 

lrwin 12692, Brazil, US; H. S. Irwin et al. 9507, Brazil, NY 

& us. 

Vellozia fibrosa Goethart & Henrard.-D. Azevedo 5592 

(1943) Serra do Jatoba, Minas Gerais, Brazil, US. 

Vellozia pavicans Martius ex Schultes f.-H. S. Irwin, J. W. 

Grear, Jr., R. Souza, R. Reis dos Santos 12938 (1966) 

Chapada dos Veadeiros, Goias, Brazil, NY & US; J. M. 

Pires, N. T. Silva, & R. Souza 9052 (1963) Parque do Gama, 

Brasilia, Brazil, US; L. B. Smith 4670 (1956) Anapolis, 

Goias Brazil, CS; E. P. Heringer 8162-256 (1961) Plan0 

Piloto, Brasilia, Brazil, US; A. Andrade 35L-M. Emmerich 

348 (1960) Cabeca do Veado, Brasilia, Brazil, US; H. S. 

Irwin, R. Souza, R. Reis dos Santos 9379 (1965) ca. 12 km 

KIV of Veadeiros, 1200 m, Goias, Brazil, NY & US; H. S. 

Irwin, J. W. Grear, Jr., R. Souza, & R. Reis dos Santos 15520 

(1966) ca. 3 km S of Sobradinho, Federal District, Brazil, 

SY & US; G. Gardner 4370, Brazil, US; H. S. Irwin, R. 

Souza, & R. Reis dos Santos 8073 (1965) Chapada da Contagem, ca. 25 km NE of Brasilia, 1000 m, Federal District, 

Brazil, NY & US; H. S. Irwin & T. R. Soderstrom 5100 

(1964) Chapada da Contagem, ca. 20 km E of Brasilia, 

Federal District, Brazil, NY & US; H. S. Irwin, J. W. Grear, 

Jr., R. Souza, & R. Reis dos Santos 12574 (1966) Chapada 

dos Veadeiros, Goias, Brazil, NY & US; H. S. Irwin et al. 

11399, Brazil, NY & US. 

Vellozia glabta Mikan.-B. Maguire, G. M. Magalhaes, C. K. 

Maguire 49048 (1960) Serra Cipo, 1200 m, Minas Gerais, 

Brazil, NY & US; B. Maguire, C. K. Maguire, J. M. Pires 

44622 (1959) Serra Rola Moca, 4400 ft, Minas Gerais, 

Brazil, US; Duarte 8355-Mathos 464, Brazil, NY. 

Vellozia glandulifera Goethart & Henrard.-L. B. Smith & 

Ayensu 15998, Brazil, NY. 

Vellozia glauca Poh1.-Maguire, Maguire & Pires 44793 

(1959) 6 mi before Cristalina, Goias, Brazil, NY & US; 

Irwin et al. 21395 (1968) 10 km S of Guara, Goifis, Brazil, 

NY. 

Vellozia glochidea Poh1.-A. P. Duarte 8340-A. Mathos 677 

(1964) Goiania, Goias, Brazil, NY; H. S. Irwin, J. W. 
NUMBER 15 3 

Grear, Jr., R. Souza & R. Reis dos Santos 14584 (1966) 

Posse, Serra Geral de Goias, Goias, Brazil, NY & US; E. P. 

Heringer 10923 (1966) Serra Dourada, Goias, Brazil, US. 

Vellozia granulata Goethart & Henrard.-Riedel s.n. (1824) 

Serra da Lapa, Brazil, K; Glaziou 19934 (1892) Diamantina, 

hfinas Gerais, Brazil, NY. 

Vellozia grihea Goethart & Henrard.-Weddell 3006, Goyaz & 

Cujabo, Brazil, NY. 

Vellozia hatschbachii L. B. Smith & Ayemu.-L. B. Smith, 

Ayensu & Hatschbach 16002 (1972) 33 km NE Diamantina, 

Minas Gerais, Brazil, US. 

Vellozia hemisphaerica Seubert in Martius.-Blanchet 2544 

(1838) Jacobina, Bahia, Brazil, NY. 

Vellozia hirsuta Goethart & Henrard.-Hatschbach 28093, 

Brazil, NY; L. B. Smith, Ayensu & Hatschbach 15999 (1972) 

Serra do Espinhaco, Diamantina, Minas Gerais, Brazil, US. 

Vellozia hypoxoides L. B. Smith.-Irwin et al. 34120 (1972) 

Serra dos Pireneus, Goias, Brazil, US & NY. 

Vellozia incuroata Martius ex Schultes f.-Maguire, Maguire 

& Pires 44732, (1959) 49 mi from Diamantina, Minas 

Gerais, Brazil, NY & US; Pereira 2959-Pabst 3795 (1963) 

Serra de Ouro Branco, Minas Gerais, Brazil, US. 

Vellozia intermedia Seubert in Martius.-L. B. Smith 6854 

(1952) Serra do Cipo, Minas Gerais, Brazil, NY & US. 

Vellozia irwinii L. B. Smith.-Irwin, Maxwell & Wasshausen 

20998, Brazil, NY. 

Vellozia lanata Poh1.-Pires 9895 (1963) Parque do Gama, 

Federal District, Brazil, NY & US; Veralucia & Graziella 

s.n. (1968) Serra Dourado, Goiis, Brazil, NY. 

Vellozia lappa L. B. Smith & Ayensu.-L. B. Smith & Ayensu 

15988 (1972) Guinda, Diamantina, Minas Gerais, Brazil, US. 

Vellozia leptopetala Goethart & Henrard.-Maguire et al. 

49184 (1960) road between Diamantina and Gouveia, 

Minas Gerais, Brazil, NY & US. 

Vellozia lithophila Schu1tes.-R. E. Schultes & I. Cabrera 

18317 (1952) Rio Kubiyu, Cerro Kanenda, Vaupes, Colombia, US; J. Cuatrecasas 7700 (1939) San Jone del Guaviare, 

Rio Guaviare, Colombia, US. 

Vellozia machrisiana L. B. Smith.-Irwin, Grear, Souza & 

Reis dos Santos 12454 (1966) Chapada dos Veadeiros, 

Goias, Brazil, NY & US; Irwin 12697 (1966) Chapada dos 

Veadeiros, Goias, Brazil, NY: Prance & Silva 58275, Brazil, 

NY. 

Vellozia maculata Goethart & Henrard.-Glaziou 22218-a 

(1895) Cachoeiras da Vargem, Brazil, NY. 

Vellozia magdalenae L. B. Smith & Ayensu.-Lima & Brade 

13298 (1934) Alto do Desengano, 2000 m, Rio de Janeiro, 

Brazil, NY; Lima & Brade 14288 (1935) Serra da Furquilha, 

Rio de Janeiro, Brazil, NY. 

Vellozia metzgerae L. B. Smith.-W. Egler sen. (1947) 

Diamantina, Minas Gerais, Brazil, NY. 

Vellozia minima Poh1.-Duarte 2379 (1945) Serro do Cipo, 

Conceicao 1250 m, Brazil, US; Maguire et al. 49099 (1960) 

Serra de Cipo, 1200-1300 m, Minas Gerais, Brazil, US; H. S. 

Irwin, H. Maxwell & D. C. Wasshausen 20630 (1968) Serro 

do Cip6, Minas Gerais, 1500 m, Brazil, US. 

Vellozia modesta L. B. Smith & Ayensu.-L. B. Smith, Ayensu 

& Hatschbach 15997 (1972) Serra de Espinhaco, Diamantina, Minas Gerais, Brazil, US. 

Vellozia nuda L. B. Smith & Ayensu.-Hatschbach & Pelanda 

28029, Brazil, NY. 

Vellozia ornata Martius ex Schultes f.-L. B. Smith, Ayensu 

& Hatschbach 16001 (1972) 33 km NE of Diamantina, 

Minas Gerais, Brazil, US. 

Vellozia phalocarpa Poh1.-Anderson et al. 35855 (1972) Serra 

do Espinhaco, Minas Gerais, Brazil, NY & US. 

Vellozia pilosa Goethart & Henrard.-Glaziou 19933, (1892) 

Diamantina, Minas Gerais, Brazil, NY. 

Vellozia piresiana L. B. Smith.-Duarte 2068, Brazil, NY. 

Vellozia plicata Martius.-Porto 29572, Brazil, NY; FrazLo 

s.n., Brazil, NY; Duarte 3635-Gomes 420 (1953) Minas 

Gerais, Brazil, US: Lima 192, Brazil, NY; Brade 15947, 

Brazil, NY. 

Vellozia pumila Goethart & Henrard.-Glaziou 22215 (1894) 

Goias, Brazil, NY; H. S. Irwin, R. Souza, & R. Reis dos 

Santos 9428 (1965) ca. 12 km NW of Veadeiros, 1200 m, 

Goias, Brazil, NY & US; H. S. Irwin, R. Souza, R. Reis dos 

Santos 10922 (1963) Serra dos Pirineus, 1100 m, Goias, 

Brazil, NY & US; H. S. Irwin et al. 9230, Brazil, NY & US. 

Vellozia punctulata Seubert in Martius.-Blanchet 2561, 

Serra Gardina, Brazil, NY. 

Vellozia resinosa Martius ex Schultes f.-B. & C. K. Maguire 

& J. M. Pires 44694 (1959) Serra Cipo, 3800 ft, Minas 

Gerais, Brazil, NY & US. 

Vellozia riedeliana Goethart & Henrard.-L. B. Smith k 

Ayensu 15983, (1972) 27 km W of Serro, Minas Gerais, 

Brazil, NY. 

Vellozia sellovii Seubert in Martius.-Pereira 2677-Pabst 3513 

(1957) Serra da Piedade, Minas Gerais, Brazil, US. 

Vellozia squalida Martius ex Seubert f.-W. A. Archer & M. 

Barreto 4937 (1936) Serra do Cipo, Minas Gerais, Brazil, 

US; Duarte 6401 (1961) Serra do Cipo, Minas Gerais, 

Brazil, NY. 

Vellozia swallenii L. B. Smith.-J. M. Pires, N. T. Silva, R. 

Souza 9712 (1936) Brasilia, Brazil, US; J. M. Pires, N. T. 

Silva, R. Souza 9655 (1963) Brasilia, Brazil, US; G. Fonjecs 

11, Brazil, NY. 

Vellozia 'taxifoh Martius ex Schultes f.-Ynes Mexia 4289, 

Brazil, US; Pereira 1290 (1955) Sta. Maria Madalena, 

Pedra Dubois, 1200 m, Rio de Janeiro, Brazil, NY. 

Vellozia tenella Martius ex Schultes f.-Schwacke 6470, 6749, 

Brazil, NY; Pereira 2549-Pabst 3385 (1957) Caraca, Minas 

Gerais, Brazil, US; Damazio 91192, Brazil, NY; Damazio 

91185, Brad, NY. 

Vellozia teres L. B. Smith & Ayensu.-Irwin, Maxwell k 

TVasshausen 21000, Brazil, NY. 

Vellozia tragacantha Martius ex Schultes f.-Claussen 13, 

Brazil, NY; Claussen 1794, Brazil, NY; Irwin 2495 (1959) 

72 km SLY of Diamantina, Minas Gerais, Brazil, US. 

Vellozia triquetra Poh1.-G. Gardner 2735, Formigas, near 

Minas Nopas, Brazil, US. 

Vellozia tubiflora A. Richard.-Maguire et al. 30835, Amazonas, Guayana, NY; Maguire et al. 29159, Amazonas, 

Guayana, NY; Maguire et al. 30941, Amazonas, Guayana, 

NY; Maguire et al. 29245, Amazonas, Guayana, NY; Maguire et al. 30955, Amazonas, Guayana, NY; Maguire et al. 

30703 Amazonas, Guayana, NY; Maguire et al. 31003, 

Amazonas, Guayana, NY; Maguire et al. 31679, Amazonas, 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Guayana, NY; Maguire et al. 31777, Amazonas, Guayana, 

NY; Maguire et al. 32307, Amazonas, Guayana, NY; Maguire et al. 33532, Amazonas, Guayana, NY; Maguire et al. 

35131, Amazonas, Guayana, NY; Maguire et al. 35136, 

Amazonas, Guayana, NY; Maguire et al. 35248, Amazonas, 

Guayana, KY; Maguire et al. 36051, Amazonas, Guayana, 

NY; Maguire et al. 43855, Amazonas, Guayana, NY; Maguire et al. 45918, Amazonas, Guayana, KY; E. Foldats 

3526 (1960) Cerca del Puerto Ayacucho, Terr. Fed. Amazonas, Venezuela, US; L. Williams 13459 (1940) Pto. 

Ayacucho, Terr. Amazonas, 150 m, Venezuela, US; L. 

it’illiams 12974 (1940) Pto. Ayacucho, Terr. Fed. Amazonas, 

100 m, Venezuela, US; L. Williams 16022 (1942) Sanariapo, 

150 m, Venezuela, US; J. A. Steyermark & L. Aristeguieta 

54 (1962) NE of Luepa, 1200 m, Venezuela, NY & US; 

J. A. Steyermark & S. Nilsson 554 (1960) NE of Luepa, 

1200 m, Venezuela, NY & US; J. A. Steyermark 60326 

(1944) forest of Rio Karuai, 1220 m, Bolivar, Venezuela, 

US; Tainayo 3105 (1946) Gran Sabana, Bolivar, Venezuela, 

US; G. H. H. Tate 1196 (1938) Mount Auyan-tepui, 1100 

m, Venezuela, US; T. Lasser 1777 (1946) Guayana venezalana, Venezuela, US; J. A. Steyermark 93208 (1964) 

Auyan-tepui, Bolivar, Venezuela, US. 

Vellozia variabilis Martius ex Schultes f.-B. Maguire, G. M. 

Magalhaes & C. K. Maguire 49262 (1960) Serro Grao 

Mogul, 900-1100 m, Minas Gerais, Brazil, US; H. S. Irwin, 

R. Souza, & R. Reis dos Santos 9809 (1965) Serra dos 

Cristais, Goias, 1125 m, Brazil; L. B. Smith & Ayensu 15981 

(1972) 23 km from Serro, Minas Gerais, Brazil, US; H. S. 

Irwin, R. Souza & R. Reis dos Santos 9915 (1965) Serra dos 

Cristais, Goias, 1250 m, Brazil, US; H. S. Irwin, R. Souza Fi 

R. Reis dos Santos 11709 (1966) Serra Dourado, Goias, 800 

m, Brazil, US; H. S. Irwin, R. Souza 8s R. Reis dos Santos 

9726 (1965) Serra dos Cristais, Goias, 1175 m, Brazil, US: 

E. Pereira 2857-Pabst 3693 (1957) Minas Gerais, Brazil, US; 

Brade 13893, Brazil, NY; Lima 58-3003, Brazil, NY; Irwin 

et al. 9867, Brazil, NY. 

Vellozia verruculosa Martius ex Schultes f.-L. 0. Williams 

8112 (1943) Casa Branca, 1200 m, Minas Gerais, Brazil, US. 

Vellozia virgata Goethart & Henrard.-Schwacke 5889, Brazil, 

SY; Porto 543 (1915) Minas Gerais, Brazil, NY. 

Vellozia wasshausenii L. B. Smith.-Irwin et al. 20318, Brazil, 

NY. 

Barbacenia blackii L. B. Smith.-Macedo 2962 (1951) Cipo, 

1600 m, Minas Gerais, Brazil, US. 

Barbacenia celiae Maguire.-Maguire et al. 40025, Guayana, 

NY; Maguire et al. 40154, Guayana, NY; Maguire et al. 

40298, Guayana, NY. 

Barbacenia conicostigma Goethart & Henrard.-L. B. Smith, 

Ayensu & Hatschbach 15982 (1972) 27 km W of Serra, 

Datas, Minas Gerais, Brazil, US. 

Barbacenia delicatula, L. B. Smith & Ayensu.-L. B. Smith, 

Ayensu & Hatschbach 15975 [1972] Conceicao do Mato 

Dentro, Rio Santo Antonio, Minas Gerais, Brazil, US. 

Barbaceniu exscapa Martius.-Anderson et al. 35831 (1972) 

Serra do Espinchaco, Minas Gerais, Brazil, NY. 

Barbacenia flava Martius ex Schultes f.-L. B. Smith 6856 

(1959) Serra do Cipo, 1200 m, Minas Gerais, Brazil, US; 

B. Maguire, G. M. Magalhaes, C. K. Maguire 49116 (1960) 

20 km from Concepcao on road to Diamantina, Minas 

Gerais, Brazil, US; L. B. Smith 6695 (1952) Serra do Cip6, 

1100 m, Minas Gerais, Brazil, US; R. M. & A. F. Tryon 

6836 (1965) Serra do Cipo, 1120 m, Minas Gerais, Brazil, 

US; B. hlaguire, G. M. Magalhaes & C. K. Maguire 49011 

(1960) Serra do Cipo, 1200 m, Minas Gerais, Brazil, US; 

A. Macedo 3747 (1952) Serra do Cipo, Minas Gerais, 

Brazil, US; B. & C. K. Maguire & J. M. Pires 44691 (1959) 

Serra Cipo, 3700 ft, Minas Gerais, Brazil, US; SegadasVianna & Loredo 110 (1953) Serra do Cipo, Minas Gerais, 

Brazil, US. 

Barbacenia fragrans Goethart & Henrard.-Dueke s.n. (1929), 

Brazil, NY; Widgren 1267 (1845) Minas Gerais, Brazil, 

US; A. F. Regnell 1239 (1867) Minas Gerais, Brazil, US. 

Barbacenia gardneri Seubert in Martius.-Pereira 2849-Pabst 

3685 (1957) Minas Gerais, Brazil, US. 

Barbacenia gaveensis Goethart & Henrard.-L. B. Smith 6445 

(1954) Pedra da Gavea, Federal District, 850 m, Brazil, US. 

Barbacenia gentianoides Goethart & Henrard.-SegadasVianna & Loredo 1066 (1953) Serra do Cip6, Minas Gerais, 

Brazil, US. 

Barbacenia gounelleana Beauverd.-L. Emygdio 1428 (1957) 

Itatisia, Rio de Janeiro, Brazil, US; E. Pereira 7050 (1962) 

Itatisia, Rio de Janeiro, Brazil, US. 

Barbacenia hatschbachii L. B. Smith & Ayensu.-Hatschbach 

24284, Brazil, NY. 

Barbncenia ignea Martius ex Schultes f.-L. 0. Williams & 

V. Assis 6817 [1945] Serro de Monjolo, Minas Gerais, 

Brazil, US. 

Barbacenia involucrata L. B. Smith.-Irwin et al. 20973, 

Brazil, KY. 

Barbacenia irwiniana L. B. Smith.-Porto 1140, Brazil, NY. 

Barbacenia longifbra Martius.-H. S. Irwin 2486 (1959) 10 

km N of Guida, Minas Gerais, Brazil, US; W. A. Archer 

4089 (1936) between Diamantina and Guinda, Minas 

Gerais, Brazil, US. 

Barbacenia longiscapa Goethart & Henrard.-Irwin et al. 

22186 (1969) Serra do EspinhaGo, 1370 m, Minas Gerais, 

Brazil, NY. 

Barbacenia luzulifolia Martius ex Schultes f.-Pereira 2664- 

Pabst 3500 (1957) Serra da Piedade, 1800 m, Minas Gerais, 

Brazil, NY. 

Barbacenia macran tha Lem.-Irwin, Maxwell & Wasshausen 

20071, Brazil, NY. 

Barbacenia nana L. B. Smith & Ayensu.-Smith, Ayensu & 

Hatschbach 15973 (1972) Conceicao do Mato Dentro, Rio

Santo Antonio, Minas Gerais, Brazil, US. 

Barbacenia paranuensis L. B. Smith.-G. Hatschbach 15715 

(1967) Serra dos Mulatos, Brazil, US; G. Hatschbach 29212, 

Brazil, US. 

Barbacenia purpurea Hook.-A. L. Schott 151098, Brazil, US. 

Barbacenia riedeliana Goethart & Henrard.-Anderson et al. 

35123 (1972) Serra do Espinhaco, Minas Gerais, Brazil, 

NY; G. Hatschbach & P. Pelanda 27806 (1971) Serra do 

Espinhaco, Gouveia, Minas Gerais, Brazil, US. 

Barbacenia rubro-virens Martius.-B. Maguire et al. 49177 

(1960) between Diamantina and Gouveia, Minas Gerais, 

Brazil, US; B. Maguire et al. 49260 (1960) Serra Grao 

Mogul, 900-1100 m, Minas Gerais, Brazil, US. 
NUMBER 15 5 

Barbacenia schwachei Goethart & Henrard.-E. P. Heringer 

5263 (1956) Serra do Cipo, Minas Gerais, Brazil, US. 

Barbacenia sellouii Goethart & Henrard.-Menezes & Continho 5, Brazil, NY. 

Barbacenia seubertiana Goethart & Henrard.-J. P. Lanna 

Sobro 4488 (1956) Guanabera, Tijuca, Rio de Janeiro, 

Brazil, US; Strang 239, Brazil, NY; Strang 523, Brazil, NY; 

Goethart & Henrard 4332, Brazil, NY. 

Barbacenia stenophylla Goethart & Henrard.-Irwin, Souza 

& Reis dos Santos 9901 (1965) 6 km S of Cristalina, 1175 

m, Serra dos Cristais, Goias, Brazil, US. 

Barbacenia tomentosa Martius.-H. S. Irwin, R. M. Harley 

& E. Onishi 29559 (1971) Serra do Espinhaco,' Minas 

Gerais, 1650 m, Brazil, US; N. L. Menezes 1, Jardim 

Botanic0 do Rio de Janeiro, Brazil, NY. 

Barbacenia uandellii Pohl ex Seubert in Martius.-Gardner 

5224, hlinas Gerais, Brazil, US. 

Barbacenia uiscosissima Goethart & Henrard.-A. Macedo 

2826 (1950) 1500 m, Minas Gerais, Brazil, US. 

Barbacenia williamsii L. B. Smith.-Pereira 2460-Pabst 3296 

(1957) Serro do Curral, Minas Gerais, Brazil, US; L. W. 

IYilliams & V. Assis 6386 (1945) Serra do Curral, 1300 m, 

Minas Gerais, Brazil, US; Dueke s.n. (1929) Brazil, NY. 

METHODS.-Apart from the few pickled materials, 

most of the leaf fragments upon which this study 

was based were obtained from herbarium specimens. The specimens were soaked in a mixture of 

Aerosol OT solution and distilled water (cf. 

Ayensu, 1967) until maximum expansion was attained. After a quick wash in tap water, free hand 

sections were prepared. Microscopic details were 

studied in glycerine mounted unstained sections as 

well as sections stained in safranin and Delafield's 

Haematoxylin. Line drawings and photomicrographs were prepared, using a Wild M20 microscope with camera lucida and photographic 

attachments. The scanning electron micrographs 

were obtained with a Cambridge Steroscan Mark 

2A. The specimens were mounted on cover glass, 

and the stub holding the cover glass was rotated 

under pressure in high vacuum and coated with 

gold plating approximately 200-400 angstroms 

thick. Maximum pictorial effect was obtained by 

tilting the samples at a 45" angle. 

lowing characters were found to be significant in 

the diagnosis of the species examined. (a) The 

epidermal cells are generally somewhat rectangular. 

Others are almost square shaped. (b) The stomata 

are generally paracytic and tend to be in longitudinal rows. They occur mostly on the abaxial surface, 

and in the furrows when these are present. (c) 

The cuticle is thin layered but often it is thicker 

CHARACTERS USED IN THE DESCRIPTIONS.-The f01- 

around stomata and on the abaxial epidermis. (d) 

Trichomes occur in the form of hairs which may 

be unicellular or multicellular: often they appear 

in tufts. The kinds of hair may differ in the same 

species either on both surfaces or on the same surface. Protrusions in furrows may be in the form 

of hairs or epidermal extensions. The taxanomic 

importance of hairs in Velloziaceae must be 

assessed very critically since changes in the kinds 

of hairs occur with age. (e) The vascular bundles 

are of fairly equal size in each species except those 

at the margins and in the midvein that tend to 

be small. The number of bundles varies from 

species to species. It is important to note that leaf 

sections are always taken at the same level in each 

case (Lea, from the midportion). (f) The number 

of phloem units lying along the arms of the abaxial 

U- or Y-shaped sclerenchyma girder is usually two 

in a mature leaf. In juvenile leaves the phloem 

units are often united. (9) The vascular bundles 

are surrounded by distinct bundle sheaths which 

extend to the abaxial and adaxial epidermis in the 

case of Barbacenia species or the sheath is connected to the translucent layers radially arranged 

adaxially on vascular bundles of Vellozia species. 

(h) The central and marginal bundles are associated with distinct sclerenchyma. (k) Commissural 

bundles are often present. The absence of them 

in any particular species does not necessarily mean 

that they are totally lacking within the leaf, for 

this may merely reflect their absence at the midportion of the leaf from which sections are taken 

for examination. 

The descriptive terms which I have employed 

are essentially those published by Metcalfe and 

Gregory (1964), Ayensu (1969a), and Metcalfe 

(1971). In these publications several diagrammatic 

representations of sclerenchyma patterns in leaves 

of Cyperaceae and Velloziaceae were given. 

of the American species of Vellozia by Smith 

(1962), the species were placed into two sections 

on the basis of the following characters: 

Tepals free or obscurely short-connate above the ovary .......... 

............................................................... Section I: Vellozia 

Tepals forming a distinct tube above the ovary ........................ 

.......................................................... Section 11: Radia 

The species of Barbacenia were classified further 

as follows: 

CLASSIFICATION OF THE SPECIES.-h the Synopsis 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a b I I 

e t 

FIGURE 1.-Diagrams showing the presence of paracytic type of stomata on adaxial and abaxial 

surfaces of leaf epidermis: a, Barbacenia gounelleana, adaxial surface; b, B. gounelleana, abaxial 

surface (Pereira 7050) ; c, B. flaua, adaxial surface; d, B. paua, abaxial surface (Maguire 49116); 

e, B. vandeilii, adaxial surface; f, B. vandellii, abaxial surface (Gardner 5224) ; g, B. conicostigma, 

adaxial surface; h, B. conicostigma, abaxial surface (L. B. Smith and Ayensu 15982); i, B. 

stenophylla, adaxial surface; j, B. stenophylla, abaxial surface (Irwin et al. 9901); k, B. 

involucrata, adaxial surface; 1, l3. invoiucrata, abaxial surface (Irwin et al. 20973); m, B. rubrouirens, adaxial surface; n, B. rubro-uirens, abaxial surface (Maguire 49177) ; 0, B. luzulifolia, 

adaxial surface; p, B. luzulifolia, abaxial surface (C Pereira 2664-Pabst 3500). 

L 0 P 
NUMBER 15 7 

- C 

FIGURE 2.-Diagrams showing type and distribution of hairs and stomata on epidermis: a, VelZozia bicolor (Maguire 49047) ; b, V. lithophila (Schultes and Cabrera 18317); c, V. glandulifem 

(L. B. Smith & Ayensu 15998) ; d, Rarbacenia fragrans (Dueke s.n.) ; e, B. nana (L. B. Smith 

and Ayensu 15973); f, B. macrantha (Irwin et al. 20017). 

1. Perianth-tube not more than twice as long as the ovary 

2. Ovary section of the perianth partially or wholly 

glaborous, the costae evident (lacking in B. beauverdii) ..................................................................... Subkey I 

of a new genus, Aylthonja. The characters upon 

which she based her judgment agree with those 

of the section Radia of Vellozia established by 

2. Ovary section of .the perianth wholly vestite ............... Smith Furthermore there to be Some 

1. Perianth-tube more than twice as long as the ovary ...... ‘‘corona.” In his description of VelEozia flower, 

............................................................................. subkey 11 confusion regarding the meaning of the term 

................................................................................... Subkey 111 

Menezes (1971b) recently proposed new combinations in Velloziaceae, including the recognition 

Maguire (1969) used the term “stamina1 corona” 

to refer to the structure between the ovary and 

the anthers. Menezes on the other hand considered 
a SMITHSONIAN CONTRIBUTIONS TO BOTANY 

ae f 

I I 

m n 

0 P 

FIGURE 3.-Diagrams showing stomata, types of epidermal cells and hairs: a, Vellozia virigata, 

adaxial surface; b, V. virgata, abaxial surface (Porto 543); c, V. cinerascens, adaxial surface; 

d, V. cinerascens, abaxial surface (Lehnkner 323); e, V. glauca, adaxial surface; f, V. glauca, 

abaxial surface (Maguire et al. 44793) ; g, V. dawsonii, adaxial surface; h, 6'. dawsonii, abaxial 

surface (Irwin et al. 12622); i, V. grisea, adaxial surface; j, V. grisea, abaxial surface (Weddell 

3006) ; k, V. flavicans, adaxial surface; 1, V. flavicans, abaxial surface (Andrade 356-Emmerich 

348); m, V. cachirnbensis, adaxial surface; n, V. cachimbensis, abaxial surface (Pires et al. 

6111); 0, V. rnaculata, adaxial surface; p. V. rnaculata, abaxial surface (Glaziou 222-18A). 
NUMBER 15 9 

b e 

C f 

FIGURE 4.-Diagrams showing variation in hair types on epidermis: a, Vellozia punctata 

(Blanchet 2561); b, V. grisea (Weddell 3006); c, V. nuda (Hatschbach & Pelanda 2829); 

d, V. hirsuta (Hatschbach 2803); e, V. riedeliana (L. B. Smith and Ayensu 15983); f, V. plicata 

(Duarte 3635). 

the same structure as a tissue of the perianth which For the present I have subdivided the species 

mimicks dilated filaments. The study of Noher de of the New World members according to Smith’s 

Halac and Cococci (1971 ) stressed the stamina1 classification. The descriptions of the species are 

origin of this “corona” with the lateral lobes being arranged alphabetically within each section and 

essentially staminodes. These differences of opinion subkey. 

will naturally have to await detailed studies of ACKNOWLEDCMENTS.-~ wish to dedicate this confloral morphology in the family. tribution to Dr. Lyman B. Smith, Senior Botanist, 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Smithsonian Institution, and the foremost authority on the taxonomy of Velloziaceae, on the 

occasion of his seventieth birthday. 

I am most grateful to Ms. Cynthia Bailey, Research Assistant, for her continuous assistance in 

the preparation of the manuscript. Her interest and 

enthusiasm are much appreciated. Mr. Eduardo 

Barros was very helpful in the preparation of the 

anatomical slides upon which this work was based. 

hls. Diane Robertson prepared the camera lucida 

drawings for which I am most grateful. My thanks 

are also extended to the Scanning Electron Microscope staff of the National Museum of Natural 

History for helping with the production of the 

SEM photos. I wish to thank Dr. Basset Maguire 

for making available to me several leaf samples 

from his collections and for his continuous interest 

in my work. I also wish to thank Dr. Thomas R. 

