Manihot Manihotoides (Euphorbiaceae) - CNCFlora

Organization for Flora Neotropica 

Manihot Manihotoides (Euphorbiaceae) 

Author(s): David J. Rogers and S. G. Appan 

Reviewed work(s): 

Source: Flora Neotropica, Vol. 13, Manihot Manihotoides (Euphorbiaceae) (Jun. 22, 1973), pp. 1- 

272 

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FLORA NEOTROPIC 

fC^ 

Monograph No. 13 

MANIHOT 

MANIHOTOIDES 

(Euphorbiaceae) 

by 

David J. Rogers 

and 

- 

_ ^ 

TROPIC 

S. G. Appan 

TROPIC OF 

CANCER 

FLORA 

NEOTROPICCA 

OF 

CAPRICORN 

Published For 

Organization for Flora Neotropica 

by 

Hafner Press 

New York 

June 22, 1973

Copyright ? 1973 

HAFNER PRESS 

Published by 

Hafner Press 

A Division of Macmillan Publishing Co., Inc. 

866 Third Avenue 

New York, New York 10022 

Library of Congress Catalog Card Number 72-88251 

Printed in U.S.A. by 

NOBLE OFFSET PRINTERS, INC. 

New York, N.Y. 10003 

40-

MANIHOT AND MANIHOTOIDES (EUPHORBIACEAE)1 

A COMPUTER-ASSISTED STUDY 

DAVID J. ROGERS2 AND S. G. APPAN3 

INTRODUCTION 

This monograph is an outgrowth of our primary objective, the systematics and 

evolution of cultivated plants. The senior author chose Manihot esculenta Crantz as an 

object of study because this cultivated species, which is distributed in most tropical areas, 

had received little attention from any scientific aspect and because this crop, grown 

primarily for its roots, differed in many cultural and evolutionary aspects from grain 

crops. In the course of the study of the variation within the cultigen, it became evident 

that much of the intraspecific variation must be attributed to hybridization with other 

species in the genus Manihot, and that no progress with evolutionary studies could be 

made until all the species of the genus were embedded in a modern classificatory system. 

Once the species are properly classified, one has a better opportunity to proceed with 

more precise studies of the evolution of the cultigen. Our own concepts of the biological 

problems are given in a separate section, the biology of speciation. 

A modern classification is one in which a conscious attempt is made to employ a 

consistent biological species concept (the species is a closed gene pool). There are two 

stages in deriving the biological species concept for a group of organisms, the first of 

which is embodied in the following pages, and the second requires that hypotheses of the 

closed gene pool be tested by biosystematic means (Rogers and Appan, 1969). Clearly the 

second stage is beyond our capacities at the moment, for many pragmatic reasons, the 

greatest being the fact that this genus is largely tropical, in areas beyond our ability to 

raise and test the plants for their genetic compatability or lack thereof. We believe that 

this monograph can serve as a guide for biosystematic studies, and as a source of 

information for agricultural work that is vitally needed for improvement of the cultivars 

in Manibot esculenta. 

The many decisions necessary to establish this classification were supported by three 

large computer programs developed in the Taximetrics Laboratory under the direction of 

the senior author. Descriptions of the programs and their applicability to this type of 

taxonomic work are to be found in two companion publications, Rogers and Fleming, 

1973, Rogers and Appan, 1969, and Rogers and Appan, in press. Further details of the 

programs may be found in a series of publications from the Taximetrics Laboratory, as 

follows: Bisby, 1970; Estabrook, 1966, 1967; Estabrook and Rogers, 1966; Estabrook 

and Brill, 1969; Hawkesworth et al, 1968; Irwin and Rogers, 1967; Prance et al, 1969; 

Rogers, 1963, 1970; Rogers et al, 1967; Rogers and Appan, 1969, in press 1972; Rogers 

and Fleming, in press, 1973; Wirth et al, 1966. The three programs are TAXIR, an 

information retrieval system, CHARANAL, for study of the significance and value of 

characters, and GRAPH, a clustering method. Each of these programs is designed to 

accomplish much of the necessary data manipulation for monographic work, and each 

provides the monographic worker with long sought tools to aid in more objective 

decision-making processes. We are indebted to our colleague, Henry S. Fleming, who 

Submitted to Flora Neotropica 26 June 1972. 

2Professor, Department of Biology, University of Colorado, Boulder, Colorado 80302. 

3 Research Associate, Gulf Universities Research Consortium, Mississippi Test Facility, Bay St. 

Louis, Mississippi 39520. 

1

2 Flora Neotropica 

played a fundamental role in the design and development of the computer programs, and 

has been a never-ending source of botanical and general biological knowledge. 

We wish to extend our thanks to the keepers and curators of the following herbaria 

for the loan of their valuable collections: Arnold Arboretum of Harvard University, 

Cambridge, Massachusetts, (A)4; University of Arizona, Tucson, Arizona, (ARIZ); 

Instituto de Botanica Agricola del I.N.T.A., Buenos Aires (BAB); Suriname Forest 

Service, Paramaribo, Suriname (BBS); British Museum (Natural History), London, (BM); 

Museo Botanico, Facultad de Ciencias E.F. y Naturales, Cordoba, Argentina (CORD); 

Dept. of Botany, University of California, Davis, (DAV); Dudley Herbarium, Stanford 

University, Stanford, California (DS); Field Museum of Natural History, Chicago, Illinois, 

(F); Florida State University, Tallahassee, Florida, (FSU); Conservatoire et Jardin 

botaniques, Geneve, Switzerland, (G); Gray Herbarium of Harvard University, Cambridge, 

Massachusetts, (GH); the Herbarium, Kew (K); Rijksherbarium, Leiden, Netherlands (L); 

Instituto Miguel Lillo, Tucuman, Argentina, (LIL); Botanische Staatssammlung, Munich, 

Germany, (M); Museu Paraense Emilia Goeldi, Belem, Brasil, (MG); University Herbarium, 

University of Michigan, Ann Arbor, Michigan, (MICH); Missouri Botanical Garden, St. 

Louis, Missouri, (MO); United States National Arboretum, Washington, D.C., (NA); New 

York Botanical Garden, Bronx, New York, (NY); Fielding Herbarium, Dept. of Botany, 

Oxford University, (OXF); Laboratoire de Phanerogamie, Museum National d'Histoire 

Naturelle, Paris, France (P); Herbarium of Pomona College, Claremont, California, (POM); 

Kriebel Herbarium, Dept. of Biol. Sci., Purdue University, Lafayette, Indiana, (PUL); 

Jardim Botanico, Rio de Janeiro, (RB); Naturhistoriska Riksmuseum, Botanical Depart- 

ment, Stockholm, Sweden, (S); Instituto de Botanica, Sao Paulo, Brasil, (SP); University 

of Texas Herbarium, Austin, Texas, (TEX); Botanical Museum and Herbarium, Utrecht, 

Netherlands, (U); Herbarium of the University of California, Berkeley, California, (UC); 

U.S. National Herbarium, Smithsonian, Washington, D.C., (US); Instituto Botanico, 

Caracas, Venezuela, (VEN); Naturhistorisches Museum, Wien, Austria, (W). 

HISTORY OF CLASSIFICATION 

Pre-Linnaean. 

Bauhin (1651) was the first European botanist to recognize and describe the generic 

term Manihot. He studied the plants collected by Andre Thevet in Brasil, and honored 

Thevet by designating plants, now recognized as Manihot esculenta Crantz, M. theveti. 

This epithet was recognized and used by a number of pre-Linnaean workers, such as 

Tournefort (1700), and was continued in at least one post-Linnaean work, that of Miller 

(1754). 

Post-Linnaean. 

1753 Linnaeus placed the only species of Manihot known at the time in the genus 

Jatropha, and designated the species Jatropha Manihot. He cited earlier workers, 

including Bauhin (1658) Sloane (1696), and Plukenet (1700), but did not cite a 

specimen which could be designated the type. 

1754 Miller, in the 4th Edition of the Gardener's Dictionary, provided the first valid 

post-Linnaean description of the genus Manihot. Miller did not employ consistent 

binomial nomenclature, but followed earlier workers in designation of M. theveti. 

1758 Loefling, in Iter Hispanicum, described a plant Janipha fructescens, and this 

epithet was later attributed to him. (For example, 1771, Linnaeus, Mantissa, p 

4Acronyms of herbaria follow Lanjouw and Stafleu, 1964. Index Herbariorum, Part I, 5th ed. 

Regnum Vegetabile 31.

Introduction 3 

126 as a synonym of Jatropha Janipha). However, on examination it is clear that 

"frutescens" was part of the description, and not intended to be a specific 

epithet. The same mistake was made in interpretation of the German edition of 

his work, published in 1766. 

1763 

1763 

Adanson was long considered the author of the genus, Manihot, before the work 

of Miller was recognized. 

Jacquin was the first to recognize and validly describe a species other than M. 

esculenta. He described Jatropha carthaginensis, from Cartago, in present-day 

Colombia. Jacquin also listed Janipha frutescens of Loefling under his Jatropha 

carthagenesis. 

1766 Crantz, in Institutiones Rei Herbariae, provided the first post-Linnaean validly 

published species epithet for the cultigen, M. esculenta. He cites no specimens, 

but refers to a plate in Merian, Dissertatio de generatione et metamorphosibus 

insectorum Surinamensium (1726), which we have designated as lectotype of the 

species, and of the genus Manihot. 

1771 Linnaeus redesignated Jacquin's Jatropha carthaginensis as Jatropha janipha, 

apparently using Loefling's epithet "janipha." Linnaeus may have recognized that 

Loefling had not intended to employ a binomial, although he cites Janipha 

frutescens of Loefling in synonymy under Jatropha janipha, p 126, Mantissa, vol 

1. 

1773 J. F. Gmelin, in Onamatologia Botanica Complete, vol 5, p 7, provided the first 

designation of Jatropha dulcis, apparently citing a pre-Linnaean author, J. Gesner, 

but without a botanical description, Pohl (1827) cited J. dulcis as a synonym of 

his species M. aipi. Pax 1794 

(1910) transferred Gmelin's name to Manihot and 

considered M. aipi Pohl as a synonym. 

Sesse, M. and Mociio, J. M., Spanish botanists who collected in Mexico, intended 

to write a Flora Mexicana, but apparently their manuscript was destroyed. Sesse 

employed (or worked with) Vincente de Cervantes. Cervantes published 8 new 

taxa, apparently using Sesse's descriptions, in a supplement to La Gazeta de 

Literatura vol 3, July 2, 1794, two of which are supposedly Manihot, Jatropha 

triloba and Jatropha palmata. Since the descriptions of these two are very general, 

it is impossible for us to assign these names to any particular taxon. See McVaugh 

(1945) for an excellent survey of this historical 1817 

problem. 

Humboldt, Bonpland and Kunth, in Nov. Gen. Spec. II, p 107, listed five species 

from Mexico and northern South America as members of the genus Janipha, 

including J. foetida from Mexico, J. loeflingii, northern South America, (these 

authors also list Jatropha frutescens, attributed to Loefling, as a synonym of J. 

loeflingii), Janipha yuquilla from northern South America (Novae Andalusia), J. 

aesculifolia from Campeche, Mexico, and J. manihot, transferring Linnaeus' 

Jatropha manihot to Janipha. 

1825 Vellozo, Fr. J. Marianus A. Conceptione, Flora Fluminensis. See Stafleu (1967) 

for history of publication of text and tables. The Atlas (correct date of 

publication 1835) contained four illustrations of Jatropha species, J. manihot, J. 

palmata, J. stipulata, and J. silvestris. Although the illustrations are generalized, 

without adequate detail, and certainly with some artist's license, there is little 

doubt that the first three are species of Manihot, J. manihot = M. esculenta, J. 

palmata = M. leptopoda (Muell.-Arg.) Rogers & Appan; Jatropha stipulata 

probably is M. 1826 

esculenta, but the fourth Vellozo plate, Jatropha silvestris, is either 

a true Jatropha species, or something else which we cannot determine. 

Sprengel, in Syst. Veg. 3: 76. Named one species attributable to Manihot, namely 

Jatropha tripartita. Pohl transferred Sprengel's species to Andenoropium, and 

Mueller von Argau transferred the species to Manihot in 1886.


1827 Pohl, J. E., in Plantarum Brasiliae Icones et Descriptions, pages 17 to 56, provided 

the first monographic study of the genus Manihot. Pohl spent some time in Brasil, 

collecting and recording notes about the species. His species concepts are usually 

robust, and most of his species are recognized today. Pohl divided the cultigen M. 

esculenta into two separate species, M. utilissima and M. aipi, with the intent of 

differentiating between those cultivars with high concentrations of HCN (M. 

utilissima) and those with low concentrations of the cyanogenic glucoside (M. 

aipi). All together, he listed 48 species, most of them described for the first time. 

Pohl's original set of collections are found in the Vienna herbarium along with 

many of his 1840original 

plates. 

Steudel, E. T., in Nomenclator Botanicus, Part I (1840) and Part II (1841) placed 

1841 all previous names of Manihot, including those of Pohl, into Linnaean 

nomenclature, transferring all Manihot species to the 1840 

genus Jatropha of Linnaeus. 

He did not describe any new species, nor did he advance the knowledge of the 

classification of Manihot by his cataloging. 

Graham, in Edin. Philos. Jour., erected the variety multifida assigned to Janipha 

loeflingii. 

1843 Hooker, in Icon. PI., raised Graham's var. multifida to species rank and gave the 

species name M. grahami. 

1859 Torrey, in Emory's Rept. U. S. Mex. Bound. Surv. established the variety 

angustiloba, assigned to Janipha manihot. 

1863 Baillon in Adansonia, established four new species in Manihot, M. weddeliana, M. 

amaroleitensis, M. hilariana, and M. stricta. 

1864 Wawra, in Flora XLVII, established M. pohlii. 

1865 Mueller von Argau, in Linnaea, established M. rhomboidea. 

1866 Mueller von Argau, in de Candolle Prodromus 15(2): 1056-1075, produced the 

second monographic treatment of the genus Manihot establishing 43 species. He 

first used the concept of sectional divisions within the genus, and employed a 

much broader species definition than Pohl. Mueller von Argau maintained Pohl's 

M. ultilissima for the bitter varieties of M. esculenta, but placed Pohl's M. aipi as a 

variety in M. palmata, along with several other varieties, which gave more of an 

idea of the relationship among the cultivated varieties of M. esculenta. 

1872 Mueller von Argau established the species M. microcarpa. 

1874 Mueller von Argau, in Martius' Flora Brasiliensis XI, Part II, pages 438-486, 

accepted 72 species for the genus, nearly all confined to Brasil. For the first time 

a key (conspectus) was provided, but in this work, Mueller did not divide the 

genus into sections. It is evident that Mueller (as we) employed mostly vegetative 

characters to distinguish the species. 

1874 Grisebach erected the species Janipha anisophylla. 

1894 Watson, Proc. Am. Acad. 25, established M. pringlei. 

1896 Taubert, Bot. Jahrb. Syst. 21, established M. mossamedensis. 

1903 Greenman, Proc. Amer. Acad. 39, established M. caudata. 

1905 Chodat and Hassler, Bull. Herb. Boissier 2, named M. guaranitica and M. 

tweediana f. nana. Their work was based on collections made by Hassler in 

Paraguay. These collections are some of the finest specimens collected of the 

genus Manihot. 

1907- Ule, in a series of publications, described many new species. 

1908- It is obvious that his concept of a species was particularly narrow, and very few of 

1914 the species he erected have been maintained. 

1908 Huber, Bol. Mus. Paraense Hist. Nat. 5, established M. marajoara. 

1910 Brandegee, Univ. Calif. Publ. Bot 4, described Manihot pauciflora.


1910 F. Pax in Engler, Pflanzenreich IV. 147. II, Heft 44, 21-111, presented the last 

full monographic treatment of Manihot. Pax brought together the known 

taxonomic literature, examined and cited most of the classical collections, and 

provided a truly valuable reference to the genus. He recognized a total of 128 

species, in 11 sections, and provided diagnostic keys to all taxa. Pax's treatment 

seems to have had the objective of pragmatism rather than a true reflection of the 

biological relationships amongst the taxa. Pax did no field work, and his 

knowledge of biogeography and ecology was rather confused. As a result, the taxa 

bear little relationship to the geographic distribution nor to ecological relationships. 

Pax maintained the concept of two species in the one cultigen, M. 

esculenta, following the concept initially established by Pohl. He maintained 

Pohl's name (M. utilissima) for those cultivars with high concentrations of HCN, 

but, like Mueller von Argau before him, changed the epithet for those cultivars 

with low concentrations of HCN. Pax did not, however, accept Muell.-Arg.'s 

epithet, M. palmata, but designated the "sweet" cultivars M. dulcis. 

It is a tragedy that most of the specimens upon which he based his work were 

destroyed during World War II. Perhaps his decisions would have been more 

meaningful had all these valuable collections been available for study. 

1911 Standley and Goldman, Contr. U.S. Nat. Herb. 13, established M. chlorosticta. 

1914 Pax and K. Hoffmann, Pflanzenreich IV. 147 (Heft 63) establishedM. pittierii and 

M. boliviana. 

1923 Johnston, Contr. Gray Herb. 68, established M. rubricaulis. 

1924 Pax and K. Hoffmann, Pflanzenreich IV. 147 (Heft 85) Additamentum VII, 

described eleven new species, and two new varieties. 

1930 

1937 

1938 

Pittier, Jour. Wash. Acad. Sci. 20, described M. filamentosa. 

Standley, Publ. Field Mus. Bot. 17, describedM. isoloba. 

Ciferri, Relaz. e Mon. Agrario-Colonial 44 recognized the priority of Crantz's 

epithet, M. esculenta, and established the modern concept which places all 

cultivars together under one broad designation, whether the cultivars have high or 

low concentrations of HCN. 

1939 Lanjouw, Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 1942 

69, described M. saxicola. 

Croizat, Jour. Arnold Arb. 23, described four new species from North America. 

1946 

1961 

Standley, Am. Midi. Nat. 36, described M. tomatophylla. 

McVaugh, Brittonia 1964 

13, described M. auriculata and M. michaelis. 

Martinez-Crovetto, Bonplandia 1, described M. bunzikeriana. 

1965 

1967 

N. D. Cruz, Bragantia 24, established M. jolyana. 

N. D. Cruz, Bragantia 26, established M. handroana. 

The senior author began studies in the classification of Manihot in 1952. The original 

objective was to study the evolution of one cultivated species, but it became evident that 

the evolutionary studies must be preceded by a modern classification of the variants 

within the cultivated complex. Herbarium samples of the cultigen, M. esculenta, were 

clearly inadequate to describe the variation and make a classification. Field studies were 

initiated in 1953, in the West Indies and in Costa Rica, during which time, methods of 

sampling, recording, and preparation of documenting herbarium samples were evolved. In 

subsequent years, additional field studies (expanded to include the wild species of the 

genus) in most of the regions where the cultigen and other species of the genus are native, 

were carried out. A total of 533 population samples, each documented with photographs, 

ethnological and ecological data, have been collected. A monograph of M. esculenta 

Crantz (Rogers and Fleming, 1973) which gives the intraspecific classification of the 

cultigen, is published separately. Some of the philosophy adopted for categorization of


cultivated plants, as well as a complete discussion of the computer methods employed in 

that study, are included with the classification. The cultivars are classified into 19 

"groups" which do not have formal taxonomic status. All the specimens which document 

the classification of the cultivars were collected by the senior author. The first set of the 

specimens representing the wild species has been deposited at The New York Botanical 

Garden, whereas the first set of the documenting specimens for the cultigen, M. esculenta 

has been deposited at the National Arboretum, Washington, D.C. 

The junior author, a native of Kerala, India, an important region for cultivation of M. 

esculenta, came to the University of Colorado in 1967 as a graduate student. His doctoral 

dissertation, completed in December, 1969, was entitled "The North American Species of 

Manihot Delimited by Computer Aided Taximetric Methods" and is housed in the 

Library of the University of Colorado, Boulder. This dissertation, which gives details of 

the computer methods employed in the study, is being published separately by the 

Colorado Associated University Press. The junior author has continued work on the 

classification of the genus as post-doctoral fellow and research associate. 

As a footnote to the history of this study, and as an attempted explanation to the 

curators and keepers of the valuable herbarium materials which have been away from 

their respective homes for much too long, we append the following notes. 

The past two decades have seen a very fundamental investigation into the methods 

and thought processes which underlie the most important biological endeavor, taxonomy. 

All human endeavor is, eventually, the making of classifications, and biological 

taxonomists have the longest history of investigation into the formal processes of 

classification of any group of scientists. The reason that the past two decades have been 

important is that a new tool, the electronic computing machine, became generally 

available for purposes other than purely military. Along with the development of the 

computing machines, a new type of mathematics dealing with finite sets and qualities of 

things became available. The merging of the concepts of taxonomy, the appropriate 

mathematics, and efficient use of the computing machine, has been one of the major 

objectives of the Taximetrics Laboratory so that the classification of Manihot could be 

accomplished with the most objective procedures available. Before we considered our 

computer methods complete and satisfactory for taxonomic work, each method was 

given a thorough testing against some real taxonomic problem found in Manihot. The 

herbarium specimen, which has been derided by so many unthinking "scientists," is the 

document which provides the acid test for any theoretical development, and we have used 

the collections from thirty-two herbaria in these tests. We thank all the curators for their 

patience during the time we were developing and proving our techniques. 

GEOGRAPHY AND ECOLOGY 

All species of the genus Manibot are native in the New World tropics. The only 

species found in other tropical regions of the world are those which have been introduced 

since the discovery by Columbus. Manihot esculenta was first carried by Portuguese from 

the east coast of Brasil to West Africa (Jones, 1959). Following the initial introduction 

into Africa in the 16th Century, this species was spread slowly across the sub-Saharan 

areas of Africa, and carried by natives to the island of Madagascar. Spanish explorers and 

tradesmen carried the species from the west coast of MesoAmerica to the Philippines, and 

thence to the mainland of southeast Asia. We believe that introduction to south India was 

from the east coast of Africa much later. The Dutch brought Manihot esculenta to its 

highest degree of culture in Indonesia. Today, nearly every island of the Pacific large 

enough to support any type of agriculture contains some cultivars of this species. This 

cultigen is also grown in south Florida, apparently first introduced by native Indians long 

before the advent of white man. It has also been grown sporadically in the state of


Mississippi. Manihot grahami, native in southern Brasil and northern Argentina, is now 

found sporadically as a small shade tree and as volunteer wildlings along the Gulf Coast of 

the United States, from Florida west to Louisiana. 

Manibot glaziovii was transported to many Old World tropical areas as a potential 

rubber crop. Today, this species has become naturalized, or at least weedy, in some areas 

of Malaya, India, and Africa. Several other species, such as M. tristis subsp saxicola, have 

been intermittently introduced into Africa, Indonesia, and to India, as potential value in 

plant breeding with M. esculenta. 

In the New World, the native range of the species is from southern Arizona in the 

United States (M. davisiae and M. angustiloba) to Argentina (M. grabami and M. 

anisophylla). Only one wild species is native to the West Indies, M. brachyloba, and this 

species, which has a wide distribution in northern South America is found only on the 

island of Hispaniola. We have no explanation for this particular distribution, and attribute 

its presence on Hispaniola to a chance introduction. 

The species of Manihot are all rather sporadic in their distribution, and never become 

dominant members of the local vegetation. As discussed in Rogers (1963), there are two 

major concentrations of species, one in Mexico, the other in Brasil. The species found in 

the two areas are remarkably disjunct, and with the exception of the cultivated species, 

none of the North American species are found in South America, and only one collection 

of one species (M. brachyloba) of South American affinity has been recorded in Central 

America (Costa Rica). The preponderant number of species (80 of the 98 species 

recognized in this monograph) are South American, and in South America, by far the 

greatest number are found on the ancient area in eastern central Brasil principally in the 

states of Goias, Minas Gerais and interior Bahia. 

Most Manihot species are found in relatively dry regions, and only a few are typically 

found in rain forest regions. Those species whose distribution coincide with rain forest are 

typically found in openings in the forest, either man-made or naturally caused, such as 

flood banks, blowdowns of senescent forests, and the like. These considerations lead us to 

the hypothesis that most species are heliophiles, capable of growth only where there is no 

shading, and that many of them are "weedy" types, capable of invasion into open areas. 

The North American species particularly, are found on limestone-derived soils, of 

relatively recent origin. As indicated by the distribution maps, most of these 

representatives are found along the edges of the Sierras Madre, Oriental and Occidental, 

or are found in regions of Karst topography, as represented by the Yucatan Peninsula and 

the Isthmus of Tehuantepec. 

All of the species of the genus are frost-sensitive. For this reason, their altitudinal 

limit in the tropics is approximately 2000 meters. Only two species are known (M. 

grahami and M. anisophylla) whose native distributions are in regions with occasional but 

predictable frost. Manihot grahami, a small tree species, has been introduced to the near 

coastal regions along the Gulf of Mexico in the southern United States. In this area the 

species is successful, (although sporadic) but may be killed by particularly severe and 

extended periods of below freezing temperatures. 

BIOLOGY OF SPECIATION 

Our species interpretation is based upon the following observations, made during the 

extensive field work of the senior author in most of the important areas of distribution, 

with the additional valuable information collected by Dr. Howard Irwin of The New York 

Botanical Garden. The basic concept developed during this study is that the genus appears 

to be quite recent in origin, and that the species (with a few exceptions) are still rapidly 

evolving. The species are, therefore, not sharply delimited. Most of the species are quite 

variable with respect to vegetative structures, but are relatively uniform in their floral


organs. This is best explained by the monoecious or dioecious inflorescences, which 

makes cross-pollination much more likely. In many species, the pistillate flowers are open 

and ready for pollination before the staminate flowers of the same inflorescence have 

opened. The usual pollination mechanism is provided by insects, and the sticky pollen 

adheres to their bodies. A wide variety of hymenopterous insects, and numerous 

foraminiferans visit the flowers. These phenomena provide an opportunity to produce a 

heterozygous gene pool, allowing great phenotypic plasticity. However, we also believe 

that, although the genetic material is heterozygous for the vegetative portions of the 

plant, some precise control is maintained over the floral mechanisms thus maintaining the 

stability of the reproductive processes, mentioned above. A number of experimental 

crosses, (Abraham, 1957; Bolhuis, 1953; Cruz, 1968; Jennings, 1957; Lanjouw, 1939; 

Magoon et al, 1966; Nichols, 1947 a, b) and observations of frequent hybridity of 

cultivars of Manihot esculenta and local wild species, seem to bear out these hypotheses. 

Hybridization has, therefore, played a large role in the development of the variations 

found. We support the hypothesis of Harlan (1961) that a number of wild species may 

have been developed as a result of chance hybridization between cultivars and local wild 

species, again with an extremely heterozygous gene pool. The stability of these wild 

species (which might be termed "feral" species) is minimal, and over a period of years, 

many phenotypic variations are found which could easily expand the species boundaries. 

We have reflected this hypothesis in the taxonomic treatment by frequent emendations of 

species established 100 years or more ago. Also, the subspecies is the only taxonomic 

category below the species level which we employ. Since the variability within 

populations is great, only geographic separation can be employed at the infraspecific level 

with any degree of certainty. 

As indicated in the section of Geography and Ecology, by far the largest numbers of 

species are found on the ancient plateau of east-central Brasil. Although this is an ancient 

area long available for growth of the angiosperms, we believe that most of the species of 

Manihot in this region are of relatively recent origin, having developed rather explosively 

after many fires started by man had burned the vegetation at frequent intervals. Most of 

the species of this Brasilian area are perennial subshrubs, with large woody roots whose 

stems die back to the root crown at frequent intervals, responding either to dry periods or 

to fires (which are usually coincident with the dry seasons). The large shrub and small 

tree species of South America occur in regions to the east and south of the central 

Brasilian plateau. 

We recognize that many of our decisions will have to be changed as more material is 

collected and more study is devoted to this significant and economically important genus. 

SYSTEMATIC CRITERIA 

General 

The criteria for classification of the Euphorbiaceae are generally in two classes, those 

that distinguish the family and the major subdivisions of the family, and those that are 

primarily for differentiation of species within the genera. In the first, flower and fruit 

morphology are of primary importance, and in the second, vegetative morphology is the 

most significant. These indicate that the classifications of the family and genera to date 

are categorized as alpha taxonomy, and this monograph is, of necessity, an alpha 

classification modified by the concepts of biosystematics. More sophisticated data from 

biosystematic and chemotaxonomic studies remain to be analyzed, but even when these 

have been analyzed, they still need to be correlated with the external morphology to 

provide a decision on the taxonomic position of the taxon. Therefore, alpha taxonomy 

provides the basis on which to design biosystematic studies.


We believe, however, that by employing well-designed computer programs, analyzing 

many characters for each taxon, and keeping the concepts of biosystematics in mind, we 

have given a more than adequate reflection of the intraspecific and interspecific relations 

in Manihot. By examination of the Table of Characters and Attributes Employed (Table 

I), one may determine precisely the basis upon which our classification rests, and the 

processes by which decisions were made. Although some of the species are represented by 

single, inadequate specimens, by comparisons of these with those species which do have 

adequate representation, and using the same objective measure for delimitation of the 

taxa in all cases, a unified classification has been produced. 

We have no doubt that when more information for some of the species becomes 

available, there will have to be new combinations, changes in taxon level, and even 

reduction of some of our taxa to synonymy. Even with the most sophisticated methods 

of analysis of the data, final judgment of the taxa to be recognized must be based on the 

knowledge and skill of the taxonomist, and we are certainly open to mistakes in these 

areas. 

Growth habit 

The species of Manihot are perennial, and vary in growth pattern from nearly 

acaulescent subshrubs to small trees with trunk diameter (DBH) of 25.0 cm and heights 

of 10 to 12 m. Decumbent subshrubs, semiherbaceous subshrubs, shrubs, and small trees 

are found in the genus. The branching pattern is typically dichotomous or trichotomous, 

the point of branching determined by the appearance at the branching point of a terminal 

inflorescence. Bark of the woody species is generally smooth, but in some cases (as in M. 

caudata) may become slightly scaly with age. Many of the species are lacticiferous, and 

two rubber-producing species, (M. glaziovii and M. caerulescens subsp caerulescens) are 

still cultivated to some extent. 

Many of the stems of subshrub and shrubby species, adapted to regions of marked 

dry periods, die back to a root crown regularly, and other larger shrubs regularly lose 

their leaves during the dry season. Adaptation to arid conditions is further indicated by 

the presence of enlarged storage roots. 

Pigmentation 

Colors vary in Manihot species from dark brown to light tan in the roots, from 

brown to gray to yellow or red in the stems, from red to yellow to green on the petioles, 

and from red to green in the flowers. Since perception of color varies with the eye of the 

observer, and with age of the plant part observed, and since the observations are not 

consistent over a sufficiently large number of specimens, we have not been able to 

employ pigmentation as systematic criteria. The senior author has employed the 

Nickerson Color Fan of the American Horticultural Council in all collections made by 

him, and these are consistently reported in the species descriptions. Both the official 

color designation, and the associated numeric notation, which follows the Munsell color 

classification, are provided (where available) in the species descriptions. For example, for 

the petiole, the following notation may be found: "brilliant yellow green (5GY 8/8)." 

Although this is a very precise notation, it is recognized that it is still subject to the same 

limitation noted above. 

Roots 

Many of the species of Manihot have tuberous roots. Unfortunately, there are 

insufficient samples of this organ associated with herbarium specimens (particularly in the 

South American material) to employ characters from them as systematic criteria. Of 

those species we have examined, there are indications that many valuable characters are


TABLE I 

Characters and Attributes Employed in this Study 

1. Habit of growth 

1. compact subshrub, less than 25 cm tall Fig 98C 

2. fleshy, partially woody subshrub Fig 102B 

3. ,erect, woody subshrub, less than 1 m tall Fig 69D 

4. clambering (straggling) woody subshrub Fig 19D 

5. decumbent, herbaceous subshrub, less than 0.5 m tall Fig 21A 

6. woody vine, greater than 1.0 m long Fig 90A 

7. clambering (straggling) shrub, 1.0-3.0 m tall Fig 21A 

8. erect shrub, 1.0-3.0 m tall Fig 76B 

9. tall shrub, more than 3.0 m tall Fig 37A 

10. tree Fig 25A 

2. Pubescence of young stem 

1. lanate 

2. tomentose 

3. pubescent 

4. sparsely pubescent 

5. glabrous 

3. Pubescence of stipules 

ibid states in 2 

4. Pubescence of peduncle/pedicel 


5. Pubescence of petiole 


6. Pubescence of midrib/veins 


7. Pubescence of bracts/bractlets 


8. Pubescence of tepals 


9. Pubescence of ovary 


10. Nature of the vegetative shoot 

1. nearly acaulescent, internodes telescoped 

2. caulescent, internodes not telescoped 

11. Persistence of stipules 

1. persistent 

2. caducous 

12. Length of stipules 

1. long, foliaceous, more than 3 cm Fig 98B 

2. medium, 1-3 cm Fig 99B 

3. short, less than 1 cm Fig 94D 

13. Margin of stipules 

1. laciniate 

2. serrate 

3. entire 

14. Length of petioles 

1. subsessile, less than 1.0 cm 

2. very short, 1.0-2.5 cm 

3. short, 2.5-5.0 cm 

4. medium, 5.0-10.0 cm 

5. long, greater than 10.0 cm 

15. Shape of the leaf 

1. lobate Fig 6C 

2. nonlobed Fig 109B 

16. Lobing of leaf 

1. even numbered lobes Fig 111B 

2. odd numbered lobes Fig 6D 

3. not applicable (nonlobed) Fig 109B 

17. Number and comparative size of leaf lobes 

1. nonlobed Fig 109B 

2. 3 lobed Fig 74D


TABLE I (continued) 

3. 3 major and 2 smaller lobes Fig 71B 

4. 3 major and 2 smaller lobes, plus occasionally 2 very small lobules Fig 25B 

5. 3 major and 2 smaller lobes, plus regularly 2 larger lobules Fig 18C 

6. 5 to 9 lobes Fig 6D 

18. Outline of nonlobed leaves 

1. orbicular Fig 109B 

2. reniform Fig 112D 

3. cordate Fig 112A 

4. obovate Fig 107A 

5. linear Fig 105B 

6. stellate Fig 111B 

7. lanceolate Fig 107D 

8. elliptic Fig 106A 

9. deltoid Fig 112B 

10. not applicable (lobed leaves) Fig 6D 

19. Outline of median lobes of lobed leaves 

1. linear Fig 68C 

2. linear undulate Fig 64A 

3. gladiate Fig 63B 

4. hastate Fig 80A 

5. oblong Fig 59A 

6. oblong pandurate Fig 121D 

7. ovate Fig 60C 

8. obovate Fig 27A 

9. obovate pandurate Fig 34C 

10. broadly obovate Fig 79B 

11. lanceolate Fig 67A 

12. elliptic Fig 71B 

13. rotundate Fig 58C 

14. cuneate Fig 56A 

15. digitiform Fig 13C 

16. rhomboid Fig 18C 

17. rhomboid pandurate Fig 16B 

18. parabolic Fig 44B 

19. parabolic incised Fig 21B 

20. not applicable (nonlobed leaf) Fig 109B 

20. Size of nonlobed leaf 

1. greater than 12.0 cm long X 8-12 cm wide 

2. greater than 12 cm long X greater than 12 cm wide 

3. greater than 12 cm long X less than 3 cm wide 

4. 8-12 cm long X 8-12 cm wide 

5. 8-12 cm long X greater than 3.0 cm wide 

6. 8-12 cm long X 1-3 cm wide 

7. 8-12 cm long X less than 1.0 cm wide 

8. not applicable (lobed leaf) 

21. Dimensions of median lobe of lobed leaves 

1. 2.5-5.0 cm long X less than 1.0 cm wide 

2. 2.5-5.0 cm long X greater than 1.0 cm wide 

3. 5.0-12.0 cm long X greater than 12.0 cm wide 

4. 5.0-12.0 cm long X 5.0-10.0 cm wide 

5. 5.0-12.0 cm long X 1.0-5.0 cm wide 

6. 5.0-12.0 cm long X less than 1.0 cm wide 

7. greater than 12.0 cm long X 5.0-10.0 cm wide 

8. greater than 12.0 cm long X 1.0-5.0 cm wide 

9. greater than 12.0 cm long X less than 1.0 cm wide 

10. not applicable (nonlobed leaf) 

22. Shape of lowest lobes and comparison of size with upper lobes 

1. nonsymmetric slightly smaller than median lobe Fig 79B 

2. curved up, slightly smaller than median lobe Fig 58A 

3. curved down, slightly smaller than median lobe Fig 52A 

4. "S" shaped, slightly smaller than median lobe Fig 13A



5. same shape and size as that of median lobe Fig 70A 

6. same shape as median lobe and about 1/ its size Fig 71B 

7. tapering, and about /4 the size of the median lobe Fig 19A 

8. rounded, and about 1/4 the size of the median lobe Fig 25B 

9. not applicable (no lobes) Fig 109B 

23. Petiole attachment to lamina 

1. nonpeltate Fig 74D 

2. very narrowly peltate, less than 0.2 cm Fig 94C 

3. narrowly peltate, 0.2-0.5 cm Fig 45C 

4. widely peltate, 0.5-2.5 cm Fig 45A 

5. very widely peltate, greater than 2.5 cm Fig 111B 

24. Width of lamina at base of median lobe sinus 

1. leaf appears compound 

2. narrow, less than 0.5 cm 

3. variable (narrow to wide on the same plant) 

4. wide, 0.5-2.0 cm 

5. very wide, greater than 2.0 cm 


25. Width of median lobe base 

1. leaf appears compound 

2. narrow, less than 0.5 cm 

3. variable (narrow to wide on the same plant) 

4. wide, 0.5-2.0 cm 

5. very wide, greater than 2.0 cm 


26. Shape of sinus 

1. Fig 82C 

2. Fig 94B 

3. Fig 64D 

4. Fig 34C 

5. Fig 74D 

6. Fig 51B 

7. Fig 68A 

8. Fig 109B 

27. Shape of median lobe apex 

1. Fig 66B 

2. Fig 6D 

3. Fig 119D 

4. Fig 120C 

5. Fig 56A 

6. Fig 74D 

7. Fig 56D 

8. Fig 34C 

9. Fig 82C 

10. Fig 109B 

28. Shape of leaf apex (nonlobed leaf) 

1. Fig 112D 

2. Fig 112B 

3. Fig 109B 

4. Fig 107A 

5. Fig 105B 

6. Fig 6D 

29. Texture of lamina 

1. coriaceous 

2. medium membranaceous 

3. thinly membranaceous 

30. Gross appearance of lamina surface 

1. smooth 

2. glossy 

3. rugose



31. Position in which leaf lobes are held 

1. erect 

Fig 67C 

2. horizontal Fig 6A 

3. deflexed 

Fig 70A 

4. retrorse 

Fig 60C 

32. Bloom on the abaxial surface of lamina 

1. prominently pruinose 

2. medium pruinose 

3. non pruinose 

33. Pattern of waxy exudate on the abaxial lamina surface 

1. smooth 

Fig 1E 

2. verrucate Fig 1B 

3. reticulate Fig 1A 

4. pusticulate Fig 1C 

5. spongiosus Fig 1D 

34. Leaf venation 

1. camptodromous Fig 2B 

2. craspedodromous Fig 2A 

35. Inflorescence structure 

1. panicle, large, profusely branched, loose Fig 12B 

2. panicle, medium, loose Fig 37C 

3. panicle, medium, compact Fig 39C 

4. panicle, short Fig 65D 

5. raceme, single, long Fig 18D 

6. raceme, single, medium 

Fig 10C 

7. raceme, single, short Fig 14D 

8. raceme, cluster, long Fig 60A 

9. raceme, cluster, medium Fig 122A 

10. raceme, cluster, short Fig 38B 

11. raceme, subspicate, single, long Fig 20C 

12. raceme, subspicate, single, medium Fig 21D 

13. raceme, subspicate, single, short Fig 105C 

14. raceme, subspicate, cluster, long Fig 79C 

15. raceme, subspicate, cluster, medium Fig 76A 

16. raceme, subspicate, cluster, short Fig 75C 

17. corymboid raceme Fig 9D 

18. cluster of panicles Fig 36C 

19. cymose inflorescence Fig 98B 

36. Length of staminate tepal 

1. greater than 2.0 cm 

2. less than 2.0 cm 

37. Shape of staminate buds 

1. tubular Fig 21D 

2. slightly bifusiform Fig 96B 

3. distinctly bifusiform Fig 38C 

4. conical Fig 14D 

5. ovoid-ellipsoid Fig 53D 

38. Nature of bracteoles and bractlets 

1. foliaceous Fig 115C 

2. semifoliaceous Fig 95A 

3. setaceous Fig 77C 

39. Bracteole and bractlet margin 

1. entire Fig 61C 

2. serrate Fig 9D 

3. laciniate Fig 115C 

40. Position of pistillate flowers in inflorescence 

1. basal Fig 70B 

2. little above the base Fig 44C 

3. restricted to the upper half of the rachis Fig 112B 

4. terminal on a dioecious inflorescence Fig 98B



41. Size of fruits (base to apex) 

1. very small, less than 0.75 cm 

2. small, 0.75-1.25 cm 

3. medium, 1.25-1.75 cm 

4. large, 1.75-2.5 cm 

5. very large, greater than 2.5 cm 

42. Fruit surface 

1. smooth Fig 36D 

2. perceptably ribbed (No picture available) 

3. prominently ribbed Fig 120B 

43. Shape of fruit apex 

1. rounded 

2. slightly pointed 

3. prominently pointed 

4. depressed 

44. Seed size 

1. small, round, less than 1.0 cm 

2. small, oblong, less than 1.0 cm 

3. medium, round, 1.0-1.5 cm 

4. medium, oblong, 1.0-1.5 cm 

5. large, round, 1.5-2.0 cm 

6. large, oblong, 1.5-2.0 cm 

7. very large, oblong, greater than 2.0 cm 

to be discovered in the roots. Primarily, the species have a tap root (which may be 

enlarged with storage tissue), secondary roots, and adventitious roots either enlarged or 

slender and tapering. Species of the dryer regions frequently have woody roots, and a 

root crown from which new stems are produced when growing conditions are 

satisfactory. Adventitious roots from cuttings in the species M. esculenta provide the 

major carbohydrate food supply to millions of tropical people. All roots so far examined 

contain, in addition to large quantities of starch, varying concentrations of a cyanogenic 

glucoside which, upon autolysis, releases gaseous HCN. Refer to Rogers and Fleming 

(1973) and DeBruijn (1971) for more details on the roots of M. esculenta, and Rogers 

and Appan (in press) for descriptions of the roots of North American species. 

