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 Vello- 
ziaceae has been studied with the purpose of providing important character- 
states 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 sur- 
face 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 
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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. 
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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 
1 
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Vellozia leptopetala Goethart & Henrard ........................ 
31 
31 
Vellozia metzgerae L. B. Smith ..... 
Vellozia minima Pohl 33 
33 
Vellozia modesta L. B. Smith & Ayensu ................. 
Vellozia nuda L. B. Smith & Ayensu , , . . , . 
33 
Vellozia ornata Martius ex Schultes f. .......................... 34 
34 
Vellozia phalocarpa Pohl ...................................... 
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 Barbaceniop- 
sis (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 sur- 
rounded by sclerenchyma on the adaxial side in 
the form of an inverted crescentiform or V-shaped 
cap. The abaxial sclerenchyma is either U- or Y- 
shaped. 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 dis- 
Edward S. Ayensu, Department of Botany, Smithsonian Insti- 
tution, 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 speci- 
mens (106 species of two genera) were examined 
for this study. The herbarium vouchers are de- 
posited 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) high- 
way 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 Con- 
tagem, 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, Colom- 
bia, 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, Diaman- 
tina, 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, Ama- 
zonas, Guayana, NY; Maguire et al. 29159, Amazonas, 
Guayana, NY; Maguire et al. 30941, Amazonas, Guayana, 
NY; Maguire et al. 29245, Amazonas, Guayana, NY; Ma- 
guire 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; Ma- 
guire 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; Ma- 
guire et al. 45918, Amazonas, Guayana, KY; E. Foldats 
3526 (1960) Cerca del Puerto Ayacucho, Terr. Fed. Ama- 
zonas, 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 vene- 
zalana, 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; Segadas- 
Vianna & 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.-Segadas- 
Vianna & 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 & Con- 
tinho 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 speci- 
mens. The specimens were soaked in a mixture of 
Aerosol OT solution and distilled water (cf. 
Ayensu, 1967) until maximum expansion was at- 
tained. 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 photomicro- 
graphs were prepared, using a Wild M20 micro- 
scope 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 longitudi- 
nal 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 sur- 
face. 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 con- 
nected to the translucent layers radially arranged 
adaxially on vascular bundles of Vellozia species. 
(h) The central and marginal bundles are asso- 
ciated 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 mid- 
portion 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. rubro- 
uirens, 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, Vel- 
Zozia 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. beau- 
verdii) ..................................................................... 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 combi- 
nations 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 
a- 
e 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 con- 
floral morphology in the family. tribution to Dr. Lyman B. Smith, Senior Botanist, 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
Smithsonian Institution, and the foremost au- 
thority on the taxonomy of Velloziaceae, on the 
occasion of his seventieth birthday. 
I am most grateful to Ms. Cynthia Bailey, Re- 
search 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 Micro- 
scope 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 un- 
dulating. 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 ac- 
companied by partial adaxial girder or cap and 
abaxial girder. Vascular bundle completely sur- 
rounded by a distinct bundle sheath with exten- 
sions of two to four cells toward adaxial epidermis 
only. Crystals and tannin: not observed. 
NOTE: Although this specimen has all the Vel- 
lozia 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: adax- 
ial 'and abaxial cells rectangular. Stomata: tetra- 
cytic, 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: ad- 
axial 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 ab- 
axial 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; commis- 
sural bundles few. Large veins each with one to 
three wide vessels. Two phloem units lying lat- 
erally 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. Epi- 
dermis: 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. Epi- 
dermis: two to three rows of adaxial cells square 
to rectangular of thin walls; abaxial cells consist- 
ing of fibrous sclerenchyma except where stomata 
occur. Cuticle: thick, undulating, but conspicuous- 
ly 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 sub- 
jacent 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 bun- 
dles, furrows, and above midvein distinctly trans- 
lucent 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: ad- 
axial 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 un- 
dulating; abaxial surface furrowed about half of 
blade thickness. Epidermis: cells more or less 
rounded or square. One or two rows of thick- 
walled 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 fur- 
rows 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 lat- 
erally in abaxial or slightly partial V-shaped girder. 
Each vascular bundle always accompanied by adax- 
ial 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 ob- 
served. 
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; Oc- 
chiouni s.n. 
SURFACE Vmv.-Hairs: absent. Epidermis: adax- 
ial 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 sur- 
face 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; inter- 
mitent rows of thick-walled fibers observed sub- 
jacent to epidermis, especially toward leaf margin. 
Cuticle: uniformly thickened on adaxial and abax- 
ial surfaces but not observed in furrows. Stomata: 
present in furrows; guard cells thick walled; sub- 
stornatal chamber inconspicuous. Mesophyll: adax- 
ial palisade four to six layered, occupying about 
one-third of mesophyll and gradually changing into 
spongy tissue; two to four rows of distinctly trans- 
lucent cells arranged radially above vascular bun- 
dles 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 later- 
ally in flanges of partial abaxial U-shaped girder. 
Each vascular bundle always accompanied by ad- 
axial cap. Vascular bundles- completely surrounded 
by a distinct thin-walled bundle sheath. Crystals 
and tannin: not observed. 
NOTE: The partial abaxial U-shaped girder re- 
calls 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 rec- 
tangular, 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 sur- 
face unevenly undulating; abaxial surface fur- 
rowed, 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, occa- 
sionally three strands observed. Directly below 
fiber strands and subepidermal cells is a layer of 
large somewhat translucent parenchyma cells sep- 
arating 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 tis- 
sue; 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; com- 
missural 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 abax- 
ial 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. Adax- 
ial surface ridged; abaxial surface slightly undulat- 
ing; 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 paren- 
chyma 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 tis- 
sue. Few rectangular translucent palisade cells lar bundle. Crystals and tannins: none observed. 
arranged radially above vascular bundles and fur- 
rows. 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 vascu- 
Vellozia 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: adax- 
ial and abaxial cells, rectangular, thin walled. 