Soderstrom for reading portions of the manuscript 

and for making helpful comments. This study was 

supported in part by the Smithsonian Research 

Foundation. 

Vellozia 

Section I: Vellozia 

Vellozia abietina Martius 

FIGURE 5a-b: PLATE 24a-b 

SPECIMENS EXAMINED.-Maguire et al. 491 15. 

SURFACE VIEw.-Hairs: absent. Epidermis: 

adaxial and abaxial cells rectangular, thin walled. 

Stomata: tetracytic, 18 X 12 pm; observed on both 

surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

thickly V-shaped with significant median adaxial 

groove. Adaxial and abaxial surfaces evenly undulating. Epidermis: cells more or less square, 

rounded, or rectangular; cells thin walled with 

thicker inner tangential walls; few individual 

thick-walled fibers observed subjacent to adaxial 

epidermis. Cuticle: uniformly thickened, slightly 

thinner on cells forming median adaxial groove. 

Stomata: walls of guard cells generally thickened; 

slight development of inner ledges; substomatal 

chamber present. Mesophyll: palisade cells almost 

nonexistent except for one to three layers of cells 

on either side of adaxial median groove; spongy 

tissue making up nearly all of mesophyll. Vascular 

bundles: 12; commissural bundles not observed. 

Large veins each with a single wide vessel. Two 

phloem units lying laterally in short flanges of 

abaxial girder. Each vascular bundle always accompanied by partial adaxial girder or cap and 

abaxial girder. Vascular bundle completely surrounded by a distinct bundle sheath with extensions of two to four cells toward adaxial epidermis 

only. Crystals and tannin: not observed. 

NOTE: Although this specimen has all the Vellozia distinguishing characters, there is a close 

resemblance of it to Barbacenia because the partial 

adaxial girder is almost becoming a full girder 

except for the adaxial bundle sheath extensions. 

The mesophyll is nearly all spongy. 

Velloziu alata L. B. Smith 

FIGURE 5c-e; PLATE 24c-e 

SPECIMENS EXAMINED.-Maguire et a]. 49049. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial 'and abaxial cells rectangular. Stomata: tetracytic, 18 X 24 pm; observed on both surfaces, 

especially on adaxial surface. 

TRANSVERSE SECTION OF LAMINA.-DorSiventral; 

thinly crescentiform with upturned margins and 

median abaxial groove. Adaxial surface evenly 

ridged, especially in areas between vascular 

bundles; abaxial surface conspicuously furrowed 

about one-fourth thickness of blade. Epidermis: adaxial and abaxial cells square or rectangular, walls 

uniformly thickened; short rows of thick-walled 

fibers found subjacent to epidermis. Cuticle: 

thick, conspicuously ridged along entire length of 

lamina on both surfaces. Stomata: slightly sunken, 

walls of guard cells thickened with short outer 

ledges; substantial chamber present. Most of abaxial stomata confined to furrows. Mesophyll: 

adaxial palisade four to six layered, occupying 

about half of mesophyll and changes abruptly into 

abaxial spongy tissue. Cells between veins unlike 

rest of mesophyll, thick walled, somewhat rounded, 

and conspicuously translucent, resembling bundle 

sheath extensions. Vascular bundles: 42; commissural bundles few. Large veins each with one to 

three wide vessels. Two phloem units lying laterally in flanges of abaxial Y-shaped girder. Each 

vascular bundle always accompanied by partial ad- 
NUMBER 15 11 

FIGURES 5-24 

Transverse sections of leaves of Velloziaceae showing the distribution, types 

of sclerenchyma girders and strands, as well as the arrangement of mesophyll 

tissue. The measuring unit for each species represents 100 pm. 

- 

FIGURE 5.-a-b, Vellozia abietina (Maguire et al. 49115) ; c-e, V. data (Maguire et al. 49049) 

axial girder or cap. Abaxial girders of vascular Crystals: not observed. Tannin: few in mesophyll. 

bundles on either side of midvein and at margins NOTE: The distinguishing features of this 

often partial. Vascular bundle completely surround- species are (a) the rather wide blade with over 

ed by a distinct bundle sheath with extensions of 40 veins; (b) the two to four wide translucent cells 

three to four cells toward adaxial epidermis only. running radially from the tip of the abaxial fur- 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

row to the adaxial epidermis; and (c) the raised 

corrugated midsection of the leaf. The furrows are 

conspicuously narrow, almost touching at the ends. 

Velloria aloifolia Martius 

FIGURE 1Oa-c; PLATE 29d-f 

SPECIMENS EXAMINED.-Y. Mexia 5870. 

SURFACE VIEtv.-Hairs: unicellular, fingerlike or 

club shaped. Present on abaxial surface only. Epidermis: adaxial cells uniformly cuboidal, abaxial 

cuboidal to rectangular and distinctly thick walled. 

Stomata: paracytic, 18 X 12 pm; observed on 

abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

shallowly corrugate, and with a distinct adaxial 

median groove. Adaxial surface evenly to unevenly 

ridged; abaxial surface conspicuously furrowed 

about one-third to half thickness of blade. Epidermis: two to three rows of adaxial cells square 

to rectangular of thin walls; abaxial cells consisting of fibrous sclerenchyma except where stomata 

occur. Cuticle: thick, undulating, but conspicuously ridged near adaxial and abaxial median region. 

Stomata: present in furrows; guard cells generally 

thickened; substomatal chamber inconspicuous; 

short strands of thick-walled fibers observed subjacent to epidermis. Mesophyll: adaxial palisade 

four to six layered, occupying slightly more than 

half of mesophyll and abruptly changing into 

spongy tissue; upper two to three rows of palisade 

and those arranged radially above vascular bundles, furrows, and above midvein distinctly translucent and large; rest of palisade cells compactly 

arranged and filled with chloroplast. Vascular 

bundles: 52 commissural bundles present. Large 

veins each with one to two wide vessels, mostly 

one, Two phloem units lying laterally in flanges 

of abaxial Y-shaped girder. Each vascular bundle 

always accompanied by adaxial cap. Vascular 

bundle completely surrounded by a distinct bundle 

sheath with short extensions of two to three cells 

toward abaxial epidermis. Crystals and tannin: not 

observed. 

NOTE: The fingerlike and club-shaped hairs in 

the abaxial furrows of this species recall a similar 

feature in the Old World species of Vellozia 

dasylirioides and its varieties (cf. Ayensu, 1969). 

The thick blade (15 mm), with over 50 veins, has 

a shallowly corrugate shape. 

Vellozia angustifolia Goethart 8c Henrard 

FIGURE 7d-f; PLATE 28a-e 

SPECIMENS EXAMINED.-W. A. Archer, 4092; 

Centro Pesq, (R)01411; Riedel s.n. 

SURFACE VIEW.-Hairs: absent. Epidermis: adaxial and abaxial cells mostly rectangular, thin 

walled. Stomata: tetracylic, 30 X 15 pm; observed 

on both surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

shallowly V-shaped with downturned margins and 

median adaxial groove. Adaxial surface evenly undulating; abaxial surface furrowed about half of 

blade thickness. Epidermis: cells more or less 

rounded or square. One or two rows of thickwalled fibers forming almost a continuous strand 

subjacent to epidermis. Below strand is a layer of 

large translucent cells separating it from mesophyll. 

Cuticle: thin on both surfaces. Stomata: in line 

with epidermis; mostly confined to abaxial surface, 

especially in fibrous walls; guard cells thickened 

with short inner and outer ledges; substomatal 

chamber present. Mesophyll: adaxial palisade two 

to four layered, occupying about half of mesophyll 

and gradually changing into spongy tissue; palisade 

cells arranged radially above vascular bundles furrows and above midvein distinctly translucent and 

large; rest of palisade compactly arranged and 

filled with chloroplast. Vascular bundles: 16-27; 

commissural bundles present. Large veins each 

with one to three wide vessels, mostly two. Two 

phloem units almost coalesing into one lying laterally in abaxial or slightly partial V-shaped girder. 

Each vascular bundle always accompanied by adaxial cap. Vascular bundle completely surrounded by 

a distinct bundle sheath with extensions toward 

adaxial epidermis and short extensions toward 

abaxial epidermis. Crystals and tannin: not observed. 

NOTE: The one row translucent rectangular 

cells separating the adaxial strands of thick-walled 

fibers from the palisade cells is very distinctive. 

Vellozia asperula Martius 

FIGURE 9d-f 

SPECIMENS EXAMINED.-G. Gardner 5228; Occhiouni s.n. 

SURFACE Vmv.-Hairs: absent. Epidermis: adaxial and abaxial cells rectangular, thin walled. Sto- 
NUMBER 15 13 

mata: Mainly paracytic, some tetracytic, 15 X 18 

pm; present mainly on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove. Adaxial surface unevenly ridged; abaxial surface conspicuously 

furrowed, slightly more than one-third thickness 

of blade. Epidermis: cells more or less rectangular, 

some triangular, others almost rounded; intermitent rows of thick-walled fibers observed subjacent to epidermis, especially toward leaf margin. 

Cuticle: uniformly thickened on adaxial and abaxial surfaces but not observed in furrows. Stomata: 

present in furrows; guard cells thick walled; substornatal chamber inconspicuous. Mesophyll: adaxial palisade four to six layered, occupying about 

one-third of mesophyll and gradually changing into 

spongy tissue; two to four rows of distinctly translucent cells arranged radially above vascular bundles and furrows; rest of palisade compactly 

arranged and filled with chloroplast. Vascular 

bundles: 24, commissural bundles not observed. 

Large veins each with one to two, occasionally 

three, wide vessels. Two phloem units lying laterally in flanges of partial abaxial U-shaped girder. 

Each vascular bundle always accompanied by adaxial cap. Vascular bundles- completely surrounded 

by a distinct thin-walled bundle sheath. Crystals 

and tannin: not observed. 

NOTE: The partial abaxial U-shaped girder recalls those of the Xerophyta type described for some 

of the Old World species (cf. Ayensu, 1969). 

Vellozia bicolor L. B. Smith 

FIGURE 2a: PLATE 42a 

SPECIMENS EXAMINED.-Maguire et al. 49047. 

SURFACE VIEW.-Hairs: long and narrow, mostly 

unicellular but bicellular ones observed. Present 

on abaxial surface only. Epidermis: adaxial cells 

uniformly rectangular; abaxial cells even more rectangular, thin walled. Stomata: mainly paracytic, 

few tetracytic, 24 X 18 pm; observed on abaxial 

surface only. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove. Adaxial surface unevenly undulating; abaxial surface furrowed, about one-fourth thickness of blade. 

Epidermis: adaxial and abaxial cells more or less 

rectangularly flattened, especially of adaxial cells, 

walls uniformly thin walled. Discrete strands of 

fibers occur subjacent to adaxial epidermis, often 

two strands opposite each adaxial furrow, occasionally three strands observed. Directly below 

fiber strands and subepidermal cells is a layer of 

large somewhat translucent parenchyma cells separating epidermal and subepidermal cells from 

rest of mesophyll. Cuticle: distinctly thickened on 

both surfaces. Stomata: confined to furrows, almost 

flush with epidermis. Mesophyll: adaxial palisade 

three to four layered, occupying about half of 

mesophyll, and abruptly changing into spongy tissue; two layers of palisade cells arranged radially 

above vascular bundles distinctly translucent and 

large; rest of palisade cells compactly arranged and 

filled with chloroplast. Vascular bundles: 46; commissural bundles few. Large veins each with one 

to three wide vessels, mostly one. Two phloem 

units lying laterally in flanges of abaxial Y-shaped 

girder. Each vascular bundle always accompanied 

by adaxial inverted V-shaped sclerenchyma cap, 

both in turn completely surrounded by a distinct 

bundle sheath. Crystals: rhomboidal, square, and 

diamond-shaped present in subepidermal cells. 

Tannin: not observed. 

NOTE: The almost woolly-like hairs and the 

adaxial fiber strands subjacent to the epidermis are 

very distinctive to this species. 

Vellozia brachypoda L. B. Smith & Ayensu 

PLATE 42b 

SPECIMENS EXAMINED.-hWin 23534. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

mostly square to rectangular on adaxial and abaxial surfaces; thin walled. Stomata: paracytic, 15 X 

9 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Adaxial surface ridged; abaxial surface slightly undulating; furrowed one-third to one-half thickness of 

blade. Epidermis: adaxial cells dome shaped to 

conical; abaxial cells rounded to dome shaped; walls 

uniformly thin on both surfaces. Layer of parenchyma cells subjacent to adaxial epidermis. Cuticle: 

slightly thickened and ridged on both surfaces. 

Stomata: present on abaxial surface; almost flush 

with epidermis; substomatal chamber present. 

Mesophyll: three- to four-layered palisade with 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

b 

FIGURE 6.-a-b, Vellozia tragacantha (Irwin 2495) ; c-e, V. granulata (Riedel s.n.) . 

tissue grading into compactly arranged spongy tissue. Few rectangular translucent palisade cells lar bundle. Crystals and tannins: none observed. 

arranged radially above vascular bundles and furrows. Vascular bundles: 27; commissural bundles 

not observed. Veins with one to two large vessels, 

mostly one. Two phloem units lying laterally in 

flanges of Y-shaped abaxial girder. Adaxial inverted 

sclerenchyma cap present on each vascular bundle. 

Bundle sheath completely surrounding each vascuVellozia burle-marxii L. B. Smith 

FIGURE 16a-c 

SPECIMENS EXAMINED.-R. Burle-marx, s.n.; Irwin 

et al. 32520. 
NUMBER 15 15 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial and abaxial cells, rectangular, thin walled. 

Stomata: paracytic, 24 X 15 pm; observed on abaxial surface only. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove. Adaxial surface conspicuously ridged and more or less slightly 

furrowed; abaxial surface distinctly furrowed. Epidermis: cells more or less rectangular; thin walled. 

Cuticle: uniformly thin, slightly thicker on abaxial 

side of median groove. Stomata: present in abaxial 

furrows only; slightly sunken; somewhat protected 

by epidermal protuberances in furrows; substomatal chamber present. Mesophyll: adaxial palisade 

three to four layered, occupying half of mesophyll 

and abruptly changing into spongy tissue; palisade 

cells radially arranged above vascular bundles, 

translucent and slightly larger than rest of cells; 

other palisade cells compactly arranged and filled 

with chloroplast. Vascular bundles: of two types, 

major and minor veins; major veins 19; minor 

veins 18. Major veins each with one to two wide 

vessels, mostly one; two phloem units lying laterally in flanges of abaxial Y-shaped girder; each 

vascular bundle always accompanied by adaxial 

cap; vascular bundle completely surrounded by a 

distinct bundle sheath. Minor veins observed at 

adaxial side of abaxial furrows; both adaxial and 

abaxial sides of vascular bundle accompanied by 

sclerenchyma caps. Minor veins also surrounded 

by a distinct bundle sheath. Crystals and tannin: 

not observed. 

NOTE: The presence of major and minor veins 

in this species coupled with the bifurcations of the 

abaxial surfaces, especially with respect to the lateral epidermal projections, recalls the same conditions in Vellozia hemisphaerica. 

Vellozia caruncularis Martius 8c Seubert 

FIGURE 14a-c:; PLATES 18b, 34b-d 

SPECIMENS EXAMINED.-Maguire et al. 49013; 

Pires et al. 2721; L. B. Smith 6855; Heringer 5265; 

Macedo 2959. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial and abaxial cells mostly rectangular, some 

square cells as observed on main veins; thick 

walled. Stomata: paracytic, 21 X 12 pm; observed 

on furrows of abaxial surface only. 

TRANSVERSE SECTION OF LAMINA.-DOrSiVentral; 

inversely W-shaped with slight median adaxial 

groove. Adaxial surface slightly undulating; abaxial 

surface distinctly furrowed. Epidermis: cells more 

or less rectangular; abaxial cells much smaller than 

adaxial ones; short strands of fibers observed subjacent to epidermis; tangentially oriented translucent parenchyma cells separating the palisade 

cells from subepidermal cells. Cuticle: uniformly 

thin but significantly thicker in the midrib region. 

Stomata: present on both the abaxial surface and 

furrows; almost flush with epidermis; those in 

furrows are somewhat protected by epidermal 

protuberances; substomatal chamber present. Mesophyll: adaxial palisade three to four layered, 

occupying about half of mesophyll and abruptly 

changing into spongy tissue; two to three large 

radially oriented translucent cells arranged adaxially on vascular bundles and inner limits of furrows connecting three conspicuous parenchyma 

cells. Palisade cells compactly arranged and filled 

with chloroplast. Vascular bundles: 17-20; commissural bundles few. Large veins with one to 

three large vessels, mostly one. Two phloem units 

lying laterally in flanges of abaxial Y-shaped girder. 

Each vascular bundle always accompanied by adaxial inverted V-shaped sclerenchyma cap; both in 

turn completely surrounded by a distinct bundle 

sheath. Crystals: not observed. Tannin: present in 

mesophyll. 

NOTE: The five collections studied show a remarkable uniformity in leaf shape as well as in 

histology. This is indicative of the stability of this 

species. 

Vellozia ciliata L. B. Smith 

PLATES 5c, 36b, 49a-c 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

15995. 

SURFACE VIEw.-Hairs: small papillae on adaxial 

surface. Epidermis: adaxial cells square to rectangular; thin walled. Stomata: paracytic, 18 X 12 pm; 

present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Adaxial surface almost smooth; abaxial surface furrowed 

about one-third thickness of blade. Epidermis: 

adaxial cells square; few papillae observed on 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

adaxial surface. Abaxial epidermis mostly composed 

of sclerenchyma strands with few dome-shaped 

cells. Subjacent to adaxial epidermis are one to 

two rows sclerenchyma strands interspersed with 

parenchyma cells. Below sclerenchyma strands is 

distinct layer of parenchyma cells. Cuticle: very 

thin on adaxial and abaxial surfaces. Stomata: 

present in abaxial furrows; small substomatal 

chamber present. Somewhat protected by small 

papillae present in furrows. Mesophyll: five to six 

layers distinct palisade tissue changing abruptly 

into compact spongy tissue. Three to five layers 

large translucent palisade cells arranged radially 

above vascular bundle and furrows. Three to four 

layers large translucent palisade cells arranged 

radially above midvein; no small palisade tissue 

here. Vascular bundle: 37; few commissural bundles 

present. One large vessel present in each vascular 

bundle. Two phloem units laterally arranged in 

flanges of Y-shaped abaxial girder. Adaxial cap 

present on each vascular bundle. Abaxial sclerenchyma strands only present on epidermal layer. 

Bundle sheath completely surrounding vascular 

bundle. Crystals and tannins: none observed. 

Vellozia cinerascens Martius ex Schultes f. 

FIGURE 3c-d; PLATES 4c-d, 42c 

SPECIMENS ExAMmm.-Lehnkner 323. 

SURFACE VIEw.-Hairs: present in clusters. Epidermis: adaxial cells rectangular; abaxial cells 

mostly square, few rounded or rectangular; cell 

walls thin on both surfaces. Stomata: paracytic, 

18 X 6 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

flanged V-shaped. Adaxial surface deeply ridged; 

abaxial surface furrowed about one-third the thickness of blade. Epidermis: adaxial cells dome shaped 

to conical; interspersed with large strands of sclerenchyma positioned above vascular bundles and 

furrows. Abaxial cells rounded to dome shaped, 

thin walled; inner to cells lined with rectangular 

strands of sclerenchyma, leaving about one-half 

of furrows uncovered. Cuticle: thin and ridged on 

both adaxial and abaxial surfaces. Stomata: present 

in abaxial furrows and spaces between sclerenchyma strands; substomatal chamber present. Mesophyll: two to three layers palisade, grading into 

loosely compact spongy tissue. Translucent palisade 

radially arranged above vascular bundles. Vascular 

bundle: 39; commissural bundles not observed. 

One to two large vessels present in vascular 

bundles. Two phloem units laterally arranged in 

flanges of U-shaped abaxial girder. Each bundle 

always accompanied by an adaxial cap and an 

abaxial U-shaped sclerenchyma vascular bundle 

completely surrounded by a bundle sheath. Crystals and tannins: none observed. 

NOTE: The arrangement and disposition of 

adaxial and abaxial sclerenchyma strands are very 

distinctive for this species. 

Vellozia compacta Martius ex Schultes f. 

FIGURE 7a-c; PLATE 25b-f 

SPECIMEN EXAMINED.-hWin 2407; Maguire et al. 

49132; Segades-Vianna & Loredo 1085; L. B. Smith 

7005. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial and abaxial cells mostly rectangular but often 

square shaped, thin walled. Stomata: tetracytic, 24 

X 15 pm; present on both surfaces but highly concentrated on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

shallowly V-shaped with slight median adaxial 

groove. Adaxial surface undulating; abaxial surface furrowed abofit half blade thickness. Epidermis: cells more or less rounded, abaxial cells square 

shaped. Subjacent to adaxial epidermis occur one 

to three layers of parenchyma cells accentuated by 

short strands of sclerenchyma. Below sclerenchyma 

strands and subjacent parenchyma cells are one to 

two layers of large translucent cells separating it 

from mesophyll. Cuticle: thickened on both surfaces. Stomata: mostly confined to furrows in 

abaxial surface; substomatal chamber inconspicuous. Mesophyll: adaxial palisade three to four 

layered, occupying about one-third of mesophyll 

and gradually changing into spongy tissue; three 

to four rows of large translucent cells arranged 

radially above vascular bundles and furrows. Palisade cells and spongy mesophyll compactly arranged and filled with chloroplast. Vascular 

bundles: 49-59; commissural bundles present. Large 

veins each with three or more wide vessels. Two 

phloem units lying laterally in flanges of abaxial 

Y-shaped girder. Each vascular bundle always accompanied by adaxial inverted V-shaped scleren- 
NUMBER 15 17 

a 

FIGURE 7.-a-c, 

. 

Vellozia compacta (Irwin 2407); d-f, V. anguotifolia (Riedel s.n.) . 

chyma cap; both in turn completely surrounded Vellozia costata L. B. Smith & Ayensu 

by a distinct bundle sheath. Crystals not observed. 

Tannin: few in mesophyll. PLATE 36c 

-. NOTE: There are clear variations in the scleren- SPECIMEN EXAMINED.-Irwin et al. 20999. 

chyma patterns in the species examined. The SURFACE VIEw.---Hairs: absent. Epidermis: adaxadaxial sclerenchyma shapes range from strictly ial and abaxial cells rectangular, thick walled; occacap shaped to partial girder. The same is true of sional rows of square cells observed. Stomata: 

abaxial sclerenchyma where some are distinctly paracytic, 30 X 9 pm; present mainly on abaxial 

Y-shaped and others are somewhat partial. surface, few on adaxial side. 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TRANSVERSE SECTION OF LAMINA.-Dorsiventra1; 

V-shaped with slight median adaxial groove. Adaxial surface undulated, especially that of the right 

flange of V. Abaxial surface with deep furrows in 

which stomata are present. Epidermis: consisting 

of very thick outer walls except where stomata 

occur. Cuticle: uniformly thickened, slightly thicker 

around stomata. Almost continuous row of fibrous 

sclerenchyma occur immediately below the abaxial 

epidermis. Discrete clusters of fibers observed subjacent to adaxial epidermis. A single row of large 

cells of unequal shape, containing tanninferous 

deposits, occurs subjacent to adaxial and abaxial 

rows of fibers. Mesophyll: adaxial palisade five to 

seven layered, occupying half of mesophyll and 

gradually changing into spongy tissue. Abaxial 

palisade two to three layered, not highly differentiated from spongy mesophyll. One to three layers 

of thin-walled parenchymatous cells devoid of 

chloroplast occur in radial rows on adaxial side 

of vascular bundles and furrows. Vascular bundles: 

22-24; connected commissural bundles occasionally 

observed. Main veins each with one to three wide 

vessels, mostly two. Two phloem units lying laterally in flanges of abaxial partial girder. Each 

vascular bundle always accompanied by an adaxial 

cap and an abaxial V-shaped partial girder. Crystals 

and tannins: not observed. 

Vellozia crassicaulis Martius ex Schultes f, 

FIGURE 12d-f; PLATES lla-b, 32c-g, 33a 

SPECIMENS EXAMINED.-Archer 4079, 4076; Irwin 

9591, 9744, 9975, 10906, 11699, 13377. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial and abaxial cells mostly rectangular. Stomata: 

paracytic, 18 X 12 pm; observed on both surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove; occasionally 

abaxially two grooved. Adaxial surface evenly undulating; abaxial surface furrowed about one-half 

to two-thirds thickness of blade. Epidermis: cells 

more or less rectangularly flattened, especially 

adaxial cells; abaxial slightly larger than adaxial 

cells; 'walls uniformly thin walled. Distinct baculiform sclerenchyma strands, separated from each 

other by one radial row of parenchyma, occur subjacent to adaxial epidermis. Cuticle: uniformly 

thin, slightly thicker around margins and median 

abaxial surface. Stomata: present in abaxial furrows, few on adaxial surface. Mesophyll: adaxial 

palisade three layered, occupying about half of 

mesophyll; palisade separated from adaxial sclerenchyma strands by one layer of distinct translucent parenchyma cells; two to three radially 

oriented large translucent cells arranged adaxially 

on vascular bundles. Palisade cells compactly arranged and filled with chloroplast; spongy mesophyll loosely arranged. Vascular bundles: 24-40; 

17 in Archer 4076; commissural bundles few. Large 

veins each with one large vessel, occasionally two. 

Two phloem units lying laterally in flanges of 

abaxial V- or Y-shaped girder. Each vascular bundle 

always accompanied by adaxial inverted v-shaped 

sclerenchyma cap; both in turn completely surrounded by a distinct bundle sheath. Crystals: not 

observed. Tannin: few in mesophyll. 

NOTE: All the specimens investigated share 

anatomical and histological characteristics except 

for size of leaf blade. However, one specimen 

(Archer 4079) differs slightly from the others in 

that, whereas the furrows are almost 90" from the 

abaxial surface, Dr. Archer's specimen is thinly 

crescentiform on the abaxial surface from one furrow to the other. Furthermore, the specimen exhibits almost solid subjacent sclerenchyma band 

under the abaxial epidermis. Dr. Archer's specimen 

has sclerenchyma strands confined to the corners 

of the furrow gates, Another specimen (Irwin et 

al. 10906) lacks the distinctive abaxial furrows and 

the sclerenchyma strands which seems to distinguish this species. 

Vellozia crinita Gwthart & Henrard 

PLATE 42d 

SPECIMENS EXAMINED.-GlaziOu 16388. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial cells square to rectangular, few rounded; thin 

walled. Abaxial cells rectangular; thin walled. 

Stomata: paracytic, 21 X 12 pm; present on abaxial 

surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with margins curved slightly inversely 

but not flanged; small median adaxial groove. 

Adaxial surface slightly undulating; abaxial surface furrowed about one-half thickness of blade. 

Epidermis: adaxial cells rounded to rectangular; 
NUMBER 15 19 

I - f 

FIGURE 8.-a-c, Vellozia resinosa (Maguire et al. 44694) ; d, V. minima (Maguire et al. 49099) ; 

e, V. hernisphaerica (Blanchet 2544) ; f, V. taxifolia (Mexia 4289). 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

abaxial cells more rounded to conical; thin walled 

on both surfaces. Two to three layers parenchyma 

cells subjacent to adaxial epidermis and interspersed with sclerenchyma strands. Cuticle: very 

thin on adaxial and abaxial surfaces. Stomata: 

present on abaxial surface and in abaxial furrows. 

Substomatal chamber present: stomata almost flush 

with surface. Mesophyll: three to four layers 

palisade cells; not distinctive; blending right into 

compact spongy tissue. Row of large translucent 

palisade cells arranged radially above vascular 

bundles and furrows. Vascular bundle: 34; no cominissural bundles observed. Veins with a single 

large vessel. Two phloem units lying in flanges of 

Y-shaped abaxial girder; girder extends almost to 

abaxial epidermis. Cap or partial adaxial girder 

occurs on each vascular bundle. Bundle sheath 

completely surrounding each vascular bundle; adjacent to epidermis on abaxial surface. Epidermal 

extensions present in furrows. Few abaxial sclerenchyma strands observed on lower edge of furrows. Crystals: none observed. Tannins: few 

present. 

Vellozia dasypus Seubert 

FIGURE 13c-e; PLATES 16a, 33d-h, 34a 

SPECIMENS EXAMINE.---Belem and Pinheiro 3179;

R. de Lemos Froes 20246; Rose and Russell 19697; 

Blanchet 3558. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial and abaxial cells mostly of square cells, occasionally rectangular cells are observed. Stomata: 

paracytic, 24 X 12 pm; occur on both surfaces 

especially in furrows of abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped to nearly inversely W-shaped with median 

adaxial groove. Adaxial surface ridged; abaxial surface furrowed about one-fifth to one-seventh of 

blade thickness. Epidermis: cells more or less 

square; abaxial cells almost replaced by fibers except in furrows. Distinct cuboidal sclerenchyma 

strands, separated from each other by three to four 

radial' rows of parenchyma, occur subjacent to 

adaxial epidermis. Cuticle: uniformly thickened. 

Stomata: present on both surfaces, particularly in 

abaxial furrows. Mesophyll: adaxial palisade two 

to three layered; palisade separated from adaxial 

sclerenchyma strands by one layer of distinct translucent parenchyma cells; two to five radially 

oriented large translucent cells arranged adaxially 

on vascular bundles. Palisade cells compactly arranged and filled with chloroplast; palisade occupying about one-third of mesophyll and abruptly 

changing into almost tiered spongy cells. Vascular 

bundles: 27-33; commissural bundles present. 

Large veins each with one to two large vessels, 

mostly one. Two phloem units lying laterally in 

flanges of abaxial Y-shaped girder. Each vascular 

bundle always accompanied by adaxial inverted 

V-shaped sclerenchyma cap; both in turn completely surrounded by a distinct bundle sheath. 

Crystals: not observed. Tannin: present but few 

in mesophyll. 

NOTE: The four specimens under study share 

similar histological details except that the median 

abaxial surface in one (R. de Lemos Froes 20246) 

is much broader than the others, and the adaxial 

inverted V-shaped sclerenchyma in one of the four 

specimens (Russell 19697) is much more pronounced, especially at the middle of the intercostal region. 