Stems 

We have used three stem characters (Table I) to differentiate the taxa, the first one 

describing the growth habit, one, the pubescence of the young stem (character 2), and one, 

the nature of the vegetative shoot (character 10). In Character 1, Habit of Growth, the 

attributes are generally self-explanatory. A compact subshrub refers to plants where the 

internodes have been telescoped, and the leaves appear to form a rosette. The term 

"subshrub" is used for those species in which the stem is usually less than 1.0 m tall, 

which are herbaceous, fleshy, or semiwoody, and which have little or no continued 

cambium activity. "Clambering" refers to those in which the stems start as erect, but later 

may droop over, as contrasted with vines, in which the stem grows over, or through, other 

vegetation. The second stem character is self-explanatory. The third stem character 

(number 10), serves to differentiate the Section Stipulares from all the other sections. 

Pubescence of plants 

For convenience's sake, we have grouped the characters describing the surface hairs 

from all the parts of the plants together as characters number 2 to 9, and for each part,


the descriptive attributes of hairiness are the same. This does not indicate, however, that 

all the parts on a single specimen necessarily possess the same attribute of hairiness. 

Leaves 

Twenty-three characters (number 11-34) describe the leaves of the genus. All leaves 

are provided with stipules, most of which are caducous, but some are persistent. Some of 

the stipules are foliaceous, and quite prominent, but preponderantly they are slender, or 

filiform. All leaves are petiolate, although some of petioles are less than 1 cm long, and 

give the appearance of being sessile. Two general leaf forms are found, simple and 

nonlobed, or simple and lobed. The preponderant number of species have variously lobed 

leaves. All the species with simple, nonlobed leaves are South American, and fall into two 

sections, Brevipetiolatae and Peltatae. The names of the sections are descriptive of 

characteristics of importance in the classification of these species. All other species in the 

genus manifest some degree of lobing, but any one species may have leaves associated 

with the inflorescence which are completely nonlobed. This phenomenon has been a 

source of confusion in the past, in that some specimens of some species with 

characteristic lobed leaves have been misplaced with the species characterized by 

nonlobed leaves. Only by careful collecting techniques may we be assured that such 

confusion can be avoided in the future. Another source of confusion caused by 

inadequate sampling of populations is that the lobes of the species with lobed leaves vary 

(within limits) in shape, size, and number. Careful attention, however, indicates the usual 

number, the usual size, and usual shape can be determined. We have described these 

conditions in our species descriptions. 

The width of the lamina at the base of the sinus of the lobes varies from very minute 

widths of laminal tissue to quite broad. Some of the lobed leaves appear compound on 

casual observation, but there are no truly compound leaves in the genus. Petiolar 

attachment to the lamina may be either basal or peltate. Again, the variation is from 

almost no lamina surrounding the attachment point to leaves where the petiole is nearly 

central on the lamina. Each of the states of the character is accompanied by limiting 

dimensions, rather than employ purely qualitative states of differentiation. The shape and 

depth of the sinus between the lobes are useful differentiating characteristics. The 

differing shapes of the sinus are difficult to describe, and to aid the user in differentiating 

the states of the character, we have illustrated them. See Character 26, Table I, and 

figures accompanying this character. 

The cuticular wax pattern (Fig 1) on the abaxial surface of the lamina provides one 

of the most important characters for species differentiation. Unfortunately, this pattern 

must be observed at magnifications of X 40, with good light sources. The states of the 

character are sufficiently descriptive to require no further elaboration. In addition to this 

pattern, there are several species easily identified by the "bloom" associated with the 

leaves, on the abaxial surface only. 

The various shapes of the median lobe and the apex of this lobe are important 

differentiating characteristics. Each state of the characters is illustrated, and the figure 

representing each is given in Table I. 

The patterns of the secondary veins in the leaves are important, and two states 

describing these patterns, camptodromous and craspedodromous, are employed in 

character 34. These states, illustrated in Fig 2, follow the definitions given by Foster and 

Gifford (1959, p 450). Camptodromous veins are "pinnately arranged secondary veins 

interconnected and not terminating at the lamina margin" and craspedodromous veins are 

"pinnately arranged secondary veins extending to the lamina margin." In the latter type, 

vein ends may actually project perceptibly beyond the lamina, and in the former, the


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FIG 2. Leaf venation. A, craspedodromous type, the secondary veins extending to the lamina 

margin (Kearney & Peebles 14928, ARIZ); B, camptodromous type, the secondary veins interconnected 

and not terminating at the lamina margin (Peebles 8796, ARIZ). 

veins near the margin curve upward, and parallel the margin for a distance before joining 

the next vein above. 

Inflorescence 

The inflorescences of the species are, with one possible exception, indeterminate 

racemes or panicles. The exception is the pistillate inflorescence of M. stipularis (Fig 98B) 

which is a consistently two-flowered cyme. There is considerable variation in the size of 

the inflorescence on any species (as indicated by our own field experience) but this 

variation is difficult to detect on most herbarium collections. Therefore, it is impossible 

to give any absolute measurements with the character states describing the inflorescence, 

except by such comparative terms as "short," "medium," or "long." However, with the 

other descriptive terms employed for each character state, associated with the above 

comparative terms, the user should be able to distinguish the specific type of 

inflorescence of an unknown. The simplest inflorescence type is a raceme, and in some of 

these, the pedicels are very short, giving the general appearance of a spike. We have 

termed this type as a "subspicate raceme." In some species, several racemes may arise 

from a common base, in which case we have used the term "cluster" to accompany the 

descriptive term raceme, to distinguish these from those species whose inflorescences are 

simple racemes. We consider a cluster of racemes to be distinct from the panicle. 

Bracteoles and bractlets provide three systematic criteria: pubescence (character 7), 

nature (character 38) and margins (character 39). While the states of character 7 and 39 

are easily understood, those of character 38 require comparisons with more than one


specimen for accurate assignment. Dimensions of foliaceous, semifoliaceous and setaceous 

bracts are not significant as means of differentiation, but the qualitative terms are suf- 

ficiently descriptive to provide the user with good differentiating ability. 

Staminate flower 

The flowers of the species of Manihot are remarkably uniform in their morphology 

(refer to discussion in the section Biology of Speciation). The staminate flowers are 

generally toward the apex of any inflorescence. 

The outline of the staminate flower buds is a good differentiating character. Their 

shapes are described in character 37. The flowers are characterized by a single perianth 

whorl in all species, called tepal in this monograph. The term does not indicate whether 

the perianth whorl is either calyx or corolla, and there is some doubt as to the origin of 

the tepal. Generally the term infers calycine origin. In the staminate flower, the tepals are 

united for about one half their length, and the lobes of the tepals are either erect, 

spreading, or reflexed. The stamens are generally ten (sometimes eight) in number, in two 

whorls. The slender filaments support versatile anthers with two longitudinally dehiscing 

thecae. The large pollen grains are generally sticky, and polyforate, the sculpturing 

consisting of round or triangular verrucae. The disc of the staminate flower is lobed, with 

the filaments arising between the lobes. Generally, a small pistillode is found in the center 

of the disc. There are differences in overall size of the flowers, variations in pubescence, 

and in color. Pubescence of the flowers are adequately reflected in characters 7, 8 and 9. 

Pistillate flower 

Pistillate flowers occupy the lower portion of the inflorescence. There may be a bare 

space along the major inflorescence axis before the appearance of a pistillate flower, or 

the pistillate flowers may be inserted very near the base of the inflorescence. The relative 

positions of insertion of the pistillate flowers are employed in character 40. 

The tepals of the pistillate flower are free, 5-merous, generally linear, apices acute, 

variously glabrous to pubescent most prominently along the inner margins. The disc of 

the female flower is generally smooth, not lobed. The 3-carpellate, 3-locular ovary 

generally has one pendulous, anatropous ovule per locule. The three-lobed stigma is sessile 

or nearly so. The shape and size of the structures of the pistillate flower are generally 

quite uniform among the species. 

Fruit and seed 

The fruit is a capsule, with septicidal and (usually) loculicidal dehiscence. The 

surface of the fruit is generally smooth, sometimes more or less prominently 3-winged or 

ribbed. The shape is generally ovoid to ellipsoid, but certain species have obpyriform 

fruits. The apex of the fruits may be depressed, rounded, or acute. The size of the fruit, 

the shape of the apex, and the presence or absence of wings are the only characters 

available as systematic criteria. 

The carunculate seeds are typically euphorbiaceous. Their surface may be plain or 

mottled, but the degree of mottled pattern depends to some extent upon the maturity of 

the seeds. The caruncle and the whole seed vary in dimension and shape, and these are the 

only available characters of sufficient stability to use in the classification.

Systematic Position of the Genus 19 

SYSTEMATIC POSITION OF THE GENUS 

The 12th edition of Syllabus der Pflanzenfamilien (Melchior, 1964) places the genus 

Manihot in a separate tribe, Manihotae, primarily on the basis of the single perianth whorl 

in the species. We concur in this placement, because it removes some of the confusion 

caused by Pax's (1910) inclusion of the genus in the tribe Adrianeae along with several 

other genera of little relation to Manihot. Linnaeus' placement of the species Manihot 

esculenta in the genus Jatropha has caused considerable nomenclatorial confusion, though 

little confusion among those who have studied the two genera in any detail. The usual 

double perianth whorls of Jatropha clearly distinguish the two genera, and this distinction 

has been reflected in more recent placement of Manihot and Jatropha in different tribes. 

Miller and Webster (1962), discussing the differences which clearly separate the two 

genera Jatropha and Cnidosculus, raise the possibility of a close relation between the 

latter genus and Manihot. They suggest that because both Manihot and Cnidosculus have 

a single floral envelope, and both share a common base chromosome number (X = 9), that 

the genus Cnidosculus should be given a different assignment in the Crotonoideae, but 

state that it would be premature, with present knowledge, to make a definite placement 

other than in the Jatrophinae. We concur that it is premature, and that careful 

comparative studies need to be made over the whole suprageneric classification before 

good decisions can be made. Clearly such a study is beyond the scope of this monograph. 

1. MANIHOT P. Miller, The Gardener's Dictionary Vol. II, London, 1754, 4th Ed; 

Adanson, Fam. II. 356. 1763; Crantz, Inst. Rei Herb. I. 658. 1766; Pohl, P1. Bras. Ic. 

et Descr. 1. 17-56. 1827; Muell.-Arg. in DC. Prodr. 15(2): 1057-1075. 1866; in 

Martius Fl. Bras. 11(2): 438-486. 1874; Pax in Engler, Pflanzenr. IV. 147(Heft 44): 

21-111. 1910; Standley, Contr. U.S. Nat. Herb. 23: 1923; Pax & Hoffmann in 

Engler, Pflanzenr. IV. 147(Heft 85): 194-197. 1924; Croizat, Jour. Arnold Arb. 23: 

216-225. 1942. 

Jatropha L. Sp. PI. ed. 1. 1007. 1753 pro parte. 

Janipha H.B.K., Nov. Gen. et Spec. II. 84. t. 109. 1817. 

Mandioca Link, Handb. II. 436. 1831. 

Subshrubs, shrubs, trees or clambering vines. Latex usually, but not universally, 

produced in varying quantities. Plants typically producing a cyanogenetic glucoside, 

varying in concentrations from a few parts per million to ca 500 ppm. 

Roots tuberous or slender, the former condition predominant, frequently with large 

concentrations of stored starch; root epidermis smooth or roughened, peeling, frequently 

with lenticels clearly defined; subepidermis white, or frequently with varying intensities 

of red or yellow; phelloderm well-developed ca 0.2-0.3 cm thick; tuberous rooted species 

with cortex of white, cream or yellow parenchymatous tissues for starch storage; usually 

with a well-defined central vascular strand. 

Stems nearly acaulescent to caulescent; several of the suffruticose species with 

semiherbaceous stems dying back to a root crown; stem diameters to 30 cm at breast 

height; the internodes of the nearly acaulescent species telescoped; nodes of at least one 

species prominently enlarged; stems variously light gray, dark brown, yellowish to 

reddish; bark of the tree species generally smooth, infrequently roughened or peeling; 

branching generally dichotomous or trichotomous, the branching point determined by 

the presence of a terminal inflorescence; young stems with or without varying degrees of 

pubescence, green to dark red.


Leaves alternate, subsessile to long-petiolate, simple, nonlobed or palmately lobed in 

varying numbers and depth of lobing; young leaves green, bluish green to deep purple; 

stipules setaceous, caducous or persistent, margins entire, serrulate or dentate; petioles 

projected horizontally, at an angle, vertically, or, in some cases, drooping; pigmentation 

green, yellow-green, reddish-green to solid red; lamina either peltate or with basal 

attachment to petiole, lightly membranaceous to coriaceous, orbicular to linear, with 

varying depths of lobes from slight indentations to deep, cleft nearly to the midribs, with 

frequent variations of lobing within a species; the mature vegetative leaves of most species 

lobed, the lobes erect or recurved, linear, elliptic, or obovate, the margins entire, slightly 

indented, pandurate or falcate, the lobe number 3, 5, 7, 9 or more, infrequently 4, 6, or 

8; leaves associated with the inflorescence generally reduced in number of lobes, or 

nonlobed; the mature vegetative leaves of species in sections Brevipetiolatae and Peltatae 

not lobed or only very slightly lobed; the midrib usually prominent, the secondary veins 

either camptodromous or craspedodromous; the lamina glabrous or glaucous or with 

varying degrees of pubescence to tomentose, the abaxial lamina surface with a wax-like 

covering of varying patterns from smooth to reticulate, the adaxial surface either glossy 

or dull, but never with the same wax pattern exhibited by the abaxial surface. 

Inflorescence terminal, rarely axillary, largely monoecious, rarely dioecious, sub- 

spicate racemes, racemes or panicles, the pistillate flowers generally borne toward the 

base of the peduncles, or rarely admixed with the upper level staminate flowers, with 

setaceous, semifoliaceous or foliaceous bracts and bracteoles subtending the peduncles 

and pedicels, the bracts and bracteoles caducous or persistent, with entire, serrulate, 

serrate or laciniate margins. 

Flowers with a single perianth of free or united tepals, relatively small, from ca 0.5 

to 2.0 cm length, greenish, yellow-green to red or purplish; buds of staminate flowers 

ovoid, ellipsoid or bifusiform in outline, buds of pistillate flowers generally ovoid, 

sometimes somewhat elongate-ovoid; pistillate flowers protogynous, the tepals 5, glabrous 

or pubescent either both internally and externally, or on one surface only, free, ligulate, 

caducous; pistil with 3 carpellate, superior ovary, one anatropous ovule per carpel; the 

ovary subtended by a fleshy yellow, orange or red, smooth or lobed disc, staminodes 

sometimes present; the style short, the trifid stigma smooth or lobulate; staminate flowers 

urceolate, campanulate or infundibuliform, the tepals united for ca 1/2 their length, the 5 

lobes erect, slightly to pronouncedly reflexed, glabrous or pubescent either internally and 

externally, or on one surface only; disc of staminate flowers lobed or smooth, the 

filaments arising beneath, or at the margins of, the lobes; stamens 10 (rarely 8) in two 

whorls of 5 each, the inner longer or shorter than the outer whorl, the filaments slender, 

with anther attachment versatile; the anthers bithecate, the thecae opening longitudinally; 

pollen grains large, viscid, with prominent peg-like extensions of the exine. 

Fruit a capsule, with septicidal dehiscence, frequently with loculicidal dehiscence as 

well; the fruits globular, ellipsoid or conical, sometimes the distal end depressed; either 

winged or smooth, sometimes both within the same species. 

Seeds typically euphorbiaceous, with a more or less developed caruncle at the 

micropylar end, gray, brown, or mottled in varying degrees; the copious endosperm 

closely investing the embryo and the thin, flat cotyledons. 

Chromosome numbers in all species counted uniformly 2n=36. 

I LLUSTRATIONS. See figures associated with Table I. 

TYPE. Lectotype Fig 4 & 5 in Merian, Dissertatio de generatione et metamorphosibus 

insectorum Surinamensium. 1726. (Lectotype of Manihot esculenta Crantz 

selected by Rogers & Appan in accordance with Article 9 of the International Code of 

Botanical Nomenclature. Crantz, while describing Manihot esculenta, does not cite any 

specimen but cites Merian's figures. 

DISTRIBUTION. (Fig 3, 4).

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FIG 3. Geographical range of genus Maniot in North America. 

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FIG 4. Geographical range of genus Manibot in South America. 

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Systematic Position of the Genus 

Key to the Genera Manihot and Manihotoides 

1. Inflorescence many flowered; leaves evenly spaced on stem, leaf lobes usually more than 3.0 

cm long. 

1. Manihot. 

1. Inflorescence uniflorous, rarely 2 flowered; leaves borne at the apex of short lateral shoots, 

leaf lobes usually less than 3.0 cm long. 

2. Manihotoides. 

Key to the Sections of Manihot 

1. Taxon existing entirely under domestication, not known to be growing wild; cultivated for 

its edible roots; nodes on stem distinctly enlarged. 

Sect 1. Manihot (p. 25). 

1. Taxa existing mostly as wild populations, a few under cultivation, but grown for latex only 

and not for edible roots; nodes on stem slightly or not enlarged. 

2. Taxa native in North and Central America (USA, Mexico, Guatemala, British Honduras, 

Honduras, El Salvador, Nicaragua, Costa Rica and Panama). 

3. Shrubs or vines; fruits small (less than 1.75 cm long from base to apex), seeds small (less 

than 1.5 cm long). 

Sect 2. Parvibracteatae (p. 34). 

3. Trees; fruits large (more than 1.75 cm long from base to apex), seeds large (more than 

1.5 cm long). 

Sect 3. Foetidae (p. 65). 

2. Taxa native in South America (Colombia, Venezuela, Trinidad/Tobago, Guyana, 

Suriname, Guyane Francaise, Ecuador, Peru, Brasil, Paraguay, Uruguay, Argentina) and 

West Indies. 

4. Leaves deeply lobed; the sinus greater than half the length of the lobe. 

5. Plants caulescent; usually more than 25.0 cm tall; leaves widely spaced on stem; 

stipules mostly caducous, rarely persistent, usually less than 1.0 cm long; inflores- 

cence with pistillate and staminate flowers, indeterminate, racemose or paniculate. 

6. Leaf lobes variously shaped, but not linear or narrowly lanceolate. 

7. Erect woody shrubs, usually more than 0.5 m tall, leaves larger, median lobes 

usually more than 4.0 cm long. 

8. Abaxial surface of leaves glabrous or sparsely to densely pubescent, but not 

tomentose. 

9. Inflorescence a panicle. 

10. Erect shrubs or trees, (except M. filamentosa, a vine-like shrub in section 

6-see second 12 of this key). 

11. Low to medium tall shrubs or occasionally small trees, usually less 

than 5.0m tall; leaves membranaceous; petiole attachment mostly 

basal, rarely peltate, when peltate, lamina between petiole-midrib 

junction and basal edge of leaf is less than 0.5 cm wide; seeds oblong; 

bracts and bracteoles with entire margin. 

12. Median lobes entire or shallowly pandurate, when pandurate, the 

apical region not prominently dilated; population native in regions 

other than the Caribbean coast and the adjoining regions of 

Venezuela and Colombia. Sect 4. Heterophyllae (p. 77). 

12. Median lobes usually deeply pandurate, rarely entire, apical region 

prominently dilated and more or less triangular (Fig 47D); populations 

native on the Caribbean coast and the adjoining interior 

regions of Venezuela and Colombia. Sect 6. Carthaginenses (p. 113). 

11. Trees or tall shrubs, usually more than 5.0 m tall; leaves coriaceous; 

petiole attachment mostly peltate, rarely basal, when peltate, lamina 

between petiole-midrib junction and basal edge of leaf more than 0.5 

cm wide; seeds rounded; bracts and bracteoles if foliaceous (more than 

0.5 cm wide) with entire margin, otherwise (less than 0.5 cm wide) 

with dentate margin. 

Sect 11. Glaziovianae (p. 175). 

10. Vine-like shrubs, clambering over other vegetation. Sect 12. Peruvianae (p. 189). 

9. Inflorescence a raceme, single or several arising from a common base. 

13. Leaf lobe bases usually not extremely constricted, but if constricted the 

lobes overlapping at sinus; lamina at lobe bases usually more than 0.3 cm 

wide, lamina between base of sinus and petiole-midrib junction usually 

more than 0.3 cm wide. 

14. Low to medium shrubs; usually less than 3.0 m tall. 

15. Erect shrubs. 

16. Lamina coriaceous, adaxial surface dark green, abaxial surface 

distinctly glaucous, usually violet tinged; midribs usually purplish. 

Sect 7. Quinquelobae (p. 118). 

23


16. Lamina membranaceous; adaxial surface green, abaxial surface 

devoid of bloom and violet pigmentation; midribs yellowish green 

without purplish tinge. 

17. Inflorescence racemose, not subspicate; leaves usually 5 lobed, 

nonlobed leaves rare; plants glabrous, never pubescent. 

Sect 5. Anisophyllae (p. 105). 

17. Inflorescence subspicate; leaves usually 3 lobed, but often 

nonlobed; plants glabrous or with various degrees of pubescence. 

18. Petiole attachment basal. Sect 9. Sinuatae (p. 163). 

18. Petiole attachment peltate. 

Sect 10. Variifoliae (p. 172). 

15. Vine-like shrubs, clambering over other vegetation. Manihut peruviana 

(racemose species of section 12. Peruvianae). See also second 

10. 

14. Trees, usually more than 5.0 m tall. Manihot catingae and M. 

dichotoma, (racemose species of section 11. Glaziovianae). See also 

second 11. 

13. Leaf lobe bases extremely constricted; lobes not overlapping at sinus; 

lamina at lobe base usually less than 0.3 cm wide, lamina between base 

of sinus and petiole-lamina junction usually less than 0.3 cm wide. 

19. Bracts and bracteoles dilated, broadly lanceolate to ovate, more than 

0.5 cm wide; inflorescence usually pendent. 

20. Tall shrubs to trees, usually more than 3.0 m tall; lamina coriaceous; 

abaxial surface wax pattern showing discrete rings (at magnifications 

of X 40); petioles usually more than 8.0 cm long; bracts and 

bracteoles with entire margins, capsule with prominently raised ribs, 

apex acute; frequently cultivated as latex yielders. 

Sect 19. Caerulescentes (p. 238). 

20. Low to medium shrubs, usually less than 3.0 m tall, lamina 

membranaceous, wax on abaxial surface forming a reticulum (at 

magnification X 40); petioles usually less than 8.0 cm long; bracts 

and bracteoles with serrate or laciniate margins; capsule surface 

smooth, apex rounded; not known to be cultivated as latex yielders. 

Sect 18. Tripartitae (p. 230). 

19. Bracts and bracteoles narrow, lingulate to setaceous, less than 0.25 cm 

wide; inflorescence usually erect. Manihot xavantinensis (lobe-baseconstricted 

species of section 7. Quinquelobae). See also second 16. 

8. Abaxial surface of leaves tomentose. 

21. Leaves 5 or more lobed. 

22. Inflorescence less than 10.0 cm long; petioles less than 15.0 cm long; 

base of lamina without lobules. Sect 15. Grandibracteatae (p. 213). 

22. Inflorescence more than 10.0 cm long; petioles more than 15.0 cm long; 

lobules at base of lamina. Manihot jolyana (tomentose species of section 

4. Heterophyllae). See also second 12. 

21. Leaves 3-lobed. 

23. Raceme solitary; bracts and bracteoles laciniate; nonlobed leaves rare; 

lobe bases less than 1.0 cm wide; with lobules at base of lamina. Manihot 

tripartita subsp vestita (tomentose species of section 18. Tripartitae). See 

also second 20. 

23. Racemes in a cluster arising from a common base; bracts and bracteoles 

with entire margins, nonlobed leaves frequent; lobe bases more than 1.0 

cm wide; without lobules at base of lamina. Manihot anomala subsp 

pubescens (tomentose species of section 9. Sinuatae). See also second 

18. 

7. Procumbent, weak-stemmed, semiherbaceous shrubs, usually less than 0.5 m tall; 

leaves small, median lobes usually less than 4.0 cm long. 

Sect 13. Crotalariaeformes (p. 199). 

6. Leaf lobes distinctly linear, or narrowly lanceolate. 

25. Stipules caducous; petiole attachment basal. Sect 8. Graciles (p. 141). 

25. Stipules persistent; narrowly peltate. M. marajoara, M. surinamensis (linearlobed 

species of section 4. Heterophyllae). See also second 12. 

5. Plants acaulescent or nearly so; less than 25.0 cm tall; leaves borne more or less as a


rosette; stipules persistent, usually more than 1.0 cm long; inflorescence usually 

either pistillate or staminate, rarely of both sexes; pistillate inflorescence determinate, 

always 2 flowered. Sect 14. Stipulares (p. 206). 

4. Leaves nonlobed or very shallowly lobed, the sinus less than half the length of the 

lobes. 

26. Leaves subsessile; petioles short, less than 1.0 cm long. Sect 16. Brevipetiolatae (p. 215). 

26. Leaves petiolate; petioles long, more than 5.0 cm long. Sect 17. Peltatae (p. 225). 

Sectional Relations 

Figures 5A and 28A indicate the linear (as contrasted with the multidimensional) 

relationships of the sections of the genus Manihot. In addition, Fig 5A shows the 

relationship of the genera Manihot and Manihotoides. In terms of biological relationship, 

there are, of course, many other connections between these taxa, not indicated by this 

type of illustration. 

A critical examination of the Skylines (Figs 5, 28), and of those specific taxa 

delimited on the basis of poor specimens (or limited numbers of specimens), indicates 

that it is likely that some modification of specific taxa will be necessary when more 

specimens are available. However, we feel confident (on the basis of both morphological 

and geographical grounds) that the sectional differentiations represent stable taxonomic 

units. 

One may more readily achieve a grasp of the types of taxonomic relationship evident 

in the genus Manihot by careful comparison of the sectional taxa than by attempting to 

grasp the same concepts by examination of the individual species. 

1. Manihot sect Manihot 

Manihot sect Parvibracteatae Pax subsect Utilissimae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 66. 1910. pro parte. 

Manihot sect Heterophyllae Pax subsect Cartbaginenses Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 80. 1910. pro parte. 

Plantae caulinae, lignosae, infernae ad mediae frutices, folia petiolatae, penitae 

lobatae, inflorescentia monoecia panicula. 

Plants caulescent, woody, low to medium shrubs; leaves widely spaced on stem; 

petiolate, petiole attachment basal; lamina membranaceous, deeply lobed, lobes linear or 

obovate. Inflorescence a monoecious panicle; bracts and bracteoles setaceous, margins 

usually entire. 

TYPE. Manihot esculenta Crantz. 

DISTRIBUTION. World-wide lowland tropics, apparently native to the neotropics. 

1. Manihot esculenta Crantz, Institutiones Rei Herbariae; nutum naturae digestae ex 

habitu 1: 167. 1766; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 67. 1910 

(recognized by Pax as a synonym of M. utilissima Pohl); Ciferri, Relaz. e Mon. 

Agrario-Coloniali 44: 1-59. 1938; Archiv. Bot. Forli 18: 27-33. 1942; Rogers & 

Fleming, Econ. Bot. 27(1): 1973. 

Jatropha manibot L., Sp. PI. ed. 1: 1007. 1753. Janipha manihot Humboldt, Bonpland & Kunth, 

Nov. Gen. et Spec. 2: 84. 1817. 

Jatropha dulcis J. F. Gmelin, Onom. Bot. 5: 7. 1772-1778. (Pohl cites this name as Jatropha 

dulcis (Gesner) Gmelin). Manihot dulcis (J. F. Gmelin) Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 71. 1910. 

Jatropha mitis Rottb., Descript. Surinam. 21. 1776. 

Jatropha janipha Loureiro non L. (L. in Mantissa 1: 126. 1771). Fl. Cochinchinensis. 718. 1793. 

Jatropha stipulata Vellozo, Fl. Flum. Ic. 10: 5. t 82. 1825; Jatropha stipulata Arrabida in 

Steudel, Nomencl. ed. 2 1: 800. 1840. (This is the same as Jatropha stipulata Vellozo, but 

attributed by Steudel to Arrabida, the editor of Florae Fluminensis.) Type. Vellozo s n, n v.

~ 


o L 0

Systematic 

Position of the Genus 27 

FII. 5-1 

9 90 I5 s0 75 70 5 60 55 SO 

1 l 

I 

FIG 5-A. Dendrogram of relationships of the North American sections of Manihot (section 2 - 

Parvibracteatae, section 3 - Foetidae) and the relations of these to section 1, Manihot. The relation- 

ship of Manihotoides to the North American sections is also indicated. The similarity values are shown 

decreasing from left to right across the top of the figure. Section 1 joins section 2 at a value of 0.85, 

and sections 2 and 3 join at a similarity of 0.73. Manihotoides joins the Manihot sections at 0.54 

similarity value. 

Manihot aipi Pohl, P1. Bras. Ic. et Descr. 1: 29. 1827. Manihot palmata var aipi (Pohl) Muell.-Arg. 

in DC., Prodr. 15(2): 1062. 1866. M. dulcis (J. F. Gmelin) Pax var aipi(Pohl)Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 71. 1910. Type. Pohl 3777 (syntype, W). 

Manihot aipi Pohl var lutescens Pohl., PI. Bras. Ic. et Descr. 1: 31. 1827. Type. Pohl 3780 

(syntype, W). 

Manihot aipi Pohl var lanceolata Pohl, PI. Bras. Ic. et. Descr. 1: 31. 1827. Type. Pohl 3778 pro 

parte (syntypes, W-3 ). 

Manihot aipi Pohl var latifolia Pohl, PI. Bras. Ic. et. Descr. 1: 31. 1827. Type. Pohl 3776 

(syntype, W). 

Manihot utilissima Pohl. PI. Bras. Ic. et Descr. 1: 32. t 24. 1827. Muell.-Arg. in DC., Prodr. 15(2): 

1064. 1866; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 67. 1910; (and many others). 

Type. Pohl 3775 (syntype, W). 

Manihot utilissima Pohl var castellana Pohl, PI. Bras. Ic. et Descr. 1: 34. 1827. Type. Pohl 3778 

pro parte (syntypes, W-2). 

Manihot utilissima Pohl var sutinga Pohl, P1. Bras. Ic. et Descr. 1: 34. 1827. Type. Pohl 3781 

(syntype, W). 

Manihot flabellifolia Pohl, PI. Bras. Ic. et Descr. 1: 35. 1827; Jatropha flabellifolia Steudel, 

Nomencl. ed. 2. 1: 799. 1840; Manihot palmata var flabellifolia (Pohl) Muell.-Arg. in DC., 

Prodr. 15(2): 1062. 1866; Manihot dulcis (J. F. Gmelin) Pax var flabellifolia (Pohl) Pax in 

Engler, Pflanzenreich IV. 147(Heft 44): 72. 1910. Type. Pohl 1188 (syntypes, F, G, M-3, 

W-2). 

Manihot digitiformis Pohl, PI. Bras. Ic. et Descr. 1: 36. t 27. 1827;Jatropha digitiformis Steudel, 

Nomencl. ed. 2. 1: 799. 1840; Manihot palmata var digitiformis (Pohl) Muell.-Arg. in DC., 

Prodr. 15(2): 1063. 1866. Type. Pohl 1371 (syntypes, F, W-2); Pohl 1709 (syntypes, G, 

K-2). 

Manihot diffusa Pohl, P1. Bras. Ic. et Descr. 1: 55. 1827;Jatropha diffusa Steudel, Nomencl. ed. 

2. 1: 799. 1840; Manihot palmata var diffusa (Pohl) Muell.-Arg. in DC., Prodr. 15(2): 1062. 

1866. 

Manihot dulcis (J. F. Gmelin) Pax var diffusa (Pohl) Pax in Engler, Pflanzenreich IV. 147 (Heft 

44): 71. 1910. Type. Martius s n (syntype, G); Martius s n (2427?) (syntype, M). Jatropha 

paniculata Ruiz & Pavon ex Pax in Engler, Pflanzenreich IV. 147(Heft 44): 71. 1910 pro 

syn. 