Stomata: paracytic, 24 X 15 pm; observed on ab- 
axial surface only. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with median adaxial groove. Adaxial sur- 
face conspicuously ridged and more or less slightly 
furrowed; abaxial surface distinctly furrowed. Epi- 
dermis: 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; substoma- 
tal 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 lat- 
erally 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 lat- 
eral epidermal projections, recalls the same condi- 
tions 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: adax- 
ial 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 sub- 
jacent to epidermis; tangentially oriented trans- 
lucent 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. Mes- 
ophyll: 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 adax- 
ially on vascular bundles and inner limits of fur- 
rows connecting three conspicuous parenchyma 
cells. Palisade cells compactly arranged and filled 
with chloroplast. Vascular bundles: 17-20; com- 
missural 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 adax- 
ial 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 re- 
markable 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 rectangu- 
lar; thin walled. Stomata: paracytic, 18 X 12 pm; 
present on abaxial surface. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with small median adaxial groove. Adax- 
ial 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 scleren- 
chyma 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. Epi- 
dermis: 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 thick- 
ness of blade. Epidermis: adaxial cells dome shaped 
to conical; interspersed with large strands of scle- 
renchyma 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 scleren- 
chyma strands; substomatal chamber present. Mes- 
ophyll: 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. Crys- 
tals 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: adax- 
ial and abaxial cells mostly rectangular but often 
square shaped, thin walled. Stomata: tetracytic, 24 
X 15 pm; present on both surfaces but highly con- 
centrated on abaxial surface. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
shallowly V-shaped with slight median adaxial 
groove. Adaxial surface undulating; abaxial sur- 
face furrowed abofit half blade thickness. Epider- 
mis: 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 sur- 
faces. Stomata: mostly confined to furrows in 
abaxial surface; substomatal chamber inconspicu- 
ous. 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. Pali- 
sade cells and spongy mesophyll compactly ar- 
ranged 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 ac- 
companied 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: adax- 
adaxial sclerenchyma shapes range from strictly ial and abaxial cells rectangular, thick walled; occa- 
cap 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. Adax- 
ial 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 sub- 
jacent 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 differen- 
tiated 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 lat- 
erally 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: adax- 
ial 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 un- 
dulating; 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 baculi- 
form sclerenchyma strands, separated from each 
other by one radial row of parenchyma, occur sub- 
jacent to adaxial epidermis. Cuticle: uniformly 
thin, slightly thicker around margins and median 
abaxial surface. Stomata: present in abaxial fur- 
rows, few on adaxial surface. Mesophyll: adaxial 
palisade three layered, occupying about half of 
mesophyll; palisade separated from adaxial scle- 
renchyma strands by one layer of distinct trans- 
lucent parenchyma cells; two to three radially 
oriented large translucent cells arranged adaxially 
on vascular bundles. Palisade cells compactly ar- 
ranged and filled with chloroplast; spongy mes- 
ophyll 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 sur- 
rounded 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 fur- 
row to the other. Furthermore, the specimen ex- 
hibits 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 distin- 
guish this species. 
Vellozia crinita Gwthart & Henrard 
PLATE 42d 
SPECIMENS EXAMINED.-GlaziOu 16388. 
SURFACE VIEw.-Hairs: absent. Epidermis: adax- 
ial 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 sur- 
face 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 inter- 
spersed 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 com- 
inissural 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; ad- 
jacent to epidermis on abaxial surface. Epidermal 
extensions present in furrows. Few abaxial scle- 
renchyma strands observed on lower edge of fur- 
rows. 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: adax- 
ial and abaxial cells mostly of square cells, occa- 
sionally 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 sur- 
face furrowed about one-fifth to one-seventh of 
blade thickness. Epidermis: cells more or less 
square; abaxial cells almost replaced by fibers ex- 
cept 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 trans- 
lucent parenchyma cells; two to five radially 
oriented large translucent cells arranged adaxially 
on vascular bundles. Palisade cells compactly ar- 
ranged and filled with chloroplast; palisade oc- 
cupying 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 com- 
pletely 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 pro- 
nounced, especially at the middle of the inter- 
costal region. 
Vellozia declinans Goethart & Henrard 
PLATE 36d 
SPECIMENS EXAMINED.-Hatschbach 27842. 
SURFACE VIEw.-Hairs: few small papillae, ap- 
pearing 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 al- 
most 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 adax- 
ial epidermis occur two to three layers of scle- 
renchyma strands interspersed with rows of 
parenchyma cells of two to three layers. Below 
sclerenchyma strands is a distinct layer of paren- 
chyma cells. Cuticle: very thin and ridged on both 
surfaces. Stomata: present mainly in abaxial fur- 
rows; small substomatal chamber present. Meso- 
phyll: 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: adax- 
ial 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 sur- 
face rounded to conical, few rectangular; thin 
walled. Cells in furrows with fingerlike projections, 
subjacent to adaxial epidermis is a layer of paren- 
chyma 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 abax- 
ial 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 chang- 
ing abruptly into loosely spongy tissue; some large 
spongy tissue present. Vascular bundles: 25; com- 
missural bundles not 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; sometimes partial girders 
observed. Small adaxial cap present on each vascu- 
lar 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 abax- 
ial 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; mar- 
gins of lamina turned inversely. Adaxial surface 
slightly undulating; occasionally somewhat ridged; 
abaxial surface furrowed one-fifth to one-half thick- 
ness of blade. Epidermis: adaxial cells more or less 
dome shaped, interspersed with conical cells; abax- 
ial cells mainly rounded; cells thin walled. Sub- 
jacent 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 scle- 
renchyma. Cuticle: thick on abaxial and adaxial 
surfaces; smooth. Stomata: present in abaxial fur- 
rows; somewhat associated with epidermal pro- 
tuberances. Mesophyll: adaxial palisade cells one 
to two layers, gradually changing into spongy tis- 
sue; 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 varia- 
tions 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 char- 
acteristics. 