Vellozia declinans Goethart & Henrard 

PLATE 36d 

SPECIMENS EXAMINED.-Hatschbach 27842. 

SURFACE VIEw.-Hairs: few small papillae, appearing in tufts on adaxial surface. Epidermis: 

adaxial and abaxial cells square to rectangular; 

thin walled. Stomata: 18 X 6 pm; tetracytic; mostly 

present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-DOrSiventral; 

V-shaped, with median adaxial groove; turned 

slightly inversely at margins. Adaxial surface almost smooth; abaxial surface furrowed one-half 

thickness of blade. Epidermis: adaxial cells square; 

abaxial cells dome shaped to conical. Walls thin 

on abaxial and adaxial surfaces. Subjacent to adaxial epidermis occur two to three layers of sclerenchyma strands interspersed with rows of 

parenchyma cells of two to three layers. Below 

sclerenchyma strands is a distinct layer of parenchyma cells. Cuticle: very thin and ridged on both 

surfaces. Stomata: present mainly in abaxial furrows; small substomatal chamber present. Mesophyll: four to five layers of palisade tissue grading 

into compact spongy tissue. Two to four layers of 
NUMBER 15 21 

large translucent palisade cells arranged radially 

above vascular bundle and on furrows. Vascular 

bundle: 16; few commissural bundles present. Two 

to three large vessels present in each vascular 

bundle. Two phloem units lying laterally in flanges 

of U-shaped partial abaxial girder. Adaxial cap 

present on each vascular bundle. Distinct bundle 

sheath completely surrounding each vascular 

bundle. Two to three layers of sclerenchyma cells 

subjacent to abaxial epidermis; some sclerenchyma 

present in furrowed area. Crystals: none observed. 

Tannins: few present. 

NOTE: The partial abaxial girder recalls the 

Xerophyta type of sclerenchyma pattern. 

Velloria echinata Goethart & Henrard 

PLATE 36e 

SPECIMENS ExAMINED.-Riedel 1780. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial cells rounded, square to rectangular; abaxial 

cells rounded. Walls thin on both surfaces. Stomata: 

paracytic, 15 X 3 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAhfINA.-Dorsiventra1; 

V-shaped. Adaxial surface slightly undulating; 

abaxial surface furrowed almost three-fourths 

thickness of blade. Epidermis: cells on adaxial 

surface mostly rectangular, some rounded and 

crescent shaped; thin walled. Cells on abaxial surface rounded to conical, few rectangular; thin 

walled. Cells in furrows with fingerlike projections, 

subjacent to adaxial epidermis is a layer of parenchyma cells interspersed with sclerenchyma strands. 

Below parenchyma cells and sclerenchyma strands 

is distinct layer of large parenchyma cells. Cuticle: 

very thin and slightly ridged on adaxial and abaxial surfaces. Stomata: present in abaxial furrows, 

substomatal chamber virtually absent. Mesophyll: 

three distinct layers palisade tissue present; three 

layers of large translucent cells positioned radially 

above furrows and vascular bundles. Large palisade 

cells also arranged above midvein. Palisade changing abruptly into loosely spongy tissue; some large 

spongy tissue present. Vascular bundles: 25; commissural bundles not observed. One to two large 

vessels present in each vascular bundle, mostly one. 

Two phloem units lying laterally in flanges of Yshaped abaxial girder; sometimes partial girders 

observed. Small adaxial cap present on each vascular bundle. Bundle sheath completely surrounding 

each vascular bundle. No abaxial sclerenchyma 

strands or layers observed. Crystals: none observed. 

Tannins: present. 

NOTE: This species is distinct in that the abaxial furrows are exceptionally deep with its upper 

limit even higher than the adaxial sclerenchyma 

cap. 

Velloria epidendroides Martius ex Schultes f. 

FIGURE 9a-c; PLATE 31b-d 

SPECIMENS EXAMINED.-Achwake 19321; Brade 

19136; Duarte 6537; Lima and Brade 13297A; 

Macedo 2993; Pereira 2229; Maguire et al. 49060; 

Schwake 8354; L. B. Smith 6762; Duarte 4519. 

SURFACE VImv.-Hairs: few present. Epidermis: 

adaxial and abaxial cells mainly cuboidal with few 

rows of rectangular cells; cell walls thin. Stomata: 

paracytic, 21 X 14 pm; observed on abaxial side 

only. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with shallow median adaxial groove; margins of lamina turned inversely. Adaxial surface 

slightly undulating; occasionally somewhat ridged; 

abaxial surface furrowed one-fifth to one-half thickness of blade. Epidermis: adaxial cells more or less 

dome shaped, interspersed with conical cells; abaxial cells mainly rounded; cells thin walled. Subjacent to adaxial epidermis occur one to three 

layers of sclerenchyma bands interspersed with 

parenchyma cells. Below is one conspicuous layer 

of parenchyma cells. Subjacent to abaxial epidermis 

occurs a layer or two, sometimes strands of sclerenchyma. Cuticle: thick on abaxial and adaxial 

surfaces; smooth. Stomata: present in abaxial furrows; somewhat associated with epidermal protuberances. Mesophyll: adaxial palisade cells one 

to two layers, gradually changing into spongy tissue; spongy tissue compactly arranged. One to two 

layers of large translucent palisade cells arranged 

radially above vascular bundles and furrows; of 

some specimens quite numerous in midvein region. 

Vascular bundles: 10 to 27; no commissural bundles 

observed. Each vein large with one to three wide 

vessels. Two phloem units lying laterally in flanges 

of abaxial Y-shaped partial girder. Each vascular 

bundle accompanied by adaxial cap, sometimes 

much larger than abaxial girder. Vascular bundle 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

b 

FIGURE 9.-a-c, Vellozia epidendroides (L. B. Smith 6762); d-f, V. asperula (Gardner 5228). 
NUMBER 15 23 

completely surrounded by distinct bundle sheath. 

Crystals and tannins: not observed. 

NOTE: Notwithstanding the general uniformity 

of the specimens examined, there are certain variations in the depth of the furrows, the shape of the 

abaxial portions of the midveins (some are pointed, 

others are flattened), and the shape of the margins. 

These modifications may constitute varietal characteristics. 

Vellozia exilis Goethart 8c Hemard 

FIGURE 14d-e; PLATE 34e-f 

SPECIMENS EXAMINED.-Goethart and Henrard 

12696; Irwin 12692; Irwin et al. 9507. 

SURFACE VIEw.-Hairs: few and in tufts. Epidermis: cells rectangular on abaxial and adaxial 

surfaces; walls thin on both surfaces. Stomata: 

paracytic, 18 X 3 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with flanges inversely inrolled; abaxial 

median section laterally extended on both sides. 

Adaxial surface slightly undulating, occasionally 

ridged toward median of lamina; abaxial surface 

furrowed about one-fourth thickness of blade. Epidermis: adaxial cells more or less dome shaped; 

abaxial cells dome shaped to conical; cells thin 

walled. Subjacent to adaxial epidermis occur three 

to six layers of sclerenchyma cells interspersed 

with radial rows of two to four parenchyma cells; 

subjacent to sclerenchyma and radial rows of 

parenchyma cells is one layer of distinct parenchyma cells. Cuticle: thick on adaxial surface; 

thin on abaxial surface; slightly ridged. Stomata: 

present in abaxial furrows; small substomatal 

chamber present; associated with short epidermal 

projections in furrows. Mesophyll: two to three 

distinct layers of palisade tissue abruptly changing into compactly arranged spongy tissue. One 

to two layers of large translucent cells arranged 

radially above vascular bundles. Vascular bundles: 

9; no commissural bundles observed. One to two 

large vessels present, mostly one. Two phloem units 

lying laterally in flanges of Y-shaped abaxial girder. 

Adaxial cap more or less inverted V shaped. Lateral extensions formed on either side of median 

abaxial vein have compactly arranged spongy mesophyll and a broad band of sclerenchyma cells. 

Bundle sheath completely surrounding vascular 

bundles. Abaxial strands present. Crystals and tannins: not observed. 

NOTE: The leaf shape of this species is very 

distinctive in the Velloziaceae, thus rendering its 

identification simple. 

Vellozia fibrosa Goethart 8c Henrard 

PLATES 17b, 26d-f 

SPECIMENS EXAMINm.-Azevedo 5592. 

SURFACE VIEW.-Hairs: few papillae present. 

Epidermis: cells square to rectangular, thin walled. 

Stomata: tetracytic, 12 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMrNA.-Dorsiventral; 

V-shaped with small median adaxial groove; margins turned inversely, flanged. Adaxial surface very 

slightly undulating; abaxial surface furrowed onehalf thickness of blade. Epidermis: cells more or 

less square, rounded or rectangular; walls slightly 

thick, especially on inner tangential wall. Subjacent to adaxial epidermis is one to two layers 

of thick-walltd parenchyma cells interspersed with 

sclerenchyma strands; below parenchyma and sclerenchyma is a layer of translucent parenchyma 

cells. Cuticle: adaxial surface thicker than abaxial; both slightly ridged. Stomata: present in furrow and on abaxial surface. Substomatal chamber 

virtually absent. Mesophyll: one to three layers of 

distinct palisade tissue changing into compact 

spongy tissue; one to three layers of large translucent palisade cells arranged radially above vascular bundles and furrows. Vascular bundles: 16; 

commissural bundles not observed. Veins with one 

to two large vessels, mostly one. Two phloem units 

lying laterally in flanges of U-shaped partial abaxial 

girder. Each vascular bundle accompanied by an 

adaxial cap. Vascular bundle completely surrounded by bundle sheath. Abaxial strands present at 

open ends of furrows. Crystals: not observed. 

Tannins: few present. 

NOTE: The presence of partial abaxial girders 

recalls the Xerophyta type of sclerenchyma encountered in Madagascar species of this family. 

Vellozia flavicans Martius ex Schultes f. 

FIGURES 3k-1, lla-c; PLATE 31e-g 

SPECIMENS ExAMINED.-Andrade 356-Emmerich 

348; Heringer 8126-256; Irwin et al. 8073, 9379, 
24 SMITHSOSIAS CONTRIBUTIONS TO BOTAXY 

12574; Irwin et al. 11399; Gardner 4370; L. B. 

Smith 4670; Irwin et al. 12938; Irwin et al. 15520; 

Irwin and Soderstrom 5100; Pires et al. 9052. 

SURFACE VIEW.-Hairs: absent. Epidermis: adaxial and abaxial cells rectangular; thin walled. Stomata: paracytic, 21 X 18 pm; present mainly on 

abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-DOrSiVentral; 

V-shaped with small median adaxial groove. Adaxial surface undulating; abaxial surface furrowed 

about one-fourth to one-third thickness of blade, 

occasionally up to one-half (c.f. Irwin et al. 15520). 

Epidermis: adaxial and abaxial cells rounded to 

rectangular; walls uniformly thickened. Subjacent 

to epidermis occur three to four layers of sclerenchyma strands interspersed with three to four rows 

of translucent parenchyma. Below sclerenchyma 

strands and parenchyma cells is a distinct layer 

of translucent cells. Cuticle: thick on abaxial and 

adaxial surfaces; extends partially into furrows; 

ridged on abaxial surface. Stomata: present mainly 

in abaxial furrows, occasionally few occur on adaxial surface; almost flush with surface; small substomata1 chamber present. Mesophyll: palisade 

cells of four to five layers; large translucent cells 

arranged radially above vascular bundles. Spongy 

tissue compactly arranged, some cells somewhat 

translucent. Vascular bundles: 18 to 42; few commissural bundles observed and occasional fused 

bundles seen (c.f, Gardner 4370). One to two large 

vessels in each vein, mostly one. Two phloem units 

lying laterally in flanges of Y-shaped abaxial girder. 

Adaxial cap occurs on each vascular bundle. Abaxial strands or layers present. Distinct bundle sheath 

completely surrounds vascular bundles. Crystals: 

none observed. Tannins: some present. 

NOTE: All the specimens studied show that 

the large translucent cells occur only on vascular 

bundles, only and not above the furrows as well. 

This species is quite distinct, even at different 

stages of leaf development. 

Vellozia glabra Mikan 

PLATES 24f-g, 258 

SPECIMENS ExAMINED.-Maguire et al. 44622; 

Maguire et al. 49048; Duarte 8355-Mathos 464. 

SURFACE VIEw.-Hairs: on abaxial furrows. Epidermis: cells square to rectangular; thin walled. 

Stomata: paracytic, 21 X 12 pm; present mainly 

on abaxial surface. 

TRAMVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with very small median adaxial groove; 

very slight inversion at margins. Adaxial surface 

almost smooth: abaxial surface furrowed about 

one-half thickness of blade. Epidermis: adaxial and 

abaxial cells square to rectangular; walls thicker 

on inner tangential side. Subjacent to epidermis 

occur two to three layers of parenchyma cells interspersed with one to two layers of sclerenchyma 

strands. Below parenchyma and sclerenchyma 

layers is a distinct layer of large translucent parenchyma cells. Cuticle: slightly thicker on adaxial 

side; slightly ridged on both surfaces. Stomata: 

present in furrows and on abaxial surface; small 

substomatal chamber present; protected by hairs 

in furrows. Mesophyll: definite palisade tissue of 

three layers abruptly changing into compact 

spongy tissue; two to three layers of translucent 

palisade cells arranged radially above vascular 

bundles and furrows. Three distinct layers of large 

translucent palisade cells in midvein region. Vascular bundles: 16-46; no commissural bundles observed. One to two large vessels in each vascular 

bundle, mostly one. Two phloem units lying laterally in flanges of Y-shaped abaxial girder. Each 

vascular bundle has an adaxial cap. Some sclerenchyma strands occur between abaxial epidermir 

and abaxial parenchyma layer. Bundle sheath completely surrounds each vascular bundle; few large 

translucent cells may appear at abaxial end of 

vascular bundle. Abaxial furrows contain many 

epidermal extensions. Crystals: none observed. 

Tannins: few present. 

Vellozia glandulifera Goethart 8c Henrard 

FIGURE 2c; PLATES Ic-d, 36f 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

15998. 

SURFACE VIEW.-Hairs: multiseriate, present on 

adaxial surface in clumps, consisting of small 

fingerlike projections; abaxial surface hairs long 

and fairly thin. Epidermis: adaxial cells mostly 

square; thin walled. Abaxial cells square to rectangular, few almost rounded; thin walled. Stomata: 

pnracylic, 24 X 15 iim; present on abaxial surface, few on adaxial surface. 
NUMBER 15 25 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove; turned 

slightly inversely at margins. Very slightly undulating on adaxial surface; abaxial surface furrowed 

one-half thickness of blade. Epidermis: adaxial 

cells square; abaxial cells slightly rounded. Walls 

thin on abaxial and adaxial surfaces. Two to three 

layers of parenchyma cells subjacent to adaxial 

epidermis and interspersed with sclerenchyma 

strands. Below is a layer of parenchyma cells. 

Cuticle: very thin and slightly ridged on both surfaces. Stomata: present in abaxial furrows in association with fingerlike projections. Mesopliyll: 

three to four layers palisade tissue changing 

abruptly into compact spongy tissue. Three to four 

layers of large translucent palisade cells radially 

arranged above vascular bundles and furrows. Vascular bundles: 22; commissural bundles not observed. One to three large vessels present in each 

vascular bundle. Two phloem units laterally arranged in flanges of Y-shaped abaxial girder. Adaxial cap present on each vascular bundle. Bundle 

sheath completely surrounding each vascular 

bundle. Some abaxial sclerenchyma strands present 

at entrances of furrows. Crystals: none observed. 

Tannins: present. 

Vellozia glauca Pohl 

FIGURES 3e-f, lld-f 

SPECIMENS EXAMINED.-Irwin et al. 21395; Maguire et al. 44793. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular; thin walled. Stomata: paracytic, 21 X 18 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

open V-shaped with margins turned slightly inversely. Adaxial surface slightly undulating; abaxial surface furrowed one-fourth thickness of blade. 

Epidermis: adaxial and abaxial cells rounded and 

thin walled. Subjacent to epidermis occur four to 

five layers of sclerenchyma strands interspersed 

with two to four layered rows of parenchyma. A 

distinct layer of large parenchyma cells subjacent 

to sclerenchyma and parenchyma cells. Cuticle: 

evenly thickened on adaxial surface; thick and 

quite ridged on abaxial surface. Stomata: present 

in furrows; small substomatal chamber present. 

Mesophyll: three to five layers palisade tissue 

grading into spongy tissue. One to two layers large 

translucent palisade cells arranged radially above 

vascular bundles. Vascular bundles: 38-58; commissural bundles not observed. Veins have one to 

three large vessels, mostly one, rarely three. Two 

phloem units lying laterally in flanges of Y-shaped 

abaxial girder. Each vascular bundle has an adaxial 

cap. Abaxial strands near openings of furrows present between abaxial epidermis and a distinct layer 

of parenchyma cells. Bundle sheath extends completely around vascular bundle. Crystals: observed. 

Tannins: present. 

Vellozia glochidea Pohl-Heringer 

FIGURE 12a-c; PLATES 5a, 31h, 32a-b 

SPECIMENS EXAMINED.-Irwin et al. 14584; Duarte 

8340-Mattos 677; Heringer 10923. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular in shape; walls thin. Stomata: 

paracytic, 21 X 18 pm; and present on abaxial 

surface only in Irwin et al. 14584. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove; turned 

slightly inversely at margins. Adaxial surface 

slightly undulating; abaxial surface furrowed up 

to one-third the thickness of blade in Irwin 14584, 

one-fourth in Heringer 10923 and Duarte 8340- 

Mattos 6ii. Epidermis: cells rounded to dome 

shaped on abaxial and adaxial surfaces; thin walled. 

Subjacent are three to four layers sclerenchyma 

strands interspersed with rows of parenchyma. 

Sclerenchyma layers very distinct and interspersed 

with single parenchyma rows in Irwin et al. 14584. 

Below sclerenchyma is a distinct layer of parenchyma cells. Cuticle: thick and smooth on adaxial 

surface; bumpy on abaxial surface; very thick and 

ridged in Irwin et al. 14584. Stomata: present on 

adaxial and abaxial surfaces and in abaxial furrows; substomatal chamber present; seen only in 

abaxial furrows in Irwin et al. 14584. Mesophyll: 

definite palisade tissue present in two to four 

layers grading into compact spongy tissue. Three to 

four layers translucent palisade cells arranged 

radially above vascular bundles. Vascular bundles: 

27; 43 in Irwin et al. 14584; few commissural 

bundles observed. Usually one large vessel in each 

vein. Two phloem units lying laterally in flanges 

of Y-shaped abaxial girder. Each vascular bundle 
26 SMITHSONIAN COKTRIBUTIONS TO BOTANY 

accompanied by an adaxial cap. Abaxial sclerenchyma strands or layers present (three to four layered in Irwin et al. 14584); parenchyma layer may 

be between sclerenchyma and mesophyll. Vascular 

bundles completely surrounded by bundle sheath; 

cells at abaxial end of sheath may be enlared. 

Crystals: none observed. Tannins: few present. 

NOTE: The specimen by Irwin et al. 14584 

does not seem to be the same as the other two 

specimens. 

Vellozia granulata Goethart & Henrard 

FIGURE 6c-e 

SPECIMENS EXAMINED.-Glaziou 19934; Riedel 

s.n. 

SURFACE VIEw.-Hairs: few papillae present. 

Epidermis: cells square to rectangular, thin walled. 

Stomata: tetracytic, 18 X 12 pm; present on abaxial surface; seen on both surfaces in Glaziou 19934. 

TRANSVERSE SECTION OF LAMINA.-DorSiventral; 

V-shaped with margins turned very slightly inversely; small median adaxial groove; groove large 

in Glaziou 19934. Adaxial surface slightly undulating, ridged in Glaziou 19934. Abaxial furrows 

one-half thickness of blade, furrows almost absent 

in Glaziou 19934. Epidermis: cells on both surfaces 

dome shaped to conical. Two to three layers thickwalled parenchyma cells subjacent to epidermis 

and interspersed with sclerenchyma strands. One 

to two layers thin-walled parenchyma in Glaziou 

19934; few sclerenchyma strands. Another layer of 

large parenchyma cells above palisade tissue. 

Cuticle: slightly ridged on both surfaces; cuticle 

fairly thick. Stomata: present in furrows; substomatal chamber virtually absent. Mesophyll: three 

layered palisade grading into spongy tissue; cells 

near abaxial surface more compactly arranged; 

spongy tissue in Glaziou 19934 completely obliterated in most sections. Large translucent palisade 

tissue radially oriented above vascular bundles 

and on furrows. Vascular bundles: 19 in Riedel 

sm.; 29 in Glaziou 19934; commissural bundles not 

observed. One to two vessels larger than others. 

Two phloem units lying laterally in flanges of 

Y-shaped abaxial girder. Adaxial cap usually small, 

rather large in Glaziou 19934. Sclerenchyma strands 

and isolated fibers forming a layer between abaxial 

epidermis and a distinct layer of parenchyma cells. 

Bundle sheath completely surrounding vascular 

bundles. Crystals and tannins: not observed. 

KOTE: The two specimens examined are not 

identical-a clear indication of misidentification. 

Vellozia grisea Goethart & Henrard 

FIGURES 3i-j, 4b; PLATES 16b, 42e, 50c 

SPECIMENS EXAI\.IINED.-Weddell 3006. 

SURFACE VIEw.-Hairs: present in clunlps On 

adaxial and abaxial surfaces. Epidermis: cells 

square to rectangular, thin walled, Stomata: paracytic, 15 X 9 pm; present on abaxial and few on 

adaxial surface. 

TRANSVERSE SECTION OF LAMrNA.-Dorsiventral; 

V-shaped with deep median adaxial groove. Adaxial surface conspicuously ridged; adaxial furrows 

almost one-half thickness of blade. Epidermis: cells 

rounded to dome shaped on both surfaces; few 

conical on adaxial surface. Two to three layers of 

sclerencliyma cells subjacent to epidermis and interspersed with parenchyma cells. Below sclerenchyma is a layer of parenchyma cells. Cuticle: 

uniformly ridged and thick on both surfaces. Stomata: present in furrows; substomatal chamber 

virtually absent. Mesophyll: three to four layers 

indistinct palisade cells grading into compact 

spongy tissue; few translucent palisade cells radially arranged above vascular bundles. Vascular 

bundles: 20; commissural bundles not observed. 

Veins having one to two large vessels, mostly one. 

Two phloem units lying laterally in flanges of 

Y-shaped girder. Each vascular bundle has in addition to Y-shaped abaxial girder an adaxial cap. 

Vascular bundle completely surrounded by bundle 

sheath. Few strands of abaxial sclerenchyma observed. Crystals: not observed. Tannins: present. 

NOTE: The adaxial surface is very heavily 

ridged. In some areas the ridges are deep enough 

to qualify as small furrows. 

Vellozia hatschbachii L. B. Smith & Ayensu 

PLATES 1Oa-b, 37a 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

16002. 

SURFACE VIEtv.-Hairs: none present. Epidermis: 

adaxial cells mostly square, few rectangular; thin 
NUMBER 15 27 

f 

FIGURE 10.-a-c, Vellozia aloifolia (Mexia 5870) ; d-f, V. leptopetala (Maguire et al. 49184). 

walled, Abaxial cells rectangular; thin walled. Stomata: paracytic, 21 X 9 pm; present on abaxial 

surface. 

TRANSVEXSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Adaxial surface very slightly undulating; abaxial surface 

furrowed one-half to three-fourths thickness of 

blade. Epidermis: adaxial cells rounded; few rectangular; thin walled. Abaxial cells rounded; interspersed with small sclerenchyma strands. Subjacent 

to adaxial epidermis occur two to three layers of 

sclerenchyma strands interspersed with radially arranged rows of two to three parenchyma cells. 

Below sclerenchyma strands is a distinct layer of 

parenchyma cells. Cuticle: fairly thick and ridged 

on adaxial and abaxial surfaces, Stomata: present 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

in abaxial furrows; substomatal chamber present; 

some epidermal furrow projections covering stomata. Mesophyll: three to four, mostly three, layers 

of compactly arranged small palisade tissue changing abruptly into spongy tissue. Two to three 

layers of large translucent cells radially arranged 

above vascular bundles and furrows. Vascular 

bundles: 46; few commissural bundles present. 

Two to three large vessels present in each vascular bundle. Two phloem units arranged laterally 

in flanges of partial Y-shaped abaxial girder. Small 

adaxial cap present on each vascular bundle. Abaxial sclerenchyma strands present between abaxial 

epidermis followed by a layer of parenchyma cells. 

Each vascular bundle completely surrounded by a 

bundle sheath. Crystals: none observed. Tannins: 

some present. 

KOTE: This species resembles Vellozia lappa 

in its general anatomy. The leaf blade is smaller 

than V. lappa. Furthermore the translucent cells 

above the vascular bundles are mostly two, whereas 

those in V. lappa are mostly four. 

Vellozia hemisphaerica Seubert in Martius 

FIGURE 8e 

SPECIMENS ExAMrNED.-Blanchet 2544. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular, thin walled. Stomata: paracytic, 18 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTIOK OF LA1LrINA.-Dorsiventral; 

V-shaped. Adaxial surface conspicuously ridged; 

abaxial surface distinctly furrowed about one-half 

thickness of blade. Epidermis: adaxial cells 

rounded to dome shaped, some conical; abaxial 

cells more rectangular; cells on both surfaces thin 

walled. Subjacent to the epidermis are one to two 

layers of sclerenchyma strands interspersed with 

parenchyma cells. Below sclerenchyma strands is 

a distinct layer of parenchyma cells. Cuticle: thin 

and ridged on adaxial surface; not as ridged but 

thicker on abaxial surface. Stomata: present in 

abaxial furrows; covered by epidermal projections; 

large, distinct hooklike protuberances in furrows 

overarch some of the stomata; substomatal chamber 

present. Mesophyll: two to three layers of palisade 

grading into compactly arranged spongy tissue to 

abaxial side; one to two layers of large translucent 

palisade cells radially arranged above major and 

minor vascular bundles. Vascular bundle: two 

types present, major and minor veins; major veins 

18 to 22; minor veins 16 to 24 arranged radially 

above furrows. Major veins contain one to two 

large vessels, mostly one; two phloem units lying 

laterally in flanges of Y-shaped abaxial girder; 

each vascular bundle accompanied by an adaxial 

cap; vascular bundles completely surrounded by 

a distinct bundle sheath. Minor veins usually with 

one large vessel; abaxial and adaxial ends accompanied by sclerenchyma caps; minor veins with 

adaxial cap and abaxial U- or V-shaped girder completely surrounded by a bundle sheath. Crystals 

and tannins: not observed. 

NOTE: This species and Vellozia burle-rnarxii 

share the following common characters: (a) major 

and minor vascular bundles and (b) distinct hooklike protuberances in furrows which overarch the 

lower stomata. 

Vellozia hypoxoides L. B. Smith 

PLATES 37c, 50a 

SPECIMENS EXAMINED.-IrWiIl et al. 34120. 

SURFACE VrEw.-Hairs: few tufts of multicellular 

hairs. Epidermis: adaxial cells square to rectangular; thin walled. Abaxial cells square to rectangular, few rounded; thin walled. Stomata: paracytic, 

18 X 9 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove. Margins 

turned very slightly inversely. Abaxial surface furrowed about one-half thickness of blade. Epidermis: 

adaxial cells square to rectangular, few rounded; 

cell walls slightly thickened. Abaxial cells rounded 

to dome shaped; walls slightly thickened. Subjacent 

to adaxial epidermis occur two layers of small 

parenchyma cells followed by a layer of large 

parenchyma cells. Few sclerenchyma strands found 

in parenchyma cell layers. Cuticle: thick and 

slightly ridged on adaxial surface; thin on abaxial 

surface. Stomata: present in abaxial furrows, small 

substomatal chamber observed, flush with epidermis. Mesophyll: two to three layers of palisade 

tissue subjacent to adaxial epidermis, rather indistinct in places. Two- to three-layered large and 

almost rounded translucent cells arranged radially 

above vascular bundles and furrows. Palisade cells 

grading into compact spongy tissue of small round 
NUMBER 15 29 

cells. Vascular bundles: 23; few commissural 

bundles observed. One to two vessels present in 

each vascular bundle, mostly one. Two phloem 

units lying laterally in flanges of Y-shaped abaxial 

girder. Small adaxial cap present on each vascular 

bundle. Bundle sheath completely surrounds each 

vascular bundle. One layer of parenchyma cells 

subjacent to abaxial epidermis. Crystals: none observed. Tannins: some present. 

Vellozia incurvata Martius ex Schultes f. 

PLATE 42f 

SPECIMENS EXAMINED.--R/lagUire et al. 44752; 

Pereira 2959-Pabst 3795. 

SURFACE VrEw.-Hairs: absent, Epidermis: cells 

rectangular to cubelike on both surfaces; thin 

walled. Stomata: tetracytic, 21 X 17 pm; present 

on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove in 

Pereira 2959, Pabst 3795; margins turned slightly 

inversely and pointed. Adaxial surface slightly undulating; abaxial surface furrowed about one-half 

the thickness of blade. Epidermis: adaxial and 

abaxial cells rounded to rectangular; cell walls 

thin on both surfaces. One layer of sclerenchyma 

cells subjacent to epidermis and interspersed by 

parenchyma cells. Below sclerenchyma cells are 

two to three layers of parenchyma cells. Cuticle: 

thick on both surfaces; only slightly ridged on 

both surfaces. Stomata: present in abaxial furrows; substomatal chamber present. Mesophyll: 

two to three layers of indistinct palisade tissue 

grading into compactly arranged spongy tissue. 

Two to three layers of large translucent cells arranged radially above vascular bundles and furrows, poorly differentiated in Maguire et al. 44752. 

Vascular bundles: 25-34; few commissural bundles 

observed. One to two large vessels present in each 

vascular bundle, usually one. Two phloem units 

lying laterally in flanges of Y-shaped partial abaxial 

girder. Adaxial cap present on each vascular 

bundle. Two to three layers of sclerenchyma cells 

separating abaxial epidermis and a distinct layer 

of parenchyma cells. Bundle sheath completely 

surrounding vascular bundles. Some epidermal 

projections present in furrows. Crystals: none observed. Tannins: some present. 

NOTE: The two specimens studied are quite 

similar; however, the abaxial median portion is 

pointed in Pereira 2959, Pabst 3795, while 

of hfaguire et al. 44752 is flattened. 

Vellozia intermediu Seubert in Martius 

PLATE 28f-g 

SPECIMEKS EXAMINED.-L. B. Smith 6854. 