Manihot loureirii Pohl, P1. Bras. Ic. et Descr. 1: 55. 1827; Jatropha loureirii Steudel, Nomencl. 

ed. 2. 1: 799. 1840. Type. Loureiro s n, n v. 

Mandioca utilissima Link, Handb. 2: 436. 1831. 

Manihot edule A. Rich. in R. de la Sagra, Fl. Cub. ed. hisp. 3: 208. 1853. 

Manihot melanobasis Muell.-Arg., Linnaea 34: 206. 1865. Type. Schomburgk 694 (syntypes, F, 

NY, P); Schomburgk 426/694 (syntype, K); Schomburgk 426 syntypes, F, OXF, P, W-2). 

Mandioca dulcis Parodi, An. Soc. Ci. Argent. 4: 127. 1877. 

Manihot manihot Cockerell, Bull. Torrey Club 19: 95. 1892. 

Manihot sprucei Pax in Engler, Pflanzenreich IV. 147(Heft 44): 71. 1910. ad interim; Manihot 

esculenta var sprucei Lanjouw, Euphorb. Surinam 33. 1931. Type. Spruce 825 (syntypes, 

BM, F, K-2, NY-2, OXF, P, W). 

Manihot flexuosa Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 195. 1924. 

Type. Luetzelburg 9 (9A, 9B?) (syntypes, F, M-2); Luetzelburg 10 (syntypes, M, NY). 

The following synonyms, all named by Ciferri (using epithets provided by Pohl as the basionym) 

were not, to our knowledge, based on the examination of specimens of Pohl, but were based 

on materials from the Dominican Republic. Ciferri did not cite any specimens, and we have 

not found any specimens either annotated by him, nor collections made by him. 

5Throughout the monograph, we employ the convention of the acronym followed with a dash 

and a number to indicate that number of specimens in the herbarium.


Manihot esculenta Crantz subsp flabellifolia (Pohl) Ciferri var coalescens Ciferri, Archiv. Bot. 

Forli 18: 31. 1942 

Manihot esculenta Crantz subsp flabellifolia (Polh) Ciferri var nodosa Ciferri, Archiv. Bot. Forli 

18: 31. 1942. 

Manihot esculenta Crantz subsp flabellifolia (Pohl) Ciferri var digitifolia Ciferri, Archiv. Bot. 

Forli 18: 31. 1942. 

Manihot esculenta subsp flabellifolia (Pohl) Ciferri var debilis Ciferri, Archiv. Bot. Forli 18: 31. 

1942. 

Manihot esculenta Crantz subsp flabellifolia (Pohl) Ciferri var flavicaulis Ciferri, Archiv. Bot. 

Forli 18: 31. 1942. 

Manihot esculenta Crantz subsp flabellifolia (Pohl) Ciferri var fuscescens Ciferri, Archiv. Bot. 

Forli18: 31. 1942. 

Manihot esculenta Crantz subsp flabellifolia (Pohl) Ciferri var argentea Ciferri, Archiv. Bot. Forli 

18: 31. 1942. 

Manihot esculenta Crantz subsp grandifolia Ciferri sectio alboerecta Ciferri var ramosissima 

Ciferri, Arch. Bot. Forli 18: 31. 1942. 

Manihot esculenta Crantz subsp grandifolia Ciferri sectio alboerecta Ciferri var luteola Ciferri, 

Archiv. Bot. Forli 18: 31. 1942. 

Manihot esculenta Crantz subsp grandifolia Ciferri sectio alboerecta Ciferri var bispaniolensis 

Ciferri, Archiv. Bot. Forli 18: 31. 1942. 

Manihot esculenta Crantz subsp grandifolia Ciferri sectio alboerecta Ciferri var rufescens Ciferri, 


Manihot esculenta Crantz subsp grandifolia sectio alboerecta Ciferri var domingensis Ciferri, 


Manihot esculenta Crantz subsp grandifolia Ciferri sectio alboerecta Ciferri var mutabilis Ciferri, 


Manibht esculenta Crantz subsp grandifolia Ciferri sectio diffusa (Pohl) Ciferri var jamaicensis 


Manihot esculenta Crantz subsp grandifolia Ciferri sectio diffusa (Pohl) Ciferri var fertilis Ciferri, 


Manihot esculenta Crantz subsp grandifolia Ciferri sectio diffusa (Pohl) Ciferri var pohlii Ciferri, 


Manihot esculenta Crantz subsp grandifolia Ciferri sectio diffusa (Pohl) Ciferri var zimmermannii 


Manihot esculenta Crantz subsp grandifolia Ciferri sectio diffusa (Pohl) Ciferri var communis 


General Species Description. 

Tropical shrubs, 1 to 4 m tall; all parts of plant with varying concentrations of a 

cyanogenic glucoside. Roots from seed a tap root, with secondary roots generally slender; 

adventitious roots arising from stem cuttings tuberous, variously shaped, from long and 

slender to globose. Stems woody, glabrous, or sparsely pubescent, except at the heavily 

pubescent apex, variously branched, from low, many-branched plants to tall and 

essentially unbranched; pith usually massive; leaf and stipule scars usually raised, 

sometimes not raised; predominantly brown, but sometimes yellow or silver. Leaves 

simple, frequently of two forms, either palmately lobed or nonlobed, the former 

predominant, the latter mostly associated with the inflorescence. Leaf lobes (2-)3-10(- 

12), linear, obovate or pandurate. The upper and lower lamina surfaces slightly pubescent 

between the veins, slightly to heavily pubescent on major veins; abaxial surface of lamina 

with a farinose layer, generally in a hexagonal pattern about the stomata; margins entire 

or slightly sinuate; stipules caducous, linear or laciniate; petioles from 5 to 25 cm long, 

frequently slightly S-shaped, attached basally to the lamina or slightly peltate, glabrous or 

slightly pubescent at base, and frequently a tuft of hairs at apex; leaves associated with 

inflorescence generally smaller, frequently 3-lobed, or sometimes simple, entire; lamina 

generally dark green, sometimes with red veins; petioles green, yellow-green, red or mixed 

red and green; young foliage at stem apices green, bluish green to dark red, very heavily 

pubescent. Inflorescence a panicle, generally from 2 to 10 cm long, glabrous; bracts and 

bracteoles strap-shaped, generally inconspicuous and caducous. Flowers monoecious, the


pistillate basal, opening first, the staminate apical, opening later. Pistillate flower 

hypogynous; perianth of 5 separate, strap-shaped tepals, red, green or purplish, pubescent 

along inner margins, and frequently with a tuft of hairs at the interior apex; ovary 

subtended by a nonlobed disc, 3 carpellate, glabrous, style short, with 3 finely dissected 

stigmas. Staminate flower with a 5 lobed perianth of tepals united about half the length, 

glabrous externally, pubescent internally; stamens usually 10 (infrequently 8) in 2 whorls, 

5 short and 5 longer, the filaments slender, glabrous, arising between the lobes of the 

basal disc, supporting versatile anthers; pollen 3-colpate, large, sticky, with spine-covered 

exine, in some cultivars sterile. Fruit a schizocarp, usually winged, but sometimes 

smooth-surfaced; dehiscence septicidal and loculicidal, leaving a central stalk. Seeds 

carunculate, elongate or rounded, variously marked, mottled brown and light brown, or 

plain. 

Detailed Cultivar Description. 

Surface of tuberous roots most frequently rough but often smooth. External root 

color most frequently brown, dark brown or reddish brown but often light brown, tan or 

light tan, seldom light brown-yellow, pinkish white, light pink or pink or pinkish brown 

or pinkish tan. Root flesh usually white to cream, sometimes cream-yellow or yellow, 

rarely with some pink intermixed. Stems commonly brown, but often silver, sometimes 

yellow, seldom silver-brown. Storey length usually 9 to 20 cm, but sometimes short, 4-8 

cm or long, 21-28 cm. Scars on stem usually either slightly raised or moderately raised, 

sometimes very large, seldom not raised (stem smooth). Stems most frequently more than 

two-branched, but commonly with one or two branches (excluding any branches at the 

top), less frequently unbranched, or a single branch at the top. Leaves with basic number 

of lobes odd, but occasionally with an even number of lobes; most frequently 7 or 8 

lobed, sometimes 9 or 10 lobed, occasionally 5 or 6 lobed, rarely 3 or 4 lobed, or 

variable, 3, 4, and 5 lobed, or 4, 5, or 6 lobed. Leaf lobe shape prevalently obovate but 

sometimes linear; margins of obovate leaves usually simple, sometimes sinuous and rarely 

pandurate, of linear-shaped lobes, margins commonly sinuous, occasionally simple, rarely 

pandurate. Length of median lobe most frequently long, greater than 17 cm, often of 

moderate length, 14 to 17 cm and occasionally short, less than 14 cm. Width of median 

lobe predominantly moderate, 2.6 cm to 4.8 cm, sometimes narrow, 1.5 cm to 2.4 cm, or 

wide, 5 cm or more. Petioles mostly green, frequently red, occasionally reddish green, 

seldom greenish red. Young foliage most frequently green, often reddish blue, sometimes 

bluish green. 

Fig 6A, B, C, D; 7A, B. 

TYPE. Fig 4 & Fig 5 in Merian, Dissertatio de generatione et metamorphosibus 

insectorum Surinamensium. 1726. (Lectotype selected by Rogers & Appan in accordance 

with Art. 9, Intern. Code Bot. Nomen. Crantz, while describing Manihot esculenta, does 

not cite any specimen but cites Merian's figures). 

DISTRIBUTION. Manihot esculenta, as a cultigen, is cultivated in all tropical 

countries of the world. The species is not known in a purely wild state. The limitations 

upon its growth are largely those of low temperature. In some subtropical areas, the 

cultigen is raised where there are a few periods of frost, but in these areas, the crop is of 

little economic significance. For this reason, we have cited only specimens which were 

selected to indicate the range of variability within the species. The collections made by 

Rogers form a part of the monographic study of the species, published separately (Rogers 

and Fleming, 1973), and all are deposited in the herbarium of the National Arboretum, 

Washington, D.C. An outline of the classification of the subspecific "groups" is given at 

the end of this section. (Table II). Other specimens cited are those confined to the 

Western Hemisphere, and were chosen because they represent the more important 

variations within the species. The crop is American in origin, and specimens from other 

areas of the world do not vary significantly from those we have selected. MEXICO. Tamauli-


, ~.....i& , " 

Vi-,~~~ 

[ W 

C - . 

FIG 6. Manihot esculenta. A, the plant; B, variegated leaf (Rogers s n); C, leaf showing slightly 

pandurate lobes (Rogers 440, NA); D, leaf showing obovate lobes (Rogers 464, NA). 

i

Systematic 


A B.. 

FIG 7. Manibot esculenta. A, leaf showing linear lobes (Rogers 460, NA); B, inflorescence. 

pas: Rogers, Appan & Rogers 530 (NY) Gomez Farias, 22 Jun 1968. GUATEMALA. Santa Rosa: 

Heyde & Lux 4306 (NY, US) Santa Rosa, Nov 1892. HONDURAS. Atlantida: Standley 53767 (F) 

Triunfo, near Tela, 28 Dec 1927. COSTA RICA. Cartago: Rogers 51 (F, W, K, COLO, MO, MICH, M, 

S, A, SP) Turrialba Institute, 14 Jul 1953; Rogers 236 (NA) Turrialba Institute, 15 Jun 1955; Rogers 

237 (NA) Turrialba Institute, 17 Jun 1955; Rogers 243 (NA) Turrialba Institute, 17 Jun 1955; Rogers 

249 (NA) Turrialba Institute, 17 Jun 1955; Rogers 252 (NA) Turrialba Institute, 17 Jun 1955. WEST 

INDIES. Jamaica: Rogers 77 (NA) Bodies Experimental Station, Jun 1954; Rogers 84 (NA) Bodies 

Experimental Station, Jun 1954; Rogers 100 (NA) Bodies Experimental Station, Jun 1954; Rogers 

101 (NA) Bodies Experimental Station, Jun 1954; Rogers 106 (NA) Bodies Experimental Station, Jun 

1954; Rogers 119 (NA) Bodies Experimental Station, Jun 1954; Rogers 120 (NA) Bodies 

Experimental Station, Jun 1954; Rogers 123 (NA) Bodies Experimental Station, Jun 1954; Rogers 

128 (NA) Bodies Experimental Station, Jun 1954; Rogers 131 (NA) Bodies Experimental Station, Jun 

1954; Rogers 257 (NA) Bodies Experimental Station, Jul 1955; Rogers 261 (NA) Bodies Experimental 

Station, Jul 1955; Rogers 263 (NA) Bodies Experimental Station, Jul 1955;Rogers 264 (NY) Bodies 

Experimental Station, Jul 1955; Rogers 267 (NA) Bodies Experimental Station, Jul 1955; Rogers 268 

(NA) Bodies Experimental Station, Jul 1955; Rogers 270 (NA) Bodies Experimental Station, Jul 


Station, Jul 1955; Rogers 278 (NA) Bodies Experimental Station, Jul 1955; Rogers 286 (NA) Bodies 

Experimental Station, Jul 1955; Rogers 293 (NA) Bodies Experimental Station, Jul 1955; Rogers 295 

(NA) Bodies Experimental Station, Jul 1955; Rogers 296 (NA) Bodies Experimental Station, Jul 


Station, Jul 1955; Rogers 312 (NA) Bodies Experimental Station, Jun 1960; Rogers 313 (NA) Bodies 

Experimental Station, Jun 1960; Rogers 321 (NA) Bodies Experimental Station, Jun 1960. 

COLOMBIA. Cundinamarca: Triana 3620(E) (P)Tiromena. VENEZUELA. Bolivar: Badillo 1475 

(VEN) Roca Altas de El Carmen, 18 Apr 1946; Velez 2465 (US) Paragua, 22 Apr 1946. GUYANA. 

Schomburgk s n (F) Savanna Province Pirara, Aug 1840; Schomburgk 426 (F, OXF, P, W-2), Pirara. 

Schomburgk 426/694 (K). Schomburgk 694 (F, NY, P) Savanna Bei Pirara. SURINAME. B.W. 2778 

(U) Zanderij I, 9 Mar 1917; B.W. 2785 (U) Zanderij I, 28 Mar 1917; B.W. 2841 (U) Zanderij I, 2 May 

1917; B.W. 2845 (U) Zanderij I, 5 Feb 1917; B.W. 2873 (U) Zanderij I, 23 May 1917. PERU. 

Huanuco: Rogers 453 (NA) Tingo Maria, Grounds of Hotel Tourista, Terrace above Rio Huallaga, 4 

Feb 1962. BRASIL. Amapa Territory: Pires, Rodriques & Irvine 50815 (NY-3) Rio Araguari, Sep 

1961; Pires, Rodriques & Irvine 5100 (NY) Rio Araguari, 20 minutes downriver from Porto Platon, 17 

Sep 1961. Amazonas: Rogers 490 (NA) 8 km NE of Manaus, on Colonia Santo Antonio, 24 Feb 1962;


Ule 7645 (MG, UC) Rio Branco, Bei Der Serra Pelloda, Oct 1908. Para: Huber s n (POM) Belem, Apr 

1908; Huber 9346 (MG) Belem, Horto Botanico; Rogers 323 (NA) Belem, Instituto Agronomico Do 

Norte, Jun 1960; Spruce 825 (BM, F, K-2, NY-2, OXF, P, W) in Vicinibus Santarem, Apr 1850. 

Pernambuco: Cruz 113 (SP) Goiania, 18 Jan 1965; Falcao, Egler & Pereira 329 (PUL) Saltinho, 3 Sep 

1954; Martius s n (M); Rogers 389 (NA) Tambe Experimental Station, 28 Feb 1961; Rogers 390 (NA) 

Tambe Experimental Station, 28 Feb 1961; Rogers 405 (NA) Araripina Agricultural Experimental 

Station in Chapada, 5 Mar 1961. Goias: Pobl Icon Tab. 24 (W-2); Pohl Icon Tab. 27 (W);Pohl 1188 

(F, G, M-2, W-2) Corgo Do Jaragua; Pohl 1371 (F, W-2) Corallinho; Pohl 1709 (G, K-2); Pohl 3775 

(W) Villa Boa; Pohl 3776 (W) Villa Boa; Pohl 3777 (G.W);Pohl 3778 (W-5) Villa Boa;Pohl 3780 (W) 

Villa Boa; Pohl 3781 (W) Ad Villa Boa; Ule 471 (P-2) Serra de Santa Barbara, Jan 1893. Bahia: 

Luetzelburg 10 (M, NY) S. Bento, Dec 1911, Luetzelburg 9A (F, M) Casa de Petro, Dec 1911; 

Luetzelburg 9B (M), Dec 1911: Martius s n (G); Martius 2427 (M) Apr.; Minas Gerais: Rogers 365 

(NA) Sete Lagoas, Grounds of Federal Experimental Station, 17 Feb 1961; Rogers 370 (NA) Sete 

Lagoas, Grounds of Federal Experimental Station, 17 Feb 1961. Sao Paulo: Labouriau 186 (SP) 

Instituto de Botanica, Sao Paulo, Cultivated, 17 Dec 1965. Rio de Janeiro: Peckolt 201B (W), 

Cantagello. BOLIVIA. La Paz: Bejarano 115 (COLO, F, K, US, W) Nor Yungas, Caranavi, end of air 

strip, near Coroico River, Jan 1962; Rogers 417 (NA), Nor Yungas, Caranavi, Flood plain near Rio 

Coroico, 6 Jan 1962; Rogers 419 (NA) Nor Yungas, road between Caranavi and Al Coche, 7 Jan 1962; 

Rogers 422 (NY-3) Nor Yungas, between Caranavi and Al Coche, along Rio Coroico, 7 Jan 1962; 

Rogers 423 (G-2) Nor Yungas, Chacaltaya, first terrace above Rio Coroico near Caranavi, 7 Jan 1962. 

Santa Cruz: Rogers 432 (NA) Canto Minera, 12 Jan 1962; Rogers 442 (NA) Ibanez, Paurito, 50 km SE 

of Santa Cruz, 18 Jan 1962. Province Unknown/Uncertain: Rusby 886 (NY) Mapiri, Apr 1886. 

PARAGUAY. Province Unknown/Uncertain: Christopher s n (OXF) Rio Paraguay, Jun 1944. 

COUNTRY: Unknown/Uncertain: Tweedie 103 (K). Tweedie 104 (K). 

TABLE II 

Rogers' specimens by Groups, cited in Monograph of 

Manihot esculenta Crantz (Rogers and Fleming, 1973). 

Group 

No. Specimen Major Categories 

264, 268 

2 267, 278 

3 243, 312 

4 

5 

6 

389, 252 

101, 249, 277 

263,270 

Obovate-lobed 

- 

Leaves 

7 106,236,295 

8 100,490 

9 77, 313 

10 323 "Rough"-rooted 

11 257,321 Stems brown, 

LC12 370NAE 390AND___ yellow or red 

13 84, 261 j Linear-lobed 

---------' Leaves 

"Smooth"-rooted 

15 

17 

120, 237 

123, 453 

. . 

Obovate-lobed Stems silver (or 

18 

19 

128,271,296 

119, 293, 365, 405 

Leaves 

LOCAL NAMES AND USES. The major common names are: cassava (English speaking 

areas), manioc (French speaking areas), yuca (Spanish speaking areas), mandioca 

(Portuguese speaking areas). All these may be modified by adding the equivalent of


"bitter" (poisonous) or "sweet" (nonpoisonous) to the major name. In Brasil, aipim or 

macachiera may be used rather than "sweet" to differentiate those cultivars used as a 

fresh vegetable. All these names (or some equivalent sound) seem to have been in use by 

natives in the Western Hemisphere tropics, and may apply either to the plant or to one of 

its useful products. For example, cassava (or cassabe) generally describes a large, dried 

cake made from the ground and dried root. Beju also refers to this product in Brasil. 

The major useful part of the plant is the starch-filled tuberous root, most often 

produced as an adventitious root from stem cuttings. However, in parts of Africa, 

particularly in the former Republic of the Congo (now designated Zaire), the foliage is 

eaten as a spinach with great regularity, and in considerable quantities. Use of the foliage 

is infrequent in the tropics of the Western Hemisphere. 

The roots may be used as a fresh vegetable, in a manner similar to use of the tubers 

of Solanum tuberosum, if the cultivar is known to produce roots with low concentrations 

of HCN. There are apparently no roots completely devoid of the poisonous substance, 

but in the "sweet" variations, most of the HCN is confined to the outer layer, or 

phelloderm. Those not entirely familiar with this cultigen would be well advised to treat 

all cultivars as poisonous, and to process them sufficiently to reduce the concentrations 

of HCN to tolerable levels. 

By far the largest food use of the roots follows a series of processes intended to 

remove the HCN (though recent evidence indicates that a trace of HCN remains after the 

best processing techniques), and perhaps to impart some desirable flavor to the product. 

In Brasil, the largest consumer of M. esculenta, much of the crop is produced as farinha, a 

dry meal eaten alone or in combination with many other foods. Gari, a product found in 

west Africa, is similar to Brasilian farinha. The number of variations on methods of 

preparation of the roots is almost endless. Frequently, a fermentation, or microbial 

conversion process, is employed in food production. Apparently, the flavors added by 

fermentation make the processed food more palatable. Tapioca is one of the best known 

of cassava products reaching the temperate zone markets. 

The starch from M. esculenta roots may be used in various industrial processes, such 

as paper sizing, in corrugated paper, or boxes, in many fillers where starches are 

important. Dried chips from the roots are imported into western Euorpean nations as part 

of cattle and hog rations. 

NOMENCLATURE. The nomenclatorial history of this species reflects, to some 

extent, the development of species concepts from their inception to the present. Linnaeus 

(1753) named the cultigen Jatropha Manihot. He did not accept Manihot as a separate or 

recognizable genus although European botanists used Manihot as a distinct taxon more or 

less equivalent to a genus for nearly 100 years before 1753. Linnaeus' inclusion of the 

cultivated species in Jatropha caused considerable confusion in the literature until the 

monographic work of Pohl, who, in 1827, definitively described the genus and many 

other species of the genus. Although Pohl's work was the definitive one, Miller, in 1754, 

is credited with the first post-Linnean application of the generic term. Miller intended to 

use an epithet first applied by Bauhin in 1651, M. theveti, but since he did not clearly 

espouse a strict binomial system (frequent polynomials) his species epithet is not 

permitted, and the name esculenta, first used by Crantz in 1766, becomes the first valid 

name for the cultivated species. Crantz's name was not recognized, however, until Pax's 

monograph in 1910, where the epithet was listed in synonymy underM. utilissima Pohl, 

and was not generally accepted until 1938, when Ciferri applied the modern concept 

wherein all the cultivars, with both high and low concentrations of the cyanogenic 

glucoside, are members of the same highly variable species. 

Many of the species epithets applied to M. esculenta are mere nomenclatorial 

synonyms, but several are here placed in synonymy for the first time, because we employ 

a broader concept of the species than any previous students. Thus, M. flabellifolia Pohl, 

M. digitiformis Pohl, M. diffusa Pohl, M. loureirii Pohl, M. melanobasis Muell.-Arg., M.

34 

Flora Neotropica 

sprucei Pax, and M. flexuosa Pax, all represent variations which fit well within our 

concept of this species. It is likely that each of the above species names is based upon 

materials collected from plants not consciously grown for their roots, but come from 

"volunteer" plants, apparently wild. 

We have placed all formal subspecific taxonomic designations of M. esculenta in 

synonymy because it is impossible to apply formal subspecific taxon epithets to fleeting 

variants which are not related to some precise geography or ecological region. In a 

monograph of the cultivars of M. esculenta (Rogers and Fleming, 1973), 19 informal 

"Groups" (see Intern. Code Nomen. Cult. PI., Utrecht, 1961) are recognized as clusters of 

similar morphologies, without relation to geographic distribution. The purpose of this 

grouping is to provide agricultural workers with guides for development of new or 

improved cultivars, and to document the morphological variation encompassed by the 

species, rather than for taxonomic naming of the variants. In our interpretation, Loureiro, 

Fl. Cochinchinensis 718, 1793, could only have seen Manihot esculenta, even though he 

cites as his authority Jacquin, who clearly described Manihot carthaginensis (Jacq.) 

Muell.-Arg. Loureiro indicates that the plants described were in cultivation, and we 

seriously doubt that Manihot carthaginensis had been imported into the Eastern 

Hemisphere tropics. 

BEGINNINGS OF CULTIVATION. This discussion is concerned with the earliest domestication 

of M. esculenta which is considered to be different from the botanical origin of 

the species. As evident from the computer analyses the closest wild relative of M. 

esculenta is M. aesculifolia (H.B.K.) Pohl. Some South American species are also related 

to a lesser extent (Fig 28), and thus the botanical evidence for the beginnings of 

cultivation are not yet sufficient to designate one area over another as the place where M. 

esculenta was first domesticated. In an earlier study, Rogers (1963) called attention to 

the possibility of a Meso American origin, as contrasted to other possible points of initial 

culture. Clearly, cultivation implies human activity, and the earliest such activity, with 

respect to documented archeological evidence, points to the Tehuacan Valley in the state 

of Puebla, Mexico (Callen, 1965). Other archeological records from Colombia (Reichel- 

Dolmatoff, 1956), Peru (Sauer, 1951), Tamaulipas, Mexico (MacNeish, 1958), merely 

indicate an early use of the species. We consider the point of initial cultivation to be a 

relatively unimportant problem, because the cultigen known today is a complex species 

with many points of initial cultivation. Wherever the species occurs (or has been 

transported by man), there is evidence (putative) that the plants have hybridized with 

other locally-occurring wild species, thus changing the genetic composition of the cultigen 

in such ways that the hybrid produced becomes a cultigen essentially unique to the region 

where the hybridization occurred. Since there are no apparent genetic barriers to 

hybridization within the genus Manihot, clearly a very complex species has developed and 

each hypothesis of origin or initiation of cultivation has some validity. 

An intriguing, but unsolved problem, is the origin of cultivars with little or no HCN 

versus those with relatively high concentration of the poisonous principle. Several 

suggestions have been made, apparently originating with Nordenskiold (1924), that in the 

extreme western part of the distribution in South America there were no poisonous 

varieties, but to the east, both types were found. This seems to have no basis in fact. Both 

genetic and enviromental influences play a role in the concentrations of HCN, but no 

studies have been sufficiently precise to determine the relative roles. The best studies to 

date of HCN in M. esculenta are those of DeBruijn (1971). 

2. Manihot sect Parvibracteatae Pax emend Rogers & Appan 

Manihot sect Sinuatae Pax subsect Warmingianae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

43. 1910. pro parte. 

Manihot sect Parvibracteatae Pax subsect Elatae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

55. 1910. pro parte. 

Manibot sect Parvibracteatae Pax subsect Graciles Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

76. 1910. pro parte.

i 

ilii!~? m , 

.... 

.....t


Manibot sect Heteropbyllae Pax subsect Carthaginenses Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 80. 1910. pro parte. 

Plants caulescent, woody, low to tall shrubs; leaves widely spaced on stem; petiolate, 

petiole attachment basal or peltate; lamina usually membranaceous, deeply lobed, lobes 

variously shaped, including linear. Inflorescence a monoecious raceme or panicle; bracts 

and bracteoles setaceous to foliaceous; margins entire to laciniate. 

TYPE. Manibot aesculifolia (H.B.K.) Pohl. 

DISTRIBUTION. (Fig 8). 

Key to the Species of Sect Parvibracteatae 

1. Bracts and bracteoles foliaceous; fruit pedicels ascending, held at an angle of about 45? to 

the rachis of the infructescence; lamina deeply cleft, less than 0.5 cm wide from base of 

sinus to petiole attachment point. 

2. M. pringlei. 

1. Bracts and bracteoles setaceous or semifoliaceous, never foliaceous; fruit pedicels not 

ascending, held horizontally or curved downwards; lamina not deeply cleft, more than 0.5 

cm wide from base of sinus to petiole attachment point. 

2. Leaf venation camptodromous; apex of secondary and tertiary lobes rounded. 

3. Tall erect shrubs more than 1.0 m tall; petiole attachment basal; staminate tepal lobes 

erect or slightly reflexed at anthesis. 

4. Staminate buds auriculate; leaf lobe margin repand. 

3. M. auriculata. 

4. Staminate buds campanulate or conical; leaf lobe margin entire, or pandurate, never 

repand. 


6. Leaf lobes oblong or oblong-pandurate, never linear; wax pattern of mature 

abaxial leaf surface verrucate; inflorescence a profusely branched panicle. 

4. M, aesculifolia. 

6. Leaf lobes linear, never oblong or oblong-pandurate; wax pattern of mature 

abaxial leaf surface smooth, inflorescence a sparsely branched panicle. 

5. M rubricaulis. 

5. Inflorescence a raceme. 

7. Leaf lobes obovate or obovate-pandurate; distal end of capsules rounded or 

projected but never depressed; fruit dehiscence septicidal; seeds oblong; 

arborescent shrubs (more than 1.5 m tall), or clambering vine-like shrubs. 

8. Staminate buds conical; racemes very short, usually less than 5.0 cm long, 

never longer than 8.0 cm; arborescent shrubs more than 1.5 m tall. 6. M. oaxacana. 

8. Staminate buds campanulate; racemes usually longer than 5.0 cm; clambering 

vine-like shrubs. 7. M. cblorosticta. 

7. Leaf lobes rhomboid or rhomboid-pandurate; distal end of capsules depressed; 

fruit dehiscence loculicidal; seeds rotund; medium sized (less than 1.5 m tall) 

slender-branched shrubs. 8. M. davisiae. 

3. Sprawling subshrubs, less than 1 m tall; petiole attachment peltate; staminate tepal 

lobes prominently reflexed at anthesis. (This condition occurs rarely in M. rhomboidea 

subsp rhomboidea). This taxon predominately has craspedodromous leaf venation. See 

second 2. 

2. Leaf venation craspedodromous (with the rare exception of some forms of M. rhomboidea 

subsp rbomboidea); apex of secondary and tertiary lobes attenuate or cuspidate, never 

rounded. 

9. Leaf lobes long (more than 12.0 cm); petiole attachment basal, distal end of capsules 

depressed; fruit dehiscence loculicidal; seeds large, more than 1.25 cm long; erect 

shrubs. 9. M. angustiloba. 

9. Leaf lobes short (less than 12.0 cm); petiole attachment peltate; distal end of capsules 

rounded or projected, never depressed; fruit dehiscence septicidal; seeds small, less than 

1.25 cm long, decumbent or sprawling shrubs. 

10. Petiole attachment narrowly peltate (lamina base less than 0.5 cm wide); staminate 

buds campanulate or slightly constricted in the middle; staminate tepal lobes 

prominently reflexed at anthesis; seeds oblong; sprawling shrubs. 

11. Inflorescence a raceme; secondary leaf lobe apices attenuate; distribution mostly 

south of the Tropic of Cancer. 10. M. rhomboidea. 

11. Inflorescence a subspicate raceme; secondary leaf lobe apices often dilated; 

distribution mostly north of the Tropic of Cancer. 11. M. subspicata.


10. Petiole attachment widely peltate (lamina base more than 0.5 cm wide); staminate 

buds distinctly constricted in the middle; staminate tepal lobes only slightly reflexed 

at anthesis; seed rotund; decumbent shrubs. 12. M. walkerae. 

2. Manihot pringlei Watson, Proc. Am. Acad. 26: 148. 1894; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 45. 1910. 

More or less erect, tall shrubs 3.0-4.0 m tall; dichotomously and trichotomously 

branching. Roots numerous, prominently swollen, elongated, 0.5-1.0 m long, shallow; 

epidermis relatively smooth, brown (5 YR 4/5),6 lenticels oriented along horizontal lines; 

subepidermis light tan, somewhat yellowish; cortex whitish, soft, somewhat pulpy, with a 

very weak smell of HCN. Young stems glabrous, greenish gray-brown; mature stems 

glabrous, light gray, lenticels vertically oriented. Leaves alternate; stipules deciduous, 

glabrous; petioles about 15.0 cm long, occasionally longer, terete, glabrous, greenish, 

reddish (7.5 R 4/11) towards the base, lamina nonpeltate, ventral surface deep green, 

glabrous, dorsal surface pale green, glabrous, abaxial surface wax pattern smooth; 

venation camptodromous, veins on the ventral surface green, glabrous, on the dorsal 

surface reddish (7.5 R 4/11), glabrous, palmately 7 or 9 lobed; median lobes 

oblong-pandurate, rarely oblong, entire lobed, usually about 12.0 cm long, apex 

acuminate, the base of the lobe very narrowly constricted, less than 0.25 cm across, 

lamina between the base of sinus and petiole attachment point less than 0.5 cm wide; 

lowest lobes about half as long as median lobes, more or less similar in shape to median 

lobes. Inflorescence a monoecious, terminal, subspicate/corymbose raceme, about 30.0 

cm in length, the lower half of the rachis usually devoid of flowers, all parts glabrous; 

bracteoles prominently foliaceous, lanceolate, apex accuminate, often as long as 3.0 cm, 

0.5 cm wide, margin laciniate, veins on dorsal and ventral surfaces purplish, bractlets 

foliaceous, purplish veined, often as long as 2.0 cm, 0.3 cm wide. Pistillate flowers restricted 

to the base of the upper half of the inflorescence, borne on long, straight, ascending 

pedicels, held at an angle of ca 45? to the rachis, the lowermost pedicel often 5.0-7.0 cm 

long, subsequent pedicels decreasing gradually in length towards the apical end of the 

inflorescence, creating more or less a corymbose type of inflorescence, pistillate tepals ca 

1.6 cm long, reddish (2.5 R 4/10) externally, greenish yellow (2.5 GY 9/8) internally, 

cleft down to base into 5 lobes, lobes oblong lanceolate, tapering; disc very prominent, 

fleshy; staminodes often present around the disc, pistil ca 0.7 cm long, ovary 

ovate-ellipsoid, the trifid stigma moderately lobed and lobulate. Staminate flowers almost 

tubular, slightly constricted in the middle, tepals ca 1.8 cm long, reddish (2.5 R 4/10) 

externally, greenish yellow (2.5 GY 9/8) internally, cleft nearly 1/3 way down into 5 

lobes, lobes not prominently reflexed at anthesis, disc yellowish (5 Y 9/9), 10 lobed, 

stamens 10, in 2 whorls of 5 each, superior whorl ca 1.3 cm long, inferior whorl ca 1.2 cm 

long. Fruit pedicels long, straight, ascending, 3.0-7.0 cm long; capsules ovate-ellipsoid ca 

1.5 cm long from base to apex, disc at the base very prominent, stigma scar at the apex 

distinctly pointed, fruit surface more or less smooth, perceptibly 6 ribbed, ribs very dark 

maroon (2.5 R 3/7), greenish in between ribs; fruit dehiscence septicidal. Seeds oblong, ca 

1.25 cm long, basal end gently curved inside, caruncle moderately prominent. Seedlings at 

early stages have a characteristic swollen hypocotyl region. Fig 9B, C, D. 

TYPE. Pringle 3558: Mexico, San Luis Potosi: Limestone hills, Las Canoas, 15 Jul 

1890 (syntype, GH). We have not found Las Canoas on recent maps. Efforts to find the 

type locality on a recent trip to Mexico were unsuccessful. 

DISTRIBUTION. (Fig 9A) Mexico: Tamaulipas and San Luis Potosi, along the 

6 Refer to discussion of color designation in Systematic Criteria.


aJJg~~~~~~~~~~~~~~~~~~~~~~~~ 

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ss"\ ' yx tj 

P 

FI distiuin B,-? .Mnhtpige. h a A i (Rgr,Apn&Rgr 2,N),la 

(oes, pa oes58 ?;D nlrsec Rgr,Apn&Rgr 2,N) 

FIG 9. Manibot pringlei. A, distribution; B, habit (Rogers Appan & Rogers 528, NY); C, leaf 

(Rogers, Appan & Rogers 528, NY); D, inflorescence (Rogers, Appan & Rogers 528, NY).


southern and southwestern slopes of Sierra de Tamaulipas and the eastern slopes of Sierra 

Madre Oriental, in a more or less crescent-shaped area bordering the plains surrounding 

Tampico, at 300-2000 m. Growing in dense woods (Crutchfield & Johnston s n) and 

amidst dry scrub vegetation (Webster 11229), invariably in limestone-based soil. Rogers, 

Appan & Rogers 528 was colonizing a disturbed habitat on unconsolidated sandy red 

lateritic soil dumped after road construction in a mixed broad leaf oak forest with Dioon 

edule, Croton sp., Solanum shrubs and scattered palms. This ability to colonize disturbed 

habitats is manifested by most of the species of Manihot. MEXICO. Tamaulipas: Crutchfield 

& Johnston s n (TEX) 3 miles SE of Gomas Farias, 26 Sep 1960; Dressier 1913 (MO) Sierra de 

Tamaulipas, Rancho las Yucas, 20 Jul 1957; Palmer 266 (F-2, NY, UC, US-3) Vicinity of Victoria, 

Feb-Mar 1907; Reiche 1072 (M) S. Victoria, Sep 1926; Rogers, Appan & Rogers 528 (COLO-2, F-3, 

G-3, K-2, M-2, MEXU, MICH-2, MO-2, NY-4, US-3, W-2) 13.5 miles SW of Ciudad Victoria on Road to 

Juamave, 21 Jun 1968; Webster 11229 (DAV) 15 miles SW of Ciudad Victoria, 9 Jun 1962. San Luis 

Potosi: Kenoyer A-198 (F, MICH) Valles 3 Sep 1938; Pringle 3826 (A, F, MICH-2, MO, NY, UC, 

US-2, W) Lime Stone Hills, Las Canoas, 14 Aug 1891; Purpus 5465 (F, MO, NY, UC) Minas de San 

Rafael, May 1911. Province Unknown/Uncertain: Collector Unknown/Uncertain (MO). 