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. Epi- 
dermis: 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. Epi- 
dermis: 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 paren- 
chyma 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 chang- 
ing 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. Lat- 
eral extensions formed on either side of median 
abaxial vein have compactly arranged spongy mes- 
ophyll and a broad band of sclerenchyma cells. 
Bundle sheath completely surrounding vascular 
bundles. Abaxial strands present. Crystals and tan- 
nins: 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 abax- 
ial surface. 
TRANSVERSE SECTION OF LAMrNA.-Dorsiventral; 
V-shaped with small median adaxial groove; mar- 
gins turned inversely, flanged. Adaxial surface very 
slightly undulating; abaxial surface furrowed one- 
half thickness of blade. Epidermis: cells more or 
less square, rounded or rectangular; walls slightly 
thick, especially on inner tangential wall. Sub- 
jacent to adaxial epidermis is one to two layers 
of thick-walltd parenchyma cells interspersed with 
sclerenchyma strands; below parenchyma and scle- 
renchyma is a layer of translucent parenchyma 
cells. Cuticle: adaxial surface thicker than abax- 
ial; both slightly ridged. Stomata: present in fur- 
row 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 trans- 
lucent palisade cells arranged radially above vas- 
cular 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 surround- 
ed 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 en- 
countered 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: adax- 
ial and abaxial cells rectangular; thin walled. Sto- 
mata: paracytic, 21 X 18 pm; present mainly on 
abaxial surface. 
TRANSVERSE SECTION OF LAMINA.-DOrSiVentral; 
V-shaped with small median adaxial groove. Adax- 
ial 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 scleren- 
chyma 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 adax- 
ial surface; almost flush with surface; small sub- 
stomata1 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 com- 
missural 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. Abax- 
ial 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. Epi- 
dermis: 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 in- 
terspersed with one to two layers of sclerenchyma 
strands. Below parenchyma and sclerenchyma 
layers is a distinct layer of large translucent paren- 
chyma 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. Vas- 
cular bundles: 16-46; no commissural bundles ob- 
served. One to two large vessels in each vascular 
bundle, mostly one. Two phloem units lying lat- 
erally in flanges of Y-shaped abaxial girder. Each 
vascular bundle has an adaxial cap. Some scleren- 
chyma strands occur between abaxial epidermir 
and abaxial parenchyma layer. Bundle sheath com- 
pletely 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 rec- 
tangular, few almost rounded; thin walled. Stomata: 
pnracylic, 24 X 15 iim; present on abaxial sur- 
face, 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 undu- 
lating 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 sur- 
faces. Stomata: present in abaxial furrows in asso- 
ciation 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. Vas- 
cular bundles: 22; commissural bundles not ob- 
served. One to three large vessels present in each 
vascular bundle. Two phloem units laterally ar- 
ranged in flanges of Y-shaped abaxial girder. Adax- 
ial 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; Ma- 
guire et al. 44793. 
SURFACE VIEw.-Hairs: absent. Epidermis: cells 
square to rectangular; thin walled. Stomata: para- 
cytic, 21 X 18 pm; present on abaxial surface. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
open V-shaped with margins turned slightly in- 
versely. Adaxial surface slightly undulating; abax- 
ial 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; com- 
missural 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 pres- 
ent between abaxial epidermis and a distinct layer 
of parenchyma cells. Bundle sheath extends com- 
pletely 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 paren- 
chyma 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 fur- 
rows; 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 scleren- 
chyma strands or layers present (three to four lay- 
ered 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 abax- 
ial surface; seen on both surfaces in Glaziou 19934. 
TRANSVERSE SECTION OF LAMINA.-DorSiventral; 
V-shaped with margins turned very slightly in- 
versely; small median adaxial groove; groove large 
in Glaziou 19934. Adaxial surface slightly undu- 
lating, 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 thick- 
walled 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; substo- 
matal chamber virtually absent. Mesophyll: three 
layered palisade grading into spongy tissue; cells 
near abaxial surface more compactly arranged; 
spongy tissue in Glaziou 19934 completely oblit- 
erated 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: para- 
cytic, 15 X 9 pm; present on abaxial and few on 
adaxial surface. 
TRANSVERSE SECTION OF LAMrNA.-Dorsiventral; 
V-shaped with deep median adaxial groove. Adax- 
ial 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 in- 
terspersed with parenchyma cells. Below scleren- 
chyma is a layer of parenchyma cells. Cuticle: 
uniformly ridged and thick on both surfaces. Sto- 
mata: present in furrows; substomatal chamber 
virtually absent. Mesophyll: three to four layers 
indistinct palisade cells grading into compact 
spongy tissue; few translucent palisade cells ra- 
dially 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 addi- 
tion to Y-shaped abaxial girder an adaxial cap. 
Vascular bundle completely surrounded by bundle 
sheath. Few strands of abaxial sclerenchyma ob- 
served. 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. Sto- 
mata: paracytic, 21 X 9 pm; present on abaxial 
surface. 
TRANSVEXSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with small median adaxial groove. Adax- 
ial surface very slightly undulating; abaxial surface 
furrowed one-half to three-fourths thickness of 
blade. Epidermis: adaxial cells rounded; few rec- 
tangular; thin walled. Abaxial cells rounded; inter- 
spersed with small sclerenchyma strands. Subjacent 
to adaxial epidermis occur two to three layers of 
sclerenchyma strands interspersed with radially ar- 
ranged 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 sto- 
mata. Mesophyll: three to four, mostly three, layers 
of compactly arranged small palisade tissue chang- 
ing 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 vas- 
cular bundle. Two phloem units arranged laterally 
in flanges of partial Y-shaped abaxial girder. Small 
adaxial cap present on each vascular bundle. Abax- 
ial 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: para- 
cytic, 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 accom- 
panied by sclerenchyma caps; minor veins with 
adaxial cap and abaxial U- or V-shaped girder com- 
pletely 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 hook- 
like 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 rectangu- 
lar; thin walled. Abaxial cells square to rectangu- 
lar, 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 fur- 
rowed 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 epi- 
dermis. Mesophyll: two to three layers of palisade 
tissue subjacent to adaxial epidermis, rather indis- 
tinct 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 ob- 
served. 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 un- 
dulating; 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 fur- 
rows; 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 ar- 
ranged radially above vascular bundles and fur- 
rows, 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 ob- 
served. 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: para- 
cytic, 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 fur- 
rowed about one-half thickness of blade. Epider- 
dis: adaxial cells rounded to dome shaped; abaxial 
cells rounded. One to two layers parenchyma sub- 
jacent to epidermis; below parenchyma cells are 
two to three layers of sclerenchyma strands inter- 
spersed 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 fur- 
rows; 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. Vas- 
cular bundles: 55; few commissural bundles ob- 
served. One to two large vessels, mostly one. TWO 
phloem units lying laterally in flanges of Y-shaped 
abaxial girder. Adaxial cap present on each vas- 
cular bundle. One to two layers sclerenchyma cells 
between abaxial epidermis and a distinct paren- 
chyma layer. Vascular bundle completely sur- 
rounded by bundle sheath. Crystals: none ob- 
served. Tannins: present. 
Velloziu irwinii L. B. Smith 
PLATE 43a 
SPECIMENS EXAMINED.-Irwin, Maxwell, Was- 
shausen 20998. 
SURFACE VIEw.-Hairs: multicellular, present on 
adaxial surface. Epidermis: cells circular to rec- 
SMITHSONIAN COXTRIBUTIONS TO BOTANY 
tangular, thin walled. Some crystals present. Sto- 
mata: 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 fur- 
rowed over one-half the thickness of blade. Epi- 
dermis: 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 abax- 
ial 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. Epi- 
dermis: cells on adaxial and abaxial surfaces 
square to rectangular; thin walled. Stomata: para- 
cytic, 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 al- 
most smooth; abaxial surface furrowed about one- 
half thickness of blade. Epidermis: adaxial cells 
square to rectangular in shape; thin walled. Abax- 
ial 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 in- 
terspersed 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 sub- 
stomatal chamber present. Numerous furrow pro- 
jections present. Mesophyll: three to four layers 
palisade tissue grading into compactly arranged 
spongy tissue. Three to four layers of large trans- 
lucent 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: para- 
cytic, 15 X 12 pm; present on abaxial surface. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with small median adaxial groove. Adax- 
ial surface very slightly undulating; abaxial sur- 
face 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; sub- 
stomatal 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 com- 
pletely 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 flat- 
tened 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; pres- 
ent on abaxial surface. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with small median adaxial groove. Adax- 
ial surface slightly undulating; abaxial surface fur- 
rowed 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 paren- 
chyma 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; sur- 
rounded with epidermal projections in furrows, 
small substomatal chamber observed. Mesophyll: 
distinct two- to three-layered palisade tissue grad- 
ing 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 be- 
tween abaxial epidermis and a layer of parenchyma 
cells. Bundle sheath completely surrounding each 
vascular bundle. Crystals and tannins: none ob- 
served. 
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: para- 
cytic, 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 undu- 
lating, abaxial surface furrowed one-third to one- 
halt the thickness of blade. Epidermis: adaxial 
cells rectangular; abaxial cells rounded to rec- 
tangular; 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 fur- 
rows; substomatal chamber present; associated with 
fingerlike projections. Mesophyll: three- to four- 
layered 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 scleren- 
chyma 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 pres- 
ent. 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 por- 
tion 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: para- 
cytic, 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 one- 
half thickness of blade. Epidermis: adaxial cells 
mostly rectangular; abaxial cells rounded to dome 
shaped; cells on both surfaces thin walled. Sub- 
jacent 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 sur- 
face; substomatal chamber present; stomata asso- 
ciated 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 scleren- 
chyma 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 scle- 
renchyma 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 pres- 
ent. Epidermis: adaxial cells square to rectangular; 
abaxial cells rounded. Walls thin on both sur- 
faces. 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 pres- 
ent. hlesophyll: three- to four-layered palisade cells 
grading into compactly arranged spongy tissue. 
Two- to three-layered large translucent cells ar- 
ranged 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. In- 
verted V-shaped adaxial cap present on each vascu- 
lar bundle. Abaxial sclerenchyma strands present 
in abaxial corners of furrows and extending partly 
into furrows. Bundle sheath completely surround- 
ing 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. Ex- 
treme margins of lamina turned inversely. Adaxial 
surface slightly undulating; abaxial surface fur- 
rowed 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 abax- 
ial surfaces. Stomata: present on adaxial surface 
and in abaxial furrows, more numerous on abaxial 
surface; substomatal chamber present on both sur- 
faces: 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 trans- 
parent cells arranged radially above vascular 
bundles and furrows. Large translucent cells occur 
above entire area of midvein. A layer of paren- 
chyma 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. Tan- 
nins: few present. 
Vellozia phalocarpa Pohl 
PLATE 46a 
SPECIMENS EXAMINED.-Anderson et al. 35855. 
SURFACE VIEw.-Hairs: absent. Epidermis: adax- 
ial 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. Ex- 
treme margins of lamina turned slightly inversely. 
Adaxial surface slightly undulating; abaxial sur- 
face furrowed one-third to one-half thickness of 
blade. Epidermis: cells on adaxial and abaxial sur- 
faces rounded; wall thickened. Adaxial epidermis 
interspersed with strands of sclerenchyma. Sub- 
jacent to epidermis and strands of fibers occur two 
to three layers of parenchyma cells. Cuticle: mark- 
edly thick and slightly ridged on both surfaces. 