SURFACE VIEw.-Hairs: present. Epidermis: 

that 

cells 

square to rectangular; thin walled. Stomata: paracytic, 21 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped; small adaxial groove present. Adaxial 

surface slightly undulating; abaxial surface furrowed about one-half thickness of blade. Epiderdis: adaxial cells rounded to dome shaped; abaxial 

cells rounded. One to two layers parenchyma subjacent to epidermis; below parenchyma cells are 

two to three layers of sclerenchyma strands interspersed with radial rows of two- to three-layered 

parenchyma cells. Subjacent to sclerenchyma is a 

distinct layer of parenchyma cells. Cuticle: thick 

and slightly ridged on adaxial surface; thin on 

abaxial surface. Stomata: present in abaxial furrows; asscociated with large numbers of epidermal 

projections; small substomatal chamber. Mesophyll: 

three to four layers of translucent tissue changing 

abruptly into compact spongy tissue. Four to five 

layers of large translucent palisade cells arranged 

radially above vascular bundles and furrows. Vascular bundles: 55; few commissural bundles observed. One to two large vessels, mostly one. TWO 

phloem units lying laterally in flanges of Y-shaped 

abaxial girder. Adaxial cap present on each vascular bundle. One to two layers sclerenchyma cells 

between abaxial epidermis and a distinct parenchyma layer. Vascular bundle completely surrounded by bundle sheath. Crystals: none observed. Tannins: present. 

Velloziu irwinii L. B. Smith 

PLATE 43a 

SPECIMENS EXAMINED.-Irwin, Maxwell, Wasshausen 20998. 

SURFACE VIEw.-Hairs: multicellular, present on 

adaxial surface. Epidermis: cells circular to rec- 
SMITHSONIAN COXTRIBUTIONS TO BOTANY 

tangular, thin walled. Some crystals present. Stomata: paracytic, 24 X 12 pm; present on abaxial 

surface. 

TRANSVERSE SECTION OF LAMINA.--Dorsiventral; 

V-shaped; turned slightly inversely at margins; 

small adaxial grooves present. Adaxial surface 

slightly undulating; abaxial surface deeply furrowed over one-half the thickness of blade. Epidermis: adaxial cells rounded to dome shaped; 

abaxial cells dome shaped to conical. Cells on both 

surfaces thin walled. Subjacent to epidermis is a 

layer of parenchyma cells; below parenchyma are 

sclerenchyma strands interspersed with parenchyma 

cells; subjacent is another layer of parenchyma 

cells. Cuticle: very thin on both surfaces. Stomata: 

present in furrows, and on surface; substomatal 

chamber present. Mesophyll: three to four layers 

of palisade cells grading into compactly arranged 

spongy tissue; cells near abaxial side less compact. 

Two to three layers large translucent palisade 

cells arranged radially above vascular bundles and 

furrows. Vascular bundles: 55; commissural bundles 

not observed. One to two large vessels present in 

each vascular bundle, mostly one. Two phloem 

units arranged laterally in flanges of Y-shaped abaxial girder. Adaxial cap present on each vascular 

bundle. Vascular bundles completely surrounded 

by bundle sheath. Crystals: present. Tannins: some 

present. 

Vellozia lappa L. B. Smith & Ayensu 

PLATE 3'id 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

15988. 

SURFACE VIEw.-Hairs: short hairs present in 

clumps on adaxial surface; none on abaxial. Epidermis: cells on adaxial and abaxial surfaces 

square to rectangular; thin walled. Stomata: paracytic, 21 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAhiINA.-Dorsiventra1; 

V-shaped with small median adaxial groove; turned 

slightly inversely at margins. Adaxial surface almost smooth; abaxial surface furrowed about onehalf thickness of blade. Epidermis: adaxial cells 

square to rectangular in shape; thin walled. Abaxial cells rounded to square; thick walled; abaxial 

epidermal cells replaced by sclerenchyma cells in 

a number of places. Subjacent to adaxial epidermis 

occur two to three layers of parenchyma cells interspersed with sclerenchyma strands. Below is a 

distinct layer of parenchyma cells, sometimes two. 

Cuticle: very thin on abaxial and adaxial surfaces. 

Stomata: present in abaxial furrows; small substomatal chamber present. Numerous furrow projections present. Mesophyll: three to four layers 

palisade tissue grading into compactly arranged 

spongy tissue. Three to four layers of large translucent cells radially arranged above vascular 

bundles and furrows. Vascular bundles: 30; few 

commissural bundles observed. One to two large 

vessels present in each vascular bundle, mostly 

one. Two phloem units lying laterally in flanges 

of Y-shaped or partially Y-shaped abaxial girder. 

Small adaxial cap present on each vascular bundle. 

Abaxial sclerenchyma strands replacing some of 

abaxial epidermal cells. Row of parenchyma cells 

betiyeen abaxial epidermis and spongy tissue. 

Bundle sheath completely surrounding vascular 

bundle; occasionally two bundle sheaths around 

one bundle are observed. Crystals and tannins: 

none observed. 

NOTE: The presence of two bundle sheaths 

around one vascular bundle is quite distinctive. 

Vellozia leptopetala Goethart & Henrard 

FIGURE 10d-f; PLATE 30d-f 

SPECIMENS EXAMINED.-Maguire et al. 49 184. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular; thin walled. Stomata: paracytic, 15 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Adaxial surface very slightly undulating; abaxial surface furrowed about one-half thickness of blade. 

Epidermis: cells rounded to dome shaped on adaxial 

surface; rounded on abaxial surface; thin walled. 

Subjacent to adaxial epidermis occur one to two 

layers of sclerenchyma strands interspersed with 

rows of parenchyma tissue of two to three cells 

thick. Below sclerenchyma is a distinct layer of 

parenchyma cells. Cuticle: thin and slightly ridged 

on both surfaces. Stomata: present in furrows; substomatal chamber virtually absent. Mesophyll: 

three- to four-layered palisade tissue changing 

abruptly into compact spongy tissue. One to two 

layers of large translucent cells arranged radially 
NUMBER 15 

above vascular bundles and furrows. Vascular 

bundles: 18; no commissural bundles observed. 

One to two large vessels, mostly one. Two phloem 

units lying laterally in flanges of U- or V-shaped 

partial abaxial girder. Adaxial cap present on all 

vascular bundles. Distinct layer of parenchyma 

cells separating abaxial epideqmis and one to two 

layers abaxial sclerenchyma. Bundle sheath completely surrounding vascular bundle. Crystals and 

tannins: none observed. 

NOTE: The partial abaxial sclerenchyma recalls 

the Xerophyta type of sclerenchyma girders found 

in some Old World Velloziaceae species. The flattened abaxial portion of the median vein is also 

distinctive. 

Vellozia magdaleme L. B. Smith 8c Ayensu 

PLATES 14a, 43b 

SPECIMENS EXAMINED.-Lima and Brade 13298, 

14288. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

rounded, square to rectangular on both surfaces; 

thin walled. Stomata: paracytic, 18 X 15 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Adaxial surface slightly undulating; abaxial surface furrowed about one-third the thickness of blade. 

Epidermis: adaxial cells dome shaped to square and 

rectangular; walls slightly thickened. Abaxial cells 

rounded to dome shaped; thin walled. Subjacent to 

adaxial epidermis occur one to two layers of parenchyma cells interspersed with sclerenchyma strands. 

Below parenchyma layers is a distinct layer of 

parenchyma cells. Cuticle: fairly thick on both 

adaxial and abaxial surfaces; slightly ridged on 

abaxial. Stomata: present in abaxial furrows; surrounded with epidermal projections in furrows, 

small substomatal chamber observed. Mesophyll: 

distinct two- to three-layered palisade tissue grading into fairly compact spongy tissue; cells less 

compact toward abaxial surface. One to two layers 

of large translucent cells arranged radially above 

vascular bundles and on furrows. Vascular bundles: 

18 to 19; no commissural bundles observed. One 

to two large vessels present in vascular bundles, 

mostly one. Two phloem units arranged laterally 

in flanges of Y-shaped, sometimes partial abaxial 

girder. Adaxial caps present on all vascular 

bundles. Abaxial sclerenchyma strands present between abaxial epidermis and a layer of parenchyma 

cells. Bundle sheath completely surrounding each 

vascular bundle. Crystals and tannins: none observed. 

NOTE: The two specimens are identical except 

for the adaxial portion of midvein that is flattened 

in Lima & Brade 13298, but pointed in Lima and 

Brade 14288. 

Vellozia metzgerae L. B. Smith 

PLATES 5b, 43c 

SPECIMENS EXAMINED.-W. Egler s.n. 

SURFACE VrEw.-Hairs: present on both adaxial 

and abaxial surfaces; usually fairly short and in 

tufts, cells multicellular. Epidermis: adaxial cells 

square, few rounded; thin walled. Abaxial cells 

square to rectangular; thin walled. Stomata: paracytic, 21 X 15 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove; margins 

turned inversely. Adaxial surface slightly undulating, abaxial surface furrowed one-third to onehalt the thickness of blade. Epidermis: adaxial 

cells rectangular; abaxial cells rounded to rectangular; cells on both surfaces are thin walled. 

Two to three layers of parenchyma cells subjacent 

to adaxial epidermis and interspersed with one to 

three layers of sclerenchyma strands. Below is a 

distinct layer or two of large parenchyma cells. 

Cuticle: fairly thick on both surfaces; only slightly 

ridged on both surfaces. Stomata: present in furrows; substomatal chamber present; associated with 

fingerlike projections. Mesophyll: three- to fourlayered palisade grading into fairly compact spongy 

tissue. Two to four layers of large translucent cells 

arranged radially above vascular bundles and on 

furrows; large translucent cells quite numerous in 

median region of lamina. Vascular bundles: 16; 

few commissural bundles observed. One to three 

large vessels observed in each vascular bundle. 

Two phloem units laterally arranged in flanges 

of Y-shaped or partial U- or V-shaped abaxial 

girder; each bundle with an inverted V-shaped 

adaxial sclerenchyma cap. Some abaxial sclerenchyma strands present. Bundle sheath completely 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

- 

FIGURE 11.-a-c, Vellozia flauicans (Gardner 4370) ; d-f, V.-glauca (Maguire et al. 44793) . 
NUMBER 15 33 

surrounding each vascular bundle. Crystals: not 

observed. Tannins: few present. 

Vellozia minima Pohl 

FIG~IRE 8d; PLATE l5b 

SPECIMENS ExAMIi%Eo.-Duarte 2379; Maguire et 

al. 49099. 

SURFACE VIEw.-Hairs: absent. Epidermis; cells 

rounded and rectangular; thin walled. Stomata: 

paracytic, 15 X 18 pm; present on abaxial and 

adaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

widely V-shaped with small median adaxial groove. 

Both surfaces slightly undulating; no furrows present. Epidermis: adaxial and abaxial cells rounded 

to dome shaped; thin walled. Subjacent to adaxial 

epidermis is a layer of parenchyma cells. Cuticle: 

very thin on both surfaces. Stomata: present on 

adaxial and abaxial surfaces; substomatal chamber 

present. Mesophyll: fairly indistinct palisade tissue 

of two to three layers grading into a loose spongy 

tissue. Translucent cells radially arranged above 

some vascular bundles. Vascular bundles: 6-8; no 

commissural bundles observed. Two phloem units 

lying laterally in flanges of U-shaped abaxial 

girder; adaxial cap present on each bundle. Bundle 

sheath completely surrounding each vascular 

bundle. Crystals and tannins: none observed. 

NOTE: As its name implies, this species is 

perhaps the smallest in the New World vellozias. 

The number of bundles is few but firmly organized. 

The abaxial portion of the midvein is slightly 

concave. The translucent cells in the adaxial portion of the midvein are quite distinct. The palisade 

cells are somewhat indistinct and loosely arranged. 

Vellozia modesta L. B. Smith & Ayensu 

PLATE 43d 

SPECIMEKS EXAMINED.-L. B. Smith, Ayensu, 

Hatschbach 15997; Hatschbach 27869. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

rounded to rectangular, thin walled. Stomata: paracytic, 21 X 15 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

widely V-shaped with small median adaxial groove; 

margins turned slightly inversely. Adaxial surface 

undulating; abaxial surface furrowed about onehalf thickness of blade. Epidermis: adaxial cells 

mostly rectangular; abaxial cells rounded to dome 

shaped; cells on both surfaces thin walled. Subjacent to adaxial epidermis is one to two layers of 

parenchyma cells interspersed with sclerenchyma 

strands. Below is a layer of parenchyma cells. 

Cuticle: fairly thin and smooth on both surfaces. 

Stomata: present in furrows and on abaxial surface; substomatal chamber present; stomata associated with numerous epidermal projections in 

furrows. Mesophyll: three- to five-layered palisade 

cells changing abruptly into loosely spongy tissue. 

Large, translucent cells arranged radially above 

vascular bundles and on furrows. Vascular bundles: 

38; no commissural bundles observed. One to two 

large vessels present in each vascular bundle, 

mostly one. Two phloem units arranged laterally 

in flanges of thin Y-shaped or partial U- or V-shaped 

abaxial girder. Small adaxial cap present on all 

bundles. Sclerenchyma strands present on abaxial 

side, fiber often replacing epidermal cells. Layer 

of parenchyma cells separating abaxial sclerenchyma strands from spongy tissue. Bundle sheath 

completely surrounding each vascular bundle. 

Crystals: not observed. Tannins: present. 

NOTE: This species is very distinctive because 

of the arrangements of adaxial and abaxial sclerenchyma strands as well as the fusiform epidermal 

cells at the outer portions of the furrows. 

Vellozia nuda L. B. Smith & Ayensu 

FIGURE 4c; PLATE 37e 

SPECIMENS ExAhfMrNEo.-Hatschbach and Pelanda 

28029. 

SURFACE VIEw.-Hairs: numerous papillae present. Epidermis: adaxial cells square to rectangular; 

abaxial cells rounded. Walls thin on both surfaces. Stomata: paracytic, 7 X 15 pm; present on 

abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove; turned 

slightly inversely at extreme margin of lamina. 

Adaxial surface only very slightly undulating; 

some papillae present. Abaxial surface furrowed 

about one-third thickness of blade. Epidermis: 

adaxial cells slightly rounded; few dome shaped; 

walls thin. Abaxial cells dome shaped; walls thin. 
34 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

Subjacent to adaxial epidermis occur five to six 

layers of sclerenchyma strands interspersed with 

radially arranged rows of four to five parenchyma 

cells. Below sclerenchyma strands is a distinct layer 

of parenchyma cells. Cuticle: adaxial and abaxial 

cuticle very thin and ridged. Stomata: present in 

abaxial furrows; small substomatal chamber present. hlesophyll: three- to four-layered palisade cells 

grading into compactly arranged spongy tissue. 

Two- to three-layered large translucent cells arranged radially above vascular bundles. Vascular 

bundles: 24; few commissural bundles observed. 

One to two large vessels present in each vascular 

bundle, mostly one. Two phloem units arranged 

laterally in flanges of Y-shaped abaxial girder. Inverted V-shaped adaxial cap present on each vascular bundle. Abaxial sclerenchyma strands present 

in abaxial corners of furrows and extending partly 

into furrows. Bundle sheath completely surrounding each vascular bundle. Crystals: none observed. 

Tannins: present. 

Vellozia ornata Martius ex Schultes f. 

PLATE 37f 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

16001. 

SURFACE VIEw.-Hairs: few tufts of multicellular 

hairs present. Epidermis: adaxial cells square to 

rectangular; thin walled. Abaxial cells square to 

rectangular; thin walled. Stomata: paracytic, 18 X 

9 pni; present on adaxial and abaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Extreme margins of lamina turned inversely. Adaxial 

surface slightly undulating; abaxial surface furrowed one-fourth to one-third thickness of blade. 

Epidermis: adaxial cells rounded to rectangular, 

small in size; walls slightly thickened. Abaxial cells 

rounded to dome shaped; walls slightly thickened. 

Adaxial epidermis interspersed with strands of 

sclerenchyma fibers, especially arranged radially 

above furrows and vascular bundles; subjacent to 

this occurs a layer of parenchyma cells. Cuticle: 

thick and quite ridged on both adaxial and abaxial surfaces. Stomata: present on adaxial surface 

and in abaxial furrows, more numerous on abaxial 

surface; substomatal chamber present on both surfaces: stomata flush with epidermis. Mesophyll: 

three to four layers of palisade cells changing 

abruptly to small, rounded, compactly arranged 

spongy tissue. Two to three layers of large transparent cells arranged radially above vascular 

bundles and furrows. Large translucent cells occur 

above entire area of midvein. A layer of parenchyma cells on abaxial side occurs between spongy 

mesophyll and abaxial epidermis. Vascular bundles: 

23; few commissural bundles observed. One to 

three vessels present in each vascular bundle. Two 

phloem units lying laterally in flanges of thick 

Y- or U-shaped abaxial girder. Adaxial cap present 

on each vascular bundle. Abaxial sclerenchyma 

strands present usually at corners of outer area 

of furrows. Bundle sheath completely surrounding 

each vascular bundle. Crystals: none observed. Tannins: few present. 

Vellozia phalocarpa Pohl 

PLATE 46a 

SPECIMENS EXAMINED.-Anderson et al. 35855. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial cells rounded to square, few almost triangular; 

thin walled; abaxial cells rectangular, few square 

shaped, thin walled. Stomata: paracytic, 24 X 12 

pm; present on abaxial surface. 

TRAKSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Extreme margins of lamina turned slightly inversely. 

Adaxial surface slightly undulating; abaxial surface furrowed one-third to one-half thickness of 

blade. Epidermis: cells on adaxial and abaxial surfaces rounded; wall thickened. Adaxial epidermis 

interspersed with strands of sclerenchyma. Subjacent to epidermis and strands of fibers occur two 

to three layers of parenchyma cells. Cuticle: markedly thick and slightly ridged on both surfaces. 

Stomata: present in abaxial furrows; virtually no 

substomatal chamber observed; somewhat associated with small projections in furrow. Mesophyll: three to four layers of palisade cells 

changing abruptly into a compactly arranged 

spongy tissue of small, rounded cells. Two to three 

layers of large translucent cells radially arranged 

above vascular bundles and furrows. Large translucent cells occur above midvein. Vascular bundles: 

26; few commissural bundles observed. One to two 

large vessels present in each vascular bundle. Two 
NUMBER 15 35 

phloem units lying laterally in flanges of Y-shaped 

abaxial girder. Adaxial cap present. One to two 

layers of sclerenchyma strands between abaxial epidermis and a layer of parenchyma cells subjacent 

to spongy mesophyll. Bundle sheath present on 

each vascular bundle. Crystals: present. Tannins: 

some observed. 

Vellozia pilosa Goethart & Henrard 

PLATE 43e 

SPECIMENS EXA;\IINED.-Glaziou 19933. 

SURFACE VrEw.-Hairs: some present. Epidermis: 

cells rectangular, thin walled on both surfaces; 

papillae present. Stomata: paracytic, 18 X 9 pm; 

present on abaxial surface. 

V-shaped with small median adaxial groove. Adaxial surface distinctly ridged; small furrows on 

abaxial surface. Epidermis: abaxial and adaxial 

cells rounded to dome shaped; cells on both surfaces thin walled. Subjacent to adaxial epidermis 

is a layer of parenchyma cells. Cuticle: very thin 

on adaxial and abaxial surfaces. Stomata: present 

on abaxial surface; no substomatal chamber observed. hfesophyll: three layers of palisade tissue 

blending into compactly arranged spongy tissue. 

Vascular bundles: 15; no commissural bundles 

observed. One to two large vessels present in each 

vascular bundle. Two phloem units lying laterally 

in flanges of U-shaped abaxial girder. Adaxial cap 

present on each vascular bundle. No abaxial sclerenchyma present. Bundle sheath surrounding each 

vascular bundle. Crystals and tannins: none observed. 

TRAKSVERSE SECTION OF LAMINA.-Dorsiventral; 

Vellozia piresiana L. B. Smith 

PLATE 43f 

SPECIMESS ExAMINED.-Duarte 2068. 

SURFACE VrEw.-Hairs: absent. Epidermis: cells 

square to rectangular on both surfaces, thin walled. 

Stomata: paracytic, 12 X 6 pm; present on abaxial 

surface. 

TRANSVERSE SECTION OF LAMINh.-DOrSiventral; 

V-shaped with small but distinct median adaxial 

groove. Adaxial surface undulating; abaxial surface furrowed about one-third the thickness of 

blade. Epidermis: adaxial and abaxial cells rectangular to dome shaped; cell walls thin on both 

surfaces. Subjacent to adaxial epidermis occur 

sclerenchyma strands interspersed with one to two 

layers of parenchyma cells. Cuticle: adaxial cuticle 

thin and slightly ridged; abaxial cuticle thin but 

deeply ridged. Stomata: present in abaxial furrows; no substomatal chamber observed. Mesophyll: three layers of indistinct palisade tissue 

grading into fairly compactly arranged spongy 

tissue. Large translucent cells arranged radially 

above vascular bundles and furrows; adaxial translucent cells in midvein very distinct. Vascular 

bundles: 37; no commissural bundles observed. 

Each vascular bundle contains one to two large 

vessels, mainly one. Two phloem units lying laterally in flanges of U-shaped abaxial girder. Large 

adaxial cap on each bundle, often larger than 

abaxial girders. Abaxial sclerenchyma strands 

present; some parenchyma cells between abaxial 

sclerenchyma and 3pongy tissue. Crystals: none 

observed. Tannins: present. 

NOTE: The adaxial sclerenchyma girders which 

are often larger than the abaxial girders are very 

distinctive in Vellozia. 

Vellozia plicata Martius 

FIGURE 4f; PLATE 44a 

SPECIMENS EXAMINED.-POrtO 29572; Frazao sen.; 

Duarte 3635-Gomes 420; Lima 192; Brade 15947. 

SURFACE VIEw.-Hairs: present in clusters as 

well as in fingerlike multicellular projections. Epidermis: adaxial cells almost rounded; few square to 

rectangular; abaxial cells square to rectangular, 

few rounded; walls thin on adaxial and abaxial 

surfaces. Stomata: paracytic, 15 X 6 pm; present 

on abaxial and adaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

W-shaped with margins turned inversely. Adaxial 

surface conspicuously undulating in some specimens; abaxial furrows very large in places. Entire 

lamina distinctly corrugated. Epidermis: adaxial 

and abaxial cells rounded to dome shaped; cells on 

both surfaces very thick walled. Subjacent to adaxial epidermis occur one to two layers of sclerenrhyma strands interspersed with parenchyma cells. 

Below sclerenchyma strands is a layer of parenchyma cells. Cuticle: thin and slightly ridged on 
36 SMITHSONIAS CONTRIBUTIONS TO BOTANY 

U d 

e 

FIGURE 12.-a-c, Vellozia glochidea (Irwin et al. 14584); d-f, V. crassicaulis (Irwin et al. 9744) 
NUMBER 15 

both surfaces. Stomata: present on abaxial surface 

as well as furrows; almost flush with epidermal 

surface; present on adaxial surface; substomatal 

chamber present. Xlesophyll: three layers of palisade tissue grading into compactly arranged spongy 

tissue. Vascular bundles: consisting of two types, 

major and minor veins; major veins 11-15; minor 

veins 30-63; no commissural bundles observed. 

Major veins have three to four large vessels. Two 

phloem units arranged laterally in flanges of 

U-shaped abaxial girder; adaxial girder present on 

each major vein, often extending close to adaxial 

epidermis; each major vascular bundle completely 

surrounded by a bundle sheath. Sclerenchyma 

strands present between abaxial epidermis and a 

layer of parenchyma cells of some specimens 

(Duarte 3635-Gomes 420; Lima 192; Porto 29572; 

Frazao s.n.; Brade 15947). Crystals: not observed. 

Tannins: few present. 

NOTE: This species is very distinctive because 

of the plicate habit of its leaves, the presence of 

major and minor vascular bundles, and its striking resemblance to Vetlozia triquetra (Gardner 

2735). 

Vellozia pumila Goethart & Henrard 

PLATES 5d. 44b 

SPECIMENS ExAMINED.-Irwin et al. 9230; Irwin 

et al. 9428; Irwin et al. 10922; Glaziou 22215. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular on both surfaces; thin walled. 

Stomata: paracytic, 27 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMrNA.-Dorsiventral; 

V-shaped; adaxial surface undulating, sometimes 

conspicuously ridged; abaxial surface furrowed 

about one-fifth to one-third thickness of the blade. 

Epidermis: cells on both surfaces rounded or dome 

shaped; few conical; thin walled. Subjacent to 

adaxial epidermis occur sclerenchyma strands interspersed with parenchyma cells. Below sclerenchyma strands is a distinct layer of parenchyma 

cells. Cuticle: adaxial cuticle thin and slightly 

ridged; abaxial cuticle thicker and very ridged. 

Stomata: present in abaxial furrows; substomatal 

chamber present. Mesophyll: two to three layers 

of palisade grading into a compactly arranged 

spongy tissue. One to two layers large translucent 

palisade cells arranged radially above vascular 

bundles. Vascular bundles: 2 1; no commissural 

bundles observed. Usually one large vessel present 

in each vacular bundle. Two phloem units arranged laterally in flanges of Y-shaped abaxial 

girder. Adaxial cap present on each vascular 

bundle. Bundle sheath completely surrounding 

each vascular bundle. Crystals: none observed. 

Tannins: present. 

Vellozia punctulata Seubert in Martius 

FIGURE 4a; PLATE 44c 

SPECIMENS ExAMINED.-Blanchet 2561. 

SURFACE VIEW.-Hairs: present in tufts. Epidermis: cells square to rectangular on both surfaces; 

thin walled. Stomata: paracytic, 15 X 9 pm; present 

on abaxial surface. 

TRANSVERSE SECTION OF LAmiw.-Dorsiventral; 

V-shaped with small median adaxial groove; margins turned slightly inversely. Adaxial surface 

slightly ridged; abaxial surface furrowed about 

one-half thickness of blade. Epidermis: cells on 

both surfaces rounded to dome shaped; thin walled. 

Subjacent to adaxial epidermis are sclerenchyma 

strands interspersed with parenchyma cells. Below 

sclerenchyma strands is a layer of parenchyma cells. 

Cuticle: thick on adaxial surface and slightly 

ridged; thinner on abaxial surface and slightly 

ridged. Stomata: present in abaxial furrows; no 

substomatal chamber observed. Mesophyll: three 

layers of palisade cells grading into fairly compact 

spongy tissue. Two to three layers of large translucent cells arranged radially above vascular 

bundles and furrows. Vascular bundles: 25; no 

commissural bundles observed. One to three large 

vessels present. Two phloem units lying laterally 

in flanges of Y-shaped abaxial girder. Adaxial cap 

present on each vascular bundle. Some abaxial 

sclerenchyma strands present between abaxial epidermis and a layer of parenchyma. Bundle sheath 

completely surrounding vascular bundle. Crystals: 

none observed. Tannins: present. 

Vellozia resinosa Martius ex Schultes f. 

FIGURE 8a-c; PLATES 30g-h, 31a 

SPECIMENS EXAMINED.-Maguire et al. 44694. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

square to rectangular on both surfaces; thin walled. 

Stomata: paracytic, 21 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove; margins 

turned slightly inversely. Adaxial surface slightly 

undulating; abaxial surface furrowed about onehalf thickness of blade. Epidermis: adaxial cells 

rectangular to dome shaped; abaxial cells rounded 

to dome shaped. Cells on both surfaces thin walled. 

Subjacent to adaxial epidermis are two to three 

layers of parenchyma cells interspersed with sclerenchyma strands. Below is a distinct layer of 

parenchyma cells. Cuticle: thick on adaxial surface; about one-half as thick on abaxial surface; 

smooth on both surfaces. Stomata: present in furrows; associated with hairlike protrusions; substomata1 chamber present. Mesophyll: three to 

four layers of palisade cells grading into compactly 

arranged spongy tissue. Two to three layers of 

large translucent cells arranged radially above 

vascular bundles and furrows. Vascular bundles: 

33; few commissural bundles observed. Two 

phloem units laterally arranged in flanges of 

Y-shaped abaxial girder. Adaxial cap on each vascular bundle. Few abaxial sclerenchyma strands 

present. Bundle sheath completely surrounding 

each vascular bundle. Crystals and tannins: none 

observed. 

Vellozia sellovii Seubert in Martius 

PLATE 46c 

SPECIMENS ExAMINED.-Pereira 2677-Pabst 35 13. 

SURFACE VrEw.-Hairs: absent. Epidermis: adaxial cells rounded; thin walled. Abaxial cells square 

to rectangular; thin walled. Stomata: paracytic, 12 

X 6 pm; present on abaxial surface and adaxial 

surface. 

TRAKSVERSE SECTION OF LAMINA.-Dorsiven tral; 

V-shaped with median adaxial groove. Both adaxial 

and abaxial surfaces slightly undulating. Epidermis: cells on adaxial surface mostly rounded to 

dome shaped, few conical; thin walled. Cells on 

abaxial surface rounded to dome shaped; thin 

walled. Cuticle: thick on adaxial and abaxial surfaces, slightly ridged. Stomata: present on adaxial 

and abaxial surfaces; large substomatal chamber 

present on both surfaces. Mesophyll: four to five 

layers of indistinct palisade tissue grading into 

compactly arranged spongy tissue. Two to three 

layers of large translucent cells arranged in midvein region and radially above each vascular bundle. Vascular bundles: 19; few commissural bundles 

observed. One to two large vessels present in each 

vascular bundle, mostly one. Two phloem units 

lying laterally in flanges of Y-shaped abaxial girder. 

Inverted V-shaped adaxial cap present on each 

vascular bundle. No abaxial sclerenchyma strands 

or layers observed. Distinct layer of parenchyma 

cells immediately subjacent to abaxial epidermis. 

Bundle sheath completely surrounding each vascular bundle. Crystals and tannins: none observed. 

NOTE: The mesophyll in this species is poorly 

differentiated and therefore resembles the mesophyll pattern encountered in Barbacenia species. 

However, all other characteristics indicate that it 

is in the correct genus. 

Vellozia squalida Martius ex Schultes f. 

PLATE 29a-c 

SPECIMENS EXAMINED.-W. A. Archer and M. 