The roots of M. pringlei seem to contain very little HCN. The authors' collecting 

experience indicated almost no odor when the fresh roots (Rogers, Appan & Rogers, 528) 

were dug. Since these plants are found in the same general region and same general 

habitat as the caves reported in archeological digs by MacNeish (1958), where seeds of M. 

dulcis (=M. esculenta) were found, this species may have contributed some genetic 

material to the cultivated complex. The structure of its inflorescence and the size of the 

flowers differentiate M. pringlei very well from all other species in Mexico. 

Cuttings and seeds of this species have been grown successfully 

at the Fairchild 

Tropical Gardens, Miami, Florida. Reports that this species had been introduced into 

Madagascar are apparently unfounded. 

3. Manihot auriculata McVaugh, Brittonia 13: 190. 1961. 

Arborescent shrubs/low trees, ca 6.0 m tall, diam at base ca 6.0 cm, with long weak 

trailing branches. Stems glabrous, grayish brown. Leaves alternate; stipules deciduous, 

glabrous; petioles 15.0-20.0 cm long, terete, glabrous; lamina nonpeltate, dorsal and 

ventral lamina surfaces glabrous, abaxial surface wax pattern smooth; venation camptodromous, 

midribs glabrous; palmately 5 lobed, 3 major and 2 slightly smaller lobes; 

median lobes oblong, ca 14.0 cm long and ca 5.0 cm wide, margin broadly sinuate, apex 

acuminate; lowest lobes ca half as long as median lobes, slightly reflexed downwards. 

Inflorescence a long raceme, monoecious, ca 15.0 cm in length, all parts glabrous; 

bracteoles setaceous, margin smooth; bractlets setaceous. Pistillate flowers restricted to 

the base of the inflorescence, borne on short (ca 1.0 cm long) pedicels, tepals and pistils 

not seen. Staminate buds ovoid-ellipsoid, tepals pale yellowish, ca 1.7 cm long, 5 lobed, 

lobes auriculate at sinus, disc 10 lobed; stamens 10, in 2 whorls of 5 each, longer ones ca 

1.3 cm in length, shorter ones ca 0.8 cm, anthers 0.30-0.35 cm long. Fruits and seeds non 

vidi. Fig 10B, C. 

TYPE. McVaugh 15283: Mexico, Nayarit: Mirador del Aguila, ca 14 miles N of Tepic 

10 Jul 1957 (holotype, MICH). 

DISTRIBUTION. (Fig 10A). Mexico, Nayarit, western slopes of Sierra Madre 

Occidental, alt ca 500 m. Lower forested slopes of steep, rocky ravine. 

This species is characterized by its unique auriculate staminate tepal, a feature not 

occurring in any other Manibot species. 

4. Manihot aesculifolia (Humbolt, Bonpland & Kunth) Pohl, P1. Bras. Ic. et Descr. 1: 55. 

1827; Muell.-Arg. in DC., Prodr. 15(2): 1065. 1866; Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 58. 1910; Standley, Contr. U. S. Nat. Herb. 23: 645. 1923; Croizat, 

Jour. Arnold Arbr. 23: 218. 1942.


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Systematic 


Janipha aesculifolia Humboldt, Bonpland & Kunth, Nov. Gen. & Sp. 2: 85. PI. 109. 1817; 

Jatropha aesculifolia Steudel, Nomencl. ed. 2.1: 799. 1840. 

Manihot intermedia Weatherby, Proc. Am. Acad. 45: 427. 1910. (Contr. Gray Herb. 2: 427 

1910); Pax in Engler, Pflanzenreich IV. 147(Heft 44): 101. 1910; Standley, Contr. U. S. Nat. 

Herb. 23: 643. 1923; Croizat, Jour. Arnold Arb. 23: 221. 1942. Type. Pringle 13938 

(syntypes, ARIZ-2, F, GH, MICH, MO, UC, US). 

Manihot olfersiana Pax, Pflanzenreich IV. 147(Heft 44): 55. 1910; McVaugh, Brittonia 13: 193. 

1961. Type. Ubde 1176 nv. 

Manihot gualanensis Blake, Contr. U. S. Nat. Herb. 24: 13. 1922. Type. Blake 7688 (holotype 

US, isotypes F, US). 

Erect tall shrubs to 7.0 m, trunk at base ca 10.0 cm in diam. Roots prominently 

swollen, large; epidermis brown, rough and flaky; subepidermis creamy white. Young 

stems green, glabrous, internally white; mature stems grayish brown, glabrous; all parts 

with white latex. Young apical foliage yellowish green to dark reddish purple (10 P 3/9). 

Leaves alternate; stipules deciduous, glabrous; petioles ca 20.0 cm long, terete, glabrous, 

pure green or with various degrees of pigmentation such as green with bluish cast at base 

and apex, moderately reddish (2.5 R 4/10) throughout, or dark red throughout; lamina 

nonpeltate, silvery beneath, deep green above (2.5 G 5/9), abaxial surface wax pattern 

verrucate with a characteristic microscopic pulverulence; venation camptodromous; 

palmately 5-9 lobed; median lobes 15.0-30.0 cm in length, 4.0-10.0 cm in width, 

oblong, entire lobed, or oblong-pandurate, the primary constriction of pandurate lobes 

differing considerably in depth and width, apex acute to acuminate, bristle pointed, base 

of lobe usually 0.75-2.0 cm across, rarely less; lowest lobes ca half as long, and more or 

less similar in shape, to median lobes. Inflorescence monoecious, usually a large profusely 

branched terminal panicle, often as long as 45.0 cm, occasionally shorter and less 

profusely branched, all parts glabrous; bracteoles setaceous or semifoliaceous, in the latter 

case as long as 2.0 cm, linear, margin smooth; bractlets setaceous. Pistillate flowers not 

restricted to the base of the inflorescence but occurring from the base up to almost the 

apex, interspersed with staminate flowers, tepals ca 1.2 cm long, cleft down to base into 5 

lobes, lobes lanceolate-oblong, ca 0.4 cm wide; disc fleshy, entire, pistil ca 0.6 cm long, 

ovary subglobose, glabrous, the trifid stigma much lobed and lobulate. Staminate flowers 

campanulate, often pendulous, tepals ca 1.3 cm long, color varies from pure greenish 

yellow to various degrees of purple pigmentation, different types occurring together often 

in the same local population, cleft 1/3 way down into 5 lobes, lobes oblong, obtuse, not 

prominently reflexed at anthesis; disc 10 lobed, depressed and fleshy; stamens 10, in 2 

whorls of 5 each, superior whorl ca 0.9 cm long, inferior whorl ca 0.7 cm long. 

Infructescence usually a long cluster of fruits, characteristic of this species, fruit pedicles 

ca 1.0 cm long, slightly curved downwards; capsules almost globular, ca 1.5 cm long from 

base to apex, surface green, slightly tuberculate, without ribs, apex rounded; fruit 

dehiscence septicidal. Seeds oblong, ca 1.25 cm long, relatively thin and flat, dorsal side 

with conspicuous horizontal stripes; caruncle moderately prominent, broad. Fig 10-D; 

12-A, B. 

TYPE. Humboldt & Bonpland s n: Mexico, Campeche: Campeche (photo of syntype, 

P). 

DISTRIBUTION. (Fig 11). Mexico: Sinaloa, Nayarit, Jalisco, Colima, Michoacan, 

Guerrero, Mexico, Vera Cruz, Oaxaca, Chiapas, Yucatan and Quintana Roo Territory; 

British Honduras; Guatemala; Honduras; El Salvador; Nicaragua; Costa Rica and Panama. 

Occurring in the interior region of the west and east facing coastal belts, usually along the 

foothills and lower slopes of the mountains, to ca 1300 m, rarely in the plains. In the 

Yucatan area it flourishes on steep banks of cenotes (limestone sinks with water at 

bottom, Karst topography), localized and not spreading far beyond the cenote margins. 

Towards the north with lower rainfall, it grows in dry thorn forest vegetation dominated 

by Acacia, Cnidoscolus, Coccoloba, Randia, etc.; or at the edge of savannas in grassy

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second growth thickets. MEXICO. Sinaloa: Brandegee s n (GH, UC) Culiacan, 30 Aug 1904; 

Gentry 6101 (ARIZ, MICH, MO, NY) Palmar, 50-70 miles N of Guamuchil, 22 Aug 1941; Palmer 

1536 (F, NY, US-2) Culiacan, Aug-Sep 1891; Rogers 520 (BM, COLO, F, MEXU, MICH, MO, US) 18 

km N of Culiacan, 10 Jul 1966; Rogers 521 (COLO, F-2, G-2, K, MEXU, MO, NY-3, US, W) 30 miles 

N of Mazatlan, 11 Jul 1966; Rose 1611 (GH) between Rosario and Colomas, 12 Jul 1897; Rose 3204 

(GH) Foot Hills of Sierra Madre, near Colomas, Jul 1897. Nayarit: Feddema 1007 (DS, US) Circa 15 

km SE of San Bias, Along Road to Miramar, 26 Aug 1959;McVaugh 15355 (MICH) 2 miles SE of Las 

Varas, Road to Mazatlan, 12 Jul 1957. Jalisco: Rogers 510 (A, COLO, F-2, G, M, MEXU, MICH, MO, 

NY-3, S, US-2, W) 16 km SW of Autlan, 29 Jul 1963; Rogers 515 (G, K, MICH, NY, US-2) Circa 22 

km W of Tecaltitlan, 31 Jul 1963. Colima: Jancey 334 (A, ARIZ, MEXU, NY, P, SP, UC) 3 miles N of 

Manzanillo, Jul 1965; Jancey 335 (ARIZ, BM, DAV, MEXU) 6 miles N of Manzanillo, Jul 1965. 

Michoacan: Hinton 13972 (GH, NA, NY, US) Coalcoman, 21 Jul 1939. Mexico: Hinton 1207 (BM, 

MO) Chorrera, 30 Jul 1932; Hinton 4374 (BM, GH) Ypericones, 26 Jul 1933; Hinton 4555 (BM, GH) 

Bejucos, 19 Aug 1933. Vera Cruz: Purpus 6112 (UC) Banos Del Carrizal, Aug 1912; Purpus 8448 (GH, 

NY, UC, US) Baranca de Panoaya, Jul 1919; Purpus 8484 (MO-2) Baranca de Panoaya, Jul 1919. 

Guerrero: Crisman & Willis 206 (TEX) 5 miles N of Ocotito, 13 Jun 1954; Hinton 10376 (F, GH, NY, 

UC, US) Cundanchico, 2 Jul 1937; Hinton et al 6264 (BM, GH) Pungarabato, 9 Jul 1934; Hinton et al 

6468 (BM, F, MO, NY, US) Pungarabato, Coyuca, 20 Aug 1934; Mexia 8778 (ARIZ, F, MICH, MO-2, 

NY, U, UC-2, US) Temisco, Barranca de La Suriana, 9 Nov 1937; Pringle 13938 (ARIZ-2, F, GH, 

MICH, MO, UC, US) Iguala Canyon, 29 Jul 1907; Rusby 2 (NY, US) Limon Mountain, 28 Jul 1910. 

Oaxaca: Nelson 2852 (F, US-2) between Topana, Oaxaca and Tonala, Chiapas, Jul-Aug 1895; Williams 

9867 (F, MICH) Almoloya, Jul 1937. Chiapas: Jancey 327 (A-2, S-2, SP) 20 miles S of Tonala, Jul 

1965; Jancey 328 (NY) 20 miles S of Tonala, Jul 1965; Jancey 329 (NY) 20 miles S of Tonala, Jul 

1965; Laughlin 1125 (NY) El Chorreadero, 21 Jun 1966; Matuda 16375 (F, MICH) Cacaluta, 

Escuintla, 14 Jun 1947; Matuda 4813 (F, NY) between Mazapa and Motozintla, 19 Jul 1941. 

Yucatan: Gaumer 1142 (BM, F-2, GH, MICH, MO, NY) Progresso; Gaumer & Sons 23394 (F, MO) 

Orchidia, Dec 1916; Jancey 320 (NY) 10 miles NE of Piste, Jun 1965; Jancey 321 (DAV-2, MEXU) 

10 miles NE of Piste, Jun 1965; Jancey 322 (ARIZ, MEXU, NY) 10 miles NE of Piste, Jun 1965; 

Jancey 323 (SP, UC) 10 miles NE of Piste, Jun 1965;Jancey 324 (M, MICH, P) 10 miles NE of Piste, 

Jun 1965; Jancey 325 (NY) 10 miles NE of Piste, Jun 1965; Jancey 326 (BM, COLO, F, G, K, MO, 

US, W) 10 miles NE of Piste, Jun 1965; Lundell & Lundell 7473 (A, DS, F, MICH, US) Chichen Itza, 

off Kaua Road, Jun-Jul 1938; Novelo 114 (NA) Merida, Aug 1939; Rogers 504 (A, BM, F-2, G-2, K, 

M, MO, NY-3, P, S, UC-2, US, W-2) E Edge of Valladolid, Steep Banks of Cenote, 18 Jul 1963; Schott 

518 (BM, F, MO, US-2) Merida, Jun-Jul 1865; Steere 1673 (F, MICH) Chichen Itza, 29 Jun 1932; 

Steere 1947 (F, MICH) Champoton, Campeche, Jul 1932. Quintana Roo Territory: Lundell & Lundell 

7737 (MICH) Quintana Roo, Coba, Jun-Jul 1938; Lundell & Lundell 7848 (MICH) Quintana Roo, 

Coba, Along Dzitnup Trail, Jun-Jul 1938. Province Unknown/Uncertain: Galeotti 3735 (P) Torullo, 

1840. GUATEMALA.,Huehuetenango: Steyermark 51316 (F-2, US) Trail between Democracia and 

Santa Ana, 25 Aug 1942. Zacapa: Standley 73669 (F) Vicinity of Zacapa, Oct 1940; Standley 73683 

(F) Vicinity of Zacapa, Oct 1940; Standley 73783 (F) between Zacapa and Chiquimula, 9 Oct 1940. 

Chiquimula: Standley 73736 (F) near Divide on Road from Zacapa to Chiquimula, 9 Oct 1940; 

Standley 74305 (F) Quebrada Shusho, 14 Oct 1940. El Quiche: Aguilar 1373 (F), 1942. Izabal: Blake 

7688 (F, US-2) Gualan, 26 May 1919. Suchitepequez: Steyermark 47778 (F, NY) S of Alotenango, 19 

Jun 1942; Steyermark 47779 (F) S of Alotenango, 19 Jun 1942. Santa Rosa: Standley 79452 (F) S of 

Guazacapan, 6 Dec 1940. Province Unknown/Uncertain: Cook & Doyle 311 (US), 6 Jun 1904. 

BRITISH HONDURAS. El Cayo: Gentle 2539 (A, F, MICH) Vaca, 1 May 1938. HONDURAS. Santa 

Barbara: Thieme 5473 (US) San Pedro Sula, Jun 1887. Cortes: Bangham 300 (A, F) San Pedro Sula, 

10 Aug 1929; Johansen 38 (F, US) La Lima, 31 Aug 1929. EL SALVADOR. San Salvador: Calderon 

776 (GH, NY, US) La Chaera, San Salvador, 1922. Province Unknown/Uncertain: Choussy 73 (US) 

Finca San Nicolas, 1923. NICARAGUA. Chinandega: Maxon, Harvey & Valentine 7107 (US-2) 

Ameya, Jun 1923; Maxon, Harvey & Valentine 7192 (NY, US-3) Ameya, Jun 1923;Standley 11481 

(F, US) vicinity of Chichigalpa, Jul 1947. Jinotega: Standley 9657 (F) vicinity of Jinotega, Jun-Jul 

1947. Managua: Gamier s n (F) Sierra de Managua. Granada: Levy 41 (P-2) environs of Granada, Jun 

1869. Zelaya: Standley 19836 (F) vicinity of El Recreo, Apr-May 1949. COSTA RICA. Guanacaste: 

Cook & Doyle 648 (US), 22 May 1903; Tonduz 13977 (BM, US) dans Les Broussailes a Nicoya, May 

1900. Puntarenas: Holm & Iltis 250 (BM, F, MO, P) vicinity of Cascajal, 5 Jul 1949. Province 

Unknown/Uncertain: Brenes 3993 (F) San Ramon, Rio Cacao, 24 Jun 1924; Echeverria 4158 (F, UC), 

Jul 1945; Pittier & Durand 6579 (US). PANAMA. Canal Zone: Tyson 4108 (MO) Farfan Beach area, 

29 May 1966. Province Unknown/Uncertain: Hayes 717 (BM, K) common about Paraiso, 4 Jun 1862. 

LOCAL NAMES. Yuca sylvestre. (Tonduz 13977) Yuca de monte (Tonduz 13977); 

Yuca (Standley 11481); Yuca cimarrona (Maxon, Harvey & Valentine 7107); Yquilla 

(Steyermark 51316); Chichput (Lundell & Lundell 7737); Batul (Novelo 114); Tacote 

(Rogers 510).


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FIG 12. Manihot aesculifolia. A, leaf (Rogers 504, NY); B, inflorescence (Rogers 515, NY). M. 

rubricaulis. C, distribution-dots represent subsp isoloba and the letter E represents subsp rubricaulis; 

D, leaf of subsp rubricalus (Palmer 224, US).


USES. Some collections such as Steyermark 51316 and Lundell & Lundell 7737 are 

reported to have edible roots. 

Pohl correctly indicated the relations of this species to Manihot, and transferred it 

from Janipha, where it was originally described by H.B.K. Later synonyms are mere 

failures to understand the wide-ranging ecotypic variability of the species. Many 

misidentifications of specimens have been made, most frequently confusing M, 

aesculifolia with M. carthaginensis, from which this species differs in the much stouter 

inflorescences in M. aesculifolia, the larger general dimensions, and in distribution. M. 

carthaginensis is confined to the relatively dry coastal regions of northern South America. 

Manibot aesculifolia is the closest wild relative of M. esculenta. Its varied, 

wide-ranging habitats, general morphology, and complex ecotypic variability make its 

relation to M. esculenta most striking. It is most easily differentiated from M. esculenta 

by the verrucate abaxial lamina condition, by its stout inflorescence, large numbers of 

flowers per inflorescence, the larger size of the staminate flowers, and by its relatively 

smooth stem without swollen nodal regions. 

Plant breeders should find M. aesculifolia a rich source of genetic material for a 

variety of purposes. They should be careful to recall, however, the ecotypic variability of 

the species in selection of their material. 

5. Manihot rubricaulis I. M. Johnston, Contr. Gray Herb. 68: 90. 1923; Croizat, Jour. 

Arnold Arb. 23: 222. 1942. 

Erect shrubs, 1.0-5.0 m tall with many slender stems arising from a woody base; 

foliage borne at the tip of the twigs. Young stems glaucous; mature stems glaucous, 

reddish brown; latex present. Leaves alternate; stipules deciduous, glabrous; petioles 

6.0-12.0 cm long, occasionally longer, glabrous, terete; lamina nonpeltate, deep green or 

ashy blue-green in color, dorsal and ventral lamina surfaces and midribs glabrous, abaxial 

surface wax pattern smooth; venation camptodromous; palmately 5-9 lobed; median 

lobes 6.0-10.0 cm long, occasionally as long as 17.0 cm, 0.75-2.0 cm wide, narrowly 

linear with almost parallel sides, occasionally a shallow constriction present near the apex, 

apex acute to acuminate, bristle tipped, sinuses between lobes prominently calloused, 

giving a "rooster's foot" appearance to leaves; lowest lobes about half as long as median 

lobes with a characteristic 'S' shaped bend. Inflorescence a monoecious, axillary, 

few-flowered, panicle with sparse branching, sometimes raceme-like 5.0-7.0 cm long, 

occasionally as long as 15.0 cm, all parts glabrous; bracteoles setaceous, or semifoliaceous, 

in the latter case, to 1.25 cm in length, margin smooth; bractlets setaceous. Pistillate 

flowers restricted to the base of the inflorescence, borne usually on branched peduncles, 

pedicels 2.0-4.0 cm long, tepals yellowish green, sometimes with an ashy blue-green 

tinge, 0.8-1.2 cm long, cleft to the base into 5 lobes, lobes 0.3-0.4 cm wide, 

lanceolate-oblong, disc fleshy, entire, pistils 0.4-0.6 cm long, ovary subglobose, glabrous, 

the 3 divisions of the stigma moderately lobed and lobulate. Staminate flowers 

campanulate, tepals greenish yellow, sometimes with an ashy blue-green tinge, 0.9-1.3 cm 

long, cleft 1/3 way down into 5 lobes, lobes oblong, obtuse, not prominently reflexed at 

anthesis, disc fleshy, 10 lobed; stamens 10, in 2 whorls of 5 each, the longer ones ca 0.9 

cm long, shorter ones 0.7 cm long, filaments and anthers cream colored. Fruit pedicels 

2.0-4.0 cm long, almost straight; capsules ovate-ellipsoid, 1.2-1.75 cm long from base to 

apex, surface slightly tuberculate, ribs not prominent, apex rounded, fruit dehiscence 

septicidal. Seeds oblong, 0.9-1.2 cm long, surface finely pitted; caruncle moderately 

prominent, trapeziform. 

DISTRIBUTION. (Fig 12C). Mexico, States of Sonora, Chihuahua, Sinaloa and 

Durango. The plants of this species are sporadically distributed from Durango north and 

west in the high, relatively dry ranges of the Sierra Madres Occidental. It apparently sheds


its leaves during the dry periods, and some of the younger stems may die back during the 

same periods. Its altitudinal distribution, one of the highest of any species of the genus, 

makes it a potential donor of useful genetic materials for breeders interested in increasing 

the tolerance of Manihot esculenta to drought and cool temperatures. 

Key to the Subspecies of Manihot rubricaulis 

1. Fruits smaller, less than 1.5 cm in length; seeds shorter, less than 1.0 cm in length. 

5a. M. rubricaulis subsp rubricaulis. 

1. Fruits larger, more than 1.5 cm in length; seeds longer, more than 1.0 cm in length. 

5b. M. rubricaulis subsp isoloba. 

5a. Manihot rubricaulis I.M. Johnston subsp rubricaulis 

Manihot rubricaulis I. M. Johnston, Contr. Gray Herb. 68: 90. 1923; Croizat, Jour. Arnold Arb. 

23: 222. 1942. 

This subsp differs from subsp isoloba with respect to the size of fruits and seeds. The 

fruits are relatively smaller, less than 1.5 cm in length from base to apex. The seeds are 

also shorter, less than 1.0 cm in length. The leaf lobes are generally narrower, mostly less 

than 1.0 cm in width. Fig 12D; 13A, B. 

TYPE. Palmer 224: Mexico, Durango: City of Durango and vicinity (holotype, GH; 

isotypes, (BM, F, GH, NY, UC, US-2). 

DISTRIBUTION. (Fig 12C). Mexico, States of Sinaloa and Durango. MEXICO. Sinaloa: 

Brandegee s n (UC) vicinity of Culiacan, Cerro Colorado, 1 Nov 1904. 

5b. Manihot rubricaulis I. M. Johnston subsp isoloba (Standley) Rogers & Appan, stat 

nov 

Manihot isoloba Standley, Publ. Field Mus. Bot. 17: 197. 1937; Croizat, Jour. Arnold Arb. 23: 

223. 1942. 

This subspecies differs from the subsp rubricaulis with respect to the size of fruits 

and seeds. The fruits are relatively larger, more than 1.5 cm in length (from base to apex). 

The seeds are also bigger, over 1.0 cm in length. The leaf lobes are generally wider, mostly 

not less than 1.0 cm in width. Fig 13C, D. 

TYPE. Gentry 2372: Mexico, Chihuahua: Guasaremos, Rio Mayo 10 Aug 1936 

(holotype, F; isotypes, ARIZ, MO, UC, US). 

DISTRIBUTION. (Fig 12C). Mexico, States of Sonora, Chihuahua and Sinaloa. Alt 

1000 to 2750 m. MEXICO. Sonora: Gentry 1468 (ARIZ, F, MO, UC, US) Bakachaka, Rio Mayo, 6 

Jul 1935; Muller 3696 (NA, UC) Nacore Chico, Slopes above El Rio Bonito, 7 Oct 1939; Shreve 

6753A (ARIZ, MICH) Santa Margarita Mountains, 6 miles E of Mina Verde, 23 Sep 1934; White 2758 

(MICH) Cation de Los Apaches, N of Aribabi, 6 Sep 1939; White 2930 (GH, MICH, NA) La Palmita 

between Granados and Bacadehuachi, 7 Jul 1940; White 3103 (ARIZ, GH, MICH) Cation de Bavispe, 

Jul-Aug 1940; White 3647 (DS, GH, MICH) Piedra Parada, Sep 1940. Chihauhua: Gentry, Correl & 

Arguelles 18083 (US) Rio San Juan Valley, 29 Oct 1959; Jones s n (POM-2) Guayanopa Canyon, 

Sierra Madre Mountains, 24 Sep 1903; Knobloch 1335 (MICH) vicinity of Areponapuchie, Slopes of 

Barranca de Urique, 23 Aug 1954; Lesueur 1249 (F, GH, MO, TEX, UC, US) Rio Aros, 6 Aug 1937; 

Townsend & Barber 404 (BM, F-2, NY) Sierra Madres, near Seven Star Mine, 5 Sep 1899. Sinaloa: 

Gentry 6583 (ARIZ, GH, MICH, MO, NY) Quebrado de Mansana, Sierra Surotato, Sep 1941. 

VERNACULAR NAME. pato de gallo (Gentry 2372). 

6. Manihot oaxacana Rogers & Appan, sp nov. 

Frutices 2.0-5.0 m altae, glabrae; folia septemloba, lobus medianus ca 10.0 cm 

longus, ca 4.0 cm latis; inflorescentia racemosa, brevis, ca 5.0 cm longa; flores staminati 

conici; capsulae ca 1.5 cm longae; semina ca 0.9 cm longa. 

Arborescent shrubs, 2.0-5.0 m tall, trunk diam ca 10.0 cm, branching begins above


.- 3 

r 

'.t ....~ . . ......... 

FIG 13. Manihot Manibot rubricaulis subsp rubricaulis. A, basal lobe of leaf showing the characteristic 

'S' shaped curve (Palmer 224, UC); B, inflorescence (Palmer 224, UC). M. rubricaulis subsp isoloba. C, 

leaf (Townsend & Barber 404, BM); D, inflorescence (Gentry 1468, ARIZ).


the middle, dichotomously and trichotomously branching, branches with a tendency to 

spread out. All parts of the plant with strong odor of HCN. Roots slightly swollen; 

epidermis rough, strong brown (5 YR 4/5) subepidermis white. Young stems glabrous, 

strong brown (5 YR 4/5), internally light yellow-green (7.5 GY 9/4); mature stems 

brownish gray; latex cream colored. Young foliage at apex strong yellow-green (7.5 GY 

6/8). Leaves alternate, stipules deciduous, glabrous; petioles generally 10.0-15.0 cm long, 

terete, dull red-green with some olive shading; lamina nonpeltate, ventral surface deep 

green, glabrous; dorsal surface pale green, glabrous, midribs and major veins prominent, 

abaxial surface wax pattern reticulate; venation camptodromous; palmately 7 lobed, 3 

major, 2 smaller, and often 2 minute lobes; median lobes usually ca 10.0 cm long and ca 

4.0 cm wide, obovate-pandurate, occasionally obovate, entire lobed, the primary 

constriction of pandurate lobes shallow to deep, apex varies from almost truncate with an 

acute conical tip in the middle, to acuminate. Plants begin to flower when ca 1.0 m tall; 

inflorescence a short, moneocious, axillary raceme, usually ca 5.0 cm long, never longer 

than 8.0 cm, pedicels comparatively longer at base, gradually decreasing in length towards 

apex, giving the inflorescence a more or less conical shape, all parts glabrous; bracteoles 

setaceous, less than 0.3 cm in length, margin smooth; bractlets setaceous. Pistillate 

flowers restricted to the base of the inflorescence, borne on 1.0-2.0 cm long pedicles, 

tepals externally brilliant yellow-green (2.5 GY 9/8), cleft to the base into 5 lobes, lobes 

lanceolate-oblong, disc fleshy, entire, brilliant yellow-green (2.5 GY 9/8), ovary 

subglobose, entire, the three divisions of the stigma moderately lobed and lobulate. 

Staminate flowers characteristically conical, often as long as 1.8 cm, tepals brilliant 

yellowish green externally (2.5 GY 9/8), cleft 1/3 of the way down into 5 lobes, lobes 

tapering, acuminate, disc 10 lobed, brilliant yellowish green (2.5 GY 9/8), stamens 10, in 

2 whorls of 5 each, filaments and anthers white. Fruit pedicels 1.0-2.0 cm long, slightly 

curved downwards; capsules more or less globular, ca 1.5 cm long from base to apex, 

surface slightly rugose, with 6 very small wings, apex rounded, fruit dehiscence septicidal. 

Seeds oblong, ca 0.9 cm long, carunculate and bluntly acute, caruncle moderately 

prominent. Fig 14B, C, D. 

TYPE. Rogers 505: Mexico, Oaxaca: 17 km ENE of Juchitan, 22 Jul 1963 

(holotype, NY; isotypes, COLO, F, G, K, MEXU, MO, P, S, SP, UC, US, W). 

DISTRIBUTION. (Fig 14A). Mexico: Oaxaca. Flourishes in the plains and foothills 

surrounding Juchitan and Tehuantepec, and upstream on both sides of Rio Tehuantepec; 

to 150 m. It appears that the range of this species extends upwards as high as 1000 m, 

towards the headwaters of Rio Tehuantepec. Webster, Miller & Miller 11641, collected at 

an alt of 870 m, may represent an ecotypic form of this taxon, but the poor condition of 

the specimen does not render critical comparisons possible. Similarly Conzatti 2413, 

collected from Quiotepec, Cuicatlan at an alt of 700 m resembles M. oaxacana, but 

positive identification has not been possible due to the poor condition of the specimen. 

Further collections from these areas will confirm the possibility of the range of this 

species extending along the northeastern foothills of the Sierra Madre del Sur, up to 

Cuicatlan which is on the periphery of the range of Manibotoides pauciflora. 

In the typical habitat along the foothills near Juchitan and Tehuantepec, large 

self-seeding colonies are common. Manihot oaxacana grows under partial shade in a 

relatively undisturbed vegetation composed of thorny leguminous trees, shrubs, cacti, etc. 

Appears to be adapted to grow in several soil types such as sandy (King 1213), loose 

limestone with coarse friable reddish brown soil mantle (Rogers 507), metamorphic rock 

(Webster, Miller & Miller 12955), and lateritic soil on noncalcareous rocks (Webster, 

Miller & Miller 11641). MEXICO. Oaxaca: Conzatti 2413 (F) Quiotepec, Cuicatlan, 21 Jun 1909; 

King 1213 (NY, TEX, UC, US) Isthmus of Tehuantepec, 2-4 km E of Tehuantepec, 1 Jul 1959; King 

366 (US) Isthmus of Tehuantepec, 5.5 km E of Juchitan, 2 Jul 1958; King 737 (US) Isthmus of 

Tehuantepec, 10-15 miles NW of Tehuantepec, 26 Jul 1958; Purpus 7353 (UC) San Geronimo, Jul 

1914; Rogers 506 (G, M, MICH, NY, S) 7-8 km WNW of Tehuantepec, 23 Jul 1963; Rogers 507


ir t 

NY); D, inflorescence (Rogers 505, NY).


(COLO-2, F-2, G, K, MEXU, MO, P, NY-2, US-3, W) 10 km WNW of Tehuantepec, 23 Jul 1963; 

Webster, Miller & Miller 11641 (DAV) 70 miles SE of Oaxaca, 22 Jun 1962; Webster, Miller & Miller 

12955 (DAV) 12 miles E of Juchitan, 11 Aug 1962; Webster, Miller & Miller 13007 (DAV) 8 miles 

W of Tehuantepec, 12 Aug 1962. 

This species was confused in the past with Manihot chlorosticta. The former is an 

arborescent shrub, while the latter is a clambering vine. There are several other features 

which characterize M. oaxacana such as the very short raceme, conical brilliant yellow 

flowers, smaller fruits and seeds, etc. Besides, the geographical ranges of these two species 

are disjunct and consequently there is little likelihood of interbreeding. This Oaxaca 

population represents a closed gene pool in itself and as such a distinct biological species. 

Rogers (1965) discovered and defined this population as an undescribed species, based on 

field study observations. The specific epithet denotes the distribution which is confined 

only to the state of Oaxaca. 

In the computer analysis Manihot oaxacana cluster joins M. chlorosticta cluster at a 

"C-value" of .857 and soon both the clusters integrate tightly. The general ecological 

habitat of both these species is very similar, both flourishing along the coastal belt. All 

these suggest that these two species are very closely related. Without cytogenetic data it is 

unsound to hypothesize; nevertheless, it seems obvious that M. oaxacana and M. chlorosticta 

once belonged to the same ancestral gene pool, eventually splitting and diverging 

into two closed gene pools. The M. oaxacana population, occupying a pocket confined by 

mountain barriers, is strongly suggestive of the possibility of a segment of the ancestral 

gene pool being restrained to evolve independently in isolation. 

7. Manihot chlorosticta Standley & Goldman, Contr. U. S. Nat. Herb. 13: 375. 1911. 

Manibot colimensis Croizat, Jour Arnold Arb. 23: 221. 1942. Type. Holotype. Ferris 6140 (A), 

isotypes (F, US). 

Manihot mobilis Standley, Am. Midi. Nat. 36: 177. 1946 Type. Holotype Leavenworth & 

Hoogstraal 1556 (F). 

Clambering vines or vinelike decumbent shrubs, to 5.0 m, the trunk at the base about 

10.0 cm in diam. soon dividing into slender, smooth, long branches ca 3.0 m in length, 

forming rather heavy masses on other vegetation. Young stems greenish gray, glabrous; 

mature stems grayish with reddish brown tinge, glabrous; latex present. Young foliage at 

apex yellowish green. Leaves alternate; stipules deciduous, glabrous; petioles ca 8.0 cm 

long, terete, glabrous, dark reddish; lamina nonpeltate, deep green above, pale green 

beneath, abaxial surface wax pattern reticulate; venation camptodromous; palmately 5-7 

lobed, 3 major, 2 smaller, often 2 more minute lobes; median lobes with considerable 

variation in size and shape, length from 5.0-12.0 cm, width 2.0-6.0 cm, obovate entire 

lobed or obovate pandurate, the primary constriction of pandurate lobes varying in 

depth, width, and sinuosity, apex acute to more or less acuminate; lowest lobes ca 1/4 as 

long as median lobes. Inflorescence a monoecious, few-flowered raceme, mostly axillary, 

ca 8.0 cm in length; all parts glabrous; bracteoles nonfoliaceous, less than 0.3 cm in 

length, purplish, margin smooth; bractlets nonfoliaceous. Pistillate flowers restricted to 

the base of the inflorescence; pedicels usually long, up to 5.0 cm, tepals lemon-yellow, 

without any purplish pigmentation, cleft to the base into 5 lobes, lobes oblong-lanceolate, 

ca 1.0 cm long and ca 0.3 cm wide; disc fleshy, entire, lemon-yellow; pistil about 0.6 cm 

long; ovary subglobose, glabrous, the three divisions of the stigma moderately lobed and 

lobulate, cream colored. Staminate flowers campanulate, tepals ca 1.4 cm long, 

lemon-yellow without any purplish pigmentation, cleft 1/3 way down into 5 lobes, lobes 

oblong-obtuse, not prominently reflexed at anthesis; disc fleshy, 10 lobed, lemon-yellow, 

stamens 10, in 2 whorls of 5 each, longer whorl ca 1.0 cm long, shorter whorl ca 0.8 cm 

long, filaments and anthers cream colored. Fruit pedicels usually long, to 5.0 cm, robust, 

very slightly curved. Capsules ovate ellipsoid, 1.25-1.75 cm long from base to apex,


surface coarsely tuberculate, perceptibly 6 ribbed, apex slightly pointed, fruit dehiscence 

septicidal. Seeds oblong, 1.25-1.5 cm long, caruncle moderately prominent, trapeziform. 

Fig 15B, C, D. 

TYPE. Nelson & Goldman 7401: Mexico, Baja California: San Jose del Cabo, 6 Jan 

1906 (holotype, US; Photo of type, F). 