Stomata: present in abaxial furrows; virtually no 
substomatal chamber observed; somewhat asso- 
ciated with small projections in furrow. Meso- 
phyll: 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 trans- 
lucent 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 epi- 
dermis 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. Adax- 
ial surface distinctly ridged; small furrows on 
abaxial surface. Epidermis: abaxial and adaxial 
cells rounded to dome shaped; cells on both sur- 
faces 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 ob- 
served. 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 scle- 
renchyma present. Bundle sheath surrounding each 
vascular bundle. Crystals and tannins: none ob- 
served. 
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 sur- 
face furrowed about one-third the thickness of 
blade. Epidermis: adaxial and abaxial cells rec- 
tangular 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 fur- 
rows; no substomatal chamber observed. Meso- 
phyll: three layers of indistinct palisade tissue 
grading into fairly compactly arranged spongy 
tissue. Large translucent cells arranged radially 
above vascular bundles and furrows; adaxial trans- 
lucent 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 lat- 
erally 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. Epi- 
dermis: 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 speci- 
mens; 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 adax- 
ial epidermis occur one to two layers of scleren- 
rhyma strands interspersed with parenchyma cells. 
Below sclerenchyma strands is a layer of paren- 
chyma 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 pali- 
sade 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 strik- 
ing 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 abax- 
ial 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 in- 
terspersed with parenchyma cells. Below scleren- 
chyma 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 ar- 
ranged 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. Epider- 
mis: 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; mar- 
gins 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 trans- 
lucent 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 epi- 
dermis 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 abax- 
ial surface. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with median adaxial groove; margins 
turned slightly inversely. Adaxial surface slightly 
undulating; abaxial surface furrowed about one- 
half 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 scle- 
renchyma strands. Below is a distinct layer of 
parenchyma cells. Cuticle: thick on adaxial sur- 
face; about one-half as thick on abaxial surface; 
smooth on both surfaces. Stomata: present in fur- 
rows; associated with hairlike protrusions; sub- 
stomata1 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 vas- 
cular 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: adax- 
ial 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. Epider- 
mis: 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 sur- 
faces, 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 mid- 
vein region and radially above each vascular bun- 
dle. 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 vas- 
cular bundle. Crystals and tannins: none observed. 
NOTE: The mesophyll in this species is poorly 
differentiated and therefore resembles the meso- 
phyll 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. Sto- 
mata: paracytic, 27 X 15 pm; present on the abax- 
ial surface. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with small median adaxial groove; mar- 
gins turned very slightly inversely. Adaxial surface 
slightly undulating; abaxial surface furrowed 
about one-half thickness of blade. Epidermis: adax- 
ial 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. Sto- 
mata: present in abaxial furrows; small substomatal 
chamber present; furrows lined with epidermal 
projections. Mesophyll: three layers of palisade 
cells grading into compactly arranged spongy tis- 
sue. 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 vas- 
cular 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 thick- 
ness 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 inter- 
spersed 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. Crys- 
tals 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 cham- 
ber virtually absent. 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 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. Adax- 
ial surface slightly undulating; abaxial surface 
furrowed about one-half thickness of blade. Epi- 
dermis: 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 abax- 
ial 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. Epider- 
mis: cells square to rectangular on both surfaces; 
thin walled. Stomata: paracytic; 21 X 12 pm; pres- 
ent 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 one- 
half 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 ob- 
served. 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. Vas- 
cular bundles: 28; no commissural bundles ob. 
served. One to two large vessels present in each 
vascular bundle. Two phloem units arranged lat- 
erally 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; Claus- 
sen 13, 1794. 
SURFACE VIEw.-Hairs: few present in small 
tufts. Epidermis: cells mostly square to rectangular 
on both surfaces, few rounded, thin walled. Sto- 
mata: paracytic, 15 X 9 pm; present on abaxial 
and adaxial surfaces. 
TRANSVERSE SECTION OF LA;LIINA.-Dorsiventral; 
V-shaped with small median adaxial groove. Adax- 
ial and abaxial surfaces only very slightly undu- 
lating. 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 sur- 
face; 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 ob- 
served. 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 com- 
missural 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 vas- 
cular bundle and extending almost to adaxial 
epidermis. Each vascular bundle completely sur- 
rounded 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. Tan- 
nins: 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. Adax- 
ial 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 fur- 
rows; 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 vas- 
cular 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 bun- 
dle. 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. Adax- 
ial surface slightly undulating; abaxial surface 
furrowed about one-half thickness of blade. Epi- 
dermis: cells on both surfaces rounded to rectangu- 
lar; 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 epi- 
dermis 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 paren- 
chyma 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; sub- 
stomata1 chamber not observed. Mesophyll: three- 
to four-layered palisade cells changing abruptly 
into compactly arranged spongy tissue. Two to 
three large translucent palisade cells radially ar- 
ranged 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 ar- 
ranged in flanges of Y-shaped abaxial girder. Abax- 
ial 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. Adax- 
ial 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 epi- 
dermis 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 adax- 
ial surface; fairly thick and slightly ridged on 
abaxial surface. Stomata: present in abaxial fur- 
rows 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: adax- 
ial and abaxial cells mostly rectangular, generally 
thin walled; slightly thick on veins. Stomata: para- 
cytic, 18 X 12 pm; observed on both surfaces. 
TRANSVERSE SECTION OF LAMINA.-Dorsiventral; 
V-shaped with slight median adaxial groove. Adax- 
ial surface evenly undulating; abaxial surface 
furrowed about one-third of blade thickness. Epi- 
dermis: 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 continu- 
ous band of sclerenchyma. Cuticle: thin on both 
surfaces. Stomata: almost flush with epidermis; few 
on adaxial surface, mostly confined to abaxial sur- 
face, 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 bun- 
dles: 57-59; commissural bundles present. Large 
veins each with one to two large vessels, occasion- 
ally three or more. Two phloem units lying lat- 
erally 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; pres- 
ent 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 scle- 
renchyma strands interspersed with parenchyma 
cells. Below is a distinct row of parenchyma cells. 