Barreto 4937; Duarte 6401. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

rectangular and thin walled on both surfaces. Stomata: paracytic, 27 X 15 pm; present on the abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove; margins turned very slightly inversely. Adaxial surface 

slightly undulating; abaxial surface furrowed 

about one-half thickness of blade. Epidermis: adaxial cells rounded, thin walled. Abaxial cells dome 

shaped to conical, thin walled. Subjacent to adaxial 

epidermis are two to three layers sclerenchyma 

strands interspersed with two to three rows of 

parenchyma cells. Below is a distinct layer of 

parenchyma cells. Cuticle: thin and smooth on 

adaxial surface; ridged on abaxial surface. Stomata: present in abaxial furrows; small substomatal 

chamber present; furrows lined with epidermal 

projections. Mesophyll: three layers of palisade 

cells grading into compactly arranged spongy tissue. One to two layers of large translucent cells 

arranged radially above vascular bundles. Vascular 

bundles: 27; few commissural bundles observed. 
NUMBER 15 39 

One to two large vessels, mostly one, present in 

each vascular bundle. Two phloem units lying 

laterally in flanges of Y-shaped abaxial girder. 

Adaxial cap present on each vascular bundle. 

Bundle sheath completely surrounding each vascular bundle. Abaxial sclerenchyma strands present 

at entrances to furrows. Crystals and tannins: none 

observed. 

Vellozia swallenii L. B. Smith 

FIGURE 13a-b; PLATE 33b-c 

SPECIMENS EXAMINED.-G. Fonjecs 11; Pires et 

al. 9712, 9655. 

SURFACE VIE\+'.-Hairs: absent. Epidermis: cells 

rectangular on both surfaces, thin walled. Stomata: 

paracytic, 15 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped, turned slightly inversely at margins; 

lamina thickens about halfway to margins, then 

tapers off. Adaxial surface slightly undulating; 

abaxial surface furrowed about one-fourth thickness of blade. Epidermis: abaxial and adaxial cells 

dome shaped to rectangular; cells on both surfaces 

thin walled. Four to five layers of sclerenchyma 

cells subjacent to adaxial epidermis and interspersed with four to five layers of parenchyma 

cells. Cuticle: thin and slightly ridged on both 

surfaces. Stomata: present in abaxial furrows; 

small substomatal chamber present. Mesophyll: 

two to three layers of large palisade cells present; 

four to five layers of translucent cells arranged 

radially above vascular bundles. Two to three 

layers of palisade tissue subjacent to translucent 

cells and grading into compactly arranged spongy 

tissue. One to two layers of large translucent cells 

arranged radially below vascular bundles. Vascular 

bundles: 13-33; no commissural bundles observed. 

One to three large vessels present in each vascular 

bundle. Two phloem units arranged laterally in 

flanges of Y-shaped abaxial girder. Adaxial cap 

present on each vascular bundle. Three to four 

layers of abaxial sclerenchyma strands interspersed 

with three to four layers of parenchyma cells. Crystals and tannins: none observed. 

The thickening of the lamina in the 

intercostal portion of the leaf is very characteristic 

of all the specimens examined. 

NOTE: 

Vellozia taxifolia Martius ex Schultes f. 

FIGURE 8f: PLATE 30a-c 

SPEcihrLNa ExAmhm.-Pereira 1290; Ynes Mexia 

4289. 

SURFACE VIE\+'.-Hairs: absent. Epidermis: cells 

rounded; few rectangular on both surfaces; walls 

thin. Stomata: paracytic, 21 X 12 pm; present on 

abaxial surface. 

TRAMVERSE SECTION OF LA1CrINA.-Dorsiventral; 

V-shaped with shallow median adaxial groove; 

margins turned slightly inversely. Adaxial surface 

slightly undulating; abaxial surface furrowed 

about one-third thickness of blade. Epidermis: 

adaxial cells dome shaped to rectangular; abaxial 

cells rounded to dome shaped; cells on both surfaces 

have thin walls. One layer of sclerenchyma strands 

subjacent to adaxial epidermis interspersed with 

few parenchyma cells. Below sclerenchyma is a 

layer of distinct parenchyma cells. Cuticle: fairly 

thick and ridged on adaxial and abaxial surfaces. 

Stomata: present in abaxial furrows; associated 

with small hairlike projections; substomatal chamber virtually absent. Mesophyll: three- to fourlayered palisade tissue grading into compactly 

arranged spongy tissue. One to two layers of large 

translucent cells arranged radially above vascular 

bundles and furrows. Vascular bundles: 15-17; no 

commissural bundles observed. One to two large 

vessels present in each vascular bundle. Two 

phloem units lying laterally in flanges of U- or 

Y-shaped abaxial girder. Adaxial cap present on 

each vascular bundle. Bundle sheath completely 

surrounding each vascular bundle. Crystals: none 

observed. Tannins: present. 

NOTE: The abaxial sclerenchyma recalls the 

type in Madagascan species which I originally 

termed the Xerophyta Type (cf. Ayensu 1969). 

Vellozia tenella Martius ex Schultes f. 

PLATE 44d 

SPECIMENS EXAMINED.-Damazio Herb. No. 

91 185, 91 192; Pereira 2549-Pabst 3385; Herb. 

Schwache 6470, 6749. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular on both surfaces; thin walled. 

Stomata: paracytic, 15 X 9 pm; present on abaxial 

surface. 
40 ShIITHSONIAX CONTRIBUTIONS TO BOTANY 

FIGURE 13.-a-b, Vellozia sruallenii (Pires 9712) ; c-e, V. dasypus (Rose and Russell 19697). 
NUMBER 15 41 

- 

FIGURE 14.-a-c, Velloziu carunculuris (Heringer 5265) ; d-e, V. exilis (Irwin et al. 9507). 
42 ShIITHSONIAN CONTRIBUTIONS TO BOTANY 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small adaxial median groove. Adaxial surface slightly undulating; abaxial surface 

furrowed about one-half thickness of blade. Epidermis: cells on both surfaces rounded to dome 

shaped and thin walled. Subjacent to adaxial 

epidermis is a layer of sclerenchyma cells, below 

which is a distinct layer of parenchyma. Cuticle: 

slightly thickened and ridged on adaxial and abaxial surfaces. Stomata: present in abaxial furrows; 

substomatal chamber virtually absent. Mesophyll: 

two layers of palisade tissue grading into loosely 

arranged spongy tissue. One to two large palisade 

cells arranged radially above vascular bundles and 

furrows of Damazio H. N. 91 192. Vascular bundles: 

12-29; no commissural bundles observed. One to 

two large vessels in each vascular bundle. Two 

phloem units lying laterally in flanges of U-shaped 

abaxial girder. Adaxial cap present on all vascular 

bundles. Bundle sheath completely surrounding 

each vascular bundle. Crystals: none observed. 

Tannins: present. 

NOTE: Abaxial sclerenchyma girder recalls the 

Xei-ophyta Type (cf. Ayensu 1969). 

Vellozia teres L. B. Smith & Ayensu 

PLATE 44e 

SPECIMENS EXAJIINED.-Irwin, Maxwell, and 

Wasshausen 21000. 

SURFACE VIEw.-Hairs: papillae present. Epidermis: cells square to rectangular on both surfaces; 

thin walled. Stomata: paracytic; 21 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove; margins 

turned slightly inversely. Adaxial surface slightly 

undulating; abaxial surface furrowed about onehalf thickness of blade. Epidermis: adaxial cells 

dome shaped to conical, thin walled. Abaxial cell? 

rounded, interspersed with some sclerenchyma 

strands. Subjacent to adaxial epidermis are one to 

two layers parenchyma cells interspersed with 

sclerenchyma strands. Below is a distinct layer of 

parenchyma cells. Cuticle: thin on both adaxial 

and abaxial surfaces; ridged. Stomata: present on 

abaxial furrows; substomatal chamber not observed. Mesophyll: palisade tissue not very distinct, 

grading into loosely arranged spongy tissue. One 

to two layers of large translucent cells arranged 

radially above vascular bundle and furrows. Vascular bundles: 28; no commissural bundles ob. 

served. One to two large vessels present in each 

vascular bundle. Two phloem units arranged laterally in flanges of large U-shaped abaxial girder. 

Small adaxial cap present on each vascular bundle. 

A layer of abaxial sclerenchyma band present. 

Crystals and tannins: none observed. 

Vellozia tragacantha Martius ex Seubert 

FIGURE 6a-b 

SPECIMENS EXAMINED.-H. S. Irwin 2495; Claussen 13, 1794. 

SURFACE VIEw.-Hairs: few present in small 

tufts. Epidermis: cells mostly square to rectangular 

on both surfaces, few rounded, thin walled. Stomata: paracytic, 15 X 9 pm; present on abaxial 

and adaxial surfaces. 

TRANSVERSE SECTION OF LA;LIINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Adaxial and abaxial surfaces only very slightly undulating. Epidermis: adaxial cells rounded to dome 

shaped, few conical; cells larger in median adaxial 

groove. Xbaxial cells rounded to dome shaped. 

Cuticle: thin and slightly ridged on adaxial surface; thick and slightly ridged on abaxial surface. 

Stomata: present on abaxial and adaxial surfaces; 

substomatal chamber present; distinctly sunken. 

Mesophyll: four to five layers of palisade cells 

grading into compactly arranged spongy tissue. 

Two layers of large translucent tissue arranged 

radially above midvein and some vascular bundles. 

Vascular bundles: 8; no commissural bundles observed. One to two large vessels present in each 

vascular bundle. Two phloem units lying laterally 

in flanges of Y-shaped abaxial girder. Adaxial cap 

present on each vascular bundle, sometimes as 

large as abaxial girder. Bundle sheath completely 

surrounding each vascular bundle. Crystals and 

tannins: none observed, 

Vellozia triquetra Pohl 

PLATE 44f 

SPECIMENS ExAhfINED.-Gardner 2735. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

round to rectangular on both surfaces; walls thin. 
NUMBER 15 43 

C 

- f 

FIGURE 15.-a-c, Vellozia candida (Carauta 1868); d-f, V. cachimbensis (Pires et al. 6235). 

Stomata: paracytic, 18 X 12 pm; present on abax- sclerenchyma strands interspersed with parenchyma 

ial surface. cells. Below sclerenchyma strands is a distinct layer 

TRANSVERSE SECTION OF LAMINA.-DOrSiventral; of parenchryma cells. Cuticle: thin and slightly 

roughly V-shaped with lamina highly corrugated. ridged on adaxial and abaxial surfaces. Stomata: 

Both surfaces deeply undulating. Epidermis: adax- present in abaxial surface; flush with epidermis; 

ial and abaxial cells rounded to dome shaped, substomatal chamber present. Mesophyll: two to 

thick walled. Subjacent to adaxial epidermis are three layers of indistinct palisade tissue grading 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

into compactly arranged spongy tissue. Some large 

translucent cells present between major veins. 

Large lacunae present in mesophyll especially 

toward abaxial side. Vascular bundles: two types 

present; major veins 15; minor veins 86; no commissural bundles observed. Major veins with one 

to two large vessels; two phloem units lying 

laterally in flanges of abaxial crescentiform or 

U-shaped girder; adaxial cap present on each vascular bundle and extending almost to adaxial 

epidermis. Each vascular bundle completely surrounded by a bundle sheath. Minor veins have 

one to two larger vessels and a single phloem unit; 

few sclerenchyma strands, completely surrounded 

by bundle sheath. Some abaxial strands present 

between abaxial epidermis and a single layer of 

parenchyma cells. Crystals: none observed. Tannins: present. 

NOTE: This species is in every respect similar 

to Vellozia plicata. 

Vellozia variabilis Martius ex Schultes f. 

PLATES 12a-b, 27a-f 

SPECIMENS ExAMINm-Pereira 2857-Pabst 3493; 

Brade 13893; Lima 58-3003; Irwin et al. 9867; 

Irwin et al. 9915, 11709, 9809, 9726; Maguire 

49262. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular on both surfaces, thin walled. 

Stomata: paracytic, 15 X 12 pm; present on abaxial 

surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with slight median adaxial groove. Adaxial surface slightly undulating: abaxial surface 

furrowed about one-third to one-half thickness 

of blade. Epidermis: adaxial cells dome shaped to 

rectangular; abaxial cells rounded to dome shaped; 

cells on both surfaces are thin walled. Subjacent 

to adaxial .epidermis are two layers of parenchyma 

cells interspersed with sclerenchyma strands. Below 

parenchyma and sclerenchyma is a distinct layer 

of translucent parenchyma cells. Cuticle: thick and 

ridged on adaxial surface; thin and ridged on 

abaxial surface. Stomata: present in abaxial furrows; small substomatal chamber present, small 

hairs present in furrows. Mesophyll: distinctly 

three-layered palisade cells grading into compactly 

arranged spongy tissue. Two to three layers of 

large translucent cells arranged radially above vascular bundles and on furrows. Vascular bundles: 

21-46; few commissural bundles observed. One to 

two large vessels present, mostly one. Two phloem 

units lying laterally in flanges of Y-shaped abaxial 

girder. Adaxial cap present on each vascular bundle. Bundle sheath completely surrounding each 

vascular bundle. A layer of abaxial sclerenchyma 

present between abaxial epidermis and a distinct 

layer of parenchyma. Crystals: none observed. 

Tannins: present. 

Vellozia verruculosa Martius ex Schultes f. 

PLATES 19a-b, 26a-c 

SPECIMENS EXAMINED.-L. 0. Williams 8 112. 

SURFACE VrEw.-Hairs: few papillae present. 

Epidermis: cells square to rectangular on both 

surfaces; few rounded on abaxial surface; walls 

thin on both surfaces. Stomata: paracytic, 12 X 6 

pm; present on adaxial and abaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Adaxial surface slightly undulating; abaxial surface 

furrowed about one-half thickness of blade. Epidermis: cells on both surfaces rounded to rectangular; thin walled. One to two layers of parenchyma 

cells interspersed with sclerenchyma strands occur 

between adaxial epidermis and a distinct layer of 

parenchyma cells. Cuticle: fairly thick on both 

surfaces and slightly ridged. Stomata: present in 

abaxial furrows as well as on abaxial and adaxial 

surfaces. Substomatal chamber present; stomata 

almost flush with surface. Mesophyll: three- to 

four-layered palisade tissue grading into compactly 

arranged spongy tissue. One to two layers of large 

translucent cells arranged radially above vascular 

bundles and on furrows. Vascular bundles: 21; 

few commissural bundles observed. One to two 

large vessels present in each vascular bundle, 

mostly one. Two phloem units arranged laterally 

in flanges of v- or U-shaped abaxial girder. Adaxial 

cap present on each vascular bundle. Some abaxial 

sclerenchyma strands present between abaxial epidermis and a layer of parenchyma cells. Bundle 

sheath conipletely surrounding each vascular 

bundle. Crystals: none observed. Tannins: present. 
NUMBER 15 45 

Vellozia virgata Goethart & Henrard 

FIGURE 3a-b; PLATE 46d 

SPECIMENS ExAMINED.-schwacke 5889; Porto 

543. 

SURFACE Vw.w.-Hairs: few papillae present. 

Epidermis: cells square to rectangular on both 

surfaces; some rounded on adaxial surface; walls 

thin on both surfaces. Stomata: paracytic, 12 X 3 

pm; present on abaxial and adaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

widely V-shaped with large median adaxial groove. 

Adaxial and abaxial surfaces slightly ridged; very 

small furrows in Porto 543. Epidermis: cells 

rounded to dome shaped on both surfaces; thin 

walled. Cuticle: slightly thickened and ridged on 

both surfaces; a little thicker on abaxial surface. 

Stomata: present on abaxial and adaxial surfaces; 

substomatal chamber present. Mesophyll: palisade 

tissue indistinct; spongy tissue compact. Vascular 

bundle: 7-10; few commissural bundles observed. 

One to two large vessels present in each vascular 

bundle. Two phloem units laterally arranged in 

flanges of Y-shaped abaxial girder. Adaxial cap 

present on each vascular bundle. Bundle sheath 

completely surrounding each vascular bundle. 

Crystals and tannins: not observed. 

Velloziu wasshausenii L. B. Smith 

PLATE 45d 

SPECIMENS EXAhfINED.-IrWin et al. 20318. 

SURFACE ViEw.-Hairs: few papillae present on 

abaxial surface. Epidermis: cells rectangular on 

adaxial surface; cells square to rectangular, 

rounded on abaxial surface; walls thin on abaxial 

and adaxial surfaces. Stomata: paracytic, 15 X 9 

pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMISA.-Dorsiventral; 

widely V-shaped, intercostal region significantly 

expanded. Adaxial surface undulating; abaxial 

surface furrowed about one-half thickness of blade. 

Epidermis: cells on both surfaces rounded to dome 

shaped. Subjacent to adaxial epidermis is a layer 

of sclerenchyma strands interspersed with parenchyma cells. Below sclerenchyma is a distinct layer 

of parenchyma cells. Cuticle: thick and smooth on 

adaxial surface; thinner and smooth on abaxial 

surface. Stomata: present in abaxial furrows; substomata1 chamber not observed. Mesophyll: threeto four-layered palisade cells changing abruptly 

into compactly arranged spongy tissue. Two to 

three large translucent palisade cells radially arranged above vascular bundles and on furrows. 

Vascular bundles: 17; no commissural bundles 

observed. One to two large vessels present in each 

vascular bundle. Two phloem units laterally arranged in flanges of Y-shaped abaxial girder. Abaxial sclerenchyma layer present. Bundle sheath 

completely surrounding each vascular bundle. 

Crystals and tannins: none observed. 

Section 11: Radia 

Vellozia cachimbensis L. B. Smith 

FIGURES 3m-n, 15d-f 

SPECIMENS EXAMINED.-PireS et al. 6235; PireS 

6111. 

SURFACE VIEw.-Hairs: small papillae present in 

clusters on both surfaces. Epidermis: cells square 

to rectangular; few adaxial and abaxial cells 

rounded; thin walled on both surfaces. Stomata: 

paracytic, 15 X 9 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with slight median adaxial groove. Adaxial surface slightly ridged; abaxial surface furrowed 

about one-third thickness of blade. Epidermis: 

adaxial cells rectangular to rounded; abaxial cells 

rounded to dome shaped. Subjacent to adaxial epidermis are one to two rows of parenchyma cells 

interspersed with sclerenchyma strands above 

radially arranged translucent cells on each furrow; 

below parenchyma cells occurs a distinct layer of 

parenchyma cells; few abaxial sclerenchyma 

strands. Cuticle: fairly thick and smooth on adaxial surface; fairly thick and slightly ridged on 

abaxial surface. Stomata: present in abaxial furrows with small substomatal chamber. Mesophyll: 

three to four layers of indistinct palisade tissue 

grading onto compactly arranged spongy tissue. 

One to two layers of large translucent palisade 

cells radially arranged above vascular bundles and 

furrows. Vascular bundle: 2 1-36; commissural 

bundles not observed. Large veins with one to 

three large vessels, mostly one. Two phloem units 

arranged laterally in flanges of Y-shaped abaxial 

girder. Each vascular bundle accompanied by an 
46 ShfITHSONIAN CONTRIBUTIONS TO BOTANY 

abaxial girder and adaxial cap. Bundle sheath 

completely surrounding vascular bundle. Crystals: 

none observed. Tannins: few present in mesophyll. 

NOTE: The adaxial inverted caps are almost 

V-shaped and extend closely to epidermis. 

Vellozia cana Goethart & Henrard 

PLATES 18a, 4513 

SPECIMENS EXAILIINED.-hWin et al. 11734, 12556. 

SURFACE VIEw.--Hairs: absent. Epidermis: adaxial and abaxial cells mostly rectangular, generally 

thin walled; slightly thick on veins. Stomata: paracytic, 18 X 12 pm; observed on both surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with slight median adaxial groove. Adaxial surface evenly undulating; abaxial surface 

furrowed about one-third of blade thickness. Epidermis: cells more or less square shaped, adaxial 

cells slightly larger than abaxial ones; adaxial cells 

evenly interrupted by strands of sclerenchyma. 

Some of abaxial cells replaced by almost continuous band of sclerenchyma. Cuticle: thin on both 

surfaces. Stomata: almost flush with epidermis; few 

on adaxial surface, mostly confined to abaxial surface, specifically in furrows; guard cells thick walled 

with no projecting ledges; substomatal chamber 

present. Mesophyll: adaxial palisade four to six 

layered, occupying about half of mesophyll and 

abruptly changing into spongy tissue. Palisade 

layers alternate with large translucent cells radially 

arranged above vascular bundles and furrows; rest 

of mesophyll compactly arranged and filled with 

chloroplast. Translucent cells radially arranged 

above vascular bundles and furrows. Vascular bundles: 57-59; commissural bundles present. Large 

veins each with one to two large vessels, occasionally three or more. Two phloem units lying laterally in flanges of abaxial Y-shaped girder, Each 

vascular bundle always accompanied by abaxial 

inverted V-shaped sclerenchyma cap, both in turn 

completely surrounded by a distinct bundle sheath. 

Crystals: not observed. Tannins: present but not 

abundant. 

NOTE: The abaxial strands at the wide end of 

the furrows and the aImost horizontal median 

abaxial portion are very distinctive. 

Vellozia candida Mikan 

FIGURE 15a-c; PLATES 34g-h, 35a, 50a 

SPECIMENS EXAMINED.-Carau ta 160; Kuhlman 

8-XI-922; Porto 91195; Jardim Bot., Rio de 

Janeiro 2485. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

rounded, square to rectangular on both surfaces; 

thin walled. Stomata: paracytic, 24 X 15 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with margins turned inversely. Adaxial 

surface undulating; abaxial surface furrowed about 

one-third thickness of blade. Epidermis: adaxial 

cells dome shaped or rectangular; abaxial cells 

rounded to dome shaped; cell walls thin on both 

surfaces. Subjacent to adaxial epidermis are sclerenchyma strands interspersed with parenchyma 

cells. Below is a distinct row of parenchyma cells. 

Cuticle: thick and slightly ridged on adaxial surface; thinner and slightly ridged on abaxial surface. 

Stomata: present on abaxial surface especially in 

furrows; substomatal chamber present; stomata 

Rush with epidermal surface. Mesophyll: threelayered palisade tissue changing abruptly to compactly arranged spongy tissue. Large translucent 

cells arranged radially above vascular bundles. 

Vascular bundles: 25-35; commissural bundles not 

observed. One to two large vessels observed, mostly 

one. Two phloem units laterally arranged in 

flanges of Y- or U-shaped abaxial girder. Each vascular bundle always accompanied by inverted 

adaxial cap and bundle sheath completely surrounding vascular bundles. Crystals: none observed. Tannins: few. 

The large abaxial crescentiform sclerenchyma band is very distinctive. 

NOTE: 

Velloria caput-ardeae L. B. Smith & Ayensu 

PLATE 36a 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

15989. 

SURFACE VIEw.-Hairs: none present. Epidermis: 

adaxial cells mostly square, few rounded; thin 

walled. Abaxial cells rectangular; thin walled. Stomata: paracytic, 18 X 12 pm; present on abaxial 

surface. 
NUMBER 15 47 

FIGURE 16.-a-c, Vellozia burle-murxii (US. 2537159) : 

d-f, V. lithophila (Schultes and Cabrera 18317). 
48 SMITHSONIAS COSTRIBUTIONS TO BOTANY 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove present. Adaxial surface slightly undulating; abaxial 

surface furrowed one-half thickness of blade. Epidermis; cells dome shaped on abaxial and adaxial 

surface; few square on adaxial surface; walls thin 

on both surfaces. Subjacent to adaxial epidermis 

are three to four layers parenchyma cells interspersed with some sclerenchyma strands. Clusters 

of radially arranged parenchyma cells are intermittently present subjacent to adaxial epidermis. 

Below parenchyma cells and sclerenchyma strands 

is a distinct layer of parenchyma cells. Cuticle: 

thin and ridged on abaxial and adaxial surfaces. 

Stomata: present in abaxial furrows; substomatal 

chamber not observed; somewhat protected by 

projections in furrow. Mesophyll: one to two layers translucent palisade cells radially arranged 

above vascular bundles and furrows. Three to four 

layers small palisade cells present between translucent rows; small palisade grading into compact 

spongy tissue. Vascular bundle: 49; few commissural bundles present. One to two vessels present 

in each vascular bundle, mostly one. Two phloem 

units laterally arranged in flanges of thin Y-shaped 

abaxial girder. Adaxial cap present on each bundle, 

but very small. Bundle sheath completely surrounding vascular bundle. Few abaxial sclerenchyma 

strands present. Crystals: none observed. Tannins: 

few present. 

Vellozia dawsonii L. B. Smith 

FIGURE 3g-h; PLATES la-b, 45c, 51a 

SPECIMENS EXAMINED.-hWin et al. 12648. 

SURFACE VIEw.-Hairs: present, tuftlike. Epidermis: adaxial and abaxial cells mostly rectangular. 

Stomata: paracytic, 18 X 15 pm; occur on both 

surfaces but highly concentrated in abaxial furrows. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with slight median adaxial groove and 

abaxially two grooved. Adaxial surface undulating; 

abaxial surface furrowed about one-third of blade 

thickness. Epidermis: cells more or less rectangular shaped on adaxial surface and almost rounded 

on abaxial surface. Abaxial epidermis partially replaced with sclerenchyma fibers. Subjacent to 

adaxial epidermis occur one to two layers of parenchyma cells interspersed with short strands of 

sclerenchyma. Below sclerenchyma strands and 

subjacent paranchyma is a layer of large translucent cells separating subjacent cells from mesophyil. Cuticle: generally thin, but slightly 

thickened around median region. Stomata: mostly 

confined to furrows in abaxial surface, conspicuous 

in places. Mesophyll: adaxial palisade three to four 

layered, occupying a little under half of mesophyll 

and changing abruptly into spongy tissue; three 

to four rows of large translucent cells arranged 

radially above vascular bundles and furrows. Palisade and spongy cells compactly arranged and 

filled with chloroplasts. Mesophyll in median section of leaf composed of large translucent cells, 

bulliform-like. Vascular bundles: 52; commissural 

bundles present. Large veins with one to two wide 

vessels. Two phloem units lying laterally in flanges 

of abaxial Y-shaped girder. Each vascular bundle 

always accompanied by adaxial inverted V-shaped 

sclerenchyma cap; both in turn completely surrounded by a distinct bundle sheath. Crystals: not 

observed. Tannin: few in mesophyll. 

NOTE: The tuftlike appearance of unicellular 

and multicellular macro-hairs is very distinctive. 

Vellozia durnitiana R. E. Schultes 

PLATES 2a-b, 45a 

SPECIMENS EXAMINED.-Barriga 13804; Cabrera 

and Schultes 15050A. 

SURFACE VrEw.-Hairs: long and narrow, unicellular and multicellular, often with coalesced 

bases, observed on abaxial surface only. Epidermis: 

adaxial and abaxial cells uniformly rectangular. 

Stomata: tetracytic, 21 X 18 pm; present on abaxial 

surface only. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with shallow median adaxial groove but 

tending to shallowly corrugate. Adaxial surface 

gently undulating. Abaxial surface deeply furrowed about two-thirds of blade thickness. Epidermis: cells square shaped, adaxial cells slightly 

larger than abaxial ones; cells somewhat translucent. Subjacent to epidermis occur two to three 

layers of parenchyma cells followed by strands of 

fibers. Below these layers and fibers occurs a layer 

of large translucent cells separating epidermal 

and subepidermal layers of cells from mesophyll. 
NUMBER 15 49 

Cuticle: thin on both surfaces. Stomata: confined 

to abaxial furrows only; almost flush with epidermis. hlesophyll: adaxial palisade four- to fivelayered, occupying about half of mesophyll, and 

abruptly changing into spongy tissue; three to 

five rows of translucent layers radially arranged 

above vascular bundles and abaxial furrows. Vascular bundles: 18-33; commissural bundles few. 

Large veins each with one to two wide vessels, 

mostly one. Two phloem units lying laterally in 

flanges of abaxial Y-shaped girder. Each vascular 

bundle always accompanied by adaxial inverted 

V-shaped sclerenchyma cap and abaxial Y-shaped 

girder; both in turn completely surrounded by a 

distinct bundle sheath. Crystals: square shaped 

and styloid-like. Tannin: present especially in 

palisade cells. 

NOTE: The two collections are identical in 

their anatomy except that Barriga 13804 has a 

slightly narrower leaf. 

Vellozia hirsuta Goethart & Henrard 

FIGURE 4d; PLhTE 37b 

SPECIMENS ExAmNm.-Hatschbach 28093; L. B. 

Smith and Ayensu 15999. 

SURFACE VIEw'.-Hairs: some present in clumps 

on adaxial and abaxial surfaces; multicellular. 

Epidermis: cells square; few rounded and rectangular on both surfaces. Cells thin walled. Stomata: 

tetracytic, 18 X 15 pm; present on abaxial and 

adaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with median adaxial groove; turned 

slightly inversely at margins. Surface slightly undulating on adaxial surface; abaxial surface furrowed about one-fourth to one-half thickness of 

blade. Epidermis: adaxial cells dome shaped; few 

conical. Abaxial cells rounded. Walls thin on abaxial and adaxial surface. Subjacent to adaxial epidermis are two to three layers of parenchyma cells, 

interspersed with sclerenchyma strands. Below this 

is a complete layer of parenchyma cells. Stomata: 

present in abaxial furrows and on abaxial and 

adaxial surfaces; substomatal chamber present. 

Mesophyll: three layers palisade tissue grading into 

compact spongy tissue. One to two layers large 

translucent palisade cells arranged radially above 

vascular bundles; large number translucent palisade cells above midvein. Vascular bundles: 35; 

few commissural bundles observed. One to two 

large vessels present in each vascular bundle; 

mostly one. Two phloem units arranged laterally 

in flanges of Y-shaped abaxial girder. Each vascular 

bundle has an adaxial cap or partial girder. Some 

abaxial sclerenchyma strands present on abaxial 

corners of furrows. Bundle sheath completely surrounding each vascular bundle. Crystals and tannins: none observed. 

Velloria lanata Pohl 

PLATES 3b, 17b, 45e 

SPECIMENS EXAMINED.-Pires et al. 9895; Veralucia and Graziella 26/1/1968. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

rectangular, thin walled. Stomata: tetracytic, 21 X 

12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

deeply V-shaped with median adaxial groove. Adaxial surface slightly undulating; abaxial surface 

furrowed about one-half thickness of blade. Epidermis: adaxial cells rounded to rectangular; abaxial cells rounded to dome shaped. One to two layers 

parenchyma cells subjacent to epidermis and interspersed with sclerenchyma strands. Below parenchyma and sclerenchyma is a distinct layer of 

parenchyma cells. Cuticle: thick and slightly 

ridged on adaxial surface; very thin on abaxial 

surface. Stomata: present in furrows; large substomata1 chamber present. Mesophyll: three-layered 

palisade tissue grading into compact spongy tissue. 