DISTRIBUTION. (Fig 15A). Mexico, states of Baja California, Sinaloa, Jalisco, 

Colima, Michoacan and Guerrero, along the coastal belt between the sea and the 

mountain ranges; 0-250 m, occasionally up to 500 m; on cliffs facing ocean; between the 

beach and coastal lagoons; in heavy clay soils of coastal plains in thorn forest vegetation; 

etc. The range of this species is parallel to that of M. aesculifolia, along the west coast, 

and it appears that M. chlorosticta is abundant towards the ocean, and M. aesculifolia 

flourishing at the back of the coastal belt towards the mountains; the two overlapping in 

the middle common territory. MEXICO. Baja California: Brandegee s n (UC) San Jose del Cabo, 

Nov 1902; Brandegee 550 (GH, UC, US) San Jose del Cabo, Sep 1890; Grabendorfer 550 (UC) San 

Jose del Cabo, 1899; Hastings & Turner 64-355 (UC) 33 miles N of Cabo San Lucas, 17 Oct 1964; 

Purpus 517 (UC) San Jose del Cabo, Jan-Feb 1901. Sinaloa: Ferris & Mexia 5218 (A) Vicinity of 

Labrades, 21 Sep 1925; Gentry 5026 (ARIZ, MICH, MO, NA, NY) Cerro E of Culiacan, 23 Nov 1939; 

Gentry 7085 (A, F, NY, UC, US) Maraton, 12 miles W of Culiacan, 21 Sep 1944; Ortega 6345 (GH, 

UC) Mazatlan, Villa Union, Escamillas, Aug 1926; Ortega 7024 (F), 1933; Ortega 7273 (F), 1934; 

Ortega 6488 (DS) Mazatlan; Rose 3266 (US) between Rosario and Concepcion, 28 Jul 1897. Jalisco: 

McVaugh & Koelz 1695 (MICH, US) near Playa de Cuastecomate, Dec 1959. Colima: Emrich 140 (F) 

Tecoman, Nov 1906; Emrich 156 (F) Tecoman, Nov 1906; Ferris 6140 (A, F, US) vicinity of 

Manzanillo, 28 Nov 1925; Gregory & Eiten 321 (MICH, MO, NY) ca 5 miles WNW of Manzanillo, 11 

Jul 1956; McVaugh 15638 (MICH, TEX) 5 miles NW of Manzanillo, 22 Jul 1957; McVaugh 15875 

(MICH) Bahia de Santiago, 29 Jul 1957;McVaugh & Koelz 1553 (MICH, US) 11 miles SSW of Colima 

on Manzanillo Road, Dec 1959; Palmer 1027 (US) Manzanillo, Dec 1890; Palmer 1027A (BM, NY, 

US) Manzanillo, Dec 1890; Rogers 511 (COLO-2, F, G, K, MEXU, MO, NY-3, US, W) Playa de 

Santiago, 7 km W of Manzanillo, 30 Jul 1963; Rogers 512 (A, ARIZ, BM, G-2, M, MG, MICH, NY, P, 

S, SP, UC, US) 16 km SSW of Colima on Highway 110, 30 Jul 1963. Michoacan: Leavenworth 451 (F, 

GH) Bank of Rio Apatzingan, 2 miles S of Apatzingan, 5 Aug 1940; Leavenworth & Hoogstraal 1532 

(F, MO, NY) Canyon between Acahuato and Apatzingan, 14 Aug 1941; Leavenworth & Hoogstraal 

1556 (F) Tancitaro Region, between Apatzingan and La Majada, 13 Aug 1941. Guerrero: Hinton 

10345 (GH, K, NY, US) Petatlan, 20 Jun 1937;Hinton 10939 (K, MICH, MO, NY, UC, US-2) Atoyac, 

19 Nov 1937; MacDaniels 212 (F) Acapulco, 24 Aug 1935; Palmer 272 (GH, US) Acapulco and 

vicinity. 

LOCAL NAME. Cuadrado (Ortego 6345). 

Croizat correctly indicated the close relation between his M. colimensis and M. 

chlorosticta, but decided that the single character of seed size and shape and geographic 

range difference were sufficient basis for erecting a separate species. Variation in this 

character indicates the need to synonymize Croizat's species, and geographic range is 

continuous. 

8. Manihot davisiae Croizat, Jour Arnold Arb. 23: 225. 1942. 

Erect shrubs, 1.0- 3.0 m tall, cespitose, woody at the base. Stems glabrous, grayish 

brown. Leaves alternate, stipules deciduous, glabrous; petioles usually ca 10.0 cm long, 

terete, glabrous; lamina nonpeltate, dorsal and ventral surfaces glabrous, abaxial surface 

wax pattern smooth; venation camptodromous; palmately 7 lobed, 3 major, 2 smaller and 

often 2 more minute lobes; median lobes usually ca 10.0 cm long, rhomboid, entire lobed 

or rhomboid pandurate, 3.0-4.0 cm wide, rarely narrower, pandurate lobes usually with a 

prominently dilated apical lobule, apex acuminate. Inflorescence, a moneocious axillary 

raceme, usually short, ca 8.0 cm in length, occasionally longer, all parts glabrous; 

bracteoles nonfoliaceous less than 0.3 cm, margin smooth; bractlets setaceous. Pistillate 

flowers restricted to the base of the inflorescence, borne on short (ca 1.5 cm long) 

pedicels, tepals not seen. Staminate flowers campanulate ca 1.2 cm long, tepals yellowish 

green without any purplish pigmentation, cleft 1/3 way down into 5 lobes, lobes


r I 

C D 

'', 

" 

FIG 15. Manibot cblorosticta. A, distribution; B, habit (Rogers 511, NY); C, leaf (Rogers 512, 

NY); D, inflorescence 

(Rogers 512, NY).


oblong-obtuse, not prominently reflexed at anthesis; disc fleshy, 10 lobed; stamens 10, in 

2 whorls of 5 each, longer ones ca 0.8 cm long, shorter ca 0.6 cm long, filaments and 

anthers cream colored. Fruit pedicels ca 1.5 cm long, slightly curved downwards; capsules 

depressed globular, ca 1.5 cm long from base to apex, surface nearly smooth without 

wings, apex depressed; fruit dehiscence loculicidal, the commissural sutures not 

disjoining. Seeds almost rotund, ca 1.25 cm long; basal end prominently bulged in 

contrast to the tapering carunculate end, caruncle not prominent. Fig 16B, C. 

TYPE. Lemmon s n: USA, Arizona: Santa Catalina Mountains; 27 Aug 1883 

(holotype, US; isotype, BM). 

DISTRIBUTION. (Fig 16A), USA: Arizona; Mexico: Sonora, Chihuahua and Sinaloa; 

usually at high alt, in the Santa Catalina mountains, Baboquivari mountains and northern 

regions of the Sierra Madres Occidental; but at the southern end of the range this species 

descends to sea level. Manihot davisiae appears to be sympatric with M. angustiloba 

almost over its entire range with the possible exception of the Baja California area, where 

no M. davisiae specimens have been collected so far. USA. Arizona: Davis s n (A) Sabino 

Canyon Trail, Santa Catalina Mountains, 15 Aug 1942; Harris 16475 (NY) The Basin, Santa Catalina 

Mountains, 5 Aug 1916; Lemmon 70 3055 (UC) Santa Catalina Mountains, 27 Aug 1883; Livingston 

& Thornber s n (ARIZ-2, POM) Santa Catalina Mountains, Carillos Ranch, 11 Aug 1906; Millspaugh 

3055 (F), 27 Aug 1883; Peebles 8796 (ARIZ-2, F, GH, US) Baboquivari Mountains, 3 Aug 1932; 

Thornber s n (ARIZ, POM) Santa Catalina Mountains, Soldiers Canyon, 14 Aug 1910; Tisher s n (L) 

Santa Catalina Mts. Aug 1912. MEXICO. Sonora: White 542 (ARIZ) Santa Rosa Canyon, near 

Bavispe, 17 Jul 1938; Wiggins & Rollins 221 (ARIZ, MICH, MO, NY, US) 2.5 milesE of Main 

Highway between La Palma and Cienequita, 11 Sep 1941. Chihuahua: Gentry 2450 (ARIZ) Sawakoa, 

Rio Mayo, 25 Aug 1936. Sinaloa: Gentry 14293 (US) Cerros del Fuerte, 18-24 miles N of Los 

Mochis, 25 Sep 1954; Waterfall 12808 (MICH, US) 2 miles E and 14 Miles N of Los Mochis, 17 Aug 

1956. 

Leaf polymorphism is commonly prevalent in Manibot species and, as such, leaf 

outline will have to be employed with caution when used as a taxonomic character. At 

the southernmost extremity of the range of M. davisiae some forms (Gentry 14293; 

Gentry 2450) occur which not only possess completely entire lobed leaves, but also 

characteristically mottled seeds, in contrast to the normal M. davisiae seeds. Geographically, 

the area where these forms grow is adjoining the range of M. isoloba. Cytogenetic 

investigations may throw more light on the possibility of these forms constituting a 

bridging population in the ancestral history linking M. davisiae and M. isoloba. Certain 

collections, such as White 542, which have both M. davisiae specimens and M. angustiloba 

specimens, prevent ruling out the possibility that these two species may merely be two 

segregating forms of one interbreeding gene pool. Field studies of M. davisiae were not 

carried out, and it may become necessary to revise its taxonomic status when more field 

data become available. 

9. Manihot angustiloba (Torrey) Mueller von Argau emend Rogers & Appan. 

Janipha manihot H.B.K. var angustiloba Torrey in Emory, Rep. U. S. Mex. Bound. Surv. 2: 199. 

1859 pro parte; Manihot angustiloba (Torrey) Muell.-Arg. in DC., Prodr. 15(2): 1073. 1866 

pro parte; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 83. 1910. pro parte; Croizat, Jour 

Arnold Arb. 23: 223. 1942 pro parte; McVaugh, Brittonia 13(2): 189. 1961 pro parte. 

Manihot acutiloba Weatherby, Proc. Am. Acad. 45: 427. 1910. (orthographic variant). 

Erect shrubs, 1.0-3.0 m tall, frequently branched, often forming clumps from 

tuberous root stocks; very strong odor of HCN in all plant parts. Roots prominently 

swollen, ca 0.5 m long, fusiform; epidermis light brown, slightly rough to rough, 

subepidermis tan to white; cortex white; vascular strands yellow. Stems glabrous, 

moderate reddish brown (2.5 R 3/7). Leaves alternate, stipules deciduous, glabrous; 

petioles terete, glabrous; lamina nonpeltate, dorsal and ventral surfaces glabrous, abaxial 

surface wax pattern smooth; venation craspedodromous; palmately 7 lobed, 3 major, 2


_,._ ,' 

X 

I 

/'"' 

/ .. . i 1. 

C ^l _ B .... 

C-C p 

les 8796, ARIZ).,. angustiloba. -, .istribution. 

les 8796, ARIZ). M. angustiloba. D, distribution.J 

i . 

z.i. 

i._.i.; ,1 / i.. 

FIG 1.Mnhtdvsa.AditiuinB,la(Pels89,RI 

I 

1 

j 

Z) ,ifoecne(eb


smaller and often 2 more minute lobes; median lobes usually more than 12.0 cm long, 

hastate, occasionally the basal lobules poorly developed, or absent, lobe and lobule apices 

acuminate, terminating in a spine-like point, an extension of a vein, very often a smaller 

lobulet present below the basal lobule, lamina margin above the basal lobule sinuate, 

entire, or incised into small falcate lobulets; lowest lobes about 1/4 as long as median 

lobes, attenuate. Inflorescence a monoecious, axillary raceme, usually ca 12.0 cm long, 

occasionally shorter, all parts glabrous; bracteoles setaceous, margin smooth; bractlets 

setaceous. Pistillate flowers borne on branched peduncles at the base of the inflorescence; 

tepals greenish yellow without purplish pigmentation, ca 1.5 cm long, cleft to the base 

into 5 oblong-lanceolate lobes; disc fleshy, entire, yellowish; pistil ca 0.9 cm long, ovary 

subglobose to slightly elongated, glabrous, the three divisions of stigma clearly divided, 

each well-lobed and lobulate. Staminate flowers campunulate, often as long as 1.8 cm, 

tepals brilliant yellowish green (2.5 GY 8/9), cleft 1/3 way down into 5 oblong lobes, not 

prominently reflexed at anthesis; disc fleshy, yellowish, 10 lobed; stamens 10, in 2 whorls 

of 5 each, longer ones ca 1.1 cm long, shorter ones ca 0.9 cm long, filaments and anthers 

white. Fruit pedicels 1.0-2.5 cm long, slightly curved downwards; capsules depressedglobose, 

ca 1.5 cm long from base to apex, surface very slightly tuberculate, without ribs, 

apex depressed; fruit dehiscence loculicidal, the commissural sutures not disjoining. Seeds 

almost rotund, ca 1.25 cm long, the basal end prominently bulged in contrast to the 

tapering carunculate end; caruncle not prominent. Fig 17A, B, C, D. 

TYPE. Schott III. No. 8: Mexico, Sonora, Sierras oeste de Sta. Cruz y Tubac. VI. 

VII. 1855 (syntype, NY); C. Wright 1811: N. Mex., without precise locality, 1851-1852 

(syntypes GH, MO, NY, US). 

DISTRIBUTION. (Fig 16D). USA: Arizona and doubtfully in New Mexico; Mexico: 

Sonora, Chihuahua, Sinaloa and Baja California; usually at ca 1000-2000 m, in the Santa 

Catalina Mountains, Baboquivari Mountains, and the northern regions of Sierra Madres 

Occidental, but towards the southern end of the range descending to sea level. A small 

segment of the population occurs in the Sierra de la Giganta in Baja California. This 

species appears to be sympatric, almost over its entire range, with M. davisiae. In various 

habitats, on exposed rocky slopes, along rocky ridges, clinging in rock crevices, on 

basaltic hill slopes, in oak woodland, in oak grassland, etc. USA. Arizona: Darrow, Gould, 

Phillips & Pultz 1937 (ARIZ) Carr Canyon, 9 Sep 1944; Goodding 35-57 (ARIZ) Coyote Mountains, 

Quinlin Pass, Pima County, 2 Aug 1957; Goodding 5370 (ARIZ) Mule Mountains, 22 Sep 1939; 

Goodding 6548 (ARIZ) Sycamore Canyon, 17 Aug 1937; Goodding 6549 (ARIZ, NA) Sycamore 

Canyon, 16 Jul 1938; Gould & Haskell 3248 (ARIZ, UC) Baboquivari Mountains, Fresnal Canyon, 

Pima County, 1 Sep 1945;Harrison & Kearney 7983 (POM) Rincon Mountains, 2 Aug 1931; Harrison, 

Kearney & Hastings 6031 (ARIZ) Nogales, 15 Sep 1929; Harrison, Kearney & Hope 8904 (ARIZ, F) 

Florida Canyon, Santa Rita Mountains, 20 Aug 1932; Kearney & Peebles 14816 (ARIZ, POM) 

Patagonia Mountains, Santa Crus County, 24 Aug 1940; Kearney & Peebles 14928 (A, ARIZ, GH, NY, 

US) below Baboquivari Canyon, Pima County, 31 Aug 1940; Lemmon 3054 (BM, GH, UC) Santa 

Catalina Mountains, 17 Aug. 1883; Peebles, Harrison & Kearney 4584 (ARIZ, US) Nogales, 7 Aug 

1927; Peebles, Harrison & Kearney 5588 (ARIZ) Patagonia Mountains, 18 Aug 1928; Peebles & 

Kearney 8742 (GH, MICH, UC) Rincon Mountains, 31 Jul 1932;Pringle 127 (MO) foot hills of Santa 

Rita Mountains, 17 Jul 1881; Pringle 76 (GH) foot hills of Santa Rita Mountains, 8 Sep 1884; Swingle 

s n (NA) Baboquivari Mountains, Allison Dam, Pima County, 10 Jul 1931; Thackery 487 (ARIZ, DS, 

MO, NA, NY) Baboquivari Flat, Pima County, 17 Jul 1928. MEXICO. Baja California Sur Territory: 

Carter 4982 (UC) Sur Sierra de la Giganta, 19 Sep 1965; Carter 5126 (UC) Sur Sierra de la Giganta, 4 

Oct 1965. Sonora: Gentry 14282 (US) Cerro, ca 10 miles S of Ciudad Obregon, 22 Sep 1954; Gentry 

17802 (US) San Bernado, Rio Mayo, 13 Sep 1959; Palmer 233 (BM, NY, US-2) Guaymas, 1887; 

Shreve 6371 (ARIZ-2, DS, F) Bajada, S of Las Trincheras, 2 Sep 1933; White 3013 (MICH) Rio de 

Bavispe, 25 Jul 1940; White 542 (GH, MICH) Santa Rosa Canyon, near Bavispe, 17 Jul 1938; Wiggins 

7056 (DS, US) Along Rio Magdalena, 9 Sep 1934; Wiggins 7155 (A, DS, MICH, US) 20 miles SE of 

Magdalena, 12 Sep 1934; Wiggins & Rollins 221 (UC) 2.5 miles E of Main Highway between La Palma 

and Cienequita, 1 Sep 1941. Chihuahua: Gentry 2371 (ARIZ, F, MO, NA, UC, US) Guasaremos, Rio 

Mayo, 10 Aug 1936. Sinaloa: Hastings & Turner 64-134 (ARIZ, DS) near Yacht Hotel, Topolobampo, 

5 Oct 1964; Palmer 222 (ARIZ, MICH, US) Topolobampo, Sep 1897; Rogers 519 (F-2, G-2, K, 

MEXU, NY-2, US-2, W) vicinity of Topolobampo, 18 miles SW of Los Mochis, 10 Jul 1966. 

LOCAL NAMES. Pico gallo (Carter 5126).

56 

2m.al~ 

.~~~~~~~1 A .. 

"C~~~~~~~~~~~~~~~~~~~~~~~~~ 

l.. ,' 

~~~~~~~~~~~~~ ? ? 

~----" 

-,u 

er~~~~~~~~~~~~~~~~~~ 

rv..... ~...:..:"-.~i.'__;..- . 

. ..._ 

:: 


D 

i. 

~' 

C D ~~~~~~~~~~~~~~~~~~~~~~~ 

FIG 17. Manihot angustiloba. A, habit (Rogers 519, NY); B, roots (Rogers 519, NY); C. leaf 

(Carter 4982, UC); D, inflorescence (Popenoe 134, Fairchild Tropical Garden Herbarium). 

ir


We have elected to designate syntypes for M. angustiloba, rather than designate 

either a lectotype or agree with Croizat's decision (Jour Arnold Arb. 23: 223-224. 1942) 

for the designation of a holotype. In Torrey's description (Rep. U. S. Mex. Bound. Surv. 

2: 199. 1859) of Janipha Manihot, H.B.K. var angustiloba, there are three specimens 

cited, two of which apply to this species, and the third, collected by Gregg, (specimen 

seen by us at MO) clearly belongs to M. subspicata Rogers & Appan. Since Torrey did not 

designate a holotype, and since two of the specimens Schott s n and Wright 1811) served 

to designate the new variety, we see no obvious means by which a holotype can later be 

designated (see Int. Rules, Bot. Nomen, Art. 7, note 3, 1966). We further disagree with 

Croizat's interpretation of Wright's label, which he states to be "New Mexico" when in 

truth the label reads "N. Mexico": which may as easily be northern Mexico as New 

Mexico. 

10. Manihot rhomboidea Mueller von Argau, Linnaea 3: 205. 1865. 

Sprawling shrubs, ca 0.5 m tall, branching from base onwards, branches slender, 

weak. Roots prominently swollen, napiform, often as long as 25.0 cm; epidermis dark 

brown, rough; subepidermis white; with strong smell of HCN. Young stems glabrous; 

mature stems grayish brown, often with deep reddish tinge, glabrous, latex present. 

Leaves alternate, stipules deciduous, glabrous; petioles usually about 6.0 cm long, 

occasionally as short as 3.0 cm, glabrous, terete, greenish or purplish tinged; lamina very 

narrowly peltate, ventral surface glabrous, color varying from pure green to various 

degrees of purplish tinge, dorsal surface glaucous, occasionally silvery, abaxial surface wax 

pattern smooth; venation camptodromous or craspedodromous; palmately 7 lobed, 3 

major, 2 smaller and, often, 2 more minute lobes; median lobes varying in size and shape, 

length 3.0-15.0 cm, width 0.5-5.0 cm, rhomboid, entire-lobed, or rhomboid pandurate 

to hastate or gladiate, or with several combinations of these in the same local population; 

apex acute to acuminate; lowest lobes about 1/4 as long as median lobes, attenuate. 

Inflorescence a monoecious raceme, of two types: either a short axillary raceme less than 

10.0 cm in length, (flowers occurring from base to the apex, pistillate flowers restricted to 

the base), or a long terminal raceme, often as long as 25 cm, (the lower half of the rachis 

completely devoid of flowers, pistillate flowers occurring at the base of the upper half of 

rachis); peduncles and pedicels glabrous, pure green or purplish tinged, bracteoles setaceous 

less than 0.3 cm in length, glabrous, margin smooth; bractlets setaceous, glabrous. 

Pistillate flowers, when occurring at the very base of the inflorescence, borne on pedicels 

about 2.0 cm long, slightly curved downwards; when occurring at the base of the upper 

half of the rachis, borne on pedicels about 2.0 cm long with a characteristic downward 

bend, almost at a right angle; tepals ca 1.1 cm long, pure greenish yellow to various 

degrees of purplish tinged, cleft to the base into 5 oblong-lanceolate lobes; disc orange 

yellow, fleshy, entire; pistil ca 0.5 cm long, ovary greenish, glabrous, subglobose to 

slightly elongated, the 3 divisions of the stigma clearly divided to base and relatively 

long-lobed and lobulate, cream colored. Staminate flowers ca 1.5 cm long, campanulate, 

sometimes almost conical; tepals greenish yellow to various degrees of purplish tinged, 

cleft 1/3 way down into 5 oblong-obtuse or tapering lobes, prominently reflexed at 

anthesis; disc orange-yellow, fleshy, 10 lobed; stamens 10, in 2 whorls of 5 each, longer 

ones ca 1.1 cm long, shorter ones ca 0.9 cm long, fully exerted at anthesis, filaments 

greenish white, anthers pale yellow. Fruit pedicels slightly curved downwards in case of 

fruits occurring at the very base of the inflorescence, or bent down almost at a right angle 

in case of fruits occurring at the base of the upper half of the rachis; capsules almost 

globular, ca 1.0 cm or less in length from base to apex, surface more or less tuberculate, 

without ribs, apex rounded; fruit dehiscence septicidal. Seeds oblong, 0.5-0.8 cm in 

length; caruncle moderately prominent, trapeziform.


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4218 dnd 

21 

FIG 18. Manibot rbomboidea. A, distribution, dots represent subsp rbomboidea and the letter E 

represents subsp microcarpa. M. rbomboidea subsp rbomboidea. B, the plant (Rogers 508, NY); C, leaf 

(Sess4, Moci*o, Castillo & Maldonado 4218, F); D, inflorescence (Rose & Hay 6201, F).


DISTRIBUTION. (Fig 18A). Mexico, states of Sinaloa, Nayarit, Aguas Calientes, 

Guanajuato, Queretaro, Jalisco, Colima, Michoacan, Mexico, Morelos, Puebla, Guerrero, 

Oaxaca, and Chiapas; Guatemala, Huehuetenango, El Quiche, Izabal, Baja Verapas; 

Honduras, Francisco Morazan; El Salvador; Nicaragua. 

Key to the Subspecies of Manihot rhomboidea 

1. Leaf lobes rhomboid, rhomboid-pandurate, hastate or a combination of these 3 shapes, but 

never gladiate; staminate tepals green or with various degrees of reddish or purplish 

pigmentation. 

10a. M. rhomboidea subsp rhomboidea. 

1. Leaf lobes gladiate with or without falcate ascending lobules near base; staminate tepals 

totally green with no purplish or reddish tinge. 

10b. M. rhomboidea subsp microcarpa. 

1Oa. Manihot rhomboidea Mueller von Argau subsp. rhomboidea 

Manibot rhomboidea Muell.-Arg., Linnaea 3: 205. 1865 pro parte. Type. Sesse s n (Sesse, 

Mocino, Castillo & Maldonado, 4218, 4222, 4573 (F-3) 

Manibot mexicana I. M. Johnston, Contr. Gray Herb. 68: 90. 1923; Croizat, Jour Arnold Arb. 

23: 220. 1942. Type. Goldsmith 120 (holotype, GH; isotype, US). 

Manibot ludibunda Croizat, Jour Arnold Arb. 23: 219. 1942. Type. Seler & Seler 2814 

(holotype, NY; isotype, US). 

Median lobes rhomboid, rhomboid-pandurate, or hastate, with several combinations 

of these 3 forms, but never gladiate or gladiate with falcate basal lobules. Several plant 

parts pure green or with various degrees of reddish or purplish pigmentation. Fig 18B, C, D. 

TYPE. Sesse s n (in herb. Boissier) non vidi. Sesse s n (Sesse, Mocino, Castillo & 

Maldonado 4218, 4222, 4573): Mexico (F-3). 

DISTRIBUTION. (Fig 16D). Mexico. Sinaloa, Aquascalientes, Guanajuato, Queretaro, 

Jalisco, Colima, Michoacan, Mexico, Morelos and Puebla. Alt 300-2000 m, on limestone 

scrub land with legumes and organ pipe cacti, rough rocky mountain sides with dry 

shrubs including Ipomoea, Bursera, Cnidoscolus, etc., arid hillsides with shrub forest; 

steep eroded ravines in gray calcareous conglomerate soil, etc. MEXICO. Sinaloa: Rose 1608 

(US) between Rosario and Colomas, 12 Jul 1897. Aguascalientes: Rose & Hay 6201 (F, US) near city 

of Aguascalientes, 20 Aug 1901. Guanajuato: Duges 203 (GH), Guanajuato, 1900; Duges 8 (GH) 

Guanajuato, Sep 1903. Queretaro: Rose & Rose 11190 (F, NY, US) near Queretaro, Aug 1906. 

Jalisco: Faberge s n (TEX) 12 miles E of Guadalajara, Jul 1963; Goldsmith 120 (GH, US) hillside of 

Zapotlan, 1905; Holway 513 (F) Guadalajara, 12 Oct 1896;McVaugh 13298 (MICH) near El Molino, 

ca 25 miles SW of Guadalajara, 1 Oct 1952; McVaugh 17615 (MICH-2) ca 1 mile SW of San Juan de 

Los Lagos, 1 Sep 1958; Palmer 142 = 156 (BM, GH, NY, US-3) Baranca, Jun-Jul 1886;Pringle 11318 

(F, US) Slopes of Barranca of Guadalajara, 19 Jul 1902; Pringle 5159 (GH) Bluffs of Baranca, near 

Guadalajara, 12 Sep 1891; Weintraub & Roller 122 (MICH) 5 miles S of Yahualica along road to 

Tepatitan, 22 Jul 1955; Wilbur & Wilbur 1743 (DS, MICH, US) 10 miles S of Autlan towards La 

Resolana, 15 Jul 1949; Wilbur & Wilbur 2088 (DS, MICH-2, US) ca 15 miles SE of Autlan, 1 Aug 1949. 

Colima: McVaugh 15531 (MICH) 10 miles SSW of Colima on Manzanillo Road, 18 Jul 1957; McVaugh 

15571 (MICH) 11 miles SSW of Colima on Manzanillo Road, 19 Jul 1957. Michoacan: Arsene s n (F) 

Morelia, Jun 1912; Arsene 5272 (BM, MO, NY, US) vicinity of Morelia, Punguato, 18 Aug 1910; 

Galeotti 3734 (P) Cordillera, 1840; Leavenworth & Hoogstraal 1548 (F, MO) Apatzingan, Canyon 

below Acahuato, 14 Aug 1941. Mexico: Hinton 4220 (ARIZ, BM, F, MO, US) Tenayac, 14 Jul 1933; 

Hinton 4349 (BM, F, GH) Acatitlan, 21 Jul 1933; Hinton 4437 (BM, F, US) Tenayac, 17 Aug 1933; 

Hinton 6188 (BM, F, NY, US) Acatitlan, 19 Jun 1934; Hinton 940 (BM) Puerto Salitre, 2 Jul 1932. 

Morelos: Kenoyer s n (MICH) Tepoztlan, 16 Jul 1938. Puebla: Miranda 2092 (GH) Amatitlan, 25 Jul 

1942; Rogers 508 (MO, NY-3, W) 47 km NW of Acatlan on Highway 190, 25 Jul 1963; Rogers 508 A 

(G) 47 km NW of Acatlan on Highway 190, 25 Jul 1963; Rogers 508 B (K) 47 km NW of Acatlan on 

Highway 190, 25 Jul 1963; Rogers 508 C (NY) 47 km NW of Acatlan on Highway 190, 25 Jul 1963; 

Rogers 508 D (US) 47 km NW of Acatlan on Highway 190, 25 Jul 1963, Rogers 508 E (S) 47 km NW 

of Acatlan on Highway 190, 25 Jul 1963; Rogers 508 F (MICH) 47 km NW of Acatlan on Highway 

190, 25 Jul 1963; Rogers 508 G (F) 47 km NW of Acatlan on Highway 190, 25 Jul 1963; Rogers 

508 H (COLO) 47 km NW of Acatlan on Highway 190, 25 Jul 1963; Rogers 508 1 (MEXU) 47 km NW 

of Acatlan on Highway 190, 25 Jul 1963; Webster, Miller & Miller 13080 (DAV) 29.5 miles NW of 

Acatlan, 18 Aug 1962; Webster, Miller & Miller 13080A (PUL) 29.5 miles NW of Acatlan, 18 Aug


1962; Webster, Miller & Miller 13080 B (PUL) 29.5 miles NW of Acatlan, 18 Aug 1962. Province 

Unknown/Uncertain: Jurgensen 582 (BM) Sierra San Pedro Nolasco, 1843;Pavon 34163 (F); Sesse & 

Mocifo 30856 (F). GUATEMALA. Huehuetanango: Melbus & Goodman 3652 (F) below San Antonio 

Huista, 29 Sep 1944; Seler & Seler 2814 (NY, US) Uaxackanal, 13 Jul 1896; Steyermark 50614 (F) 

Cerro Pix Pix, 15 Aug 1942; Steyermark 51474 (F) between Nenton and Miramar, 29 Aug 1942. 

LOCAL NAMES. yuca cimarrona (Steyermark 51474). 

The three major phenotypic forms of Manihot rhomboidea subsp rhomboidea, the 

rhomboidea form, the mexicana form and the ludibunda form appear as three peaks in 

the "SKYLINE" (Fig 5), with several intermediate forms between the peaks. This 

indicates close phenotypic similarity between the forms. A study of the geographical 

distribution shows that the forms do not have any geographical integrity of their own, 

and all the forms occur over the entire range of this subspecies. There are several 

collections where two or more forms are represented in the same collection, (c f Rogers 

508). 

The close phenotypic similarity of subspecies rhomboidea with the other subspecies 

microcarpa is evident in Fig 5. However, there are a few morphologic characters which 

consistently differentiate these two populations (described in the morphological diagnosis 

of these subspecies) indicating that there is only limited interbreeding between these two 

populations. Nevertheless, these two populations do not seem to represent 2 closed gene 

pools. There are indications of genetic exchange between these two populations along the 

line where these two meet, as evident from the frequent occurrence of forms which are 

morphologically more or less intermediate between these two subspecies, in the region 

where they merge. These two populations are, therefore, assumed to be partially isolated 

segments of one single closed gene pool, and as such designated as two subspecies of one 

biological species. 

10b. Manihot rhomboidea Mueller von Argau subsp microcarpa (Mueller von Argau) 

Rogers & Appan, stat nov 

Manihot rhomboidea Muell.-Arg., Linnaea 3: 205. 1865 pro parte. Type. Sesse s n (Sesse, 

Mocino, Castillo & Maldonado 4224) (syntype, F). 

Manihot microcarpa Muell.-Arg., Flora 55: 42. 1872. Type. Karwinsky s n (M, photo at F, G). 

Manihot parvicocca Croizat, Jour Arnold Arb. 23: 219. 1942. Type. Matuda 1665 (holotype, A: 

isotypes, MICH, MO, NY, US). 

Median lobes gladiate, with or without falcate, ascending lobules near the base. Most 

plant parts green, without other pigmentation. Fig 19A, B. 

TYPE. Karwinsky s n: Mexico: (M, photo of type F, G). 

DISTRIBUTION. (Fig 18A). Mexico: Nayarit, Jalisco, Colima, Michoacan, Mexico, 

Guerrero, Morelos, Oaxaca and Chiapas; Guatemala; El Salvador; Honduras; and 

Nicaragua. Alt 300-2000 m. Ecologically this subspecies is very similar to Manihot 

rhomboidea subsp rhomboidea, but subsp rhomboidea is sympatric over the entire range 

of section Foetidiae, while subsp microcarpa is allopatric, not overlapping the range of 

any Manihot species. The range of subsp microcarpa is the longest among the taxa in this 

study, extending from 13 N in Nicaragua to 22 N in Nayarit, Mexico, indicating its 

adaptability to flourish within such a wide latitudinal range. MEXICO. Nayarit: Feddema 909 

(US) 3 miles NE of Puga, 22 Aug 1959; McVaugh 13386 (MICH) 10 miles SE of Tepic, 6 Oct 1952; 

McVaugh 16580 (MICH) near KM 886, ca 10 miles SE of Tepic, 30 Aug 1957. Jalisco: McVaugh 

15954 (MICH) ca 11 miles N of bridge of Rio Chihautlan, 1 Aug 1957;McVaugb 18128 (MICH, US) 

Puente San Pedro, 5 miles SW of Tecaltitlan, 22 Sep 1958; Palmer 142 = 156 (GH, NY) Rio Blanco, 

Jun-Jul 1886; Rose & Hough 4751 (US) near Tequila, Jul 1899; Zingg 10 (F) Mountains near Bolanos, 

1935. Colima: McVaugb 15486 (MICH) ca 18 miles E of Colima at km 189, 17 Jul 1957; McVaugh 

15531 (MICH-2) 10 miles SSW of Colima on Manzanillo Road, 18 Jul 1957. Michoacan: Hinton 

15072 (NA, NY, US) Barroloso, 7 Aug 1939. Mexico: Hinton 4467 (BM, F, GH, US) Ixtapan, 1 Aug 

1933. Morelos: Kenoyer s n (ARIZ, MICH) Cuernavaca, 16 Jul 1938; Knechtel 507 (W) Cuernavaca;


>, I 

Wb 

IF ,i J 

TJ 4 

FIG 19. Manihot rbomboidea rhomboidea subsp microcarpa. A, A leaf (Knechtel (Knecbtel 507, W); B, inflorescence 

(Knechtel 507, W). W) M. M subspicata. subspcata C, C distribution; D, habit (Rogers, (Rogers Appan & Rogers 525, NY). 

O--


Lyonnet 301 (NY, US) Cuernavaca, May-Jun 1929; Martinez 15088 (MO) Cuernavaca, Aug 1946. 

Guerrero: Herald & Clark 349 (TEX) 1 mile N of Acahuizatla, 27 Jun 1954; Hinton et al 10488 (GH, 

MICH, NY, UC) Mesa Quisle, 11 Jul 1937; Hinton et al 8029 (DS, GH, MICH, NY, U, US) 

Chacamerito, 8 Jul 1935; Hinton et al 9090 (DS, MICH, NY, US) Placeras Mesa, 14 Jul 1936; Ryan & 

Floyed 11 (TEX) Aqua del Obispo, 10 Jun 1954. Oaxaca: Galeotti 3794 (P); Ghiesbreght s n (P), 

1842. Chiapas: Matuda 1665 (A, MICH, MO, NY, US) Siltepec, 8 Aug 1937; Matuda 4392 (A, NY) 

near Siltepec, 9 Jul 1941; Nelson 2899 (GH, US) top of Ridge back of Tonala, 10 Aug 1895; Purpus 

10213 (NY, UC, US) mountains near Menserrate, May 1925; Purpus 9322 (UC) Hacienda Menserrate, 

Sep 1923. Province Unknown/Uncertain: Sessd, Mocino, Castillo & Maldonado 4224 (F). GUATE- 

MALA. Huehuetenango: Steyermark 50614 (F) Cerro Pix Pix, 15 Aug 1942; Steyermark 50866 (F) 

along Rio Cuilco, 18 Aug 1942. El Quiche: Aguilar 1546 (F), 1942. Izabal: Deam 6097 (F) 1 Jun 

1909. Baja Verapas: Pittier 132 (NY, US) Cuesta de Cachil, Apr 1905. HONDURAS. Francisco 

Morazan: Rodriques 1528 (F) Santa Ines, 4 Nov 1943: Williams & Molina 10116 (F, MICH, MO) Las 

Mesas, 21 Jul 1946. EL SALVADOR. Province Unknown/Uncertain: Calderon 1023 (GH, NY) Cerro 

de la Olla, 1922. NICARAGUA. Province Unknown/Uncertain: Morley 745 (F, UC, US) 43 miles 

towards Managua, 1 Aug 1946. 

The justifications for designating this taxon as one of the subspecies of Manihot 

rhomboidea are elaborated in the discussions of M. rhomboidea subsp rbomboidea. 

Manihot microcarpa and M. parvicocca are two minor morphologic variants. 

McVaugh (1961) recognized this and synonymized these two species. A critical study of 

the geographical distribution of these two forms indicates that these two forms are 

sympatric. 

11. Manihot subspicata Rogers & Appan, sp nov. 

Janipha manihot H.B.K. var angustiloba Torrey in Emory, Rep. U. S. Mex. Bound. Surv. 2: 199. 

1859. pro parte. Type. Gregg s n (198) (syntype, MO). 

Frutices ca 1 m altae, ramis laxis, glabrae; folia septemloba lobis medianibus 

6.0-10.0 cm longibus; inflorescentia subspicato-racemosa ca 25 cm longa; capsulae ca 1.5 

cm longae; semina ca 1.0 cm longa. 

Weak stemmed, sprawling shrubs, ca 1.0 m tall, many branched from base, branches 

lax, slender, irregularly and slightly geniculate, often leaning on other vegetation. Roots 

prominently swollen, penetrating 0.3-1.0 m deep, of various shapes, napiform, fusiform, 

or irregularly swollen; epidermis more or less smooth, dark grayish brown subepidermis 

tan to white; cortex white, spongy, with strong smell of HCN. Young stems glabrous, 

greenish brown; mature stems glabrous, dark grayish brown, lower stems with prominent 

lenticels. Leaves alternate; stipules deciduous, glabrous, petioles terete, 5.0- 10.0 cm long, 

rarely longer, glabrous, dark red (10 R 3/4) above, greenish beneath; lamina very 

narrowly peltate, rarely widely peltate; dorsal and ventral lamina surfaces and midribs 

glabrous, abaxial surface wax pattern smooth; venation craspedodromous; palmately 7 

lobed, 3 major, 2 smaller and often 2 more minute lobes; median lobes 6.0-10.0 cm long, 

rarely longer, hastate, the apex of the basal lobules sometimes dilated, not narrowly 

attenuate, terminating in a spine-like point, lamina margin above the basal lobules nearly 

entire, sinuate, or incised into small pointed lobulets, the apical region of median lobes 

often slightly dilated, apex acuminate, bristle-pointed; lowest lobes about 1/4 as long as 

median lobes, attenuate. Inflorescence a monoecious, subspicate terminal raceme, tall and 

erect, about 25.0 cm in length, the lower half of the rachis devoid of flowers, all parts 

glabrous; bracteoles setaceous, less than 0.3 cm in length, margin smooth or serrate; 

bractlets setaceous. Pistillate flowers restricted to the base of the upper half of the 

inflorescence, borne on pedicels 1.0-2.0 cm in length with a characteristic bend 

downwards almost at a right angle, tepals ca 1.2 cm in length, greenish yellow externally 

and internally, cleft to the base into 5 oblong-lanceolate lobes; disc fleshy, entire, bright 

orange; pistil about 0.6 cm long, ovary subglobose, yellowish green, the trifid stigma 

moderately lobed and lobulate, cream colored. Staminate flowers campanulate to almost


conical, more or less subsessile, pedicels less than 0.5 cm in length, tepals ca 1.3 cm long, 

reddish (5 R 4/12) externally, greenish white internally, cleft 1/3 way down into 5 more 

or less tapering lobes, moderately reflexed at anthesis; disc bright orange, 10 lobed, 

fleshy; stamens 10, in 2 whorls of 5 each, longer ones ca 0.9 cm, shorter ones 0.7 cm, 

filaments and anthers cream colored. Fruit pedicels 1.0-2.0 cm long, with a characteristic 

downward bend at almost a right angle; capsules ca 1.5 cm long from base to apex, 

ovate-ellipsoid, surface fairly smooth, ribs not prominent, apex rounded; fruit dehiscence 

septicidal. Seeds oblong, ca 1.0 cm long, basal and carunculate ends smoothly rounded; 

caruncle prominent, trapeziform. Fig 19D; 20A, B, C. 

TYPE. Rogers, Appan & Rogers 529: Mexico, Tamaulipas: Circa 5 miles E of Casas, 

33 miles E of Ciudad Victoria, 21 Jun 1968 (holotype, NY; isotypes, F, K, MEXU, MO, 

US, W). 

DISTRIBUTION. (Fig 19C). Mexico: Coahuila, Nuevo Leon and Tamaulipas; along 

eastern slopes of Sierra Madres Oriental. Usually growing at ca 200- 700 m, but towards 

the southern end of its geographical range the species grows in low lands in the coastal 

plains, adjacent to or possibly slightly overlapping the range of M. walkerae. 

This species characteristically grows close to and in the shade of other shrubs, thus 

protected by the taller shrubs. The savannah type habitat supports tall shrubs and 

medium sized trees such as Cordia boissieri, lechuguilla, thorny legumes, etc. This species 

is a relatively infrequent member of the vegetation. Towards the southern extremity of 

the range it grows in prairies with very shallow reddish sandy loam. Usually grows in 

limestone based, loose and rocky soil. Like many other Manihot species this one exhibits 

the potential to colonize disturbed areas, such as in soil dumped after road construction, 

eroded ravines, etc. MEXICO. Coahuila: Marsh 1163 (F, TEX) Muzquiz, Apr 1938. Nuevo Leon: 

Crutchfield & Johnston 5460A (TEX) Mamulique Pass, 23 miles S of Sabinas Hidalgo, 4 May 1960; 

Edwards 411 (F, MO, TEX) Rancho Resendez, Lampazos, 29 Jun 1937; Gregg 198 (MO) Monterrey, 

24 Jun 1848; Perkins & Hall 3565 (F) Nuevo Leon, Road from Monterrey to Nuevo Laredo, 12 Apr 

1939; Rogers, Appan & Rogers 525 (G-2, MEXU, NY-3, US) Mamulique Pass, just off old Highway, 

Milestone 1051, 18 Jun 1968; Rogers, Appan & Rogers 526 (COLO, F-2, G, K, M, MICH, NY-2, S, 

US, W) Summit of Mamulique Pass, 38 miles N of Monterrey, 19 Jun 1968. Tamaulipas: Bartlett 

10613 (MICH) Summit of Cerro de La Tamaulipeca, 26 Jul 1930; Bartlett 13695 (MICH-2) Summit of 

Cerro de La Tamaulipeca, 4 Jul 1931; Crutchfield & Johnston 5523 (TEX) 16 miles E of San 

Fernando-Santander Jiminez Highway on Road to Lareto, 15 Sep 1960; Crutchfield & Johnston 

5784E (TEX) 5 miles E of Casas on Victoria-Sotolamarina Highway, 26 Sep 1960; Graham & 

Johnston 4721B (TEX) 26 miles S of Victoria, 15 Nov 1959; Kenoyer C 142 (GH) S of Victoria, 14 

Aug 1940; Lesueur 246 (ARIZ, F, TEX) Mountains W of Bustamante, Aug 1938; Rogers, Appan & 

Rogers 533 (A, BM, COLO, K, MEXU, NY, P, SP, UC), 26 mi S of Ciudad Victoria, 26 Jun 1968. 

LOCAL NAME. Palo mulato. (Edwards 411). 

Manihot subspicata was confused in the past with M. mexicana, which is merely one 

of the morphologic variants of M. rhomboidea subsp rhomboidea. The range of M. 

subspicata is widely disjunct from the range of M. rhomboidea subsp rhomboidea, and 

consequently there is little likelihood of interbreeding. This disjunct taxon is, therefore, 

assumed to be a closed gene pool and, as such, described as a new species. The specific 

epithet describes the characteristic subspicate inflorescence. 

12. Manihot walkerae Croizat, Bull. Torrey Club 69: 452-457. 1942. 

Decumbent shrubs, ca 0.5 m tall, profusely branching, branches slender, prostrate, 

long, weak. Roots prominently enlarged, carrot shaped, ca 10.0 cm long; epidermis very 

dark brown, rough with bands of laterally oriented corky strips; cortex white with strong 

smell of HCN; tuberous roots reported to possess adventitious buds (used as propagating 

material, Walker 1003). Young stems glabrous, greenish brown; mature stems glabrous, 

grayish brown. Leaves alternate; stipules setaceous, deciduous, glabrous; petioles slender,


'- 

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i 

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_ F: 

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;~~~~~~~~~~~~~~~~~~~~~~~~~, ' Y\/,~ 

E- 

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~ 

~ 

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FIG 20. Manibot subspicata. A, roots (Rogers, Appan & Rogers 533, NY); B, leaf (Rogers, Appan 

& Rogers 525, NY); C, inflorescence (Perkins & Hall 3565, F). M. woalkerae. D, distribution.


usually 7.0-10.0 cm long, glabrous, terete, ventral surface dark dull red, dorsal surface 

greenish; lamina peltate, over 0.5 cm wide at the region between the petiole attachment 

point and the lamina margin at the base of the leaf, ventral lamina deep green, dorsal 

lamina markedly glaucescent, abaxial surface wax pattern smooth; venation craspedodromous; 

palmately 5 lobed, 3 major and 2 smaller lobes; median lobes ovate-ellipsoid to 

more or less parabolic, pandurate, rarely entire-lobed, usually 5.0-10.0 cm long, the basal 

lobule apex abruptly dilated, cuspidate, terminating in a spine-like point; lamina margin 

above and below the basal lobule incised, often deeply, into lobulets which also terminate 

in spine-like extensions, apex acute to acuminate; the sinuses on either side of the median 

lobe with a characteristically flat base forming an almost 45? angle to the median lobe 

midrib; lowest lobes about half as long as median lobes, apex cuspidate. Inflorescence a 

monoecious, axillary, subspicate, raceme, 5.0-10.0 cm long, all parts glabrous; bracteoles 

setaceous, less than 0.3 cm long, glabrous, margin serrate; bractlets setaceous, glabrous. 

Pistillate flowers restricted to the very base of the inflorescence, borne on ca 1.5 cm long 

pedicels (which appear attached directly to stem); tepals ca 1.1 cm long, light purplish 

streaked externally, cleft to the base into 5 oblong-lanceolate lobes; disc fleshy, entire, 

with several staminodes; pistil ca 0.6 cm long, ovary subglobose, the trifid stigma well 

lobed and lobulate. Staminate flowers almost tubular, distinctly constricted in the 

middle, tepals ca 1.2 cm long, light purplish streaked externally, cleft 1/3 of the way 

down into 5 lobes, the lobes oblong, tapering, not prominently reflexed at anthesis; disc 

fleshy, 5 lobed; stamens 6-10 (6, 7, and 8 observed, 10 reported in original description), 

superior whorl ca 0.9 cm long, inferior whorl ca 0.7 cm long, filaments and anthers cream 

colored. Fruit pedicels slightly curved downwards ca 1.5 cm long, pistillate pedicels 

attached to the very base of the inflorescence; capsules almost globular, ca 1.0 cm long 

from base to apex, surface sparingly but manifestly verrucose and rugose, ribs not 

prominent; fruit dehiscence septicidal. Seeds rotund, ca 0.8 cm long; caruncle very 

prominent, large in comparison to seed size, ca 0.2 cm long and ca 0.4 cm broad, cream 

colored, slightly cleft and 2 lipped in front. Fig 21A, B, C, D. 

TYPE. Walker (Parks, H. B.) s n: USA, Texas: Mission, 1940 (holotype, A; isotypes, 

A-2). 

DISTRIBUTION. (Fig 20D). USA: Texas; Mexico: Tamaulipas. USA. Texas: Rogers 522 

(F, G, NY, US) University of Texas, Austin, 31 Aug 1967;Schott 52 (F-2, NY-2) Ringgold Barracks, 2 

Jun 1853; Walker 1003 (GH) La Joya, Apr 1940; Wornock & Barkley 147 (MO, UC, US) University of 

Texas Campus, 7 Jun 1943. MEXICO. Tamaulipas: Crutchfield & Johnston 5572 B (TEX) Papalote de 

Mirandena, 16 Sep 1960; Johnston 5363B (TEX) Rancho Loreto, 26 Apr 1960; Pringle 2243 (GH) 

near Matomoros, 31 Jul 1888. 

This distinct species may be very close to extinction in the localities along the Rio 

Grande in Texas and adjoining Mexico due to intensive cultivation in this area. The 

illustrations, and much of the detailed description of Manihot walkerae were taken from 

the small but vigorous colony established on the south side of the Department of Botany 

building at the University of Texas, Austin, Texas. 

3. Manihot sect Foetidae Rogers & Appan, sect nov 

Manihot sect Grandibracteatae Pax subsect Papillosae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 28. 1910. 

Plantae caulinae, lignosae arbores. Folia petiolatae, penitae lobatae inflorescentia 

monoecia panicula. 

Plants caulescent; leaves widely spaced on stem; petiolate, petiole attachment basal 

or peltate; lamina membranaceous to coriaceous, deeply lobed, lobes variously shaped but 

not linear. Inflorescence a monoecious panicle; bracts and bracteoles setaceous or 

semifoliaceous, margins usually entire. 

TYPE. Manihotfoetida (H.B.K.) Pohl. 

DISTRIBUTION. (Fig 22).


C 

. 

,i .~ ,s~ 

%.... .. 

i_ 

FIG 21. Manibot walkerae. A, habit (Rogers 522, NY); B, leaf showing the outline of median

FIG~o 22 Maio sec Fotde Gegapiarne 

4~~~


Key to the Species of Sect Foetidae 

1. Leaves with more than 3 lobes; capsules subglobose; inflorescence a panicle, (Manihot 

tomatophylla and M. websterae inflorescences not seen). 

2. Petiole attachment peltate; bases of leaf lobes more than 0.25 cm across; lamina not 

deeply cleft, more than 0.5 cm wide from base of sinus to petiole attachment point; leaf 

lobes obovate or obovate-pandurate; fruit pedicels more than 1.0 cm long; seeds oblong. 

3. Leaf lobes entire, never pandurate; fruit dehiscence septicidal. 

4. Leaf apices prominently caudate; petioles, peduncles, pedicels, bracteoles and 

bractlets glabrous; lenticels on mature stem dense and pronounced. 13 M. caudata. 

4. Leaf apices acute; petioles, peduncles, pedicels, bracteoles and bractlets sparsely 

pubescent; lenticels on mature stem sparse and not pronounced. 14. M. michaelis. 

3. Leaf lobes pandurate; fruit dehiscence loculicidal. 15. M. tomatophylla. 

2. Petiole attachment basal; bases of leaf lobes less than 0.25 cm across; lamina very deeply 

cleft, less than 0.5 cm wide from base of sinus to petiole attachment point; leaf lobes 

oblong; fruit pedicels less than 1.0 cm long; seeds rotund. 16. M. websterae. 

1. Leaves constantly 3 lobed; capsules conical, distal end tapering; inflorescence a raceme. 

5. Ovary tomentose. 17. M. foetida. 

5. Ovary glabrous. 

18. M. crassisepala. 

13. Manihot caudata Greenman, Proc. Am. Acad. 39: 82. 1903. 

Trees, ca 10.0 m tall, the main trunk with a diam of ca 30.0 cm supporting an 

umbrella-like spreading, thick foliaged canopy; bark grayish brown. Young stems light 

reddish brown, shiny, sparsely pubescent; mature stems glabrous, reddish brown, 

epidermis peeling off. Young foliage strong yellow-green (5 GY 6/8). Leaves alternate; 

stipules deciduous, sparsely pubescent; petioles ca 15.0 cm long, glaucous, terete, 

occasionally very slightly canaliculate, dark red (2.5 R 3/7); lamina narrowly peltate, 

ventral surface dark green (5 G 3/4), abaxial surface wax pattern reticulate; venation 

camptodromous; palmately 5 lobed, 3 major and 2 smaller lobes; median lobes obovate, 

entire, usually ca 13.0 cm long, 6.0 cm wide, occasionally as long as 25.0 cm and as wide 

as 10.0 cm, apex abruptly caudate-acuminate, terminating in a bristle, occasionally 2.0 

cm long; lowest lobes slightly smaller than median lobes, conspicuously nonsymmetric, 

curved up. Inflorescence a monoecious, axillary panicle, usually many branched, 

15.0-25.0 cm long; peduncles, pedicels and young buds coated with ashy-blue 

glaucescence; bracteoles setaceous, occasionally semi-foliaceous and as long as 4.0 cm, 

glaucous, margin smooth; bractlets setaceous, glaucous. Pistillate flowers restricted to the 

base of the inflorescence, borne on 2.0-4.0 cm long pedicels; tepals ca 1.4 cm long, 

greenish yellow, often glaucous externally, cleft to base into 5 oblong-lanceolate, tapering 

lobes; disc prominent, fleshy, entire, staminodes often present around the disc; pistil ca 

0.9 cm long, ovary subglobose to slightly elongated, the trifid stigma moderately lobed 

and lobulate. Staminate flowers campanulate, tepals greenish yellow, often glaucous, ca 

1.6 cm long, cleft 1/3 way down into 5 oblong-obtuse lobes, not prominently reflexed at 

anthesis; disc fleshy, shallowly 10 lobed; stamens 10, in 2 whorls of 5 each, superior 

whorl ca 1.1 cm long, inferior whorl ca 0.9 cm long, filaments and anthers pale yellowish. 

Fruit pedicels 2.0-4.0 cm long, almost straight; capsules ovate-ellipsoid, ca 2.25 cm long 

from base to apex, slightly pointed; fruit dehiscence septicidal. Seeds oblong, ca 1.5 cm 

long, flattened and conspicuously ribbed along the lateral edges; caruncle not prominent. 


TYPE. Pringle 8687: Mexico, Michoacan: Volcanic hills, Monte Leon, 21 Aug 1902 

(syntypes, F-2, GH, MO, NY, UC, US, W); Palmer 201: Mexico, Chihuahua: Hacienda San 

Miguel, near Batopilas, Sep 1885 (syntype, US). 

DISTRIBUTION. (Fig 23A). Mexico: Michoacan, Guanajuato, Zacatecas and Chihuahua. 

Abundant on the rocky slopes above the south shore of Lake Chapala, at ca 1500 m; 

in the volcanic hills near Monte Leon, west of Lake Chapala; and also towards the north 

in Guanajuato and Zacatecas. MEXICO. Zacatecas: Rose 2499 (GH) San Juan Capistrano, Aug


C- 

FG2 o da A dr 

D inflorescence (Rogers 509, NY). 

- 

7 4 U'I~~~~~~~~~~~ 

P~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i' 

Jri 