Cuticle: thick and slightly ridged on adaxial sur- 
face; thinner and slightly ridged on abaxial surface. 
Stomata: present on abaxial surface especially in 
furrows; substomatal chamber present; stomata 
Rush with epidermal surface. Mesophyll: three- 
layered palisade tissue changing abruptly to com- 
pactly 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 vas- 
cular bundle always accompanied by inverted 
adaxial cap and bundle sheath completely sur- 
rounding vascular bundles. Crystals: none ob- 
served. Tannins: few. 
The large abaxial crescentiform scle- 
renchyma 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. Sto- 
mata: 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 pres- 
ent. Adaxial surface slightly undulating; abaxial 
surface furrowed one-half thickness of blade. Epi- 
dermis; 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 inter- 
spersed with some sclerenchyma strands. Clusters 
of radially arranged parenchyma cells are inter- 
mittently 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 lay- 
ers translucent palisade cells radially arranged 
above vascular bundles and furrows. Three to four 
layers small palisade cells present between trans- 
lucent rows; small palisade grading into compact 
spongy tissue. Vascular bundle: 49; few commis- 
sural 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 surround- 
ing 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. Epider- 
mis: adaxial and abaxial cells mostly rectangular. 
Stomata: paracytic, 18 X 15 pm; occur on both 
surfaces but highly concentrated in abaxial fur- 
rows. 
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 rectangu- 
lar shaped on adaxial surface and almost rounded 
on abaxial surface. Abaxial epidermis partially re- 
placed with sclerenchyma fibers. Subjacent to 
adaxial epidermis occur one to two layers of paren- 
chyma cells interspersed with short strands of 
sclerenchyma. Below sclerenchyma strands and 
subjacent paranchyma is a layer of large translu- 
cent cells separating subjacent cells from meso- 
phyil. 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. Pali- 
sade and spongy cells compactly arranged and 
filled with chloroplasts. Mesophyll in median sec- 
tion 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 sur- 
rounded 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, uni- 
cellular 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 fur- 
rowed about two-thirds of blade thickness. Epi- 
dermis: cells square shaped, adaxial cells slightly 
larger than abaxial ones; cells somewhat trans- 
lucent. 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 epider- 
mis. hlesophyll: adaxial palisade four- to five- 
layered, 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. Vas- 
cular 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 rectangu- 
lar 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 un- 
dulating on adaxial surface; abaxial surface fur- 
rowed about one-fourth to one-half thickness of 
blade. Epidermis: adaxial cells dome shaped; few 
conical. Abaxial cells rounded. Walls thin on abax- 
ial and adaxial surface. Subjacent to adaxial epi- 
dermis 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 pali- 
sade 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 sur- 
rounding each vascular bundle. Crystals and tan- 
nins: none observed. 
Velloria lanata Pohl 
PLATES 3b, 17b, 45e 
SPECIMENS EXAMINED.-Pires et al. 9895; Vera- 
lucia 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. Adax- 
ial surface slightly undulating; abaxial surface 
furrowed about one-half thickness of blade. Epi- 
dermis: adaxial cells rounded to rectangular; abax- 
ial cells rounded to dome shaped. One to two layers 
parenchyma cells subjacent to epidermis and inter- 
spersed with sclerenchyma strands. Below paren- 
chyma 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 sub- 
stomata1 chamber present. Mesophyll: three-layered 
palisade tissue grading into compact spongy tissue. 
One to two layers of large translucent cells ar- 
ranged radially above vascular bundles and fur- 
rows. 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 vascu- 
lar 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, Epi- 
dermis; cells square to rectangular; thin walled. 
Stomata: paracytic, 21 X 15 pm; present on abax- 
ial 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 rec- 
tangular to slightly dome shaped. Walls thin on 
both surfaces. One to two layers of parenchyma 
cells subjacent to adaxial epidermis and inter- 
spersed with sclerenchyma strands. Below scleren- 
chyma 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 ar- 
ranged 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 bun- 
dle. Bundle sheath completely surrounding each 
vascular bundle. Crystals: none observed. Tannins: 
present. 
NOTE: This species exhibits rather wide abax- 
ial 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 paren- 
chyma cells. Cuticle: thick on both surfaces; 
slightly ridged on abaxial surface; smooth on 
adaxial surface. Stomata: present in abaxial fur- 
rows; 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 trans- 
lucent cells arranged radially above vascular bun- 
dles 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; sep- 
arated from spongy mesophyll by a distinct layer 
of parenchyma cells. Each vascular bundle is com- 
pletely surrounded by a bundle sheath. Crystals 
and tannins: none observed. 
NOTE: The two specimens are identical; how- 
ever, 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 sur- 
face distinctly undulating especially in median 
region; two small adaxial grooves present laterally 
to midvein. Abaxial surface with fairly wide fur- 
rows about one-third to one-half thickness of blade. 
Epidermis: adaxial cells dome shaped to rectangu- 
lar; abaxial cells rounded to dome shaped. Cells 
on both surfaces thin walled. Subjacent to adaxial 
epidermis occur layers of parenchyma cells inter- 
spersed 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 sur- 
faces 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 vas- 
cular 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 lat- 
erally in flanges of Y-shaped abaxial girder. Adax- 
ial girder present on all vascular bundles and 
almost extended to adaxial -epidermis. Some abax- 
ial 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 en- 
casing 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 rec- 
tangular on both surfaces; some rounded on abax- 
ial 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 abax- 
ial surface furrowed about one-third thickness of 
blade. Epidermis: adaxial cells rounded to rec- 
tangular; 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 fur- 
rows; 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 ar- 
ranged 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 scle- 
renchyma 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; Ta- 
mayo 3105; Tate 1196; Lasser 1777. 