One to two layers of large translucent cells arranged radially above vascular bundles and furrows. Vascular bundles: 37; few commissural 

bundles observed. One to three large vessels, mostly 

one. Two phloem units lying laterally in flanges 

of Y-shaped abaxial girder. Each vascular bundle 

accompanied by an adaxial cap. Abaxial strands 

present, sometimes displacing abaxial epidermis. 

Bundle sheath completely surrounding each vascular bundle. Crystals: none observed. Tannins: few 

present. 

Velloria lithophila Schultes 

FIGURES 2b, 16d-f; PLATE 2~-d 

SPECIMENS EXAMINED.-sChUlteS and Cabrera 

18317. 
50 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

SURFACE VIEw.-Hairs: present in tufts, Epidermis; cells square to rectangular; thin walled. 

Stomata: paracytic, 21 X 15 pm; present on abaxial surface. 

TRAMVERSE SECTION OF LAhrINA.-Dorsiventral; 

V-shaped with margins turned slightly inversely. 

Adaxial surface slightly undulating, more so near 

midvein; abaxial surface furrowed one-half the 

thickness of blade; furrows wide and arch shaped. 

Epidermis: adaxial cells mostly dome shaped, with 

outer tangential wall concave. Abaxial cells rectangular to slightly dome shaped. Walls thin on 

both surfaces. One to two layers of parenchyma 

cells subjacent to adaxial epidermis and interspersed with sclerenchyma strands. Below sclerenchyma and parenchyma is a distinct single layer 

of parenchyma cells. Cuticle: very thick and 

slightly ridged on adaxial surface; one-half as thick 

on abaxial surface and slightly ridged. Stomata: 

present in abaxial furrows, substomatal chambers 

not observed. Mesophyll: two to three layers of 

palisade cells grading into compact spongy tissue. 

One to two layers of large translucent cells arranged radially above vascular bundles and on 

furrows. Vascular bundles: 49; few commissural 

bundles observed. One to two large vessels present 

in each vascular bundle, mostly one. Two phloem 

units lying laterally in flanges of Y-shaped abaxial 

girder; adaxial cap present on each vascular bundle. Bundle sheath completely surrounding each 

vascular bundle. Crystals: none observed. Tannins: 

present. 

NOTE: This species exhibits rather wide abaxial furrows resembling archs of large gates. 

Vellozia machrisiana L. B. Smith 

FIGURE 17d-f; PLATES 4a-b, 35b-d 

SPECIMENS EXAMINED.-Irwin et al. 12454; Irwin 

et al. 12697. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

rounded to rectangular on both surfaces; thin 

walled. Stomata: paracytic, 15 X 15 pm; present 

on abaxial surface. 

TRANSVERSE SECTION OF LAAxINA.-Dorsiventral; 

V-shaped with small median adaxial groove; turned 

slightly inversely at margins. Adaxial surface 

slightly undulating; abaxial surface furrowed 

about one-half thickness of blade and narrow. 

Epidermis: adaxial and abaxial cells rounded to 

dome shaped; thin walled, Subjacent to adaxial 

epidermis occur one to two layers of parenchyma 

cells interspersed with one- to two-layered strands 

of sclerenchyma. Below parenchyma layers and 

sclerenchyma strands is a distinct layer of parenchyma cells. Cuticle: thick on both surfaces; 

slightly ridged on abaxial surface; smooth on 

adaxial surface. Stomata: present in abaxial furrows; almost flush with surface; small substomatal 

chamber observed. Mesophyll: three to four layers 

of indistinct palisade tissue grading into compact 

spongy tissue. Two to three layers of large translucent cells arranged radially above vascular bundles and furrows. Vascular bundles: 28-31; some 

coinmissural bundles observed. One to three large 

vessels present in each vascular bundle, mostly 

one. Two phloem units arranged laterally in flanges 

of Y-shaped abaxial girder, adaxial cap present on 

each vascular bundle. Abaxial strands present; separated from spongy mesophyll by a distinct layer 

of parenchyma cells. Each vascular bundle is completely surrounded by a bundle sheath. Crystals 

and tannins: none observed. 

NOTE: The two specimens are identical; however, the abaxial portion of the midvein is flattened 

in Irwin et al. 12697, while it is pointed in Irwin 

et al. 12454. 

Vellozia maculata Goethart & Henrard 

FIGURE 30p; PLATES 3c-d, 45f 

SPECIMENS EXAMINED.-Glaziou 22218-a. 

SURFACE VIEw.-Hairs: few present. Epidermis: 

cells square to rectangular on both surfaces; thin 

walled. Stomata: paracytic, 18 X 18 pm; present on 

abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped; margins turned inversely. Adaxial surface distinctly undulating especially in median 

region; two small adaxial grooves present laterally 

to midvein. Abaxial surface with fairly wide furrows about one-third to one-half thickness of blade. 

Epidermis: adaxial cells dome shaped to rectangular; abaxial cells rounded to dome shaped. Cells 

on both surfaces thin walled. Subjacent to adaxial 

epidermis occur layers of parenchyma cells interspersed with schlerenchyma strands. Below these 

layers is a distinct layer of parenchyma cells. 
NUMBER 15 51 

Cuticle: thick on adaxial surface; about one-half 

as thick on abaxial surface. Cuticle on both surfaces slightly ridged. Stomata: present in furrows; 

substomatal chamber virtually absent. Mesophyll: 

two to three layers of indistinct palisade tissue 

grading into compact spongy tissue. One layer 

large translucent cells arranged radially above vascular bundles; one to two layers above furrows. 

Vascular bundles: 21; no commissural bundles 

observed. One to two large vessels present in each 

vascular bundle. Two phloem units arranged laterally in flanges of Y-shaped abaxial girder. Adaxial girder present on all vascular bundles and 

almost extended to adaxial -epidermis. Some abaxial sclerenchyma strands present. Bundle sheath 

completely surrounding each vascular bundle. 

Crystals: none observed. Tannins: present. 

NOTE: The lateral extensions of the outer 

edges of the abaxial furrows serve as valves encasing the areas where stomata are present. The 

extensions also recall similar structures in Vellozia 

hemispherica and V. burle-marxii. 

Vellozia riedeliana Goethart & Henrard 

FIGURE 4e; PLATE 46b 

SPECIMENS EXAMINELL-L. B. Smith and Ayensu 

15983. 

SURFACE VIEw.-Hairs: numerous, -present in 

clusters. Epidermis: adaxial cells square to rectangular on both surfaces; some rounded on abaxial surface. Cells thin walled. Stomata: paracytic, 

15 X 3 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-DOrSiVentral; 

V-shaped with median adaxial groove. Adaxial 

surface undulating; sometimes deeply ridged abaxial surface furrowed about one-third thickness of 

blade. Epidermis: adaxial cells rounded to rectangular; thin walled; some papillae present on 

adaxial surface. Abaxial cells dome shaped; thin 

walled. Subjacent to adaxial epidermis occur two 

to three layers of parenchyma cells interspersed 

with sclerenchyma strands. Below is a distinct layer 

of parenchyma cells. Cuticle: very thick and fairly 

smooth on adaxial surface; thin and ridged on 

abaxial surface. Stomata: present in abaxial furrows; small substomatal chamber present. Stomata 

flush with or slightly above epidermal surface. 

Mesophyll: three layers of distinct palisade tissue 

grading into compactly arranged spongy tissue. 

Two to three layers of large translucent cells arranged radially above vascular bundles. Vascular 

bundles: 30; few commissural bundles observed. 

One large vessel present in each vascular bundle. 

Two phloem units arranged laterally in flanges of 

Y-shaped abaxial girder. Fairly large adaxial cap 

present on each vascular bundle. Few abaxial sclerenchyma strands present on abaxial corners of 

furrows. Bundle sheath completely surrounding 

each vascular bundle. Crystals and tannins: none 

observed. 

Vellozia tubiflora A. Richard 

FIGURE 1Ta-c; PLATES Sje-h, 38a-b 

SPECIMENS ExAMINED.-Foldats 3526; Maguire et 

al. 29156, 29245, 30703, 30835, 30941, 30955, 31003, 

31679, 31777, 32307, 33532, 35131, 35136, 35248, 

36051, 43855, 45918; L. Williams 12974, 13459, 

16022; Steyermark 93208, 60326; Steyermark and 

Aristiquita 54; Steyermark and Nilsson 554; Tamayo 3105; Tate 1196; Lasser 1777. 

SURFACE VIEw.-Hairs: present on abaxial surface of most specimens. Epidermis: cells square to 

rectangular on both surfaces, thin walled. Stomata: 

paracytic, 21 X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventra1; 

V-shaped with margins turned slightly inversely; 

small median adaxial groove present. Adaxial surface slightly undulating; abaxial surface furrowed 

about one-half thickness of blade; furrows may be 

wide or narrow. Epidermis: adaxial cells rectangular; abaxial cells rounded to dome shaped. Walls 

thin on both surfaces. Subjacent to adaxial epidermis are one to three layers of parenchyma cells 

interspersed with sclerenchyma strands. Below is 

a distinct layer of parenchyma cells. Cuticle: thick 

on both adaxial and abaxial surfaces. Stomata: 

present in abaxial furrows; substomatal chamber 

present. Mesophyll: three to four layers of palisade 

cells grading into compactly arranged spongy 

tissue. One to two layers of large translucent cells 

arranged radially above vascular bundles and on 

furrows. Vascular bundles: 13-55; no commissural 

bundles observed. One to two large vessels present. 

Two phloem units lying laterally in flanges of 

Y-shaped abaxial girder extending to epidermis. 

Adaxial cap present on each vascular bundle. Few 
52 SMITHSOXIAS COSTRIBUTIONS TO BOTANY 

e 

FIGURE lT.-a-c, Vellozia tubiflora (hlaguire et al. 35131) ; 

d-f, T’. rnachrisiann (Irwin et al. 12464). 

abaxial strands present. Bundle sheath completely 

surrounding each vascular bundle. Crystals: present. Tannins: few observed. 

NOTE: The diagnostic features that characterire this species were found to be very uniform in 

all the specimens examined. This observation is 

indicative of the reliability of the leaf characters 

in the identification of species. 
NUMBER 15 

Barbacenia 

Subkey I 

Barbacenia delicatula L. B. Smith & Ayensu 

PLATE 46f 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

15975. 

SURFACE VIEw.-Hairs: none present. Epidermis: 

adaxial cells mostly rectangular, few square and 

rounded; thin walled. Abaxial cells rectangular; 

thin walled. Stomata: paracytic, 15 X 16 pm; 

present on abaxial surface. 

TRANSVERSE SECTION OF LA.\IINA.-Isolateral; 

V-shaped; margins of lamina turned slightly inversely. Adaxial and abaxial surfaces uniformly 

smooth with slight grooves separating each vascular 

bundle. Epidermis: adaxial cells rounded to dome 

shaped; thick walled. Abaxial cells rounded; thin 

walled. Subjacent to adaxial epidermis is a distinct 

layer of parenchyma cells. Following parenchyma 

cell layer are clusters of sclerenchyma strands surrounded by one-layered parenchyma cells; usually 

occur above vascular bundles. Other smaller sclerencliyina strands observed. Cuticle: thin on both 

surfaces and slightly ridged. Stomata: present on 

abaxial surface; small substomatal chamber present; flush with surface but appearing in grooves. 

Rfesophyll: spongy tissue uniformly arranged with 

some large translucent cells especially in midvein 

region as well as midportion of mesophyll. Vascular bundles: 27; many commissural bundlei observed. One to two large vessels present in each 

vascular bundle, mostly one. Two phloem units 

lying laterally in flanges of Y-shaped abaxial girder. 

Adaxial cap strictly nonexistent. Each vascular 

bundle completely surrounded by a bundle sheath. 

Crystals and tannins: none observed. 

NOTE: This species is anatomically unique 

because of the lack of direct contact of sclerenchyma adaxial cap on the vascular bundles. The 

sclerenchyma caps that appear above the vascular 

bundles are separated by two to four layers of 

mesophyll cells. 

Barbacenia gaveensis Goethart & Henrard 

FIGURE 18a-c 

SPECIMENS EXAMINED.-L. B. Smith 6445; 

Menezes 2. 

SURFACE VIEW.-Hairs: absent. Epidermis: cells 

are mostly rectangular, few square; thin walled. 

Stomata: paracytic, 21 X 12 pm; present on abaxial 

surface. 

TRANSVERSE SECTION OF LAMINA.-hOkiteral; 

V-shaped with small median adaxial groove, Adaxial and abaxial surfaces slightly ridged, especially 

that of Smith 6445. Epidermis: cells on both surfaces rounded to dome shaped; thin walled. Subjacent to adaxial epidermis is a layer of parenchyma cells, some of which form part of bundle 

sheath. Cuticle: thin on adaxial surface, slightly 

thicker on abaxial surface; both cuticles slightly 

ridged. Stomata: present on abaxial surface; almost 

flush with epidermal surface; small substornatal 

chamber present. Mesophyll: three to four layers 

of indistinct palisade-like tissue grading into 

spongy tissue. Large translucent cells present in 

midportion of mesophyll. Vascular bundles: 38-40; 

no commissural bundles observed. One to two large 

vessels present in each vascular bundle, mostly one. 

Two phloem units arranged laterally in flanges of 

Y-shaped abaxial girder. Inverted Y-shaped adaxial 

girder present on each vascular bundle. Bundle 

sheath completely surrounding each vascular bundle. Crystals: none observed. Tannins: present in 

me sop h y 11. 

Barbacenia gounelleana Beauverd 

FIGURES la-b, 20d-f; PLATES 6d, 38f-h, 51b 

SPECIMENS ExAhfINED.-Emygdio 1428. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular; thin walled. Stomata: paracytic, 27 X 9 pm; present on abaxial surface. 

dorsiventral; V-shaped with slight median adaxial 

groove; turned slightly inversely at margins. Adaxial and abaxial surfaces slightly undulating. Epidermis: adaxial cells dome shaped to rectangular; 

abaxial cells rounded to dome shaped. Cells on both 

surfaces thin walled. Parenchyma layer subjacent 

to adaxial epidermis. Cuticle: thick on adaxial 

surface; thinner on abaxial surface; both slightly 

ridged. Stomata: present on abaxial surface; substomata1 chamber present; stomata almost flush 

with epidermal surface. Mesophyll: three to four 

layers of palisade-like cells grading into looselv 

arranged spongy tissue. Some translucent cells 

TRANSVERSE SECTION OF LAMINA.-somewhat 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

a 

FIGURE 18.-a-c, Barbacenia gaveensis (L. B. Smith 6445); d-f, B. purpurea (US. 151098) 

present in spongy tissue. Vascular bundles: 80; 

few commissural bundles observed. One to two 

vessels present in each vascular bundle, mostly one. 

Two phloem units lying laterally in flanges of 

Y-shaped abaxial girder. Inverted Y-shaped adaxial 

girder present on each bundle. Bundle sheath 

completely surrounding each vascular bundle. 

Crystals: none observed. Tannins: present. 

Barbacenia hatschbachii L. B. Smith & Ayensu 

PLATES Zla, 47b 

SPECIMEKS ExAmmD.-Hatschbach 24284. 

SURFACE VIEw.-Hairs: none present. Epidermis: 

adaxial and abaxial cells square to rectangular; 

thin Tvalled. Stomata: paracytic, 30 X 15 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-ISOlateral; 

V-shaped with slight median adaxial groove. Margins of lamina turned slightly inversely. Both 

adaxial and abaxial surfaces slightly undulating. 

Epidermis: adaxial cells dome shaped; few conical; 

thin walled. Abaxial cells rounded to dome shaped; 

thin walled. Subjacent to adaxial epidermis is a 

distinct layer of parenchyma cells. Cuticle: thick 

and ridged on adaxial and abaxial surfaces. Sto- 
NUMBER 15 55 

mata: present on abaxial surface; small substomatal chamber present. Mesophyll: palisade cells 

nonexistent; mesophyll almost entirely made up 

of spongy tissue with especially large thin-walled 

translucent cells. Vascular bundles: 35; few commissural bundles observed. One large vessel present 

in each vascular bundle. Two phloem units lying 

laterally in flanges of Y-shaped abaxial girder. 

T-shaped adaxial girder present on each vascular 

bundle. Bundle sheath completely surrounding 

each vascular bundle. Crystals: none observed. 

Tannins: few present. 

Barbacenia irwiniana L. B. Smith 

PLATE 48a 

SPECIMENS EXAMINED.-POrtO 1140. 

SURFACE VIEw.-Hairs: few fingerlike projections 

present on abaxial surface. Epidermis: adaxial 

cells square to rectangular; abaxial cells square to 

rectangular; cells thin walled. Stomata: paracytic, 

15 X 6 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAmNA.-Dorsiventral; 

V-shaped with small median adaxial groove; margins of lamina turned inversely. Adaxial surface 

slightly undulating; abaxial surface with very small 

furrows. Epidermis: adaxial cells rounded to rectangular in shape; abaxial cells mainly dome 

shaped; cells on both surfaces thin walled. Subjacent 

to adaxial epidermis is a single layer of parenchyma cells. Cuticle: very thick and slightly ridged 

on adaxial surface; thin and slightly ridged on 

abaxial surface. Stomata: present in abaxial furrows; no substomatal chamber observed. Mesophyll: two to three layers palisade cells grading 

into loosely arranged spongy tissue. Few translucent cells in spongy tissue. Vascular bundles: 39; 

few commissural bundles observed. Usually one 

large vessel present in each vascular bundle. Two 

phloem units laterally arranged in flanges of 

Y-shaped abaxial girder. Adaxial girder present on 

each vascular bundle. Each vascular bundle completely surrounded by a bundle sheath. Crystals 

and tannins: none observed. 

Barbacenia longiscapa Goethart & Henrard 

PLATE 47d 

SPECIAIEKS ExAMINED.-Irwin et al. 22 186. 

SURFACE VIEw.-Hairs: absent. Epidermis: adaxial cells rectangular, few square to rounded; thin 

ivalled. Abaxial cells rectangular, few square to 

rounded; thin walled. Stomata: paracytic, 21 X 12 

pm; present on both abaxial and adaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped with small median adaxial groove. Both 

adaxial and abaxial surfaces slightly undulating. 

Epidermis: cells on adaxial and abaxial surfaces 

rounded to dome shaped; thin walled. Layer of 

parenchyma cells subjacent to adaxial epidermis. 

Cuticle: thin and smooth on both adaxial and 

abaxial surfaces. Stomata: present on adaxial and 

abaxial surfaces; mainly flush with epidermis; substomata1 chamber present. Mesophyll: two to three 

layers of large palisade tissue changing into spongy 

tissue. Vascular bundles: 3 1-37; commissural bundles observed. One to two large vessels present in 

each vascular bundle, mostly one. Two phloem 

units lying laterally in flanges of Y-shaped abaxial 

girder. Inverted Y-shaped adaxial girder present 

on each vascular bundle. Bundle sheath completely 

surrounding each vascular bundle. Crystals: none 

observed. Tannins: present. 

Barbacenia purpurea Hooker 

FIGURE 18d-f; PLATE 39a-c 

SPECIMENS EXAMINED.-A. L. Schott s.n. US 

15 1098. 

SURFACE VIEw.-Hairs: none present. Epidermis: 

adaxial cells rounded; abaxial cells square to rectangular; cells thin walled on both surfaces. Stomata: paracytic, 21 X 6 pm; present on abaxial 

surface. 

TRANSVERSE SECTION OF LAiwNA.-Isolateral; 

V-shaped with margins turned inversely. Adaxial 

and abaxial surfaces furrowed about one-fourth 

thickness of blade. Epidermis: cells on adaxial surface dome shaped to rectangular; cells on abaxial 

surface rounded to dome shaped; thick walled 

on both surfaces. Cuticle: thin and slightly ridged 

on adaxial and abaxial surfaces. Stomata: present 

in abaxial furrows; substomatal chamber present. 

Xlesophyll: undifferentiated. Some large translucent cells radially arranged above the midvein; 

few translucent cells present in mesophyll. Vascular bundles: 37; commissural bundles not observed. One to two large vessels present in each 
56 

vascular bundle, mostly one. Two phloem units 

arranged laterally in flanges of Y-shaped adaxial 

girder. Inverted Y-shaped adaxial girder present 

on each vascular bundle. Both girders extend laterally below epidermis on both surfaces. Vascular 

bundles surrounded by bundle sheath. Crystals 

and tannins: none observed. 

S\IITHSOSI.AN COKTRIBUTIONS TO BOTANY 

Barbacenia seubertiana Goethart & Henrard 

FIGURE 19a-c; PLATE 38c-e 

SpEcIxin's ExAaiINED.--Lanna-Sobro 4488; Strang 

239, 523; Goetliart and Henrard 4332. 

SURFACE Vmv.-Hairs: absent. Epidermis: adaxial cells rounded; abaxial cells square to rectangua d 

FILURE 19.-a-c, Bnrbncenin seubwtiann (Strang 239) ; d-f, B. stenophylla (Irwin et al. 9901) 
NUMBER 15 57 

lar; walls thin on both surfaces. Stomata: paracytic, 

21 X 9 pm; present on abaxial surface. 

V-shaped. Adaxial and abaxial surfaces slightly undulating. Epidermis: cells on adaxial surface 

rounded to dome shaped; cells on abaxial surface 

dome shaped to conical; cell thin walled. Layer of 

parenchyma cells subjacent to epidermis. Cuticle: 

thin and ridged on abaxial and adaxial surfaces. 

Stomata: present on abaxial surface; small substomatal chamber present. Mesophyll: three to 

four layers of palisade-like cells grading into spongy 

tissue. Three to four layers of large mesophyll 

cells in midportion of lamina. Some translucent 

spongy tissue present on adaxial side of midvein. 

Vascular bundles: 19-57; commissural bundles not 

observed. One to two large vessels present in each 

vascular bundle; mostly one. Two phloem units 

laterally arranged in flanges of Y-shaped abaxial 

girder. Inverted Y-shaped adaxial girder present on 

each vascular bundle. Bundle sheath completely 

surrounding each vascular bundle. Crystals: none 

observed. Tannins: few in mesophyll. 

TRANSVERSE SECTION OF LAMINA.-ISOhteral; 

Barbacenia stenophylla Goethart & Henrard 

FIGURES li-j, 19d-f 

SPECIMEKS EXA;\rINFD.-Irwin et al. 9901. 

SURFACE VrEw.-Hairs: few present in tufts. Epidermis: cells square to rectangular, few rounded; 

thin walled. Stomata: paracytic, 30 X 12 pm; 

present on adaxial and abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Isolateral; 

V-shaped with small median adaxial groove. Adaxial and abaxial surfaces slightly undulating. Epidermis: cells on adaxial and abaxial surfaces 

rounded to dome shaped; walls on both surfaces 

thin. Layer of parenchyma cells subjacent to adaxial epidermis. Cuticle: very thin and ridged on 

both surfaces. Stomata: present on abaxial and 

adaxial surfaces; substomatal chamber present; 

flush with epidermal surface. Mesophyll: undifferentiated. Some translucent cells scattered in spongy 

mesophyll. Two to three layers translucent palisade 

cells arranged radially above midvein. Vascular 

bundles: 23; no commissural bundles observed. 

One large vessel present in each vascular bundle. 

Two phloem units lying laterally in flanges of 

Y-shaped abaxial girder. Inverted Y-shaped adaxial 

sclerenchyma present on each vascular bundle. 

Bundle sheath completely surrounding each vascular bundle. Crystals: none observed. Tannins: 

present mostly in mesophyll. 

Subkey I1 

Barbacenia celiae Maguire 

PLATE 48b 

SPECIMEKS ExAMIhED.-&laguire et al. 40025, 

SURFACE VrEw.-Hairs: absent. Epidermis: cells 

square to rectangular on both surfaces, walls thin. 

Stomata: paracytic, 27 X 12 pm; present mainly 

on abaxial surface. 

dorsiventral; widely V-shaped. Adaxial surface undulating; abaxial surface slightly ridged. Epidermis: cells on adaxial and abaxial surfaces rounded 

to dome shaped; thin walled. Subjacent to adaxial 

and abaxial epidermis a layer of parenchyma cells, 

some of which also form part of the bundle sheath. 

Cuticle: fairly thick and slightly ridged on adaxial 

surface; thinner and slightly ridged on abaxial 

surface. Stomata: present on abaxial surface: substomatal chamber present; usually occur in depression of small ridges. Mesophyll: three-layered 

palisade-like tissue grading into fairly compact 

spongy tissue; most of spongy tissue composed of 

large translucent cells. Vascular bundles: 39-47; 

no commissural bundles observed. One to two 

large vessels present in each vascular bundle, 

mostly one. Two phloem units lying laterally in 

flanges of Y-shaped abaxial girder. Inverted 

Y-shaped adaxial girder present on each vascular 

bundle. Bundle sheath completely surrounding 

each vascular bundle. Crystals: none observed. 

Tannins: present, numerous and occupying most 

of mesophyll. 

40 154, 40298. 

TRANSVERSE SECTION OF LAMINA.-somewhat 

Barbacenia flava Martius ex Schultes f. 

FIcuRrs lc-d, 23a-c; PLA~ES 6a-b, 40e-g 

SPECIMENS EXAMINED.-L. B. Smith 6695, 6856; 

Segadas-Vianna and Loredo 1100; Maguire et al. 

49116; Tryon and Tryon 6836; Maguire et al. 

49011; Macedo 3747; Maguire et al. 44691. 
SMITHSONIAS CONTRIBUTIONS TO BOTANY 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

circular to rectangular on both surfaces; thin 

walled. Stomata: paracytic, 21 X 15 pm; present 

on both surfaces. 

TRANSVERSE SECTION OF LAMINA.-ISOhteral; 

V-shaped with margins turned inversely. Both surfaces slightly undulating (except for Tryon 8c 

Tryon 6836 which has large ridges on both surfaces). Epidermis: adaxial cells rounded to dome 

shaped; abaxial cells dome shaped, slightly conical. 

Subjacent to adaxial epidermis is a distinct layer 

of parenchyma cells. Cuticle: thick and ridged on 

both surfaces. Stomata: present on adaxial and 

abaxial surfaces; small substomatal chamber present; usually flush with epidermis or slightly sunken. 

Mesophyll: undifferentiated spongy tissue largely 

composed of translucent cells. Vascular bundle: 

25-91; commissural bundles observed. One to two 

large vessels present in each vascular bundle. Two 

phloem units laterally arranged in flange of 

Y-shaped abaxial girder. Inverted Y-shape adaxial 

girder present on each vascular bundle. Bundle 

sheath completely surrounding each vascular bundle. Crystals: not observed. Tannins: present, 

numerous, filling most of mesophyll. 

NOTE: The specimen Tryon & Tryon 6836 

does not seem to belong to this species. 

Barbacenia ignea Martius ex Schultes f. 

FIGURE 21d-f; PLATES 39e-f, 40a 

SPECIMENS EXAMINED.-L. 0. Williams and Assis 

6817. 

SURFACE VIEw.-Hairs: absent. Epidermis: cells 

square to rectangular; thin walled. Stomata: paracytic, 24 X 12 pm; present on abaxial and adaxial 

surfaces. 

TRANSVERSE SECTION OF LAiwNA.--Isolateral; 

V-shaped; turned slightly inversely at margins of 

lamina. Adaxial and abaxial surfaces only very 

slightly undulating. Epidermis: cells dome shaped 

to rectangular on adaxial and abaxial surfaces. 

Layer of parenchyma cells subjacent to adaxial 

and abaxial epidermis. Cuticle: thin and slightly 

ridged on both surfaces. Stomata: present on abaxial 

and adaxial surfaces. Substomatal chamber present; 

stomata almost flush with epidermal surface. Mesophyll: two to three layers of small cells grading 

into large spongy tissue. Large number of translucent cells present in center of spongy mesophyll. 

Vascular bundles: 49; few commissural bundles 

observed. One to two large vessels observed, mostly 

one. Two phloem units lying laterally in flanges 

of Y-shaped abaxial girder. Inverted Y-shaped adaxial girder present on all vascular bundles. Bundle 

sheath completely surrounding each vascular bundle. Crystals: none observed. Tannins: numerous, 

present in mesophyll especially toward adaxial 

and abaxial surface. 

Barbacenia inuolucrata L. B. Smith 

FIGURE lk-1; PLATES 20b, 47c 

SPECIMENS EXAMINED.-Irwin et al. 20973. 

SURFACE VIEw.-Hairs: none present. Epidermis: 

adaxial cells rounded; abaxial cells square to rectangular; cells on both surfaces thin walled. Stomatal: paracytic, 30 X 15 pm; present on adaxial 

and abaxial surfaces. 

dorsiventral; V-shaped. Both adaxial and abaxial 

surfaces slightly undulating. Epidermis: abaxial 

and adaxial cells rounded; few dome shaped. Thin 

walled on both surfaces. Distinct layer of parenchyma cells subjacent to adaxial and abaxial epidermis. Cuticle: thick and slightly ridged on 

adaxial and abaxial surfaces. Stomata: present on 

both adaxial and abaxial surfaces; large substomatal chamber present. Mesophyll: three to four 

layers of palisade-like cells grading into spongy 

tissue. Large translucent cells present in midregion 

of mesophyll. Large translucent spongy cells present in adaxial portion of midvein. Vascular bundles: 69; few commissural bundles present. One to 

two large vessels present, mostly one. Two phloem 

units lying laterally in flanges of Y-shaped abaxial 

girder. Inverted Y-shaped adaxial girder present 

on each vascular bundle. Bundle sheath completely 

surrounding each vascular bundle. Crystals and 

tannins: none observed. 

TRANSVERSE SECTION OF LAMrNA.-somewhat 

Barbacenia schwackei Goethart & Henrard 

FIGURE 21a-c; PLATE 39d 

SPECIMENS EXAMINED.-Heringer 5263. 

SURFACE VIE!*.-Hairs: absent. Epidermis: adaxial cells rounded; abaxial cells rectangular, few 
NUMBER 15 59 

a 

e 

FIGURE 20.-a-c, Barbacenia gentianoides (Segades-Vianna and Loredo 1066); 

d-f, B. gounelleana (Emygdio 1428). 

rounded; cell walls thin on both surfaces. Stomata: cells thin walled. Layer of parenchyma cells subparacytic, 27 X 12 pm; present on abaxial surface. jacent to adaxial epidermis. Cuticle: thin and 

TRANSVERSE SECTION OF LArvmvA.-Isolateral; ridged on adaxial and abaxial surfaces. Stomata: 

V-shaped. Abaxial and adaxial surfaces slightly present on abaxial surface; small substomatal 

undulating. Epidermis: cells on abaxial and adax- chamber present. Mesophyll: undifferentiated. 

ial surface rounded to dome shaped, few conical; Compactly arranged spongy tissue interspersed 
60 SMITHSOSIAK CONTRIBUTIONS TO BOTANY 

with some translucent cells. Vascular bundles: 41; Tannins: numerous, virtually filling most mesofew commissural bundles observed. One to two phyll tissue. 

large vessels present in each vascular bundle, 

mostly one. Two phloem units lying laterally in 

flanges of Y-shaped abaxial girder. Inverted 

Y-shaped adaxial girder present on each vascular 

bundle. Bundle sheath completely surrounding SPECIMENS ExAMINED.-Menezes and Continho 5. 

each vascular bundle. Crystals: none observed. SURFACE VrEw.-Hairs: few papillae present. 