~~~~~~C., 

FIG 23. Manibot 

caudata. A, dis(triuinB,hbt(oes59NY;ClafPrne867


1897. Guanajuato: Rusby 178 (NY, US) Empalme de Gonzales, 18 Jun 1910. Michoacan: McVaugh 

15037 (MICH-2) S Shore of Lake Chapala, 5 miles E of Jalisco Line, 24 Jun 1957; Rogers 509 (F, G-2, 

K, MEXU-2, NY-3, US, W) S Shore of Lago Chapala, 28 Jul 1963. 

The specimens of McVaugh and Rogers, collected at the same locality on the south 

shore of Lake Chapala, indicate that this species is used in hedgerows, and thus 

propagated by man. This fact may account for the disjunct localities (Palmer 201, in 

Chihuahua) where the species has been collected. 

Manihot caudata is most closely related to another tree species, M. michaelis, and is 

distinguished from the latter by its long, attenuated leaf apex, and by the glabrous 

condition of several of its parts. 

14. Manihot michaelis McVaugh, Brittonia 13: 190. 1961. 

Trees, ca 10.0 m tall, trunk diam at base ca 20.0 cm, frequently branched above the 

middle from a single trunk, branches usually ascending, sometimes reclining. Roots not 

prominently swollen (Jancey 331); epidermis rough, brownish gray; subepidermis creamy 

white. Young stems pubescent, light reddish brown; mature stems sparsely pubescent, 

reddish brown to grayish brown, internally creamy white. Leaves alternate, stipules 

deciduous, sparsely pubescent; petioles generally ca 15.0 cm long, occasionally longer, 

terete, green, occasionally reddish, sparsely pubescent; lamina narrowly peltate, ventral 

surface green (2.5 G 5/9), glabrous, dorsal surface glaucous, light green (7.5 G 8/6), 

abaxial surface wax pattern reticulate; venation camptodromous; palmately 5 lobed, 3 

major and 2 smaller lobes; median lobes obovate, entire, generally ca 12.0 cm long, ca 5.0 

cm wide, juvenile plants with much larger foliage, apex acute; lowest lobes slightly smaller 

than median lobes, conspicuously nonsymmetric, curved up. Inflorescence a monoecious, 

axillary panicle, sparsely branched, ca 10.0 cm long, peduncles and pedicels sparsely 

pubescent; bracteoles greenish white, setaceous, sparsely pubescent; bractlets setaceous, 

sparsely pubescent. Pistillate flowers restricted to the base of the inflorescence, borne on 

pedicels 2.4-4.0 cm long; tepals ca 1.1 cm long, yellowish green, cleft down to base into 

5 oblong-lanceolate lobes; disc fleshy, entire; pistil ca 0.6 cm long; ovary subglobose, 

glabrous, the trifid stigma moderately lobed and lobulate. Staminate flowers campanulate;, 

tepals ca 1.2 cm long, yellowish green, cleft 1/3 way down into 5 oblong-obtuse 

lobes; disc fleshy, 10 lobed; stamens 10, in 2 whorls of 5 each, the superior whorl ca 0.8 

cm long, the inferior whorl ca 0.7 cm long, filaments and anthers cream colored. Fruit 

pedicels 2.4-4.0 cm long, almost straight; capsules ovate-ellipsoid, ca 2.0 cm long from 

base to apex, surface more or less smooth, often glaucous, perceptibly 6 ribbed, apex 

slightly pointed; fruit dehiscence septicidal. Seeds oblong, ca 1.5 cm long, flattened and 

conspicuously ribbed along the lateral edges, caruncle not prominent. Fig 24B, C. 

TYPE. McVaugb 15502: Mexico, Colima: 18 miles E of Colima, near Rio Tuxpan, 17 

Jul 1957 (holotype, MICH). 

DISTRIBUTION. (Fig 24A). Mexico: Colima and Jalisco. Narrowly localized in the 

region around Colima city, alt 300-1750 m, in flood plains and bluffs of wooded ravines, 

on both sides of Rio Tuxpan, Rio de Colima and Rio de Armenia. MEXICO. Jalisco: Rogers 

514 (A, COLO, G-2, K, MEXU, NY-4, S, US-2) 32 km E of Colima, 31 Jul 1963. Colima: Jancey 330 

(P, B, M) near Summit of First Mountain Pass W of Colima, Jul 1965;Jancey 331 (NY) near Summit 

of First Mountain Pass, 12 miles W of Colima, Jul 1965; Jancey 332 (NY) near Summit of Mountain 

Pass, 12 miles W of Colima, Jul 1965; Jancey 333 (NY) near Summit of Mountain Pass, 12 miles W of 

Colima, Jul 1965;Jones 127 (POM) Colima, 2 July 1892; Rogers 513 (A, F, M, MICH, S, SP) 30 km E 

of Colima, 30 Jul 1963. 

15. Manihot tomatophylla Standley, Am. Midi. Nat. 36: 178. 1946. 

Trees, ca 10.0 m tall, trunk diam at base ca 30.0 cm, successive dichotomous and 

trichotomous branching beginning at a height of ca 1.5 m, branches many, erect, forming


'xI, 

I 

I ' - Y E S-', 

A!~~~I ~ ~ 

m,-# 

' ' 

FIG 24. Manibot micbaelis. A, distribution; B, leaf (Rogers 514, NY); C, inflorescence (Jones 

127, POM). M. tomatopbylla. D, distribution. 

127, POM). M. tomatophylla. D, distribution. 

-'C


a more or less obovate crown supported by a main trunk, latex present. Young stems 

glabrous, grayish brown with reddish tinge; mature stems glabrous, reddish brown, usually 

with white dot-like lenticels; bark on older stems light gray to silvery gray, verrucate. 

Leaves alternate; stipules deciduous, glabrous; petioles ca 15.0 cm long, occasionally 

longer, terete, sometimes slightly canaliculate, glabrous, bright red; lamina narrowly 

peltate, ventral surface deep green, glabrous, dorsal surface with distinct ashy-blue 

glaucescence, abaxial surface wax pattern reticulate; venation camptodromous; palmately 

5-7 lobed, 3 major, 2 slightly smaller and often 2 more smaller lobes; median lobes 

obovate-pandurate, ca 13.0 cm long, ca 4.0 cm wide, primary constriction short, to as 

long as ca 4.0 cm, margin within the constriction usually rounded without any secondary 

constrictions, apex obtuse to truncate, mucronate; lowest lobes, in case of 5 lobed leaves, 

slightly smaller than median lobes, conspicuously nonsymmetric, curved up; in case of 7 

lobed leaves, ca 1/4 as long as median lobes, obtuse. Inflorescence not seen. Fruit pedicels 

2.0-5.0 cm long, more or less straight, glaucous; capsules fleshy, ovate-ellipsoid, ca 2.5 

cm long from base to apex, prominently 6 ribbed, ribs wavy, platelike, usually with a 

sharp edge, fruit surface prominently verruculose, glaucous, apex pointed; dehiscence 

loculicidal, the 3 commissural sutures not disjoining. Seeds oblong, ca 1.75 cm long, 

carunculate end acutely pointed, ventral surface with a prominent furrow along the 

middle, lateral edges prominently ribbed, surface shiny, brownish orange with fine black 

mottlings; caruncle not prominent, very small in contrast to the seed size. Fig 25A, B. 

TYPE. Leavenworth & Hoogstraal 1402: Mexico, Michoacan: Tancitaro region, 

Apatzingan, near La Majada, 9 Aug 1941 (holotype, F; isotypes, F, MICH, MO, NY). 

DISTRIBUTION. (Fig 24D). Mexico: Michoacan. Localized in the region of Apatzingan, 

alt ca 400 m, on edges of dry canyons, on banks of dry arroyos, on low hills 

supporting sparse woodlands. MEXICO. Michoacan: McVaugh 17951 (MICH, US) between San 

Juan de Los Platanos and Amatlan, 17 Sep 1958; Rogers 516 (F, G, K, MEXU, NY, W) 6 km W of 

Apatzingan, 1 Aug 1963; Rogers 517 (ARIZ, G, NY, UC) ca 5 miles N of Nueva Italia, 1 Aug 1963. 

16. Manihot websterae Rogers & Appan, sp nov7 

Arbores ca 5.0 m alti, pubescentes; folia penta vel septemloba lobis medianibus ca 

11.0 cm longibus, ca 3.0 cm latibus; inflorescentia non visa; capsulae ca 2.25 cm longae; 

semina ca 1.5 cm longa. 

Trees, ca 5.0 m tall. Young stems light reddish brown, pubescent; mature stems dark 

reddish brown, pubescent. Leaves alternate; stipules deciduous, pubescent; petioles ca 

10.0 cm long, terete, sometimes slightly canaliculate, pubescent; lamina nonpeltate, 

ventral surface green, sparsely pubescent, dorsal surface light green, sparsely pubescent, 

abaxial surface wax pattern smooth; venation camptodromous, midribs sparsely pubescent; 

palmately 5-7 lobed; median lobes ca 11.0 cm long, ca 3.0 cm wide, 

oblong-obovate, broadest ca 2/3 way up from the base, tapering into an acuminate apex, 

terminating in a bristle ca 0.75 cm long; lamina margin entire, base of lobe narrowly 

constricted, less than 0.25 cm across, sinuses between lobes very deep, lamina between 

the base of sinuses and petiole attachment point less than 0.5 cm wide; lowest lobes 

slightly smaller than, and more or less similar in shape, to median lobe. Inflorescence not 

seen. Fruit pedicels very short, less than 1.0 cm long, almost straight; capsules globose, ca 

2.25 cm long from base to apex, surface glaucous green, slightly tuberculate, ribs not 

prominent, apex rounded; fruit dehiscence septicidal. Seeds oblong, ca 1.5 cm long, 

carunculate end acutely pointed, ventral side with a prominent ridge along the middle, 

7This species is named for Dr. Grady Webster, Prof. of Botany, University of California, Davis. 

He is recognized internationally for his taxonomic studies in the Euphorbiaceae. He also collected the 

type specimen of this species.


>,4 ~ 

E~~ ;'0*2it'e 

[.?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~. 

----r- ~~ ~~~~~~~ ......-~ 

z ' 

M; ' 

' 

FIGe%' 

:- 

I 0 ~ ~ ~ \ 

/ --- _ - 

_ 

I is 'J?' _1/I - 

_? _ffid 

_ | ~~~,e 

--G 2. Mnh tmtpyl. A,hbt(oes56, N) ,la Rger 1 'Y. _w w = 

FIG 

,la Rgr 1,N).wb 

25. 

sterae. C, distribution; Manitruihot;D tomatophylla.A, 

D, leaf (Webster, Miller haite & (Roglers Miller 13074, 5160, NYV); DAV).


surface not shiny, occasionally verrucate, light brownish orange, with fine black 

mottlings, caruncle very small. Fig 25D. 

TYPE. Webster, Miller & Miller 13074: Mexico, Puebla: 8 miles SE of Izucar de 

Matamoros, 18 Aug 1962 (holotype, DAV; isotype, PUL). 

DISTRIBUTION. (Fig 25C). Known only from the type locality, alt 1500m, on 

limestone soil. 

The single collection representing this species is very distinct from all the other taxa. 

Further field studies are needed to outline its geographic boundaries and ecological 

adaptations and limits. Plants of Manihot websterae should be looked for in the region 

between Matamoros and Colima. This species can be easily distinguished from its close 

relatives M. michaelis and M. caudata by its characteristic nonpeltate leaf, extremely 

constricted lobe bases, and the short fruit pedicel. 

17. Manihot foetida (Humbolt, Bonpland & Kunth) Pohl, P1. Bras. Ic. et Descr. 1: 55. 

1827; Muell.-Arg. in DC., Prodr. 15(2): 1067. 1866; Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 28. 1910. 

Janipha foetida H.B.K., Nov. Gen et Spec. PI. 2: 84. 1817; Jatropha foetida Steudel, Nomencl. 

ed. 2. 1: 799. 1840. 

Tall trees. Young stems light reddish brown, pubescent; mature stems reddish brown, 

glaucous. Leaves alternate; stipules deciduous, pubescent; petioles ca 9.0 cm long, terete, 

pubescent, petiole attachment to lamina basal; lamina surfaces green to light green, pubescent, 

abaxial surface wax pattern smooth; venation camptodromous, 3 lobed; median 

lobes obovate, margin entire, ca 8.0 cm long, ca 0.5 cm wide, base of lobe more than 3.0 

cm across, apex abruptly tapering into an acute point, sinuses on either side of the 

median lobes narrowly cuneate; lowest lobes slightly smaller than median lobes, nonsymmetric, 

gently curved up, a major vein (in addition to the midrib) present along the base 

of the lowest lobe. Inflorescence a monoecious, axillary few flowered raceme, 4.0-7.0 cm 

long; peduncles and pedicels tomentose; bracts and bracteoles non vidi. Pistillate flowers 

restricted to the base of the inflorescence, borne on short pedicels (less than 1.0 cm long); 

tepals yellowish green, tomentose, about 1.0 cm long, cleft to base into 5 oblong tapering 

lobes; disc fleshy, entire; pistil ca 0.6 cm long, ovary tomentose, the trifid stigma short 

and moderately lobed and lobulate. Staminate flowers campanulate; tepals ca 1.1 cm 

long, yellowish green, tomentose, cleft 1/3 into 5 oblong obtuse lobes; disc fleshy, 10 

lobed; stamens 10, in 2 whorls of 5 each, superior whorl ca 0.7 cm long, inferior whorl ca 

0.6 cm long, filaments and anthers cream colored. Fruit pedicels ca 1.0 cm long; capsules 

3.0-4.0 cm long from base to apex, distinctly conical, narrowly tapering toward the apex. 

Seeds large, oblong, ca 2.5 cm long, surface shiny, dark brown, with inconspicuous black 

mottlings; caruncle very small. Fig 26B, C. 

TYPE. Humboldt s n Regni Mexicani, prope Mescala, Canada de Sopilote, Estolta, 

non vidi. 

DISTRIBUTION. (Fig 26A). MEXICO. Mexico: Hinton 3562 (GH) Limones, 6 Mar 1933; 

Hinton 4149 (A) Naranjo, Temascaltepec, 20 Jun 1933. Province Unknown/Uncertain: Sessd, Mocifo, 

Castillo & Maldonado 4229 (F). 

LOCAL NAME. Mercy marona (fide Pax, 1910). 

Manihot foetida has the largest seeds of any taxon in the genus. It is reported to have 

a foul odor. 

18. Manihot crassisepala Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 44): 

28. 1910.


1 

FI 

\ 

M 26 anBot fotd.A B, leaf" itiui (e atilo&Mlond29 

s n s oiio d , 

FIG 26. Manibot foetida. A, distribution; B, leaf (Sesse', Mocinzo, Castillo & Maldonado 4229, 

F). M. crassisepala D, distribution.


Trees. Young stems pubescent, grayish brown with reddish tinge; mature stems 

reddish brown, glaucous. Leaves alternate, stipules deciduous, pubescent; petioles ca 10.0 

cm long, terete, glabrous, in some, pubescent, petiole attachment to lamina basal; lamina 

green or pale green, glabrous, abaxial surface wax pattern smooth; venation camptodro- 

mous, 3 lobed; median lobes obovate, margin entire, ca 10.0 cm long, ca 5.0 cm wide, 

base of lobes more than 2.5 cm across, apex abruptly tapering to an acute point, sinuses 

on either side of the median lobes narrowly cuneate; lowest lobes slightly smaller than 

median lobes, nonsymmetric, gently curved up, a major vein (in addition to the midrib) 

present along the base of the lowest lobe. Inflorescence a monoecious, few flowered, 

axillary raceme, 4.0-8.0 cm long, peduncles and pedicels pubescent; bracteoles setaceous, 

pubescent, margin smooth; bractlets setaceous, pubescent. Pistillate flowers restricted to 

the base of the inflorescence, borne on short pedicels, less than 1.5 cm long; tepals 

yellowish green, very sparsely pubescent, ca 1.0 cm long, cleft to base into 5 oblong 

tapering lobes; disc fleshy, entire; pistil ca 0.6 cm long, ovary glabrous, the trifid stigma 

short, moderately lobed and lobulate. Staminate flowers campanulate; tepals about 1.1 

cm long, yellowish green, very sparsely pubescent, cleft 1/3 into 5 oblong-obtuse lobes; 

disc fleshy, 10 lobes, stamens 10, in 2 whorls of 5 each, superior whorl ca 0.7 cm long, 

inferior whorl ca 0.6 cm long, filaments and anthers cream colored. Fruit pedicels ca 1.5 

cm long; capsules ca 2.25'cm long from base to apex, distinctly conical, narrowly tapering 

towards the apex, surface finely rugose, apex pointed; fruit dehiscence loculicidal, the 3 

commissural sutures not disjoining. Seeds non vidi. Fig 27A, B. 

TYPE. Kerber 185: Mexico: 1880 (photos of syntype, F, GH, NY). 

DISTRIBUTION. (Fig 26D). Mexico, in Mexico and Morelos, alt ca 900-1300m. MEXI- 

CO. Mexico: Hinton 3564 (GH) Limones, Temascaltepec, 3 June 1933. Morelos: Pringle 11339 (GH) 

near Cueravaca, Lime Stone Hills, Yautepec, 24 Oct 1902. 

A \. . J ? 

ji L 

A I 

2 IB 3 

64,GH. 

FIG 27. Manihot crassisepala. A, leaf (Pringle 11339, GH); B, inflorescence (Hinton 3564, GH). 

/,


5 

95 90 15 10 ?5 70 6 6GO 5 50 

-------- 

--~~~1~~~ ~ F16. 211- 

10I 

11 1e _ 

12_ 

13 

11 

15 

18- 

FIG 28-A. Dendrogram of relationships of the South American sections of Manibot. The simi- 

larity values are shown decreasing from left to right across the top of the Figure. Sectional numbers 

are given on the vertical axis. 

Manihot crassisepala is very similar to M. foetida. The insufficient representation of 

M. crassisepala and M. foetida in this study precludes confident delimitation of these two. 

It may be necessary to re-examine the taxonomic status of M. crassisepala, especially its 

relationship with M. foetida, when sufficient material and field data become available. 

In the original description of this species, the type local is reported as Colima. 

Photographs of the type are available, but it has not been possible to verify the label data 

with respect to the locality of collection. It seems highly unlikely that the range of this 

species extends to Colima, however further field studies would be necessary to delineate 

its complete range. 

The note in the original description of this species indicates that the mature seeds are 

edible. 

4. Manihot sect Heterophyllae Pax emend Rogers & Appan. 

Manibot sect Grandibracteatae Pax subsect Coerulescentes Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 29. 1910 pro parte 

Manibot sect Grandibracteatae Pax subsect Rigidulae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 35. 1910 pro parte. 

Manibot sect Parvibracteatae Pax subsect Elatae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

55. 1910 pro parte. 

Manihot sect Parvibracteatae Pax subsect Tristes Pax in Engler, Pflanzenreich IV. 147(Heft 44): 


Manibot sect Parvibracteatae Pax subsect Langsdorffianae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 65. 1910 pro parte. 

Manibot sect Heterophyllae Pax subsect Carthaginenses Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 80. 1910 pro parte. 

Plants caulescent; low to medium shrubs, occasionally small trees; leaves widely 

spaced on stem; petiolate, petiole attachment mostly basal, rarely peltate; lamina

78 


membranaceous, deeply lobed, lobes usually entire or shallowly pandurate. Inflorescence 

a monoecious panicle, bracts and bracteoles foliaceous to setaceous, margins entire. 