SURFACE VIEw.-Hairs: present on abaxial sur- 
face 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 sur- 
face slightly undulating; abaxial surface furrowed 
about one-half thickness of blade; furrows may be 
wide or narrow. Epidermis: adaxial cells rectangu- 
lar; abaxial cells rounded to dome shaped. Walls 
thin on both surfaces. Subjacent to adaxial epi- 
dermis 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: pres- 
ent. Tannins: few observed. 
NOTE: The diagnostic features that character- 
ire 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 in- 
versely. 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 sur- 
rounded by one-layered parenchyma cells; usually 
occur above vascular bundles. Other smaller scle- 
rencliyina strands observed. Cuticle: thin on both 
surfaces and slightly ridged. Stomata: present on 
abaxial surface; small substomatal chamber pres- 
ent; 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. Vascu- 
lar bundles: 27; many commissural bundlei ob- 
served. 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 scleren- 
chyma 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, Adax- 
ial and abaxial surfaces slightly ridged, especially 
that of Smith 6445. Epidermis: cells on both sur- 
faces rounded to dome shaped; thin walled. Sub- 
jacent to adaxial epidermis is a layer of paren- 
chyma 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 bun- 
dle. 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: para- 
cytic, 27 X 9 pm; present on abaxial surface. 
dorsiventral; V-shaped with slight median adaxial 
groove; turned slightly inversely at margins. Adax- 
ial and abaxial surfaces slightly undulating. Epi- 
dermis: 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; sub- 
stomata1 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; pres- 
ent on abaxial surface. 
TRANSVERSE SECTION OF LAMINA.-ISOlateral; 
V-shaped with slight median adaxial groove. Mar- 
gins 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 substo- 
matal 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 com- 
missural 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; mar- 
gins of lamina turned inversely. Adaxial surface 
slightly undulating; abaxial surface with very small 
furrows. Epidermis: adaxial cells rounded to rec- 
tangular in shape; abaxial cells mainly dome 
shaped; cells on both surfaces thin walled. Subjacent 
to adaxial epidermis is a single layer of paren- 
chyma cells. Cuticle: very thick and slightly ridged 
on adaxial surface; thin and slightly ridged on 
abaxial surface. Stomata: present in abaxial fur- 
rows; no substomatal chamber observed. Meso- 
phyll: two to three layers palisade cells grading 
into loosely arranged spongy tissue. Few translu- 
cent 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 com- 
pletely 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: adax- 
ial 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; sub- 
stomata1 chamber present. Mesophyll: two to three 
layers of large palisade tissue changing into spongy 
tissue. Vascular bundles: 3 1-37; commissural bun- 
dles 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 rec- 
tangular; cells thin walled on both surfaces. Sto- 
mata: 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 sur- 
face 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 translu- 
cent cells radially arranged above the midvein; 
few translucent cells present in mesophyll. Vas- 
cular bundles: 37; commissural bundles not ob- 
served. 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 lat- 
erally 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: adax- 
ial cells rounded; abaxial cells square to rectangu- 
a 
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 un- 
dulating. 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 sub- 
stomatal 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. Epi- 
dermis: 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. Adax- 
ial and abaxial surfaces slightly undulating. Epi- 
dermis: cells on adaxial and abaxial surfaces 
rounded to dome shaped; walls on both surfaces 
thin. Layer of parenchyma cells subjacent to adax- 
ial epidermis. Cuticle: very thin and ridged on 
both surfaces. Stomata: present on abaxial and 
adaxial surfaces; substomatal chamber present; 
flush with epidermal surface. Mesophyll: undiffer- 
entiated. 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 vas- 
cular 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 un- 
dulating; abaxial surface slightly ridged. Epider- 
mis: 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: sub- 
stomatal chamber present; usually occur in depres- 
sion 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 sur- 
faces slightly undulating (except for Tryon 8c 
Tryon 6836 which has large ridges on both sur- 
faces). 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 pres- 
ent; 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 bun- 
dle. 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: para- 
cytic, 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. Meso- 
phyll: two to three layers of small cells grading 
into large spongy tissue. Large number of trans- 
lucent 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 adax- 
ial girder present on all vascular bundles. Bundle 
sheath completely surrounding each vascular bun- 
dle. 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 rec- 
tangular; cells on both surfaces thin walled. Sto- 
matal: 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 paren- 
chyma cells subjacent to adaxial and abaxial epi- 
dermis. Cuticle: thick and slightly ridged on 
adaxial and abaxial surfaces. Stomata: present on 
both adaxial and abaxial surfaces; large substo- 
matal 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 pres- 
ent in adaxial portion of midvein. Vascular bun- 
dles: 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: adax- 
ial 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 sub- 
paracytic, 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 meso- 
few 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 
C 
U 
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 sur- 
faces. Stomata: paracytic, 12 X 3 pm; present on 
abaxial and adaxial surface. 