Barbacenia sellovii Goethard & Henrard 

PLATES 21b, 47h 

U 

C 

FIGURE 21.-a-c, Barbacenia schwackei (Heringer 5263); d-f, B. ignea (Williams and Assis 6817). 
NUMBER 15 61 

Epidermis: adaxial cells rounded; abaxial cells 

square to rectangular; cell walls thin on both surfaces. Stomata: paracytic, 12 X 3 pm; present on 

abaxial and adaxial surface. 

TRANSVERSE SECTION OF LAMINA.-ISOlateral; 

V-shaped. Margins of lamina turned slightly inversely. Adaxial and abaxial surfaces only gently 

undulating. Epidermis: cells on adaxial and abaxial surfaces rounded to dome shaped, few conical; 

thin walled. Cuticle: very thin, but smooth on 

both surfaces. Stomata: present on abaxial and 

adaxial surfaces; large substomatal chamber present; flush with or slightly above epidermal surface. Mesophyll: undifferentiated. Spongy tissue 

composed of small rounded cells, some large and 

translucent. Two to three layers of large translucent cells above midvein changing abruptly into 

compactly arranged spongy tissue. Vascular bundles: 29; few commissural bundles observed. One 

to three large vessels present in each vascular bundle. Two phloem units lying laterally in flanges 

of Y-shaped abaxial girder. Inverted Y-shaped adaxial girder present on each vascular bundle. Each 

vascular bundle completely surrounded by bundle 

sheath. Crystals: none observed. Tannin: numerous, 

filling most of mesophyll cells. 

Barbacenia vandellii Pohl ex Seubert in Martius 

FIGURES le-f, 22a-c 

SPECIMENS EXAMINED.-Gardner 5224. 

SURFACE VrEw.-Hairs: absent. Epidermis: cells 

square to rectangular; thin walled. Stomata: paracytic, 24 X 15 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.--Isolateral; 

widely V-shaped. Adaxial and abaxial surfaces 

deeply undulating. Epidermis: adaxial cells mostly 

dome shaped, few conical. Abaxial cells largely 

conical. Cells on both surfaces thin walled. Cuticle: 

thin and ridged on adaxial surface; thicker and 

ridged on abaxial surface. Stomata: present on 

abaxial surface; substomatal chamber not observed. 

Mesophyll: undifferentiated; mesophyll composed 

almost entirely of spongy tissue interspersed with 

few translucent cells. Vascular bundles: 89; commissural bundles observed. One to two large vessels present in each vascular bundle, mostly one. 

Two phloem units lying laterally in flanges of 

Y-shaped abaxial girder. Inverted Y-shaped adaxial 

girder present on each vascular bundle. Bundle 

sheath completely surrounding each vascular bundle. Crystals: none observed. Tannins: present in 

mesophyll. 

Barbacenia williamsii L. B. Smith 

FIGURE 22d-f; PLATE 40b-d 

SPECIMENS ExAMINED.-Dueke s.n.; Williams and 

Assis 6386; Pereira 2460-Pabst 3296. 

SURFACE VIEW.-Hairs: absent. Epidermis: adaxial cells square to rectangular; abaxial cells more 

rounded. Walls thin on both surfaces. Stomata: 

paracytic, 18 X 12 pm; present on abaxial and 

adaxial surfaces. 

TRANSVERSE SECTION OF LAMmA.-Isolateral; 

V-shaped. Adaxial and abaxial surfaces only slightly 

undulating. Epidermis: rounded to dome-shaped 

cells on abaxial and adaxial surfaces; few conical 

abaxial cells. Cells thin walled on both surfaces. 

Layer of parenchyma cells subjacent to epidermis. 

Cuticle: thin on adaxial and abaxial surfaces; 

slightly thicker on midvein. Stomata: present on 

abaxial and adaxial surfaces; small substomatal 

chamber present; flush with epidermal surface. 

Mesophyll: undifferentiated, spongy tissue compactly arranged. Two to three layers of large translucent palisade cells arranged radially above midvein. Some translucent cells present in midportion 

of spongy tissue. Vascular bundles: 33-49; commissural bundles few. One large vessel present in each 

vascular bundle. Two phloem units arranged laterally in flanges of Y-shaped abaxial girder. Inverted Y-shaped adaxial girder on each vascular 

bundle. Bundle sheath completely surrounding 

each vascular bundle. Crystals: none observed. 

Tannins: numerous, mostly in mesophyll. 

Subkey I11 

Barbacenia blackii L. B. Smith 

PLATES 9c-d, 48c 

SPECIMENS EXAMINED.-Macedo 2962. 

SURFACE VIEw.-&iirs: numerous multicellular 

hairs on abaxial and adaxial surfaces in variable 

length. Epidermis: cells square to rectangular on 

both surfaces; thin walled. Stomata: paracytic, 21 
62 SMITHSONIAN COSTRIBUTIONS TO BOTANY 

e 

- C f 

FIGURE 22.-a-c, Barbacenia vandellii (Gardner 5224) ; 

d-f, B. williarnsii (L. 0. Williams and Assis 6386). 

X 18 pm; present on adaxial and abaxial surfaces. 

dorsiventral; V-shaped with margins turned slightly 

inversely. Both adaxial and abaxial surfaces slightly 

undulating. Epidermis: adaxial cells dome shaped, 

few conical; walls slightly thickened. Abaxial cells 

rounded to dome shaped, few conical; thin walled. 

Multicellular hairs present on adaxial and abaxial 

surfaces. Cuticle: very thin on both surfaces. Stomata: present on adaxial and abaxial surfaces; 

small substomatal chamber present. Mesophyll: 

TRANSVERSE SECTION OF LAMINA.-somewhat 

two to three layers of palisade-like tissue grading 

into compactly arranged spongy tissue; translucent 

cells present in spongy layer. Vascular bundles: 31; 

few commissural bundles observed. One to three 

large vessels present in each vascular bundle. Two 

phloem units lying laterally in flanges of Y-shaped 

abaxial girder. Inverted Y-shaped adaxial girder 

present on each vascular bundle. No abaxial sclerenchyma strands observed. Each vascular bundle 

completely surrounded by a bundle sheath. Crystals: present. Tannins: none observed. 
NUMBER 15 63 

Barbacenia conicostigma Goethart & Henrard 

FIGURE lg-h; PLATES 23a-b, 46e 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

15982. 

SURFACE VrEw.-Hairs: none present. Epidermis: 

cells square to rectangular on abaxial and adaxial 

surfaces, few rounded; thin walled. Stomata: paracytic, 21 X 15 pm; present on adaxial and abaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-hOlateral; 

V-shaped with median abaxial sclerenchyma quite 

prominent below abaxial surface. Both adaxial 

and abaxial surfaces very slightly undulating, almost smooth. Epidermis: adaxial cells rounded to 

dome shaped, thin walled. Abaxial cells more 

rounded to rectangular, thin walled. Layer of 

parenchyma cells subjacent to adaxial epidermis. 

Cuticle: slightly thickened and ridged on adaxial 

and abaxial surfaces. Stomata: present on both 

adaxial and abaxial surfaces; small substomatal 

chamber present. Mesophyll: palisade tissue nonexistent. Fairly loosely arranged spongy tissue 

present especially toward adaxial surface; large 

part made up of large translucent spongy cells. 

Cells larger in midsection of mesophyll. Vascular 

bundles: 42; few commissural bundles observed. 

One large vessel present in each vascular bundle. 

Two phloem units lying laterally in flanges of 

Y-shaped abaxial girder. Inverted Y-shaped adaxial 

girder present on each vascular bundle. Bundle 

sheath completely surrounding each vascular bundle. Crystals and tannins: none observed. 

Barbacenia exscapa Martius 

PLATE 46g 

SPECIMENS ExAMINED.-Anderson et al. 3583 1. 

SURFACE VIEw.-Hairs: present mostly on abaxial surface. Epidermis:. adaxial cells mostly square 

to rectangular, few rounded; thin walled. Abaxial 

cells rectangular, thin walled. Stomata: paracytic, 

27 X 21 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 

V-shaped. Both adaxial and abaxial surfaces gently 

undulating. Epidermis: cells on adaxial and abaxial surfaces rounded to dome shaped, walls slightly 

thickened. Subjacent to adaxial epidermis are one 

to two layers of parenchyma cells; parenchyma 

layers in midvein region much larger. Cuticle: 

fairly thick and slightly ridged on adaxial and 

abaxial surfaces. Stomata: present on abaxial surface, small substomatal chamber observed; flush 

with epidermal surface. Mesophyll: three to four 

layers of palisade tissue occupying about one-half 

of the mesophyll. Abrupt change to spongy tissue. 

Elongated palisade cells not usually found in the 

midvein region. Vascular bundles: 25 to 27; of 

small sizes; few commissural bundles observed. 

Usually one large vessel present in each vascular 

bundle. Two phloem units lying laterally in flanges 

of Y-shaped abaxial girder. Inverted Y-shaped 

adaxial girder present on each vascular bundle. 

Each vascular bundle completely surrounded by 

a bundle sheath. Crystals: none observed. Tannins: 

few present in mesophyll. 

NOTE: The adaxial and abaxial sclerenchyma 

girders are poorly developed. In some cases they 

are no more than lignified parenchyma cells. 

Barbacenia fragrans Goethart & Henrard 

FIGURES Zd, 24a-c; PLATES 9a, 22a 

SPECIMENS EXAMINED.-DUeke s.n.; Regnell 1239; 

Widgren s.n. US 933895. 

SURFACE VIEW.-Hairs: numerous, spreading 

over entire surfaces. Epidermis: adaxial cells 

square to rectangular; abaxial cells rounded to 

square, few rectangular; cell walls thin. Stomata: 

paracytic, 12 X 12 pm; present on abaxial and 

adaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-DorSiventral; 

widely V-shaped with margins turned inversely. 

Adaxial surface slightly undulating; abaxial surface quite ridged. Epidermis: cells on adaxial and 

abaxial surfaces are dome shaped to conical; thin 

walled. Subjacent to adaxial epidermis is a layer 

of parenchyma cells. Cuticle: thin and ridged on 

both surfaces. Stomata: present on abaxial and 

adaxial surfaces; substomatal chamber present. 

Mesophyll: two- to three-layered palisade cells 

grading into loosely arranged spongy tissue containing large translucent cells. Vascular bundles: 

37-47; no commissural bundles observed. One to 

two large vessels present in each vascular bundle, 

mostly one. Two phloem units lying laterally in 

flanges of Y-shaped abaxial girder extending to 

abaxial epidermis. Inverted Y-shaped adaxial girder 
64 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

present on each vascular bundle. Bundle sheath 

completely surrounding each vascular bundle. 

Crystals: none observed. Tannins: numerous, 

mostly in mesophyll. 

Barbacenia gardneri Seubert in Martius 

PLATES 7a-b, 47a 

SPECIMENS _C,xAMIsED.-Pereira 2849-Pabst 3685. 

SURFACE VIEw.-Hairs: long, thin, multicellular 

hairs present on adaxial surface. Epidermis: cells 

square to rectangular on both adaxial and abaxial 

surfaces; few rounded; thin walled. Stomata: paracytic, 18 X 12 pm; present on both adaxial and 

abaxial surfaces. 

TRANSVERSE SECTION OF LAMINA.-DOrsiventral; 

V-shaped with extreme margins of lamina turned 

slightly inversely. Both abaxial and adaxial surfaces slightly undulating; abaxial a little more 

than adaxial. Epidermis: adaxial cells rounded, 

dome shaped and conical, few crescent shaped; 

walls thin. Abaxial cells dome shaped to conical; 

walls thin. Cuticle: very thin and following contour of cells on both surfaces. Stomata: present 

on both adaxial and abaxial surfaces; substomatal 

chamber quite large on both surfaces; either flush 

with or slightly above epidermal surface. Mesophyll: two to three layers of small palisade cells 

changing abruptly to large, rounded translucent 

cells with thickened walls occurring in middle of 

mesophyll, especially next to the vascular bundles. 

Cells change abruptly to two or three layers of 

small rounded cells on the abaxial side. Very large 

midvein cells abruptly changing into small cells 

toward abaxial side. Vascular bundles: 48; many 

commissural bundles observed. One to three large 

vessels present in each bundle, mostly one. Two 

phloem units in flanges of V- or Y-shaped abaxial 

girder. Inverted Y-shaped adaxial girder present on 

each vascular bundle. Bundle sheath completely 

surrounding each vascular bundle. Crystals and 

tannins: none observed. 

Barbacenia gentianoides Goethart & Henrard 

FIGURE 20a-c; PLATE 41c-e 

SPECIMENS ExAMINED.-Segadas-Vianna and Loredo 1066. 

SURFACE VIEw.-Hairs: multicellular; some papillae present. Epidermis: cells rounded, square 

to rectangular. Stomata: paracytic, 27 X 12 pm; 

present on abaxial surface. 

TRANSVERSE SECTION OF LAMI,uA.-Dorsiventral; 

widely V-shaped with slight median adaxial groove. 

Both adaxial and abaxial surfaces undulating. Epidermis: cells on adaxial and abaxial surfaces dome 

shaped; thin walled. Distinct layer of parenchyma 

cells subjacent to adaxial epidermis. Sclerenchyma 

strands occur below parenchyma layer, usually extending on either side of adaxial side of vascular 

bundle. Cuticle; very thin on adaxial and abaxial 

surfaces. Stomata: present on abaxial surface. Substomata1 chamber present. Mesophyll: three to 

four layers of palisade cells present. Abruptly 

changing into loosely arranged spongy tissue. Some 

translucent cells present in spongy layer. Vascular 

bundles: 30; commissural bundles observed. One 

to two large vessels, mostly one. Two phloem units 

lying laterally in flanges of V- or Y-shaped abaxial 

girder. Inverted Y-shaped adaxial girder present on 

each vascular bundle. Bundle sheath completely 

surrounding vascular bundles. Crystals: none observed. Tannins: few present. 

Barbacenia longiflora Martius 

FIGURE 24d-f: PLATE 41f-h 

SPECIMENS ExAhfIixED.-Akcher 4089, Irwin 2486. 

SLRFACE VIEw.-Hairs: absent. Epidermis: cells 

rounded to rectangular, few squarish on abaxial 

and adaxial surfaces; walls thin. Stomata: paracytic, 18 X 15 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMmA.-Isolateral; 

widely V-shaped with slight median adaxial groove. 

Adaxial and abaxial surfaces slightly undulating. 

Epidermis: cells on both surfaces dome shaped, few 

conical; thin walled. Layer of parenchyma cells 

subjacent to adaxial epidermis. Cuticle: thin and 

ridged on adaxial and abaxial surfaces. Stomata: 

present on abaxial surface; substomatal chamber 

present; mainly flush with epidermal surface. 

RIesophyll: two to three layers of small cells 

grading into spongy tissue. Two to three layers 

of translucent cells present in median portion of 

lamina. Vascular bundles: 33; few commissural 

bundles observed. One to two large vessels in each 

vascular bundle, mostly one. Two phloem units 
NUMBER 15 65 

arranged laterally in flanges of Y-shaped abaxial 

girder. Inverted Y-shaped adaxial girder present 

on each vascular bundle. Each vascular bundle 

completely surrounded by a bundle sheath; two 

to four bundle sheath cells in midportion of vascular bundles conspicuously enlarged. Crystals and 

tannins: none observed. 

Barbacenia luzulifolia Martius ex Schultes f. 

FIGURE lo-p; PLATES 6c, 47e 

SPECIMENS EXAMINED.-Pereira 2664-Pabst 3500. 

SURFACE VrEw.-Hairs: none present. Epidermis: 

cells square to rectangular, few rounded on abaxial 

and adaxial surfaces; thin walled. Stomata: paracytic, 24 X 9 pm; present on both adaxial and 

abaxial surfaces. 

dorsiventral; V-shaped with margins turned slightly 

inversely. Both adaxial and abaxial surfaces evenly 

undulating. Epidermis: adaxial cells mostly 

rounded to dome shaped, few conical; thin walled. 

Abaxial cells rounded to dome shaped, few elongated; cell walls thicker than adaxial ones. Adaxial 

epidermal cells larger around midvein. Cuticle: 

very thin on both adaxial and abaxial surfaces. 

Stomata: present on both adaxial and abaxial surfaces; substomatal chamber quite large; usually 

flush with epidermal surface. Mesophyll: three 

layers of elongated palisade-like cells changing 

into large, rounded, translucent cells in the midregion of mesophyll. Vascular bundles: 43; no 

commissural bundles observed. One to two large 

vessels present in each bundle, mostly one. TWO 

phloem units lying laterally in flanges of V- or 

Y-shaped abaxial girder. Inverted Y-shaped adaxial 

girder present on each vascular bundle. Bundle 

sheath completely surrounding each vascular bundle. Crystals and tannins: none observed. 

TRANSVERSE SECTION OF LAMINA.-somewhat 

Barbacenia macrantha Lem 

FIGURE 2f; PLATFS 8a-b, 48d 

SPECIMENS EXAM INED.-Irwin, Maxwell, and 

IVasshausen 2007 1. 

SURFACE VIEw.-Hairs: numerous on both surfaces. Epidermis: adaxial cells rectangular; abaxial 

cells rounded, some square. Walls thin on both 

surfaces. Stomata: paracytic, 12 X 6 pm; present 

on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-DOrSiVentral; 

V-shaped with margins of lamina turned inversely. 

Both adaxial and abaxial surfaces slightly undulating; large numbers of hairs present on both 

surfaces. Epidermis: thin cell walls on abaxial and 

adaxial surfaces rounded to dome shaped; few 

conical. Cuticle: on abaxial and adaxial surfaces 

thin and ridged. Stomata: present on abaxial surface; substomatal chamber present. Mesophyll: 

two to three layers of palisade cells grading into 

spongy tissue. Some large translucent cells present 

in spongy tissue. Vascular bundles: 41; no commissural bundles observed. One to t~7o large vessels present in each vascular bundle. Two phloem 

units lying laterally in flanges or V- or Y-shaped 

abaxial girder. Inverted Y-shaped adaxial girder 

present on each vascular bundle. Crystals: none 

observed. Tannins: present and in numerous 

quantities, especially in palisade cells. 

Barbacenia nana L. B. Smith & Ayensu 

FIGURF 2e; PLATE-s 8c-d, 22b, 47f 

SPECIMENS EXAMINED.-L. B. Smith and Ayensu 

15973. 

SURFACE VrEw.-Hairs: present on adaxial and 

abaxial surtaces; long and multicellular. Epidermis: adaxial cells square; thin walled. Abaxial 

cells square to rectangular; thin walled. Stomata: 

tetracytic, 21 X 18 pm; present on abaxial surface. 

TRAMVERSE SECTION OF LAhlINA.-DOrsiVentral; 

V-shaped with margins turned slightly inversely. 

Adaxial and abaxial surfaces only slightly undulating; both surfaces covered with hairs. Epidermis: 

adaxial cells dome shaped to conical; thin walled. 

Abaxial cells more rounded to dome shaped; thin 

walled. Subjacent to adaxial epidermis is a distinct 

layer of parenchyma cells. Cuticle: thin and ridged 

on adaxial and abaxial surfaces. Stomata: present 

on abaxial surface; substomatal chamber present; 

usually flush with or slightly above epidermal surface. Mesophyll: two to three layers of palisade 

cells grading into a loosely arranged spongy tissue. 

Large number of translucent cells in spongy tissue, 

especially in midvein region. Vascular bundles: 44; 

few commissural bundles observed. One to two 

large vessels present in each vascular bundle, 
66 SMITHSOXIAN COX~RIBUTIONS TO BOTANY 

a 

b 

FIGURE 23.-a-c, Barbacenia flaua (Segadas-Vianna and Loredo 1100) : 

d-f, B. rubro-virens (Maguire et al. 49260). 

mostly one. Two phloem units lying in flanges of 

wide Y-shaped abaxial girder. Inverted Y-shaped 

adaxial girder present on each vascular bundle; 

sometimes appearing T-shaped near adaxial surface. 

Some sclerenchyma strands observed on adaxial SURFACE ViEtv.-Hairs: numerous, present on 

side of lamina replacing some palisade cells. Bun- both surfaces. Epidermis: adaxial cells rounded. 

dle sheath completely surrounding each vascular Abaxial cells square to rectangular. Walls thin. 

bundle. Crystals and tannins: none observed. Stomata: paracytic, 18 X 15 pm; present on 

Barbacenia paranuensis L. B. Smith 

PLATES 20a, 48e 

SPECIMENS ExAmmD.-Hatschbach 15715. 

f 
NUMBER 15 67 

abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-DOrSiVentral; 

widely V-shaped. Adaxial and abaxial surfaces 

undulating. Epidermis: cells on adaxial and abaxial surfaces rounded to dome shaped; thin walled. 

Multicellular hairs present on epidermal surfaces. 

Cuticle: very thin on both surfaces; ridged. 

Stomata: present on adaxial surface; substomatal 

chamber present. Mesophyll: three- to four-layered 

palisade cells changing abruptly into compactly 

arranged spongy tissue. Two to three layers of large 

translucent cells arranged radially above midvein. 

Some large translucent spongy cells present in midportion of mesophyll. Vascular bundles: 43; few 

commissural bundles observed. One large vessel 

present in each vascular bundle. Two phloem units 

arranged laterally in flanges of Y-shaped abaxial 

girder. Inverted Y-shaped adaxial girder present 

on each vascular bundle. Bundle sheath completely 

surrounding each vascular bundle. Crystals: not 

observed. Tannins: present, especially in midportion of mesophyll. 

Barbacenia riedeliana Goethart & Henrard 

PLATES 9b, 47g 

SPECIMENS ExAivlrNED.-Anderson et al. 35 123. 

SURFACE VIEw.-Hairs: short, multicellular papillae present on both surfaces. Epidermis: adaxial 

and abaxial cells rectangular, few square; thin 

walled. Stomata: paracytic, 18 X 12 pm; present 

on both adaxial and abaxial surfaces. 

TRANSVERSE SECTION OF LAMmA.-Isolateral; 

V-shaped. Adaxial and abaxial surfaces undulating 

with small furrows on each. Epidermis: adaxial 

and abaxial cells rounded to dome shaped, few 

conical; thin walled. Cuticle: very thin and smooth 

on both surfaces. Stomata: present on both adaxial 

and abaxial surfaces; flush with or slightly above 

epidermal surface; large substomatal chamber 

present. Mesophyll: two or three layers of small 

cells changing into one or two layers of large, 

rounded, translucent spongy cells. Two to three 

layers of large translucent cells above midvein 

changing into compactly arranged spongy tissue. 

Few papillae present in furrows of both surfaces. 

Vascular bundles: 17; few commissural bundles observed. One to two large vessels, mostly one, in 

each vascular bundle. Two phloem units lying 

laterally in flanges of Y-shaped abaxial girder. Inverted Y-shaped adaxial girder present on each 

vascular bundle. Bundle sheath surrounding each 

vascular bundle. Crystals and tannins: none 

observed. 

Barbacenia rubro-virens Martius 

FIGURES lm-n, 23d-f 

SPECIMENS ExAMINED.-Maguire et al. 49177, 

49260. 

SURFACE VIEw.-Hairs; absent. Epidermis: cells 

square to rectangular on both surfaces; some 

rounded on abaxial surface; thin walled on both 

surfaces. Stomata: paracytic, 27 X 15 pm; present 

on abaxial and adaxial surface. 

TRANSVERSE SECTION OF LAMmA.--Isolateral; 

V-shaped with margins of lamina turned inversely. 

Abaxial and adaxial surfaces slightly undulating. 

Epidermis: cells on abaxial and adaxial surfaces 

rounded to dome shaped; thin walled. Layer of 

parenchyma cells subjacent to both surfaces. Cuticle: thick and slightly ridged on adaxial and 

abaxial surfaces. Stomata: present on both surfaces; substomatal chamber present; more or less 

flush with epidermal surface. Mesophyll: three to 

four layers of small cells grading into compactly 

arranged spongy tissue. Some large translucent 

cells present in spongy mesophyll. Vascular bundles: 

44; few commissural bundles observed. One large 

vessel present in each vascular bundle. Two 

phloem units laterally arranged in flanges of 

Y-shaped abaxial girder. Inverted Y-shaped adaxial 

girder present on each vascular bundle. Bundle 

sheath completely surrounding each vascular bundle. Crystals: none observed. Tannins: numerous, 

especially in mesophyll cells near the epidermis. 

Barbacenia tomentosa Martius 

PLATE 7c-d 

SPECIhIENS EXAMINED.-IrWin et al. 29559; 

Menezes 1. 

SURFACE VIEw.-Hairs: present on adaxial and 

abaxial surfaces; multicellular. Epidermis: cells on 

adaxial and abaxial surfaces square to rectangular, 

few rounded; thin walled. Stomata: tetracytic, 24 

X 9 pm; present on adaxial and abaxial surfaces. 

TRANSVERSE SECTION OF LAmNA.-Dorsiventral; 
68 SMITHSOXIAN COXTRIBUTIONS TO BOTANY 

- 

FIGURE 24.-a-c, Barbncenia fragrans (IVidgren 1267); d-f, B. LongiPora (Archer 4089) . 

wide V-shaped. Both adaxial and abaxial surfaces 

.Izith ridges or small furrows. Tufts of multicellular hairs present on both surfaces. Epidermis: 

adaxial cells dome shaped, few conical, thin walled; 

abaxial cells dome shaped, few conical, thin walled. 

Subjacent to adaxial epidermis is a layer of thinwalled parenchyma cells. Cuticle: very thin on 

adaxial and abaxial surfaces. Stomata: present in 

abaxial furrows and on adaxial surface; substomata1 chamber present. Mesophyll: small, rectangular, palisade-like cells, closely packed. Abrupt 

change to loosely arranged spongy tissue with some 

large translucent cells. Vascular bundles: 43-44; few 

commissural bundles observed. Usually one large 

vessel present in each vascular bundle. Two 

phloem units lying laterally in flanges of Y-shaped 

abaxial girder. Inverted Y-shaped adaxial girder 

present on each vascular bundle. Bundle sheath 

completely surrounding each vascular bundle. 

Crystals: none observed. Tannins: present in 

mesophyll. 
NUMBER 15 69 

Barbacenia uiscossissima Goethart & Henrard 

PLATES 40h, 41a-b 

SPECIMENS ExAMINED.-h~acedo 2826. 

SURFACE Vmv.-Hairs: present on both surfaces. 

Epidermis: cells mostly square to iectangular, few 

rounded; cells thick walled. Stomata: paracytic, 24 

X 12 pm; present on abaxial surface. 

TRANSVERSE SECTION OF LAMINA.-ISOlatera1; 

widely V-shaped, turned slightly inversely at margins. Adaxial and abaxial surfaces undulating. 

Epidermis: adaxial cells rounded to dome shaped: 

few crescent shaped. Abaxial cells rounded to 

dome shaped; few conical. Cells on both surfaces 

thick walled. Distinct layer of parenchyma cells 

subjacent to epidermis. Cuticle: thin on adaxial 

and abaxial surfaces. Stomata: present on abaxial 

surface; substomatal chamber present. hiesophyll: 

composed of mainly large translucent cells except 

for few rows of small cells subjacent to epidermis. 

Vascular bundles: 40; few commissural bundles 

observed. Two phloem units lying laterally in 

flanges of Y-shaped abaxial girder. Inverted 

Y-shaped adaxial girder present on each vascular 

bundle. Bundle sheath completely surrounding 

each vascular bundle. Crystals: none observed. 

Tannin: few present. 

Observations 

The following observations are focused on certain highlights in the anatomical differences which 

are found within this family. The accompanying 

line drawings, along with the light microscope 

and scanning electron photomicrographs, will 

hopefully express more clearly the anatomical 

characteristics of the species and thereby serve as 

an additional aid to identification of the species. 

Vellozia: Vellozia abietina and V. tragacantha 

have in common certain anatomical features. 

Apart from their leaf shapes that are almost identical, the sclerenchyma girders associated with the 

vascular bundles are strongly developed. The 

adaxial sclerenchyma girders extend closely to the 

epidermis except for those associated with the vascular bundles next to the midvein. The mesophyll 

differ, however. In V. tragacantha there is a distinct differentiation between the palisade cells and 

LIGHT hfICROSCOPIC OBSERVATIONS.-SeCtiOn 1: 

the spongy tissue, while that of V. abietina is not 

so clear-cut as demonstrated in V. sellovii. 

As its name implies, the leaf of V. plicata is 

folded lengthwise into plaits. The lamina exhibits 

a strong development of sclerenchyma around the 

main veins where there is lateral expansion. The 

regions between the principal veins are slender 

and exhibit minimal sclerenchyma strands subjacent to the adaxial epidermis and to a minor 

degree they are developed toward the abaxial surface. In addition to the principal veins there are 

many smaller secondary, or tertiary, veins in the 

slender regions between the large vascular bundles. Another species that exhibits the characteristics described for V. plicata is V. triquetra and, 

in fact, the two species are almost inseparable. 

Another group of species that share basically 

similar anatomical structures includes V. glabra, 

V. incurvatn, and V. compacta. The midvein regions show strongly developed translucent cells 

which occupy over two-thirds of the primary vein 

portion of the leaf. The costal regions of the blade 

are pointed, while the intercostal regions are significantly expanded and pointed adaxially. Furthermore the furrows in the region are very deep, 

in some cases occupying over two-thirds of the 

lamina. Most of the specimens examined under 

these species have pointed abaxial midveins. However, in V. incurvatu (Maguire 44752) and V. 

compactn (Segades-Vianna and Loredo 1085) the 

abaxial portions of the midveins are distinctly 

flattened. These differences seem to be prevalent 

in other specimens of a particular species, while 

all other characters are practically the same. It 

seems obvious that such differences may represent 

variations within species complex and therefore 

do not have a distinct taxonomic importance. 