TYPE. Manibotjaniphoides Muell.-Arg. 


Key to the Species of Sect Heterophyllae 

1. Bracts and bracteoles setaceous, less than 0.25 cm wide. 

2. Panicle with one principal peduncle; flowers distantly positioned in inflorescence (except 

21. M. surinamensis); buds of staminate flowers ovoid-ellipsoid or slightly turbinate. 

3. Abaxial lamina surface wax pattern pusticulate or reticulate (unknown in M. zehntneri); 

panicles less than 10.0 cm long, sparsely branched; staminate buds usually less than 

0.75 cm long; petiole attachment peltate (except in 19. M. zehntneri and 22a. M. tristis 

subsp tristis). 

4. Leaf lobes oblong, more than 12.0 cm long. 

19. M. zebntneri. 

4. Leaf lobes linear or obovate, less than 12.0 cm long. 

5. Leaf lobes linear, less than 1.0 cm wide; venation most frequently craspedodromous, 

sometimes camptodromous. 

6. Inflorescence less than 5.0 cm long. 

20. M. marajoara. 

6. Inflorescence more than 5.0 cm long. 

21. M. surinamensis. 

5. Leaf lobes obovate or elliptic, more than 1.0 cm wide; venation camptodromous, 

never craspedodromous. 

22. M. tristis. 

3. Abaxial lamina surface wax pattern smooth; panicles more than 10.0 cm long; well 

branched; staminate buds usually more than 0.75 cm long; petiole attachment basal. 

7. Leaf lobes and petioles densely pubescent. 

23. M. janiphoides. 

7. All parts glabrous. 

8. Leaves more than 7 lobed, lobe bases more than 0.25 cm wide. 24. M. grahami. 

8. Leaves 3-5 lobed, rarely 7, lobe bases constricted, less than 0.25 cm wide. 

25. M. inflata. 

2. Panicle with several principal peduncles arising from one basal point; flowers clustered at 

terminal region of peduncles; staminate buds bifusiform. (Flowers of M. quinquefolia non 

vidi.) 

11. Leaf lobes more than 8.0 cm long, more than 2.5 cm wide; lobe bases usually more 

than 0.25 cm wide; most parts pubescent. 

12. Inflorescence moderately branched; flowers more than 0.75 cm long, ovary 

tomentose; leaf lobes entire. 26. M. pilosa. 

12. Inflorescence profusely branched; flowers less than 0.75 cm long; ovary glabrous; 

leaf lobes pandurate. 

27. M. corymbiflora.. 

11. Leaf lobes less than 8.0 cm long, less than 2.5 cm wide; lobe bases less than 0.25 cm 

wide; all parts glabrous. 

13. Lobe bases stiff, lobes held horizontally. 

28. M. leptopoda. 

13. Lobe bases flaccid, lobes with a tendency to droop. 

29. M. quinquefolia. 

1. Bracts and bracteoles foliaceous, more than 0.25 cm wide. 

14. Petiole attachment peltate; stipules foliaceous, more than 1.5 cm long. 30. M. condensata. 

14. Petiole attachment basal; stipules setaceous, less than 1.5 cm long. 

15. Inflorescence more than 8.0 cm long; profusely branched; leaf lobes more than 3.5 

cm wide; petiole-midrib junction with 2 minute lobules. 

16. Leaves more than 12.5 cm long; lobe bases more than 0.5 cm wide; lamina 

densely pubescent. Capsule surface smooth. 

31. M. jolyana. 

16. Leaves less than 12.5 cm long; lobe bases less than 0.5 cm wide; lamina 

moderately pubescent; capsule winged. 

32. M. handroana. 

15. Inflorescence less than 8.0 cm long, sparsely branched. Leaf lobes less than 3.5 cm 

wide; petiole-midrib junction without lobules. 33. M. pohlii. 

19. Manihot zehntneri Ule, Bot. Jahrb. Syst. 114: 10. 1914; Pax & Hoffmann in Engler, 

Pflanzenreich IV. 147(Heft 68): 48. 1919; Pax & Hoffmann in Engler, Pflanzenreich 

IV. 147(Heft 85): 195, 196. 1924. 

TYPE. Zehntner, L. 598: Brasil, Bahia: Riacho de Santa Anna, 21 Nov 1912 (photo 

of syntype, F, NY). 

Erect shrubs, ca 1.5 m tall. Leave alternate; petioles ca 8.0 cm long; lamina 

nonpeltate; palmately 5 lobed; median lobes oblong, ca 15.0 cm long, apex acute.

m' A-CC,JKr\//z X- 

/ \ A 

=rs 

r.+- -.. ./.' 

FIG 29 Manot sect Heteroya e . 

WH f~~~~~~. t. X ,., 

~I 0-xE ;i 0 ( 

FIG 29 Man~ n~ihot sec Heterpbylla.1 Gegapia rage 

~,ir 

/t~~~~~~~~~~~~~~~ 

.--I~~~~~~~~~~ 

f ;~~~i -? 

'~~~~~~~~~f": ~ ~ ~


Inflorescence a monoecious panicle, ca 9.0 cm long, sparsely branched. Bracteoles 

setaceous. Pistillate flowers restricted to the base of the upper half of the panicle. 

Staminate buds ovoid ellipsoid. Fruits non vidi. Seeds non vidi. 

DISTRIBUTION. (Fig 30A). Growing wild near Riacho de Santa Anna, Bahia, Brasil. 

LOCAL NAMES. Manicoba (Zehntner 598). 

The only representation of this taxon in this study is a photo of the type and the 

inadequate representation precluded an accurate evaluation of this Ule species. The leaf 

and inflorescence structure closely resemble Manihot esculenta, however, the use of this 

species as a source of inferior rubber, as reported by Ule (1914) indicates that species 

may be affiliated to members of the sect Glaziovianae. 

20. Manihot marajoara Chermonte de Miranda apud Huber, Bol. Mus. Paraense Hist. Nat. 

5: 120. 1908. 

Slender shrubs, ca 0.5 m tall. Young stems glabrous, dark brown; mature stems 

glabrous, ca 1.0 cm in diam. Leaves alternate; stipules caducous; petioles ca 8.0 cm long, 

glabrous; lamina narrowly peltate, width between basal edge of lamina and petiole-lamina 

junction less than 0.25 cm, abaxial and adaxial surface of lamina glabrous, abaxial lamina 

surface wax pattern tuberculate; venation frequently craspedodromous, occasionally 

camptodromous; palmately 5 lobed; median lobes linear, ca 8.0 cm long, less than 0.6 cm 

wide, lobes frequently with 2 falcate lobules at the base, one on either side, apex acute, 

base of lobes ca 0.3 cm wide, lamina between base of sinus and petiole-lamina junction 

less than 0.5 cm wide, lowest lobes ca half as long as median lobes, more or less similar in 

shape to median lobes. Inflorescence a monoecious, short, panicle, ca 3.0 cm long, few 

flowered, all parts glabrous, bracteoles and bractlets setaceous. Pistillate flowers non vidi. 

Staminate buds ovoid ellipsoid, pedicel 0.75 cm long, tepal 0.5 cm long, cleft nearly 1/3 

way down into 5 lobes. Fruits non vidi. Seeds non vidi. Fig 30C. 

TYPE. Huber, J. 2792: Brasil, Para: Marajo, 5 Jul 1902 (syntypes, PUL, RB; photo 

of syntype, F). 

DISTRIBUTION. (Fig 30B). Brasil, State of Para, Amapa Territory. BRASIL. Amapa 

Territory: Ducke 4808 (F, MG) Campo, 27 Jun 1904. Para: Huber 9346 (MG) Jutinha, Marajo, 

Apr 1908. 

LOCAL NAMES. Mandioca dos Indios (Ducke 4808) Maniva do campo (Huber 

2792). 

This species is most closely related to M. surinamensis. The 'skyline' (Fig 28)8 

indicates that the 2 species have a high similarity. 

21. Manihot surinamensis Rogers & Appan, sp nov 

Frutices gracili. Folia 5-7 lobatus; lobi mediani linearis, ca 10 cm longa, deminutus 

apicem versus. Inflorescentia panicula monoecious; ca 9.0 cm longa, flores fasciculatus 

versus pedunculorum terminalis extrema. Pistillatus flores ca 0.5 cm longae. Staminati 

gemmae ovoideus-ellipsoideus, flores parvi, minus quam 0.5 cm longae, fructi non vidi. 

Slender shrubs. Young stems glabrous, light brown; mature stems glabrous, dark 

brown. Leaves alternate; stipules persistent, setaceous, less than 0.5 cm long; petioles ca 

8.0 cm long, glabrous; lamina narrowly peltate, width between the basal edge of lamina 

and petiole-lamina junction less than 0.25 cm, abaxial and adaxial surfaces glabrous, 

abaxial surface wax pattern reticulate; venation mostly craspedodromous, occasionally 

camptodromous, veins glabrous; palmately 5 lobed, occasionally 7; median lobes linear 

with 2 falcate lobules near the base of the lobe, ca 10.0 cm long, gradually tapering 

8 Fig. 28 is a fold-out chart at end of book.


- 

FIG 30. Manihot zehntneri. A, distribution. M. marajoara. B, distribution; C, leaf (Huber 2792, 

RB). M. surinamensis. D, distribution. 

5


towards the apex, apex acuminate, base of lobes less than 0.5 cm wide, lamina between 

base of sinus and petiole-midrib junction less than 0.5 cm wide; lowest lobes alternate 

usually ca 1/4 as long as median lobes. Inflorescence a monoecious, medium sized, 

panicle, ca 9.0 cm long, flowers clustered at the terminal regions of peduncles, lower half 

of the rachis usually devoid of flowers, all parts glabrous, bracteoles and bractlets 

setaceous, ca 1.0 cm long. Pistillate flowers restricted to the base of the upper half of the 

inflorescence, pedicel ca 1.0 cm long, tepal less than 0.5 cm long, ovary subglobose. 

Staminate buds ovoid-ellipsoid, flowers small, less than 0.5 cm long, cleft ca 1/3 down 

into 5 lobes, stamens 10 arranged in 2 whorls of 5 each. Fruits non vidi. (Fig 31 A, B). 

TYPE. Rombouts 464: Suriname: upper Sipaliwini R., 9 Feb 1936 (holotype, U; 

isotypes NY, MO). 

DISTRIBUTION. (Fig 30D) Venezuela, Guyana and Suriname, in savannas. Specimens 

doubtfully assigned to Manihot surinamesis: VENEZUELA. Guayana, Cardona 1158 (F, NY, US, 

VEN), sabanas del Rio Uriman, afluente del Caroni, alt 400 m, May 1945. GUYANA. Lanjouw & 

Donselaar 789 (U), Rupununi Savanna, in directionem borealem de montibus Kanuku, 16 Feb 1959. 

SURINAME. Rombouts 409 (U-2, US), upper Sipaliwini R., Camp XVIII near 4 Gebroeders. 

Jennings (1959) mentions the use of Manihot melanobasis in breeding experiments 

with cassava in East Africa. It is clear that there was a misdetermination in case of the 

actual plants that were used in the experiments, because we have determined from the 

type material that M. melanobasis is a synonym of M. esculenta. From the description of 

the plants actually used by Jennings and from the geographic region from which the 

breeding material was collected we believe that M. surinamensis was the species used. It is 

unfortunate that documenting herbarium specimens of materials used by plant breeders 

are seldom collected and as a result the breeders work cannot be affirmed. 

22. Manihot tristis Mueller von Argau in Martius, Fl. Bras. 11(2): 449. 1874. 

Shrubs, to 3.0 m tall. Young stems pubescent or glabrous; mature stem glabrous. 

Leaves alternate; stipules persistent or caducous, margin serrate; petioles 4.0-9.0 cm long, 

glabrous or pubescent; lamina palmately 3-5 lobed, glabrous or pubescent, abaxial 

surface wax pattern reticulate; venation camptodromous, median lobes obovate, usually 

5.0-10.0 cm long, occasionally longer, ca 2.5 cm wide, apex acute, base of lobe narrow 

(less than 0.5 cm) to wide (more than 0.5 cm). Inflorescence a monoecious panicle, less 

than 8.0 cm long; bracteoles and bractlets setaceous. Pistillate flowers restricted to the 

base of the panicle, pedicel ca 2.0 cm long, tepal 0.6 cm long, deeply cleft into 5 lobes, 

ovary subglobose. Staminate buds ovoid-ellipsoid, 0.5 cm long, cleft ca 1/3 way down 

into 5 lobes, stamens 10, in 2 whorls of 5 each. 

DISTRIBUTION. (Fig' 31C). Venezuela, Amazonas Territory; Suriname; Brasil, Ama- 

pa and Roraima Territories. One subspecies, M. tristis subsp saxicola, is found on granitic 

domes, and in this respect, is similar to M. leptopoda, (species No. 28) which grows on 

granitic outcrops in the Rio de Janeiro region. 

The "skyline" (Fig 28) indicates that the plants here related as subspecies should be 

kept distinct. It is at this point, however, that the taxonomist's judgment must be 

exercised, and in this example, we overruled the computer-made relationship. Actual 

reexamination of the specimens, and the geographic distribution (Fig 31C) indicated the 

relationship here given. 

Key to the Subspecies of Manihot tristis 

1. Petiole attachment basal; leaf lobe base very narrow, less than 0.3 cm wide. 

22a. M. tristis subsp tristis. 

1. Petiole attachment peltate; leaf lobe base more than 0.3 cm wide. 

2. All parts glabrous. 

22b. M. tristis subsp saxicola. 

2. Leaf lobes, petioles, young stems, stipules, bracts, tepal etc pubescent. 

22c. M. tristis subsp surumuensis.


NI~* 

1@'I 

~^ 1 B t~ 

FIG 31. Manibot surinamensis. A, leaf (Rombouts 409, U); B, inflorescence (Rombouts 464, 

MO). M. tfistis. C, distribution, dots represent subsp tristis, the letter 0 represents subsp surumuensis, 

and the letter A represents subsp saxicola. M. tristis subsp tristis. D, leaf (Williams 15893, VEN) 

/


22a. Manihot tristis Mueller von Argau subsp tristis 

Manihot tristis Muell.-Arg. in Martius, Fl. Bras. 11(2): 449. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 59. 1910. 

Manihot orinocensis Croizat, Jour Arnold Arb. 24: 169. 1943. Type. Williams, L. 13132 

(syntypes, F, S, US, VEN). 

Erect shrubs, ca 2.0 m tall, rarely as tall as 4.0 m. All parts of the plants glabrous. 

Young stems dark brown, with a thin whitish bloom. Lamina nonpeltate, usually 3 lobed, 

sometimes 5; median lobes obovate-elliptic, rarely slightly pandurate, usually ca 7.0 cm 

long, base of the lobes narrowly constricted, less than 0.3 cm wide, lamina between base 

of sinus and petiole-midrib junction less than 0.3 cm wide, lowest lobes nonsymmetric 

and slightly curved up. Inflorescence few flowered, usually with ca 10 flowers; tepal 

purplish tinged. Fruits small, 1.0 cm long from base to apex, apex rounded, surface 

without prominent ribs, dehiscence septicidal. Seeds small, 0.8 cm long, oblong caruncle 

not prominent. Fig 31D. 

TYPE. Spruce, R. 3604: Venezuela, Amazonas Territory: Prope Maypures, ad 

flumen Orinoco, Jun 1854 (syntypes, BM, G, K-2, P, W; photo of syntype, F, NY). 

DISTRIBUTION. (Fig 31C). Venezuela, Amazonas Territory, on granitic outcrops 

near Puerto Ayacucho. VENEZUELA. Amazonas Territory: Gaillard 124 (P) plantes des Boros 

del'Orenoque, Jul 1887; Williams 13132 (F, S, US, VEN) Sobre las lajas graniticas en los alrededores 

de Puerto Ayacucho, 27 May 1940; Williams 15893 (F-2, US, VEN) en las lajas graniticas de Puerto 

Ayacucho, 25 Jun 1942. Williams 16095 (A, F, US, VEN) las lajas graniticas y espuestas de Puerto 

Ayacucho, Jul 1942. 

LOCAL NAME. Yuquilla (Gaillard 124). 

22b. Manihot tristis Mueller von Argau subsp saxicola (Lanjouw) Rogers & Appan, stat nov 

Manihot saxicola Lanjouw, Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 69: 544. 1939; Recueil 

Trav. Bot. Neerl. 36: 544. 1939. 

Erect shrubs, 1.0-3.0 m tall. All plant parts glabrous. Roots with intermittent 

globose tuberous swellings, ca 7.0 cm in diam (Shulz & Donselaar 10593) resembling 

tuberous roots of Manihot esculenta (Pires & Westra 48837). Young stems purplish 

tinged, devoid of any bloom. Stipules semifoliaceous, 1.25 cm long, 0.2 cm wide, 

persistent, margin laciniate; lamina narrowly peltate, width between basal edge of lamina 

and petiole-lamina junction 0.2 cm; abaxial lamina surface pruinose; palmately 5 lobed, 

occasionally with 2 more smaller lobes, ca 8.0 cm long, ca 2.5 cm wide, obovate, rarely 

pandurate, base of lobes more than 0.3 cm wide, lamina between base of sinus and 

petiole-midrib junction more than 0.3 cm. Inflorescence a moderately branched panicle 

with ca 25 flowers; peduncles purplish tinged; tepal yellowish green. Fruits ca 1.25 cm 

long, blackish brown, apex rounded, slightly winged, dehiscence septicidal. Seeds 1.0 cm 

long, oblong, with a moderately prominent caruncle. Fig 32A, B, C. 

TYPE. Lanjouw, J. 955: Suriname: Monte dicto voltzberg ad saxis graniticus, 23 

Sept 1923 (holotype, U; isotypes, F, MO, NY, U-2, US). 

DISTRIBUTION. (Fig 31C). Suriname and Brasil, Amapa Territory.SURINAME. B. W. 

6331 (NY, U) Voltzberg, 7 Aug 1923; Landb. 922 (U) Voltzberg, 1938; Richard s n (P) 

Guyanansi-Antillanum; Schulz & Donselaar 10593 (BBS -2, U-3), upper Coppename River, Voltzberg 

Top 2, 9 Feb 1965; Stahel 107 (DS, F, NY, U, US) Agricultural Experimental Garden, Paramaribo, 15 

Feb 1941; Versteeg 419 (U) upper Litanie River, Dec 1903; Versteeg 801 (U) upper Tapanahoni 

River, Sep 1904. BRASIL. Amapa Territory: Pires & Westra 48837 (NY) Rio Oiapoque, near Mt. 

Carupine, 14 Oct 1960. 

LOCAL NAMES. Boesi-ingi-kasabu (Schulz & Donselaar 10593) Maynoc (Richard s n) 

wilde cassava (Lanjouw 955).


f l': 

c~~~~r 

_ 

r 

"~~~~~~~~~~~~~~~~~0 

it -\ 

\ sE * ~~~~~~~~~~~~~( 

_ _rE X 

_~~~~~~~~~~~~~I 

__ W oT;t \ o _~~ 

! F- =~~~~~~~~~~~~~~e 

Ei *t 

w w\ e . 

n 

FIG 32 Maniht 

trisis subp saxcola. , habi (Schuz & Doselaa 10593 BBS).B, lea (Lan 

jow95 ) ,ifoecneScuz&Dnearl53 ) .tntssbpsrmess ,la 

(Ue794). 

FIG 32. Manihot tristis subsp saxicola. A, habit (Schulz & Donselaar 10593, BBS). B, leaf (Lan- 

jouw 955, U). C, inflorescence Schulz & Donselaar 10593, U). M. tristis subsp surumuensis. D, leaf 

(Ule 7944, K).


This subspecies shares with subspecies tristis the interesting habitat preference of 

granitic outcrops. To our knowledge, only one other species of the genus, Manihot 

leptopoda, is found on this type of substrate. These should, therefore, be important in 

breeding programs with M. esculenta to improve culture on soils of granitic origin. 

M. tristis subspecies saxicola, first described as a separate species in 1939, by 

Lanjouw, was brought into cultivation in Buitenzorg shortly after the species was 

discovered. The growth pattern under cultivation is much more erect than the typical 

pattern as a wild plant (see Lanjouw 1939 photographs). Lanjouw reported 2.3% 

"albumin" in the roots on a fresh weight basis. Bolhuis (1953) used this plant in breeding 

programs in Java in attempts to increase protein content of the roots of M. esculenta. 

Roots of F1 seedlings of the cross M. esculenta X M. tristis subsp saxicola produced 

approximately 2% protein, but in clones propagated from these seedlings, protein content 

fell back to typical levels (ca 1%). 

22c. Manihot tristis Mueller von Argau subsp surumuensis (Ule) Rogers & Appan, stat 

nov 

Manibot surumuensis Ule, Bot. Jahrb. Syst. 114: 12. 1914. 

Shrubs. Young stems and mature stems pubescent. Stipules caducous; petiole 

pubescent; lamina narrowly peltate, width between basal edge of lamina and petiolelamina 

junction 0.2 cm; abaxial lamina surface pubescent; palmately 3 lobed, ca 8.0 cm 

long, ca 2.5 cm wide, obovate, lobe base more than 0.3 cm wide, lamina between base of 

sinus and petiole-midrib junction greater than 0.3 cm. Inflorescence a sparsely branched 

panicle, usually with ca 10 flowers, peduncles, pedicels and tepals pubescent. Fruits and 

seeds non vidi. Fig 32D. 

TYPE. Ule, E. 7944: Brasil, Roraima Territory: Campos bei der Serra de Pracaua, 

Surumu, Feb 1909 (syntypes, K, L, MG, photo of syntype F, NY) 

23. Manihot janiphoides Mueller von Argau in Martius, Fl. Bras. 11(2): 480. 1874; Pax in 

Engler, Pflanzenreich IV. 147(Heft 44): 80. 1910. 

Shrubs, ca 1.5 m tall, branches with a tendency to droop. Roots slender, not tuberous. 

Young stems obtuse-angled in cross section, sparsely pubescent; mature stems 

glabrous, grayish olive green (5 GY 3/2). Leaves alternate; stipules caducous; petioles 

10.0-15.0 cm long, pubescent, strong yellow-green (7.5 GY 7/9), petiole attachment to 

lamina basal, nonpeltate; adaxial surface of lamina moderate yellow-green (7.5 GY 5/7), 

abaxial surface wax pattern smooth; venation camptodromous, veins on abaxial lamina 

surface pubescent; palmately 5 lobed, occasionally with 2 more smaller lobes; median 

lobes rhomboid, entire, rarely slightly pandurate, ca 10.0 cm long, ca 3.0 cm wide, apex 

acuminate, base angustate to brevi-angustate, base of lobes less than 0.3 cm wide, width 

of lamina between base of sinus and petiole-lamina junction less than 0.5 cm; lowest lobes 

attenuate, ca 1/3 as long as median lobes. Inflorescence a monoecious, many branched 

panicle, ca 15.0 cm long; bracteoles and bractlets setaceous, sparsely pubescent, with 

entire margin; pistillate flowers usually restricted to the base of the upper 2/3 of the 

inflorescence, pedicels sparsely pubescent, ca 1.5 cm long, tepal yellowish green or with 

purplish pigmentation, ca 1.0 cm long, ovary subglobose, glabrous. Staminate buds 

ovoid-ellipsoid, tepal ca 1.0 cm long, sparsely pubescent, yellowish green or with purplish 

pigmentation, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each, the 

superior whorl 0.8 cm long, the inferior 0.6 cm long. Fruits and seeds non vidi. 


TYPE. Warming s n: Brasil, Minas Gerais: prope Lagoa Santa (syntype, P). 

DISTRIBUTION. (Fig 3 3A). Brasil, in the states of Minas Gerais and Sao Paulo. 

BRASIL. Minas Gerais: Warming 1503 (F, G) Lagoa Santa. Sao Paulo: Cruz 116 (SP) Sao Carlos, 17


/ 'J 

an. , t _ I 

FIG 33. Manibot janiphoides. A, distribution; B, habit (Rogers 357, NY); C, leaf (Viegas s n, 

SP); D, inflorescence (Viegas s n, SP).


Nov 1965; Rogers 357 (NY) Campinas, cultivated in Grounds of Inst. Agron., 8 Feb 1961; Viegas s n 

(SP) Campo Experimental Instituto, Campinas, cultivated, 11 Dec 1941. 

The closest affinity of Manihot janiphoides is M. esculenta. The most distinctive 

difference between these two is the slender, nontuberous roots of the former. 

24. Manihot grahami Hooker, Icon. PI. 6. t. 530. 1843. 

Janipha loeflingii var multifida Graham, Edinb. Philos. Jour. 29: 172. 1840; Manihot loeflingii 

var multifida (Graham) Muell.-Arg. in DC., Prodr. 15(2): 1062. 1866; Manihot palmata var 

multifida Muell.-Arg. in DC., Prodr. 15(2): 1062. 1866; Manihot dulcis (J. F. Gmelin) Pax 

var multifida (Graham) Pax in Engler, Pflanzenreich IV. 147(Heft 44): 72. 1910. Type. 

Tweedie s n (syntypes, K-3). 

Manihot tweedieana Muell.-Arg. in Martius, Fl. Bras. 11(2): 450. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 56. 1910. Type. Tweedie s n (isotype, G). 

Manihot tweedieana Muell.-Arg. var lobata Chodat & Hassler, Bull. Herb. Boissier 2(5): 673. 

1905. Type. Hassler 5413a (syntypes, BM, F, G-3, K, MO, NY, P, S, W), Hassler 5413b 

(syntypes, BM, G-6, K, NY, P, S, W). 

Manihot tweedieana Muell.-Arg. f. nana Chodat & Hassler, Bull. Herb. Boissier 2(5): 673. 1905. 

Type. Hassler 5407 (syntypes, G-3). 

Manibot lobata (Chodat & Hassler) Pax in Engler, Pflanzenreich IV. 147(Heft 44): 82. 1910. 

Type. Hassler 5407 (syntypes, G-3), Hassler 5413a (syntypes, BM, F, G-3, K, MO, NY, P, S, 

W). Hassler 5413b (syntypes, BM, G-6, K, NY, P, S, W). 

Manihot enneaphylla Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 196. 1924. 

Type. Ekman 496 (syntypes, S-2). 

Arborescent shrubs to low trees with a strong odor of HCN in all parts, to 7.0 m tall, 

forming an umbrella-like dense canopy of foliage at the top. Roots not tuberous, 

epidermis rough, dark brown, subepidermis white. Trunk ca 30.0 cm in diameter at base; 

bark smooth, reddish brown, ca 0.6 cm thick, peeling readily from trunk, latex in small 

quantity, light yellowish white. Young stems obtuse-angled in cross section, glabrous, 

moderate olive green (7.5 GY 4/4), internal stem color of younger stems brilliant yellow 

green. Young shoots strong yellow green (5 GY 6/8). Leaves alternate, stipules long, often 

to 3.0 cm in length, filiform, glabrous, caducous; petioles usually very long, ca 20.0 cm in 

length, terete, glabrous strong yellow green (5 GY 7/10), petiole attachment to lamina 

basal, nonpeltate; lamina greenish without any purplish pigmentation, glabrous, abaxial 

surface wax pattern smooth, venation camptodromous, veins on the adaxial lamina 

surface conspicuous, bright yellowish, glabrous; palmately 7-11 lobed; median lobes 

oblong pandurate, rarely entire, gradually widening from a narrow base to a prominently 

dilated apical region which abruptly narrows down and terminates in an acuminate apex, 

usually as long as 15.0-20.0 cm, base of lobes ca 0.5 cm wide, width between base of 

sinus and petiole-lamina junction ca 1.0 cm, lowest lobes more or less similar in outline as 

median lobes but smaller. Inflorescence a monoecious profusely branched panicle, often 

as long as 30.0 cm, all parts glabrous; bracteoles and bractlets setaceous. Pistillate flowers 

restricted to base of upper 2/3 of the inflorescence, pedicels ca 2.0 cm long, tepal 1.25 

cm long, cleft to base into 5 lobes, disc orange-red, ovary subglobose, glabrous. Staminate 

buds ovoid ellipsoid, flowers large, tepal 1.25 cm long, brilliant yellow green (2.5 GY 9/8) 

with maroon dots internally at base, glabrous, disc bright orange, 10 lobed, stamens 10, in 

2 whorls of 5 each, the superior whorl 0.9 cm long, the inferior 0.7 cm long. Fruits 

round, not winged, with triangular indentations in stigmatic end, 1.8 cm long from base 

to apex, 1.9 cm from side to side, dehiscence septicidal. Seeds oblong, 1.2 cm long, with 

rib-like projections along the lateral edges, caruncle moderately prominent. Fig 34B, C, D. 

TYPE. Tweedie s n: Brasil, Parana: Woods of the Parana (syntypes, K-3). 

DISTRIBUTION. (Fig 34A). This species is native to southeastern Brasil, northern 

Argentina, Paraguay and Uruguay. It has been introduced to the southeastern United 

States, and now is found frequently naturalized from Louisiana eastward to Florida. It 

has been confused with M. esculenta in this area. It has, on occasion, been raised in


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:. '. E. ~ .... . . . :" 

(Rgr,Tirt&Ree48 Y;D nlrecne(oes bee es 9,N) 

FIG 34. Manihot grabami. A, distribution (native range only); B, habit (Rogers 498, NY); C, 

leaf (Rogers, Tbieret & Reese 498, NY); D, inflorescence (Rogers, Tbieret & Reese 498, NY).


conservatories, and there again, mislabled M. esculenta. BRASIL. Goias: Pohl Icon tab. 25 

(W). Minas Gerais: Regnell 408 (S). Sao Paulo: Castro & Kiehl s n (SP) Capoeira, Cunha, Dec 1938; 

Labouriau 187 (SP) Instituto de Botanica, Sio Paulo, cultivated, 17 Dec 1965; Viegas s n (SP) Campo 

Experimental Instituto Campinas, cultivated, 10 Dec 1941. Rio de Janeiro: Glaziou 14244 (F, K, P, 

US) Environs of Rio de Janeiro, 5 Jan 1883; Glaziou 8321 (P) Serra da Bocaina, 9 Jan 1876;Goes & 

Dionisio 560 (RB) Petropolis Correas, Sep 1943. Parana: Dusen 11158 (S) Therezina in Capoeira, 21 

Jan 1911; Dusen 16822 (S) 14 Mar 1915; Dusen 17597 (S) Tres Barras in Silva Arboribus, 30 Jan 

1916; Dusen 7812 (F, GH, MO, NY, S) Iraty, in Capoeira ad Rivulum, 28 Feb 1909; Tessmann 6012 

(SP) Rolandia, Jan 1937; Tweedie s n (BM-2, G) Santa Catarina: illegible s n (SP) Rio do Peixa, 31 Dec 

1927; Smith & Klein 11024 (US) Mun. Cacador-Curitibanos, ruderal, 14 km SE of Cacador on the 

Road to Lebbn Regis, 8 Feb 1957;Smith & Klein 11711 (NY, US) Ruderal, 41 km S of Mun. Dionisio 

Cerqueira, 23 Feb 1957; Smith & Klein 13076 (TEX) Mun. Xaxim, ruderal, 2 km N of Xaxim, 9 Nov 

1964; Smith & Reitz 9083 (F, S, US) ruderal, W of Cacador on Road to Taquara Verde, 23 Dec 1956; 

Smith & Reitz 9647 (US) 50 km W of Rio Capetinga on the Road to Dionisio Cerqueria, 30 Dec 1956; 

Smith & Reitz 9702 (US), S of Cedro, 40 km from D.C., 1 Jan 1957; Smith & Reitz 9896 (UC, US) 24 

km W of Joacaba on the Road to Jabora, 4 Jan 1957. Rio Grande Do Sul: Cruz 119 (SP) Proximo a 

Ponte Do Rio Pelotas, Divisao de Santa Catarina, 5 Jan 1965; Dutra 366 (S-2) Inter Conventos et 

Forquetinha, Dec 1899; Malme 1272 (S-2) Inter Santa Maria et Pinhal, 27 Jan 1902; Rambo 45076 

(MO, S, W) Kaffeschneis Pr. Dois Irmaos, 26 Dec 1949; Province Unknown/Uncertain: illegible s n (P) 

Aug 1872; Markgraf & Brade 3746 (RB) Itatiaia, Nov 1938. PARAGUAY. Amambay: Hassler 10680 

(A) In Altaplanitie et Declivibus, Sierra de Amambay, Nov 1907. Capital District: Rojas 4252 (A) 

Cordillera de Amambay, Feb 1922; Rojas 4753 (A) Colonia Independencia, Guaira, Feb 1924; Rojas 

6469 (A) Pedro Juan Caballero, Sierra de Amambay, Nov 1933. Guaira: Balansa 1713 (K, P-2) Villa 

Rica, 14 Feb 1876; Balansa 1713A (P) Foret de Yaguaron, Feb 1877; Balansa 1714 (K, P) Foret de 

Caaguazu, 6 Nov 1874; Jorgensen 4002 (DS, F, MO, NY, S, US) Villa Rica, Feb 1928. Misiones: 

Bertoni 47971 (NA) Puerto Bertoni, 22 Aug 1919. Province Unknown/Uncertain: Hassler 5407 (G-3) 

In Altoplanitie et Decliviis, Sierra de Maracayu, Nov; Hassler 5413 (F, NY) Sierra de Maracayu, Nov 

1900; Hassler 5413A (BM, F, G-3, K, MO, NY, P, S, W) In Altoplanitie et Decliviis, Sierra de 

Maracayu, Nov 1900; Hassler 5413B (BM, G-6, K, NY, P, S, W) In Altoplanitie et Deccliviis, Sierra de 

Maracayu, Nov 1900; Hassler 6756 (BM, F-2, MO, NY, P, S, UC, W) In Regione Cursus Superioris 

Fluminis Y-Aca, Dec 1900; Rambo 31287 (F, S) San Francisco de Paula, Vila Oliva, 20 Feb 1946. 

URUGUAY. Rivera: Wright s n (BM) Cunapiru, 1928. Tacuarembo: Rosengurtt B-4975 (MO, S) Valle 

Eden, 3 Feb 1947. Cerro Largo: Fruchard s n (P-2' Cerro Largo, Isla Zapata, Montevideo, 24 Jan 

1877. Canelones: Herter 9673 (NY) Toledo, Feb 1910. Montevideo: illegible s n (MO) Province 

Unknown/Uncertain: illegible s n, (MO) Montevideo. ARGENTINA. Salta: illegible s n (BAB) 

Department Oran, Tartagal, 3 Jan 1943; Venturi 9880 (BM, K, MO, S-2) Alemania, Department 

Guachipas, 14 Nov 1929; Victoria 8 (BAB) San Pedro, Oran, 8 Jan 1939;Zabala 15 (BAB) Cuidad de 

Salta, 13 Mar 1941. Formosa: Jorgensen 3082 (MO, US) 1919. Tucuman: Lillo 2243 (LIL) Tucuman, 

Mar 1899; Lillo 2365 (LIL) 8 Dec 1899; Lillo 7287 (F, US) Tucuman, Department Capital, 22 Dec 

1907; Meyer 12043 (LIL) Tucuman, Department Capital, 15 Apr 1947; Venturi 134 (LIL) Tafi, 7 Jun 

1925; Venturi 63 (BAB, LIL, US) Villa Lujan, Department Capital, 21 Dec 1918; Venturi 63E (US) 

Villa Lujan, Department Capital, 15 Dec 1925; Venturi 7241 (F, MO) Jeoba Buena, Department Tafi, 

3 Jun 1928. Santiago Del Estero: Argauaraz 451 (LIL) El Simbolar, 14 Dec 1942. Santa Fe: Annunzio 

55748 (BAB) Vera, District Tartagal, 21 Apr 1929; Lauro s n (BAB) Rosario, 25 May 1945.. 

Corrientes: Ibarrola 1766 (U), 20 Dec 1944; illegible 32581 (MO) Montenegro, 11 Nov 1945;Pedersen 

7 (S, US) Santa Teresa, 12 Mar 1947; Schwarz 8790 (S) Estancia Santa Teresa, 19 Nov 1949. 