TRANSVERSE SECTION OF LAMINA.-ISOlateral; 
V-shaped. Margins of lamina turned slightly in- 
versely. Adaxial and abaxial surfaces only gently 
undulating. Epidermis: cells on adaxial and abax- 
ial 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 pres- 
ent; flush with or slightly above epidermal sur- 
face. Mesophyll: undifferentiated. Spongy tissue 
composed of small rounded cells, some large and 
translucent. Two to three layers of large trans- 
lucent cells above midvein changing abruptly into 
compactly arranged spongy tissue. Vascular bun- 
dles: 29; few commissural bundles observed. One 
to three large vessels present in each vascular bun- 
dle. Two phloem units lying laterally in flanges 
of Y-shaped abaxial girder. Inverted Y-shaped adax- 
ial 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: para- 
cytic, 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; com- 
missural bundles observed. One to two large ves- 
sels 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 bun- 
dle. 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: adax- 
ial 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 com- 
pactly arranged. Two to three layers of large trans- 
lucent palisade cells arranged radially above mid- 
vein. Some translucent cells present in midportion 
of spongy tissue. Vascular bundles: 33-49; commis- 
sural bundles few. One large vessel present in each 
vascular bundle. Two phloem units arranged lat- 
erally in flanges of Y-shaped abaxial girder. In- 
verted 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. Sto- 
mata: 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 scle- 
renchyma strands observed. Each vascular bundle 
completely surrounded by a bundle sheath. Crys- 
tals: 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: para- 
cytic, 21 X 15 pm; present on adaxial and abax- 
ial 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, al- 
most 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 non- 
existent. 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 bun- 
dle. Crystals and tannins: none observed. 
Barbacenia exscapa Martius 
PLATE 46g 
SPECIMENS ExAMINED.-Anderson et al. 3583 1. 
SURFACE VIEw.-Hairs: present mostly on abax- 
ial 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 abax- 
ial 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 sur- 
face, 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 sur- 
face 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 con- 
taining 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: para- 
cytic, 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 sur- 
faces 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 con- 
tour 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. Meso- 
phyll: 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 Lo- 
redo 1066. 
SURFACE VIEw.-Hairs: multicellular; some pa- 
pillae 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. Epi- 
dermis: 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 ex- 
tending on either side of adaxial side of vascular 
bundle. Cuticle; very thin on adaxial and abaxial 
surfaces. Stomata: present on abaxial surface. Sub- 
stomata1 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 ob- 
served. 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: para- 
cytic, 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 vas- 
cular 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: para- 
cytic, 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 elon- 
gated; 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 sur- 
faces; 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 mid- 
region 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 bun- 
dle. 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 sur- 
faces. 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 undu- 
lating; 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 sur- 
face; 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 com- 
missural bundles observed. One to t~7o large ves- 
sels 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. Epi- 
dermis: 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 undu- 
lating; 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 sur- 
face. 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 abax- 
ial 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 mid- 
portion 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 mid- 
portion of mesophyll. 
Barbacenia riedeliana Goethart & Henrard 
PLATES 9b, 47g 
SPECIMENS ExAivlrNED.-Anderson et al. 35 123. 
SURFACE VIEw.-Hairs: short, multicellular pa- 
pillae 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 ob- 
served. One to two large vessels, mostly one, in 
each vascular bundle. Two phloem units lying 
laterally in flanges of Y-shaped abaxial girder. In- 
verted 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. Cuti- 
cle: thick and slightly ridged on adaxial and 
abaxial surfaces. Stomata: present on both sur- 
faces; 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 bun- 
dle. 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 multicellu- 
lar 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 thin- 
walled parenchyma cells. Cuticle: very thin on 
adaxial and abaxial surfaces. Stomata: present in 
abaxial furrows and on adaxial surface; sub- 
stomata1 chamber present. Mesophyll: small, rec- 
tangular, 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 mar- 
gins. 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 cer- 
tain 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 identi- 
cal, 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 vas- 
cular bundles next to the midvein. The mesophyll 
differ, however. In V. tragacantha there is a dis- 
tinct 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 sub- 
jacent to the adaxial epidermis and to a minor 
degree they are developed toward the abaxial sur- 
face. In addition to the principal veins there are 
many smaller secondary, or tertiary, veins in the 
slender regions between the large vascular bun- 
dles. Another species that exhibits the character- 
istics 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 re- 
gions 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 sig- 
nificantly expanded and pointed adaxially. Fur- 
thermore 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. How- 
ever, 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 character- 
istics, 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 pat- 
tern. 
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 de- 
pendent solely on environmental pressures is not 
quite clear. 
Of the array of species studied, Vellozia burle- 
marxii 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 ana- 
tomical 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 fur- 
ther 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 spe- 
cies, 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 vascu- 
lar 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 re- 
gion, 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 mor- 
phological feature. 
Three species that appear to be closely related 
from the standpoint of their leaf anatomy, espe- 
cially 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 sepa- 
rated, 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 paren- 
chyma 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 riede- 
liana 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 varia- 
tions among the specimens may be the result of 
environmental influences.- Maguire (1969: 35 ) was 
indeed correct when he noted that ?Vellozia tubi- 
flora 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 cor- 
rugation 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. An- 
other 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 there- 
fore no conclusive statement can be made to char- 
acterize 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 sub- 
key 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 character- 
istic features of this group is the presences of hairs 
on both surfaces of many species. Barbacenia ma- 
crantha, 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. gentiano- 
ides, 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 three- 
dimensional characteristics that emphasize the di- 
versity 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 con- 
siderably 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 OBSERVA- 
Trichomes: 
72 S.\fITHSONIAN CONTRIBUTIONS TO BOTANY 
of Vellozia ciliata (Plates 5c; 49a-c) and V. metz- 
gerae (Plate 5b). A vertical section through a fur- 
row of a leaf of V. ciliata (Plate 49c) shows very 
clearly the sporadic arrangement of the protuber- 
ances. Among these fingerlike projections are the 
stomata. 
Generally the epidermal cells 
are arranged in longitudinal files which run par- 
allel 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 numer- 
ous. 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 dis- 
tinctly 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 struc- 
ture 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 assess- 
ment 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 Re- 
cently Described in the Velloziaceae. American 
Journal of Botany, 553399-403. 
Leaf-anatomy and Systematics of Old 12'orld Vello- 
ziaceae. 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 Dis- 
cussion 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. Contribu- 
tions 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 Videnska- 
bernes 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 arrange- 
ment 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 scleren- 
chyma 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 enlarge- 
ment of the furrow showing the arrangement of the fingerlike projections (X 630) ; c, longi- 
tudinal 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