The Xerophyta type of sclerenchyma pattern 

which I described for some Madagascan species 

(hyensu, 1969) seems to occur in some of the New 

Tliorld species. Vellozia terella, V. verruculosa, V. 

variabilis, V. fibrosa, V. taxifolia, V. declinans, 

V. leptopetala, V. epidendroides, V. asperula, V. 

swallenii, V. cinerascens, V. costata, V. hatschbachii, 

V. lappa, V. modesta, and V. wasshausenii share 

several more or less common mesophyll characteristics, but in addition, their abaxial sclerenchyma 

girders are more or less U- or V-shaped, and they do 

not extend to the abaxial epidermis as is the case 
70 SMITHSONIAX COSTRIBUTIONS TO BOTANY 

in the Vellozia type of abaxial sclerenchyma pattern. 

The presence of fingerlike projections in thc 

abaxial furrows of such species as V. metzgerae, V. 

glandulifera, V. aloifolia, J? intermedia, and V. 

echinata, as well as the distribution of translucent 

cells above the vascular bundles and the furrows, 

indicate some form of relationship amongst these 

species. Whether these close resemblances are dependent solely on environmental pressures is not 

quite clear. 

Of the array of species studied, Vellozia burlemarxii and V. hemisphaerica share two distinct 

features that set them apart from all other vellozias. 

The development of secondary vascular bundles 

directly above the abaxial furrows and the extra 

appendages at the wider portions of the furrows 

are certainly convergent morphological and anatomical characteristics. 

Vellozia epidendroides is one of the species most 

easily identified by its anatomical characters and 

the shape of leaf in transverse section. Among the 

major distinctive features are (a) the presence of 

an almost continuous band of highly developed 

sclerenchyma fibers subjacent to both the adaxial 

and abaxial epidermis and (b) the sandal-shaped 

appearance of the sclerenchyma associated with 

the vascular bundles. Of the specimens examined, 

the abaxial portion of the midrib was found to be 

either flattened or pointed. This observation further weakens the importance of this feature in the 

taxonomy of the species. 

Vellozia minima and V. virgata are among the 

small-leaved species of the family. Their anatomy 

depicts compactly arranged palisade and spongy 

tissues. Translucent cells are absent in these species, which are among the few members of Vellozia 

that lack abaxial furrows. Other species that fall 

under this category are V. abietina, V. tragacantha, 

and V. sellovii. 

In most species of Vellozia, translucent cells are 

arranged radially on the adaxial side of the vascular bundles, as well as on the furrows. However, 

there are few species in which the translucent cells 

occur only on the vascular bundles: V. flavicans, 

V. glauca, V. glochidea, V. crassicazilis, V. swallenii, 

V. cincerascens, and V. dasypus. 

The most striking anatomical feature of Vellozia 

exilis is the lateral expansion of the midvein region, making it one of the most distinctive species 

in the family. To my knowledge there is no other 

species in the family that exhibits a similar morphological feature. 

Three species that appear to be closely related 

from the standpoint of their leaf anatomy, especially the development of fingerlike projections in 

the abaxial furrows, are Vellozia caput-ardeae, V. 

lappa, and V. phalocarpa. However, upon close 

examination it becomes obvious that whereas the 

two phloem units in V. lappa are distinctly separated, those of V. caput-ardeae and V. phalocarpa 

appear as one. Furthermore, the Y-shaped abaxial 

sclerenchyma of V. caput-ardeae and V. phalocarpa 

extends to the epidermis, while that of V. lappa 

ends about two-thirds into the spongy mesophyll. 

Another group of species that appear to be 

closely related on account of the arrangement of 

the adaxial translucent cells are Vellozia costata, 

V. ciliata, and to some extent V. hatschbachii. In 

this instant also, the species can be easily separated 

from each other by the shape of the large parenchyma cells below the adaxial sclerenchyma 

strands. 

Vellozin modesta exhibits certain features that 

set it apart from other members of this genus. 

Generally there are two abaxial sclerenchyma 

strands between two furrows. Furthermore the 

outermost cells of the furrows are made up of 

elongated parenchyma cells instead of sclerenchyma 

strands which olten occur in this region. The 

palisade cells are completely filled with tannin. 

The cells of the abaxial spongy tissue are also 

filled with tannin. 

Section IZ: Radia: In this section each species 

seems to be anatomically distinct. Vellozia riedeliana and V. candida have one major characteristic 

in common, the lack of translucent cells above the 

furrows. All other species examined in this section 

possess translucent cells on both the adaxial side 

of the vascular bundles and in furrows. 

In Vellozia maculata the vascular bundles and 

their associated sclerenchyma girders occupy more 

than two-thirds of the mesophyll space between 

two furrows. Furthermore, the abaxial sides of the 

furrows are laterally extended in a way that they 

virtually obscure the entrances into the furrows. 

Three species that share several anatomical 

characters, especially the shape of the sclerenchyma 

girders and width of their furrows, are Vellozia 

dumitiana, V. lithophila, and V. tubipora. The 
NUMBER 15 71 

examination of several specimens of V. tubiflora 

has shown conclusively that the various collections 

belong to one species-complex. The slight variations among the specimens may be the result of 

environmental influences.- Maguire (1969: 35 ) was 

indeed correct when he noted that “Vellozia tubiflora is a highly variable and polymorphic species 

with a wide ecological amplitude.” Some of the 

specimens can easily be attributed to either V. 

dumitiana or V. lathophila on anatomical grounds. 

The species examined in 

this subkey are all distinct. Barbacenia gaveensis, 

B. seubertaana, B. stenophylla, B. longiscapa, and 

B. gounelleana have slightly undulating epidermal 

surfaces while B. puipurea is distinctly corrugated 

on both surfaces. Barbacenia irwiniana shows corrugation on the abaxial surface only and a gentle 

undulation on the adaxial sclerenchyma from the 

vascular bundle. The adaxial sclerenchyma appears 

in the form of strands completely surrounded by 

distinct bundle sheaths. If the mesophyll had been 

slightly differentiated this species could easily be 

assigned to Vellozia on anatomical grounds. Another distinct feature that has not been observed 

in other species under this subkey is the virtual 

obliteration of about two-thirds of the mesophyll 

in Ba) bacenia hatschbachzi. Unfortunately only 

one specimen was available for study, and therefore no conclusive statement can be made to characterize this species. 

Each species under this 

group is very distinct anatomically; therefore, 

they do not form a distinct group that can set 

them apart from the subkeys. As observed in subkey I, the presence of marked corrugations on 

both surfaces of the leaf occur in few species. The 

only species that exhibits this character in subkey 

I1 is Barbacnza vandellii. 

One of the characteristic features of this group is the presences of hairs 

on both surfaces of many species. Barbacenia macrantha, B. viscosissima, B. paranaensis, B. fragrans, 

B. blackii, and B. nana possess hairs that are often 

longer than the width of lamina. The hairs on 

B. riedelzana are short and occur mainly on the 

furrows in the form of fingerlike projections. 

Another feature that is characteristic of some 

species of this group is the differentiation of the 

mesophyll into distinct palisade and spongy tissues. 

Such features occur in Barbacenia paranaensis, 

Barbacenia Subkey I: 

Barbacenia Subkey 11: 

Barbacenia Subkey IN: 

B. exscapa, B. gardneri, B. fragrans, B. gentianoides, and B. nana. 

It is obvious from the above observations that 

on anatomical grounds the species of the New 

IYorld Velloziaceae cannot be subdivided under 

the same sections and subkeys of Smith’s (1962). 

However, his classification of species is clearly sup 

ported by this study. 

TIon-s.-The use of the SEM in the studies of 

Velloziaceae has revealed remarkable threedimensional characteristics that emphasize the diversity in the leaf surfaces. In addition to the light 

microscopic observations, the SEM has brought to 

light anatomical details that were not previously 

obtainable with conventional microscopes. 

The two types of trichomes that 

occur generally in Velloziaceae are hairs and 

fingerlike projections or protuberances. The hairs 

come in a variety of forms. Generally they are 

either unicellular or multicellular and vary considerably in length. In general, hairs appear to 

cover most of the leaf epidermis. Usually they are 

more frequent on the abaxial surface. In fact, their 

short multicellular bases form distinct tufts (Plates, 

la,b; 3a-d; 4a-d), while the terminal cells are 

long and free. Sometimes hairs occur either in 

furrows, as in Vellozia dumitiana and V. lithophila 

(Plate 2a-d), or on the surfaces between furrows, 

as in V. dawsonii (Plate la,b). In some cases the 

hairs appear very close to the furrows as in V. 

pumila (Plate 5d). 

The hairs come in a variety of shapes. They are 

either straight (Plates 7a,b; 9a-d), curved (Plate 

5d), or twisted (Plate 1c,d). In species such as 

Ba?lracenia macrantha (Plate 8a,b) and B. nana 

(Plate 8c,d), the hairs appear to be coalesced. 

Transverse sections of hairs (Plate 8b-d ) indicate 

that in these species the four to nine hairs have 

coalesced to form a unit. In the center of Plate 8a 

is an indication of the process of coalescence of 

already coalesced hairs. It is obvious from this 

observation that there is no systematic pattern in 

the manner and the number of individual hairs 

that form a coalesced unit. After this process has 

been completed, the outer walls of the hairs stretch 

out to form a smooth membrane around the 

coalesced hairs. 

Fingerlike projections emerge from epidermal 

cells in the furrows, as shown in transverse sections 

SCAKNIZJC ELECTRON 1hfICROSCOPIC OBSERVATrichomes: 
72 S.\fITHSONIAN CONTRIBUTIONS TO BOTANY 

of Vellozia ciliata (Plates 5c; 49a-c) and V. metzgerae (Plate 5b). A vertical section through a furrow of a leaf of V. ciliata (Plate 49c) shows very 

clearly the sporadic arrangement of the protuberances. Among these fingerlike projections are the 

stomata. 

Generally the epidermal cells 

are arranged in longitudinal files which run parallel to the midvein and the long axis of the leaf. 

The cells are generally rectangular, but in some 

cases are isodianietric. The SEM has also shown 

that in some species the cells appear to be regularly 

arranged in folds (cf. Barbacenia flava, Plate 6a,b), 

or they are somewhat verrucata as in Vellozia 

glochidea (Plate 5a). At higher magnifications the 

cells appear three-dimensional and thus reveal the 

remarkable topographic details of the epidermis. 

Generally the stomata are confined to 

the abaxial epidermis. In a few species they occur 

on the adaxial epidermis, but they are less numerous. In species that possess furrows, one often finds 

that the highest concentration of stomata occurs 

in the furrows and that the stomata are often 

associated with papillae. The stomata are generally 

arranged in longitudinal rows which run parallel 

Epidermal Cells: 

Stomata: 

to the 

Ayensu, 

1967. 

1968. 

1969a. 

1969b 

19i3. 

be crolvded on certain portions of the leaf surface, 

particularly in the furrows. 

RIost species of the Velloziaceae have paracytic 

type of stomata (Plates 10-23). Few are tetracytic; 

i.e., there are two parallel and two polar subsidiary 

cells. 

One of the distinctive features of the stomata 

is the clearly defined guard cells that are fully 

extended around the pore. The lip of the stomata1 

pores may be smooth (Plates 10, 17; 21a) or distinctly pronounced (Plates 11-16, 18-21b, 22). The 

subsidiary cells are often robust, but occasionally 

they are conspicuously ridged as shown in Vellozia 

squalida (Plate 14b) and in other species (Plates 

16a, 18a, 23b). 

The SEM was used to observe the internal structure of some of the leaves. These observations 

generally confirmed the findings based on light 

microscopy. 

In summary, the scanning observations have 

added numerous factual data to aid in the assessment of the anatomy and systematics of this family. 

Hopefully this new information will enable future 

studies to expand on some of the information I 

leaf axis. Sometimes the stomata appear to have presented here. 

Literature Cited 

E. S. 

Aerosol OT Solution, an Effective Softener of 

Herbarium Specimens for Anatomical Study. Stain 

Technology, 42: 155-156. 

The Anatomy of Barbaceniopsis, a New Genus Recently Described in the Velloziaceae. American 

Journal of Botany, 553399-403. 

Leaf-anatomy and Systematics of Old 12'orld Velloziaceae. Kew Bulletin, 23 (2) :315-335. 

The Identity of T'ellozia uaifianensis: Anatomical 

Evidence. Meinoirs of the A'ew Yoyk Botanical 

Garden, 18 (2) :291-298. 

Phytogeography and Evolution of the Velloziaceae. 

Pages 105-119 in B. J. Meggers, E. S. Ayensu, IV. D. 

Duckworth, editor, Tropical Forest Ecosystems in 

Africa ai7d South America: A Comparative Reuiew. 

FVashington, D.C.: Smithsonian Institution Press. 

k'elloziaceae. In Botany of the Guayana Highland. 

Memoirs qf the A'ew York Botanical Garden, 18 (2): 

3241. 

blaguire, B. 

1969. 

Menezes, N. L. de 

1971a. Traqueides de transfuao no penero Vellozia Vand. 

Cienciae Cultura, 23 (3) :389-409. 

1971b. Sew Taxa and New Combinations in Velloziaceae. 

Ciinciae Cultura, 23 (3) :421-422. 

Sletcalfe, C. R. 

1971. Anatomy of the Monocotyledons, V: Cyperaceae. 

Clarendon Press: Oxford. 

Rletcalfe, C. R., and Gregory, M. 

1964. Comparative Anatomy of Monocotyledons. Some 

New Descriptive Terms for Cyperaceae with a Discussion of Variation in Leaf Form Noted in the 

Family. Sotes from the Jodrell Laboratory, Royal 

Botanic Gardens, Kew, 1:l-11. 

Noher, de Halac, R., and A. E. Cocucci 

Sobre la naturaleza de 10s "appendices petaloides" 

en Barbacenia purpurea (Velloziaceae) . Kurtziana, 

1971. 

6: 265-269. 

Smith, L. B. 

1962. A Synopsis of the American Velloziaceae. Contributions to the United States A'ational Herbarium, 

35: 25 1-292. 

\$'arming, E. 

1893. Note sur la biologie et l'anatomie de la feuille des 

Vellosiacees. Oversight Kongelige danske Videnskabernes Selskabs Forhandlingar, pages 57-100. 
PLATES 1-51 

NUMBER 15 

PLATE I.-Vellozia daulsonii (Irwin et al. 12622), showing the distribution and the tuftlike 

appearance of hairs: a, adaxial view; b, abaxial view (X 200). Vellozia glandulifera (L. B. 

Smith 8c Ayensu 15998) : c-d, abaxial views of the epidermis illustrating the twisted nature of 

the hairs (c, X 200; d, X 2000). 

75 
76 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 2.-Surface view of abaxial leaf epidermis showing the arrangement of long multicellular 

hairs at the entrances of the furrows: a-b, t’ellozin durnitiana (Garcia-Barriga 13804) (a, X 95; 

b, X 190); c-d, V. lithophila (Cuatrecasas 7700) (c, X 90; d, X 440). 
XUMBER 15 

PLArr 3.-VeZlozin lnnntn (\'eralucia & Graziella s.n.) : a, adaxial epidermis showing the rounded 

base and the twisted naturc of the hairs (X 240) ; b, abaxial epidermis showing the flattened 

arrangement of the clumped hair (X 260); c-d, abaxial leaf epidermis showing finlike arrangement of hairs of Vellozia maculata (Glaziou 22218a) (c, X 65; d, X 270). 
78 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 4.-a-b, Vellozia maclirisinnn (Prance and Silka 58275) ; c-d, V. cinerascens (Lehnkner 

323), showing the t)pe and distribution of hairs on the abaxial surface (a, X 65; b, X 385; 

c, X 120; d, X 290). 
NUMBER 15 79 

PL.ATE 5.-a, Veliozia glochidea (Duarte 8340-Mattos 677) , abaxial epidermal surface exhibiting 

a nodular profile (X 1330) ; b, V. meizgerne (Engler s.n.) , a view through an abaxial furrow 

showing the disposition of the fingerlike projections (X 950) ; c, V. ciliata (L. B. Smith and 

Ayensu 15395), surface view of an abaxial furrow showing the arrangement of the fingerlike 

projections (X 1100) ; d, V. pumila (Glariou 22215), surface view of the abaxial epidermis 

exhibiting the arrangement of hairs oterarching the entrances to the furrows (X 180). 
80 SMITHSOXIAS COSTRIBUTIOSS TO BOTANY 

PLATE 6.-.4baxial surface Liew showing the arrangement and types of folding of the epidermal 

cells (a, X 245; b, X 950): a-b, Barbucenia ffuzia (Tryon and Tryon 6836). Surface view of 

abaxial epidermis showing the relationship between epidermal ridges and the arrangement of 

the stomata: c, B. luzulifolta (Pereira 2664-Pabst 3500); d, B. gounelleana (Pereira 7050) (c. 

X 875; d, X 950). 
NUMBER 15 

PLATE i.-Adaxial epidcrrnal surfaces illustrating the distribution of long hairs: a-b, Barbacenia 

gardneri (Pereiia 2849-Pabst 3685) (a, X 11.5; b, X 340) ; c-d, B. tornentosa (Irwin et al. 29559) 

(c, X 80; d, X 280). 

81 
82 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLAiE 8.-Surface views of epidermis showing the presence of coalescent hairs. Barbacenia 

mncrantha (Irwin et al. 2001i): a, abaxial surface showing hairs in the process of coalescence 

(X 430) ; b, transverse section of coalesced hairs on the abaxial surface (X 2860). Barbacenia 

nana (L. B. Smith and Alensu 15973): c, adaxial surface view (X 85); d, transverse section of 

nine coalesced hairs on adaxial surface (X 2275) . 
NUMBER 15 

PLATE 9.-Adaxial surface views of leaf epidermis showing the presence of hairs and their 

arrangement: a, Uarbacenia fragl’uns (Dueke s.n.) (X 85) ; b, B. iiedeliana (Hatschbach 27806) 

(X 170); c-d, Barbaceilia biackii (Macedo 2962) (c, X 95; d, X 250). 

a3 
PLATES l0-23.-Scanning electron micrographs illustrating epidermal surfaces 

showing the type and arrangement of stomata in Velloziaceae. 

PLATF IO.-T’ellozia hatschbachii (L. B. Smith and Ayensu 16002): 

a-b, abaxial surface (a, X 1000; b, X 5000). 
SUMBER 15 

PLATE ll.--Vellozia crassicaulis (L. B. Smith & Ayensu 15965): a, 

adaxial surface (X 500); b. abaxial surface (X 5000). 

85 
86 SMITHSOSL4N COSTRIBUTIONS TO BOTANY 

PLATE 12.--T’ellozia variabilis (L. B. Smith & Ayensu 15981), adaxial 

surface (a, X 1000; b, X 5000). 
NUMBER 15 

PLATE 13.--Veb~zia tubiflora (Tamayo 3105), abaxial surface (a, X 

1000; b, X 5000). 

87 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 14.-a, Velloiia magdalenae (Santos et al. 13298), abaxial surface (X 

4700); b, V. squdida (Duarte 6401), abaxial surface (X 2890). 
NUMBER 15 

PLATE 15.-a, Vellozia modesta (L. B. Smith et al. 15997), abaxial 

surface (X 4500); b, V. minima (Irwin et al. 20630), abaxial surface 

(X 4500). 

89 
90 SMITHSOSIAN CONTRIBUTIONS TO BOTANY 

PLATE 16.-a, Vellozia dasypus (Blanchet 3558) , abaxial surface (X 3750) ; 

b, V. grisea (Weddell 3006) , abaxial surface (X 7830). 
NUMBER 15 

PLATE 17.-a, Vellozia fibrosa (Azevedo 5592), abaxial surface (X 6600) ; 

b, V. lanata (Veralucia and Graziella s.n.) , adaxial surface (X 8075). 

91 
92 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 18.-a, Vellozia candida (Rose and Russell 20669), abaxial 

surface (X 5000) ; b, V. caruncularis (L. B. Smith and Ayensu 15977) , 

abaxial surface (X 5000). 
NUMBER 15 

PLATE 19.-Vellozia verruculosa (Williams 8112) , abaxial surfaces 

(a, X 3060; b, X 5310). 

93 
94 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 20.-a, Barbacenia paranaensis (Hatschbach 29212) , 

abaxial surface (X 5000) ; b, B. inuolvcrata (Irwin et al. 20973), 

adaxial surface (X 2000) . 
NUMBER 15 

PLATE 21.-a, Barbacenia hatschbachii (Hatschbach 24284), abaxial 

(X 3400) ; b, B. sellovii (Menezes and Continho 5), adaxial surface 

(X 4200). 

95 
96 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 22.-a, Barbacenia jragrans (Dueke s.n.) , abaxial surface (X 4250) ; 

b, B. nana (L. B. Smith and Ayensu l5973), adaxial surface (X 4350). 
NUMBER 15 

PLATE 23.-Barbacenia cmicostigna (L. B. Smith and Ayensu 15982), 

adaxial surfaces (a, X 430; b, X 4300). 

97 
PLATES 24-28.-Transverse sections of leaves of Velloziaceae showing sclerenchyma patterns and mesophyll. Each species may include sections of either the 

midvein, coastal, or intercoastal regions. 

PLATE 24.-a-b, Velloria abietina (Maguire et al. 49115) 

(X 25) ; c-e, V. data (Maguire et al. 49049) (X 25) : 

f-g, V. glabra (Maguire et al. 49048) (X 25). 
NUMBER 15 

PLATE 25.-a, Vellozia glabra (Maguire et al. 49048) (X 25); b-d, V. compacta (Irwin 2407) 

(X 25) ; e-f, V. compacta (Segades-Vianna and Loredo 1085) (X 25). 

99 
100 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 26.-a-c, Vellozia verruculosa (Williams 8112) (X 25) ; d-f, V. fibrosa (Azevedo 5592) 

(X 25); g-h, V. variabilis (Irwin et al. 9809) (X 25). 
NUMBER 15 

PLATE 27.-a, Vellozia variabilis (Irwin et al. 9809) (X 25) : b-d, V. variabilis (Irwin et al. 

9915) (X 25); e-f, V. variabilis (Maguire et al. 49262) (X 25) ; g, V. angustifolia (Archer 

4092) (X 25). 

101 
102 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 28.-a-b, Vellozia angustifolia (Archer 4092) (X 25); c-e, V. angustifolia (Riedel 1824) 

(X 25) ; f-g, V. intermedia (L. B. Smith 6854) (X 25). 
NUMBER 15 

PLATE 29.-a-c, Vetlozia squalida (Archer and Barreto 4937) (X 25) : 

d-f, V. aloifolia (Mexia 5870) (X 25). 

103 
104 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

PLATE 30.-a-c, Vellozia taxifolia (Mexia 4289) (X 25) ; d-f, V. Zeptopetala (Maguire et al. 

49184) (X 25) ; g-h, V. resinosa (Maguire et al. 44694) (X 25). 
NUMBER 15 

PLATE 31.-a, Vellozia resinosa (Maguire et al. 44694) (X 25) ; b-d, V. epidendroides (Maguire 

et al. 49060) (X 25) ; e-g, V. flavicans (Gardner 4370) (X 25) ; h, V. glochidea (Irwin et al. 

14584) (X 25). 

105 
106 ShiITHSONIAh’ CONTRIBUTIONS TO BOTANY 

PLATE 32.-a-b, Vellozia glochidea (Irwin et al. 14584) (X 25) ; c-e, V. crassicaulis (Irwin et al. 

11699) (X 25) ; f-g, V. crassicaulis (Irwin et al. 13377) (X 25). 
NUMBER 15 

PLATE 33.-a, Vellozia crassicaulis (Irwin et al. 13377) (X 25) ; b-c, V. swallenii (Pires 9712) 

(X 25) ; d-f, V. dasypus (de Lemos and Froes 20246) (X 25); g-h, V. dnsypus (Rose and 

Russell (9697) (X 25). 

107 
108 SMITHSONIAS CONTRIBUTIOKS TO BOTASY 

PLATE 34.-a, Velloria dasypus (Rose and Russell 19697) (X 25); b-d, V. caruncularis (L. B. Smith 

6855) (X 25); e-f, V. exilis (Irwin et al. 9507) (X 25); g-h, u. candida (Carauta 1868) (X 25). 
NUMBER 15 

PLATE 35.-a, Vellozia candida (Carauta 1868) (X 25); b-d, V. machrisiana (Irwin 12697) 

(X 25) ; e-g, V. tubipora (Lasser 1777) (X 25) : h, V. tubiflora (Maguire et al. 35131) (X 25). 

109 
110 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 36.-a, Vellozia caput-ardeae (L. B. Smith and Ayensu 15989); b, V. ciliata (L. B. Smith 

and Ayensu 15595); c, V. COS~~~Q (Irwin et al. 20999); d, V. declinans (Hatschbach 27842) ; 

e, V. echinata (Riedel 1780); f, V. ghdulifera (L. B. Smith and Ayensu 15998) (a-f, X 100). 
NUMBER 15 111 

PLATE 37.-a, Vellozia hatschbachii (L. B. Smith and Ayensu 16002); b, V.’ hirsuta (L. B. 

Smith and Ayensu 15990); c, V. hypoxoides (Irwin et al. 34120); d, V. lappa (L. B. Smith and 

Ayensu 15988); e, V. nuda (Hatschbach and Pelanda 28029); f, V. ornata (L. B. Smith and 

Ayensu 16001) (a-f, X 100). 
112 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 38.-a-b, Velloria tubiflora (Maguire et al. 35131); c-e, Barbacenia seubertiana (Strang 

239); f-h, B. gounelleana (Emygdio 1428) (a-h, X 25). 
NUMBER 15 

PLATE 19.-a-c, Barbacenia purpurea (Schott s.n. U.S. 151098) : d, B. schwackeii (Heringer 

5263); e-f, B. ignea (Williams & Assis 6817) (a-f, X 25). 

113 
114 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 40.-a, Barbacenia ignea (Williams and Assis 6817) ; b-d, B. williarnsii (Williams and 

Assis 6386) ; e-g, B. flaw (Maguire et al. 49116) ; h, B. viscosissirna (Macedo 2826) (a-h, X 25). 
NUMBER 15 

PLATE 41.-a-b, Barbacenia uiscosissimia (Macedo 2826) ; c-e, B. gentianoides (Segades-Vianna 

and Loredo 1066); f-h, B. ZongiPora (Irwin 2486) (a-h, X 25). 

115 
116 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 42.-a, Velloria bicolor (Maguire et al. 49047) (X 25) ; b, V. brachypoda (Irwin et al. 

23534) (X 25): c, V. cinerascens (Lenhkner 323) (X 25) : d, V. crinita (Glaziou 16388) (X 25) ; 

e, V. grim (Weddell 3006) (X 150) : f, V. incuruata (Pereira 2959-Pabst 3795) (X 25). 
NUMBER 15 117 

PLATE 43.-a, Vellozia irwinii (Irwin et al. 20998): b, V. magdalenae (Lima and Brade 14288) ; 

c, V. rnetzgerae (Engler m.); d, V. modesta (Hatschbach and Pelanda 27869) ; e, V. pilosa 

(Glaziou 19933) ; f, V. piresiana (Duarte 2068) (a-f, X 25). 
118 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 44.-a, Vellozia plicata (Frazao s.n.) (X 25) ; b, V. pumih (Irwin et al. 9230) (X 150) ; 

c, V. punctulata (Blanchet 2561) (X 150); d, V. tenella (Damazio 91185) (X 25); e, V. teres 

(Irwin et al. 21000) (X 25) ; f, V. triquetra (Gardner 2735) (X 150). 
NUMBER 15 

PLATE 45.-a, Vellozia wasshausenii (Irwin et al. 20318) (X 25); b, V. cam (Irwin et al. 

12556) (X 150); c, V. dawsonii (Irwin et al. 12648) (X 150); d, V. dumitiana (Schultes and 

Cabrera 15050A) (X 150) ; e, V. lanata (Pires 9895) (X 25) ; f, V. rnaculata (Glaziou 222-18a) 

(X 25). 

119 
PLATE 46.-a, Vellozia phalocarpa (Anderson et al. 35855); b, V. riedeliana (L. B. Smith and 

Ayensu 15983); c; V. sellovii (Pereira 2677); d, V. uirgata (Schwacke 5889); e, Barbacenia 

conicostigniu (L. B. Smith and Ayensu 15982); f, B. delicatula (L. B. Smith and Ayensu 

15975) ; g, U. exscapa (Anderson et al. 35831) (a-g, X 100). 
NUMBER 15 121 

PLATE 47.-a. Barbacenia gardneri (Pereira 2849-Pabst 3685) ; b, B. hatschbashii (Hatschbach 

24284); c, B. involucrata (Irwin et al. 20973); d, B. longiscapa (Irwin et al. 22186); e, B. 

luzulifolia (Pereira 2664-Pabst 3500); f, B. nana (L. B. Smith and Ayensu 15973) ; g, B. riedeliana 

(Anderson et al. 35123) ; h, B. sellbvii (Continho 5) (a-h, X 100). 
122 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 48.-a, Barbacenia irwiniana (Porto 1140) (X 25) ; b, B. celiae (Maguire et al. 40298) 

(X 25) ; c, B. blackii (Macedo 2962) (X 25) ; d, B. macrantha (Irwin et al. 20071) (X 25) ; 

e, B. paranaensis (Hatschbach 15715) (X 150) . 
NUMBER 15 125 

PLATE 49.-SEM photomicrographs of leaf of Vellozia ciliata: a, transverse section of lamina 

showing the arrangement of sclerenchyma, mesophyll tissue, and furrow (X 210); b, an enlargement of the furrow showing the arrangement of the fingerlike projections (X 630) ; c, longitudinal section through a lamina exposing one surface of a furrow. Note the sporadic distribution 

of the fingerlike projections and stomata (X 200). 
124 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

PLATE 50.-Transverse sections of leaves viewed with the SEM to show sclerenchyma and 

mesophyll tissues: a, Vellozia hypoxoides (Irwin et al. 34120) 

and Russell 20669) (X 240); c, V. grisea (Weddell 3006) (X 315). 

(X 335) ; b, V. candida (Rose 
NUMBER 15 125 

PLATE 51.-Transverse sections of leaves viewed with the SEM to show surface and internal 

structure: a, Vellozia dawsonii (Irwin et al. 12622), illustrating the surface arrangement of 

tuft hairs, the clear delineation of palisade cells, and spongy mesophyll (X 110) ; b, Barbacenia 

gounelleana (Pereira 7050), showing a typical example of Barbacenia type of sclerenchyma 

and mesophyll patterns (X 265). 

f? U.8. GOVERNMENT PRINTING OFFICEi lB74- S46/358/8