Misiones: Bertoni 2263 (DS, LIL) Department Candelaria, Cerro Cora, 23 Oct 1945; Bertoni 2299 

(TEX) Garupa, 1 Nov 1945; Bertoni 2537 (W) Orillas Del Rio, Department San Javier, 8 Dec 1945; 

Bertoni 3499 (LIL) Department Candelaria, 11 Dec 1944; Bertoni 4638 (LIL) Perae, Department, 

Iguazu, 13 Nov 1949; Bertoni 511 (LIL) Department San Javier, 21 Dec 1944; Curran 721 (LIL, US) 

Vicinity of Puerto Leon, Jul 1914;Ekman 495 (S, US) Posadas, Santa Ana, 23 Dec 1907;Ekman 496 

(S-2) Posadas, Bonpland, 9 Jan 1908; Ibarrola 1009 (LIL, W) Department Apostoles, Pueblo (al sur), 4 

Nov 1944; illegible 164 (LIL), 8 Nov 1919;Jorgensen 31062 (BAB) Bonpland, 3 Dec 1909; Jorgensen 

& Hansen 617 (BAB) Bonpland, Department Candelaria, Nov 1910; Llamas s n (BAB) Santa Ana, 

Department Candelaria, Oct 1901; Medina 158 (NY) San Ignacio, Department San Ignacio, 20 Nov 

1946; Meyer 11773 (LIL, MO) Gobernador Rocaa Santo Pipo, Department San Ignacio; 30 Jan 1947; 

Meyer 11945 (LIL) Pinalito, 10 Feb 1947; Meyer 5421 (F, LIL, U) Posadas, 20 Feb 1944; Meyer 

5463 (U, UC) Eldorado, 6 Mar 1944;Meyer 5850 (LIL, U, UC-2) Eldorado, 6 Mar 1944;Montes 1443 

(DS) Department Candelaria, Santa Ana, 13 Nov 1945; Montes 1448 (W) Santa Ana, Department 

Candelaria, 13 Nov 1945; Montes 1896 (A) San Ignacio, Department San Ignacio, 12 Feb 1946; 

Montes 360 (LIL, W) Loreto, Department Candelaria, 3 Nov 1944; Montes 584 (BAB) Loreto, 

Department Candelaria, 3 Nov 1947; Montes 9524 (NY, W) Puerto Segundo km 17, Department 

Iguazu, 24 Oct 1950; Rodriquez 164 (GH) Santa Ana 8 Nov 1912; Rodriquez 4922 (F, GH) Santa 

Ana, 3 Nov 1912; Rojas s n (BAB) Pto. Aguirre, 27 Sep 1922; Rojas s n (BAB) N de Corrientes;


Schwarz 6629 (S) Oasis, Department San Ignacio, 22 Nov 1948; Schwarz 6738 (S) Paranay, 

Department Cainguas, 3 Dec 1948; Schwarz 7143 (LIL) Department Iguazu, Cataratas Iguazu, 22 Dec 

1948; Schwindt 1019 (S) Tabay, Department Cainguas, 27 Nov 1948; Schwindt 2759 (S) Victoria, 

Ruta 12, Department Iguazu, 18 Nov 1949; Schwindt 3091 (S, US) Ruta 14 km 252, Department 

Cainguas, 25 Jan 1950; Schwindt 3326 (S, US) Iracran a San Pedro Ruta 14, Department San Pedro, 

12 Apr 1950; Scbwindt 4839 (S) Obera (Colonia), Department San Javier, 22 Jul 1950; Schwindt 487 

(W) Puerto Rico, Department Cainguas, 24 Aug 1947; Spegazzini 18385 (BAB) Santa Ana, 4 Feb 

1907. Cordoba: Escuela Nacional de Agricultura 16203 (BAB) Escuela Nacional de Agricultura, Jun 

1906; Hunziker 10535 (US) Sierra Chico, Rio Ceballos, Department Colon, 31 Dec 1954;Sota 3341 

(W) Saldan, Department Colon, 30 Dec 1950. Buenos Aires: Cabrera 5712 (NY) Punta Lara, 28 Dec 

1939; Conde 117 (W) Isla Maciel, 19 Jan 1947;Lanfranchi 327 (BAB) Tigre, 15 Jan 1945;Miers 1240 

(US) Buenos Aires; Milano s n (BAB) Pto. Moron, Castelar, 24 Feb 1964; O'Donnell 205 (UC) 

cultivated in Institute of Darwinion, 13 Jan 1944; Palacios 23 (MO, TEX) Isla Martin Garcia, 21 Dec 

1946; Rojas 196A (BAB) Buenos Aires, Jan 1923; Spegazzini 2351 (BAB) La Plata, 22 Feb 1902; 

Spegazzini 16407 (BAB) Puerto de La Plata, 22 Dec 1906. Country Unknown/Uncertain: illegible s n 

(P). 

Collections representing introductions outside the native range. 

USA. District of Colombia: Schott s n (F) Washington, D.C. Public Gardens. Missouri: Craig s n 

(MO) Missouri Botanical Garden, cultivated No. 1/98, Jul 1911; Kellogg s n (MO) Missouri Botanical 

Garden Green House, 7 Sep 1921. Louisiana: Arsene 12263 (NY, US-2) Vicinity of Covington, Jun 

1920; Eifrig s n (F) New Orleans, 1928; Phares 11 (US) New Orleans, 1878; Rogers, Thieret & Reese 

496 (A, BM, F, S, SP, US) NW Edge of Lafayette in residence of Mr. J. Lynch, 17 May 1963; Rogers, 

Thieret & Reese 497 (F, K, US, W) in Yard of Mr. M. Huger 720 Lafayette Street, 17 May 1963; 

Rogers, Thieret & Reese 498 (G-2, NY-3) Home of Mr. J. Chretten, 1119 11th Street, Lafayette 

Parish, 17 May 1963. Mississippi: Lamorton s n (NA) Laine, Jackson County, 31 Aug 1914; Lamorton 

s n (NA-2) Laine, 24 Sept 1914; Lamorton s n (NA-2) Laine, Kreole, 29 Oct 1915; Skeban s n (MO-2) 

Ocean Springs, 9 Oct 1894. Florida: Cook & Jenkins s n (NA) Scott Place, Innerarity Point, near 

Pensacola, 7 Oct 1928; Godfrey 54844 (FSU) Tallahassee, Leon County, 21 May 1956; Godfrey 

60461 (FSU) Tallahassee, Leon County, 25 Oct 1960; Rogers & Godfrey 499 (COLO, G, M, MICH, 

MO, NY-2) Tallahassee in Vacant Lots Adams and 2 Street, 19 May 1963; West s n (NA-2) Tallahassee, 

Leon County, in Old Garden, 4 Jun 1949; Williams s n (FSU) Tallahassee, Leon County, 15 May 1955. 

INDIA. Uttar Pradesh: Raizada s n (MO) Botanical Garden, Forest Research Institute, Dehra Dun, 25 

May 1953. 

LOCAL NAMES AND USES. Guazu mandioca (Rodriguez 164), Mandioca brava (Ven- 

turi 63), Mandioca de veado (Castro and Kiehl s n), Mandioca do matto (Dutra 366), 

Mandio-Guazu (Rojas s n), Mandio-quazu (Montes 1896), Mandio-guazu (Montes 360), 

Sacha Mandioca. 

Cultivated as an ornamental plant in southern United States. 

Hooker first determined that Graham's Janipha loeflingii var multifida was a distinct 

species in the genus Manibot. Mueller von Argau (1874) apparently did not see the 

relationship between Graham's variety and his new species, M. tweedieana, although he 

had earlier (1866) transferred correctly Janipha loeflingii H.B.K. var Manihot. 

multifida Graham to 

Croizat (1944) said that M. tweedieana Muell-Arg. was incorrect because (fide 

Croizat) Pohl's M. flabellifolia was the correct species. Croizat incorrectly synonymized 

tweedieana under Pohl's flabellifolia even though Pohl's species itself is but a variation of 

M. esculenta. 

According to Article 60 of the International Rules, the varietal epithet multifida 

published by Graham has no priority as a species epithet. 

The roots of Manihot grahami are slender, never enlarged. For this reason, it would 

not seem to be a good candidate for breeding work to increase yield of M. esculenta. On 

the other hand if one is breeding for cool temperature tolerance, this species might be 

useful, particularly if representatives from the southern limits of its range were chosen. 

25. Manihot inflata Mueller von Argau in Martius, fl. Bras. 11(2): 450. 1874; Pax in 

Engler, Pflanzenreich IV. 147(Heft 44): 57. 1910.


Manihot brasiliensis Klotzsch ex Pax in Engler, Pflanzenreich IV. 147(Heft 44): 57. 1910 pro 

syn. 

Shrubs, ca 2.0 m tall. All parts glabrous. Young stems obtuse-angled in cross section. 

Leaves alternate; stipules caducous, ca 2.0 cm long, filiform; petioles ca 12.0 cm long, 

terete, petiole attachment to lamina basal, nonpeltate; abaxial lamina surface wax pattern 

smooth, venation camptodromous, palmately 3-5 lobed, rarely 7; median lobes 

oblong-elliptic, ca 10.0 cm long, ca 2.5 cm wide, margin undulate, apex acuminate, base 

angustate, base of lobes extremely constricted, leaf appears compound; lowest lobes more 

or less similar to median lobes in outline but smaller. Inflorescence a monoecious 

moderately branched panicle, ca 12.0 cm long; bracts and bracteoles setaceous. Pistillate 

flowers restricted to the base of the upper 2/3 of the inflorescence, pedicels ca 1.25 cm 

long, tepal 1.0 cm long, ovary subglobose. Staminate buds ovoid-ellipsoid, flowers 

medium sized, tepal 1.0 cm long, yellowish green externally, purplish internally, cleft 1/3 

way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Fruits and seeds non vidi. 

Fig 35B, C. 

TYPE. Riedel 190: Brasil, Rio de Janeiro: Near Rio de Janeiro (syntypes F-2, G-5, K, 

NY-2, P). 

DISTRIBUTION. (Fig 35-A). Brasil, states of Sao Paulo, Rio de Janeiro and Parana. 

BRASIL. Sao Paulo: Kublmann s n (SP) Galhetas, Guaruja, 20 Dec 1940; Mosen 3279 (S-2) 

Santos, 10 Feb 1875. Rio de Janeiro: Kublmann 06242 (RB) Serra do Cambori, 15 Dec 1942. Parana: 

Hatschbach 6404 (US) Barra Rio Ponta Gressa, Mun. Cerro Azul, 24 Oct 1959. 

06242). 

LOCAL NAMES. Mandioca braba (Hatschbach 6404), mandioca da Serra (Kuhlmann 

The closest affinity of Manihot inflata is Manihot grahami from which it differs by 

the deeper sinuses between the leaf lobes (constricted lobe bases). It also exhibits fewer 

to no pandurate lobes. 

26. Manihot pilosa Pohl, PI. Bras. Ic. et Descr. 1: 55. 1827; Muell.-Arg. in DC., Prodr. 

15(2): 1059. 1866 ex parte; in Martius, Fl. Bras. 11(2): 449. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 60. 1910. 

Jatropha pilosa Steudel, Nomencl. ed. 2. 1: 800. 1840. 

Manihot pedicellaris Muell.-Arg. in Martius, Fl. Bras, 11(2): 453 t. 64. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 62. 1910. Type. Warming s n (syntypes, F, G-2, P-2). 

Manihot hemitrichandra Muell.-Arg. in Martius, Fl. Bras. 11(2): 454. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 64. 1910;Manihot hemigynandra Muell.-Arg. in Martius, Fl. 

Bras. 11(2): 439. 1874 (orthographic variant). Type. Riedel 1848 (syntypes, F-2, G-2, K, 

NY-2, P, W). 

Manihot langsdorffii Muell.-Arg. in Martius, Fl. Bras. 11(2): 455. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 65. 1910; Type. Riedel 1530 (syntypes, F, G, NY, P); 

Manihot meyeriana Klotzsch ex Pax in Engler, Planzenreich IV. 147(Heft 44): 65. 1910 pro 

syn. 

Manihot tubuliflora Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 44); 61. 1910. 

Type. Ule 4605 (syntypes, F, NY);Moura 1018 n v. 

Manihot brevipedicellata Pax in Engler, Pflanzenreich IV. 147(Heft 44): 63. 1910. Type. Glaziou 

8322 (syntypes, F, G, K, NY, P, US). 

Tall, erect shrubs (3.0 m tall) to slender trees (10.0 m tall); lower trunk smooth, 

woody, 8.0-12.0 cm in diameter, with little pith. Roots slender, 2.5-5.0 cm in diameter, 

slightly enlarged in places; epidermis slightly rough, light tan; subepidermis tan; cortex 

cream. Young stems pubescent, obtuse-angled in cross section, grayish-brown; mature 

stems sparsely pubescent, dark brown. Leaves alternate, stipules short, less than 0.5 cm 

long, pubescent, caducous; petioles long, usually ca 20.0 cm long, sometimes as long as 

45.0 cm, obtuse-angled, pubescent, dark red, petiole attachment to lamina basal,


FIG 35. Manihot inflata. A, distribution; B, leaf (Hatschbach 6404, US); C, inflorescence 

(Hatschbach 6404, US). M. pilosa. D, distribution. 

(Hatschbach 6404, US). M. pilosa. D, distribution. 

93


nonpeltate; abaxial lamina surface wax pattern reticulate; venation camptodromous, veins 

on abaxial lamina surface pubescent; palmately 5-9 lobed, median lobes oblong, usually 

10.0-15.0 cm long, ca 4.0 cm wide, sometimes as long as 25.0 cm and as wide as 6.0 cm, 

margin entire, rarely undulate, apex acuminate, base cuneate, base of lobes 0.25-1.0 cm 

wide; lowest lobes similar to median lobes in outline but smaller. Inflorescence a 

monoecious, moderately branched, terminal panicle, flowers clustered at the terminal 

region of peduncles, ca 10.0 cm long, peduncles and pedicels pubescent; bracts and 

bracteoles short, less than 0.5 cm long, setaceous, pubescent, caducous. Pistillate flowers 

restricted to the base of the upper half of the inflorescence, pedicels 1.0-2.0 cm long, 

tepal 1.0 cm long; disc brilliant yellow (2.5 Y 9/9), ovary tomentose. Staminate buds 

bifusiform with a distinct constriction in the middle, flowers medium sized, tepal 1.0 cm 

long, cleft 1/3 down into 5 lobes, sparsely pubescent, apex of tepal lobes brilliant 

greenish yellow (7.5 Y 9/8), base of tepal brilliant yellow green (5 GY 8/8) with dark 

purplish pigmentation in the region of the sinus of tepal lobes; stamens 10, in 2 whorls of 

5 each, the superior whorl 0.6 cm long, the inferior whorl 0.5 cm. Capsules ellipsoidal, 

1.0-1.5 cm in diameter, surface smooth, without wings, dehiscence septicidal. Seeds 

oblong 1.0 cm long. Fig 36A, B, C, D. 

TYPE. Martius 942: Brasil, Minas Gerais: bei Antonio Pereira (syntypes, F, G, M). 

DISTRIBUTION. (Fig 35D). Brasil, in the states of Minas Gerais, Sao Paulo and Rio 

de Janeiro. Common in secondary woodlands, along stream banks, in sandy loam soils and 

among rocky outcrops. Altitudes to ca 1500 m. BRASIL. Minas Gerais: Barreto 5067 (A, F) 

Matta da Empreza Mun. Cambuquira, 25 Dec 1935; Barreto 9624 (F) Sentinella Mun. Diamantina, 8 

Nov 1937; Glaziou 18476 (A, F, G, K) Faria, 27 Jan 1891; Hoehne s n (SP) Caldas, 23 Jan 1919; 

Irwin, Maxwell & Wassbausen 19774 (F, G, GH, K, NY, RB, SIU, SP, UB, US, W) Serra do Itabirito, 

Circa 55 km SE of Belo Horizonte, 10 Feb 1968; Irwin, Maxwell & Wassbausen 20822 (F, IAN, MICH, 

NY, S, SIU, SP, UC, US-2) Circa 20 km N of Serro on Road (MG 2) to Diamantina, 25 Feb 1968; 

Irwin, Maxwell & Wasshausen 20862 (F, G, FH, MO, NY, RB, SIU, SP, UB, US, W) Serra do 

Espinhaco at Lapinha, Circa 21 km N of Serro on Road (MG 2) to Diamantina, 25 Feb 1968;Macedo 

2915 (MO, S, US) Escola, Vicosa, 10 Jan 1951;Magalhaes 6080 (PUL, RB) Barreiro, 29 Nov 1953; 

Mexia 4109 (A, F, GH, MO, NY, S, U, UC, US) Trail from Vicosa to Sao Geraldo, 16 Dec 1929;Mexia 

4153 (A, BM, GH, MICH, MO, NA, NY, P, S, TEX, UC, US) Road from Vicosa to Sao Miguel, km 20, 

23 Dec 1929; Regnell III 1070 (P, S-3, UC, US-2) Caldas, 27 Nov 1864; Regnell III 1070A (S); Rogers 

372 (COLO, F-2, W) Lavras, Grounds of Substation Experimental Federal, 20 Feb 1961; Rogers 373 

(G-2, K, M, MICH, MO, NY-2, US) Lavras, Grounds of Substation Experimental Federal, 20 Feb 1961; 

Warming 1519 (F, G-2, P-2) Lagoa Santa; Williams 6876 (GH) Serra da Bocania, 10 km NW of Serro, 

Mun. Serro; 4 May 1945. Sao Paulo: Castrol & Kiehl s n (SP) Cunha, 16 Feb 1939; Cruz 104 (SP) 

Mun. de Monteiro, Lobato, Na Rodovia Para Campos do Jordao, km 157, 18 Nov 1964; Cruz 105 (SP) 

Mun. Monteiro Lobato, Na Rodovia Para Campos do Jordao, km 157, 14 Jan 1965; Cruz 114 (SP) Sao 

Sebastiao da Grama, Cultivada na Fazenda Experimental Theodureto de Camargo, 19 Nov 1965; Krug 

s n (SP) Cunha, 22 Nov 1938; Kuhlmann 2751 (SP) Igarata, 12 Dec 1951; Kuhlamnn & Gehrt s n (SP) 

Serra de Cunha, 14 Mar 1939; Mosen 4386 (S) Serra de Caracol, 10 Dec 1875; Normanba s n (SP) 

Taubate, 1 Mar 1941; Novais 926 (SP) Capoeira, 20 Dec 1896; Riedel 1530 (F, G, NY, P) Lorena; 

Riedel 1848 (F-2, G-2, K, NY-2, P, W) Jundiahy; Viegas s n (SP) Aguas de Prata, 17 Jun 1940; Viegas s 

n (SP-2) Campo Experimental Instituto Campinas, cultivated, 10 Dec 1941; Viegas s n (SP) Estrada 5 

km Alem Jundiai, 12 Apr 1942. Rio de Janeiro; Glaziou 8322 (F, G, K, NY, P, US) San Jose dos 

Barreiros, 9 Jan 1876; Peckolt 95 (W-2) Cantagello; Ule 4605 (F, NY) Im Walde Bei Novo Friburgo, 

1897. 

LOCAL NAMES. Mandioca braba (or brava), maniva de veado, etc. These common 

names are not unique for this species, but are used in many areas of Brasil for many 

different species. 

The closest relative of Manihot pilosa is M. janiphoides, which in turn, is a member 

of the M. esculenta "complex" (see Fig 28). The most important distinguishing characters 

of this species are the bifusiform shape of the buds of the staminate flower, and the 

tendency 

peduncles. 

to produce the staminate flowers in clusters at the terminal regions of the


K4_s 

clk. 

FIG 36. Manibot pilosa. A, habit (Rogers 373, NY); B, leaf (Irwin, Maxwell & Wassbausen 

20822, 20822, NY); NY); C, inflorescence 

(Mexia 4109, S); D, D, fruits (Rogers 373, NY). 

I


The number of synonyms indicate our change in species concepts from the concepts 

of earlier botanists. Manihot pilosa is a variable species with several growth forms. It, like 

many other Manihot species, inhabits regions evidently disturbed by man, and apparently 

colonizes easily. The seedling plants are frequently more robust than the mature plants, a 

source of confusion to the earlier botanists. 

27. Manihot corymbiflora Pax in Engler, Pflanzenreich IV. 147(Heft 44): 80. 1910. 

Shrubs. Young stems grayish brown, pubescent; mature stems dark brown, glabrous. 

Leaves alternate; stipules caducous; petioles ca 12.0 cm long, obtuse-angled in cross 

section, pubescent, petiole attachment to lamina basal, nonpeltate; abaxial lamina surface 

wax pattern reticulate; venation camptodromous, veins on abaxial lamina surface 

pubescent; palmately 5 lobed; median lobes obovate-pandurate, ca 9.0 cm long, 

sometimes shorter, ca 3.0 cm wide, apex acute, base brevi-angustatus, base of lobes 0.5 

cm wide, width between base of sinus and petiole-lamina junction 0.5 cm; lowest lobes 

similar to median lobes in outline but 1/2 as long. Inflorescence a monoecious, terminal, 

many flowered panicle, flowers closely clustered at the terminal end of peduncles giving 

the appearance of a corymb, ca 15.0 cm long; peduncles and pedicels pubescent; bracts 

and bracteoles setaceous. Pistillate flowers restricted to the base of the upper half of the 

inflorescence, pedicels 1.0 cm long, tepal 0.5 cm long, ovary subglobose, glabrous. 

Staminate buds slightly constricted in the middle, flowers small, tepal 0.5 cm long, cleft 

1/3 way down into 5 lobes, stamens 10, in 2 whorls of 5 each. Capsules 1.25 cm long, 

dehiscence septicidal. Seeds 0.7 cm long, caruncle poorly developed. Fig 37B, C. 

TYPE. Glaziou 13203: Brasil, Rio de Janeiro: Environs of Rio de Janeiro, Feb 1882 

(syntypes, F, G, K, NY, P, US); Glaziou 14242: Brasil, Rio de Janeiro: Environs of Rio 

de Janeiro, Apr 1882 (syntypes, F, K, P); Glaziou 14243: Brasil, Rio de Janeiro: Environs 

of Rio de Janeiro, Apr 1882 (syntypes, BM, K). 

DISTRIBUTION. Fig 37A. BRASIL. Rio de Janeiro: Glaziou s n (P). 

Manihot corymbiflora, while maintained as distinct here, is a very close relative of M. 

pilosa. The only distinguishing feature is the size and number of staminate flowers on the 

inflorescence. See Fig 37C. and compare with Fig 36C. The distribution of M. 

corymbiflora coincides with the distribution ofM. pilosa. 

28. Manihot leptopoda (Mueller von Argau) Rogers & Appan, stat nov 

Jatropha palmata Vellozo, Fl. Flum. 10. t. 81. 1825 nom rejic;Jatropha palmata Arrabida in 

Steudel, Nomencl. ed. 2. 1: 799. 1840 (Arrabida is the editor of Flora Fluminensis. This 

name is the same as Jatropha palmata Vellozo); Manihot palmata (Vellozo) Muell.-Arg. var 

genuina Muell.-Arg. in DC., Prodr. 15(2): 1062. 1866; in Martius, Fl. Bras. 11(2): 459. 1874 

nom rejic; Manihot palmata (Vellozo) Pax in Engler, Pflanzenreich IV. 147(Heft 44): 55. 

1910 nom rejic. Type. Vellozo s n n v. 

Manihot palmata var leptopoda Muell.-Arg. in Martius, Fl. Bras. 11(2): 459. 1874;Manihot dulcis 

(J.F. Gmelin) Pax var leptopoda (Muell.-Arg.) Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 72. 1910. 

Erect shrubs, ca 2.0 m tall. All parts glabrous. Young stems obtuse angled in cross 

section, grayish brown; mature stems dark brown. Leaves alternate; stipules caducous; 

petioles ca 8.0 cm long, terete, dark red, petiole attachment to lamina basal, nonpeltate; 

abaxial lamina surface wax pattern reticulate, venation camptodromous; palmately 5 

lobed; median lobes lanceolate, ca 7.0 cm long, ca 2.0 cm wide, apex acuminate, base 

attenuate, base of lobes extremely constricted, less than 0.2 cm wide, width between base 

of sinus and petiole-lamina junction less than 0.2 cm; lowest lobes similar to median lobes 

in outline but slightly smaller. Inflorescence a monoecious, terminal, moderately 

branched panicle, with several principal peduncles arising from one basal point, ca 7.0 cm 

long, flowers clustered at the terminal ends of peduncles, pedicels and tepals with a bluish


/,/ ' 

.... 

(Glaziou 14243, K). M. leptopoda. D, distribution. 

C 

l . i 

@ - --- 

FIG 37. Manihot corymbiflora. A, distribution; B, leaf (Glaziou 14242, P); C, inflorescence 

(Glaziou 14243, K). M. leptopoda. D, distribution. 

i


white bloom; bracteoles and bractlets setaceous. Pistillate flowers restricted to the base of 

the upper 1/3 of the inflorescence, pedicels 1.0 cm long, staminodes present, disc bright 

yellow, ovary subglobose. Staminate buds distinctly bifusiform, flowers short, tepal 0.7 

cm long, yellowish green, cleft 1/3 way down into 5 lobes, stamens 10, arranged in 2 

whorls of 5 each. Capsules small, 1.0 cm long, surface smooth, without wings. Seeds 

small, 0.8 cm long, oblong, with a poorly developed caruncle. Fig 38A, B, C. 

TYPE. Guillemin 132: Brasil, Rio de Janeiro: Corcovado, 1839 (syntypes, A, G-2, 

K); Regnell 186 Brasil, Rio de Janeiro: Rio de Janeiro, 1841 (syntypes, G, NY, S-2); 

Riedel 191: Brasil, Rio de Janeiro: (syntypes, G, K). 

DISTRIBUTION. (Fig 37D). Brasil, in the state of Rio de Janeiro, on granitic outcrops. 

BRASIL. Rio de Janeiro: Ducke s n (PUL, RB), 19 Dec 1928; Goes & Constantino 817 (RB) 

Petropolis, Bairro Amoeda, Dec 1943; Gomes s n (K) Rio de Janeiro, 1836; Guillemin s n (P); 

Kuhlmann s n (PUL, RB) Mundo Novo, Botafogo, Nov 1920; Kuhlmann 06022 (RB), 30 Nov 1939; 

Luetzelburg 15399 (NY) Petropolis, Ipanema, Dec 1910; Machado s n (PUL, RB) Restinga da Tijuca, 

25 May 1945; Rogers 335 (NY) Just Below Summit of Pao Azucar, 1 Feb 1961. Province: 

Unknown/Uncertain: Freyreis s n (S). 

Doubtful specimens. Brasil, Bahia: Froes 20234 (US-2, K, F, NY), Catinga na Regiao de Serra de 

Sincora, 15 Feb 1943. This collection is doubtfully assigned to this species because it is distinct 

geographically from the other collections. 

LOCAL NAMES. Mandioca brava (Machado s n), Maniva (Machado s n). 

The first appearance of the epithet palmata was made by Sesse, as follows: Jatropha 

palmata Sesse, apud Cerv., Gaz. Lit. Mex. 3: suppl. 4, 1794. Vellozo's name Jatropha 

palmata, published in Fl. Flum. 10, 1825, t. 81, is clearly a later homonym. 

Mueller von Argau's transfer of Vellozo's name to the genus Manihot in DC., Prodr. 

15(2): 1062. 1866 is, therefore, not in keeping with Article 64, International Rules of 

Botanical Nomenclature. (See also Historical Section, p 4). 

An excellent review of Sesse's names is given by McVaugh, 1945. 

This species has a distinct ecological adaptation to the granitic domes of the Rio de 

Janeiro region. It would be of potential value in breeding programs with Manihot 

esculenta in adapting to granitic sandy regions. The closest relative of this species is M. 

pilosa. 

29. Manihot quinquefolia Pohl, P1. Bras. Ic. et Descr. 1: 56. 1827; Muell.-Arg. in DC, 

Prodr. 15(2): 1071. 1866; in Martius, Fl. Bras. 11(2): 472. 1874, Pflanzenreich IV. 

147(Heft 44): 32. 1910. 

Jatropha quinquefolia Steudel, Nomencl. ed. 2. 1: 800. 1840; Jatropha quinqueformis Steudel, 

Nomencl. ed. 2. 1: 800. 1840; Manihot quenqueformis Pohl ex Steudel, Nomencl. ed. 2. 1: 

800. 1840. (orthographic variant). 

Young stems glabrous. Leaves alternate, stipules caducous, petioles ca 10.0 cm long, 

terete, glabrous, petiole attachment to lamina basal, nonpeltate; abaxial lamina surface 

with a bluish bloom, wax pattern reticulate; venation camptodromous, veins glabrous; 

palmately 5 lobed; median lobes elliptic, ca 6.0 cm long, ca 2.5 cm wide, margin entire; 

apex acute, base breve-angustatus, base of lobes extremely constricted, less than 0.2 cm 

wide, flacid, with a tendency for the lobes to droop down; lowest lobes similar to median 

lobes in size and outline. Inflorescence non vidi. Capsules and seeds non vidi. 

TYPE. Martius s n: Brasil, Bahia: bei Sincora (syntypes, photos at F, G, M). 

DISTRIBUTION. (Fig 38D). Brasil, in the state of Bahia. 

This species, of which we had only photographic material, is closely related to 

Manihot leptopoda from which it is doubtfully distinguished. 

Manihot quinqueformis Pohl is cited by Steudel (1840) as a synonym of Jatropha 

quinqueformis Steudel. Pohl did not use this epithet, to our knowledge. Pohl (1827, p


A ^H ^^^^^^ ^^^^^^^HL \JT 

& ^ ^^^^^ ^^^^^^9D 

^^ 

Jr^ ^ ^^^B ^^^^rC '** 

FIG 38. Maniot leptopod. A, leaf ( o^ 33, N) ,ifoecne(oes35 Y; 

bifuifom samiate uds(Roers335 NY) M.qumueflia.D, istibuion 

~~~~~~~~~~~~~~~~~ 

=::~~~~~~~~~~~ 

FIG 38. Manibot leptopoda. A, leaf (Rogers 335, NY); B, inflorescence (Rogers 335, NY); C, 

bifusiform staminate buds (Rogers 335, NY). M. quinquefolia. D, distribution.


56) in an observation under M. quinquefolia asks what Jatropha quinquelobata is. The 

epithet in question was mentioned in Miller (1754, vol. 2: p 567) and in Gmelin 

(1772-1778, p 7). 

30. Manihot condensata Rogers & Appan, sp nov 

Folia 7 lobatus, lobi mediani oblongo-obovata, leviter panduratus, ca 15.0-20.0 cm. 

longa, bracteolae et ramuli foliaceus. Fructi manifeste alati. 

Erect shrub or small tree to 8.0 m tall, 10 cm diam. Plant parts glabrous or 

pubescent. Young stems grayish brown, obtuse-angled, in cross section, mature stems 

grayish brown. Leaves alternate; stipules foliaceous, 1.5-2.0 cm long, 0.5 cm wide, 

margins usually entire, rarely serrate, persistent; petioles ca 10.0 cm long, terete, petiole 

attachment to lamina peltate, width between basal edge of lamina and petiole attachment 

point 0.25-0.5 cm; abaxial lamina surface wax pattern verrucate; venation camptodrom- 

ous; lamina palmately 7 lobed; median lobes oblong-obovate, shallowly pandurate, ca 

15.0 cm long, ca 4.0 cm wide, apex acuminate, base acute; lowest lobes similar to median 

lobes in outline, ca half as long as median lobes. Inflorescence a monoecious, terminal 

panicle, 15.0-20.0 cm long, condensed; bracteoles foliaceous, 1.0-1.5 cm long, 0.5-0.8 

cm wide, broadly ovate, margins entire, occasionally serrate; bracteoles foliaceous, 0.6 cm 

long, 0.3 cm wide. Pistillate flowers restricted to the base of the upper 2/3 of the 

inflorescence, tepal 0.8 cm long, cleft to base into 5 lobes, ovary subglobose, winged. 

Staminate buds ovoid-ellipsoidc flowers medium sized, tepal 0.8 cm long, cleft 1/3 way 

down into 5 lobes, stamens 10, in 2 whorls of 5 each. Capsules almost globular, ca 2.0 cm 

long from base to apex, with prominent wings. Seeds rotundate, ca 1.25 cm long, 

caruncle well developed. Figs 39B, C, D. 

TYPE. White 1019: Bolivia, La Paz: Huachi, head of Beni river, 6 Sep 1921 

(holotype, NY; isotype, K, MICH). 

DISTRIBUTION. (Fig 39A). Bolivia, in the Departments La Paz and Beni. Alt ca 600 

m. BOLIVIA. Beni: Rusby 1297A (K, NY) Vicinity of Rurrenabaque, 15 Oct 1921. La Paz: Williams 

361 (BM, NY-2, US) San Buena Ventura, 20 Nov 1901. 

The distinct feature of this species is the condensed, many flowered panicle. (Fig 

39C) and the foliaceous bracteoles (Fig 39B, D). The skyline (Fig 28) indicates its unique 

set of characters, remaining as a distinct taxon until a relatively low level of similarity 

where it joins M. grahami and M. esculenta simultaneously. 

31. Manihot jolyana N. D. Cruz, Bragantia 24: 359- 368. 1965. 

Tall shrubs, ca 3.0 m tall. Young stems obtuse angled, in cross section, pubescent; 

mature stems pubescent. Leaves alternate; stipules short, less than 0.75 cm long, 

pubescent, caducous; petioles long, often as long as 35.0 cm, obtuse-angled, pubescent, 

petiole attachment to lamina basal, nonpeltate; with 2 pubescent, 0.6 cm long lobules 

inserted at the petiole-lamina junction; abaxial lamina surface smooth; venation 

camptodromous, veins on the abaxial lamina surface tomentose; lamina palmately 7 

lobed; median lobes obovate, 15.0-20.0 cm long, 6.0 cm wide, margin entire, apex 

broadly acuminate, base acute, base of lobes more than 0.5 cm wide; lowest lobes similar 

to median lobes in outline but slightly smaller. Inflorescence a monoecious, terminal, 

profusely branched, large panicle, ca 25.0 cm long; bracteoles foliaceous, 1.25 cm long, 

0.6 cm wide, elliptic, margins entire, pubescent; bractlets foliaceous. Pistillate flowers 

restricted to the base of the upper half of the inflorescence; pedicels 1.5 cm long, 

pubescent; tepal 1.6 cm long, cleft to base into 5 strap-shaped lobes with considerable 

purple pigmentation, disc prominent, ovary glabrous, elongated. Staminate buds more or 

less conical, flowers large, tepal sometimes as long as 2.2 cm, dark purplish tinged,


/ . I 


pubescent, cleft ca 1/3 way down into 5 lobes, lobe apex tapering, stamens 10, in 2 

whorls of 5 each. Fruits and seeds non vidi. Capsule surface smooth (fide original 

description). Fig 40B, C, D. 

TYPE. Cruz N.D. 103: Brasil, Sao Paulo: Eugenio Lefevre, 18 Nov 1964 (holotype, 

SP). 

DISTRIBUTION. 

(Fig 40A). Brasil, in the state of Sao Paulo. Alt ca 1250 m. BRASIL. 

Sao Paulo: Kuhlmann sn (SP) Mun. de Pindamonhangaba, Eugenio Lefevre, 23 Sep 1961. 

This recently described species is quite distinct. Its pubescence and elongated 

inflorescences give it a handsome appearance. The recency of this species, and the 

subsequent species Manibot bandroana, in addition to the other new taxa described in 

this monograph, are clear indications that more intensive investigations and collections in 

South America will very likely uncover more new taxa in the genus. N. D. Cruz (1965) 

has determined the chromosome count of this species as 2n = 36. 

32. Manihot handroana N. D. Cruz, Bragantia 26: 317-328. 1967. 

Shrubs. Young stems sparsely pubescent; mature stems glabrous. Leaves alternate; 

stipules filiform, ca 1.25 cm long, pubescent, caducous; petioles ca 15.0 cm long, 

pubescent, obtuse-angled, in cross section, petiole attachment to lamina basal, nonpeltate; 

with 2 pubescent, 0.5 cm long lobules inserted at the petiole-lamina junction; wax pattern 

of abaxial lamina surface smooth; venation camptodromous, veins on the abaxial lamina 

surface pubescent; lamina palmately 5 lobed, occasionally 7; median lobes obovate-elliptic 

8.0-12.0 cm long, 4.0 cm wide, margin entire, apex broadly acuminate, base angustate, 

base of lobes 0.4 cm wide, width between base of sinus and petiole-lamina junction less 

than 1.0 cm; lowest lobes similar to median lobes in outline, slightly smaller. 

Inflorescence a monoecious, terminal, many branched panicle, ca 15.0 cm long; 

bracteoles foliaceous, 1.0 cm long, 0.4 cm wide, pubescent; bractlets foliaceous. Pistillate 

flowers restricted to the base of the upper half of the inflorescence; pedicels 1.5 cm long, 

pubescent; tepal 1.5 cm long, cleft to base into 5 strap-shaped lobes with considerable 

purple pigmentation, disc prominent, ovary glabrous, subglobose. Staminate buds more or 

less conical, flowers large, tepal sometimes as long as 2.2 cm, dark purplish tinged, 

pubescent, cleft 1/3 way down into 5 lobes, lobe apex tapering, stamens 10, in 2 whorls 

of 5 each. Fruits and seeds non vidi. Capsule surface winged (fide original description). 

Fig 41B, C. 

TYPE. Kublmann s n: Brasil, Minas Gerais: Serra da Mantiqueira, Fazenda Corrego 

Alegre, 20 Nov 1940 (holotype, SP). 

This species and the preceding one, M. jolyana, are closely related elements. They are 

distinguished one from the other in that M. bandroana possesses smaller leaves and is less 

densely pubescent than M. jolyana. These two species are closely related to M. pohlii. 

33. Manihot pohlii Wawra, Flora 47: 252. 1864; Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 37. 1910. 

Manibot tripartita (Sprengel) Muell.-Arg. var quinqueloba Pax & K. Hoffmann in Engler, 

Pflanzenreich IV. 147(Heft 85): 194. 1924. Type. Luetzelburg 7209 (syntype, NY); 

Luetzelburg 8711 n v.; Luetzelburg 15012 n v. 

Shrubs. Branches with a tendency to droop. Young stems obtuse-angled, in cross 

section, pubescent; mature stems glabrous. Leaves alternate; stipules short, less than 0.5 

cm long, pubescent, caducous; petioles ca 10.0 cm long, obtuse-angled, in cross section, 

pubescent, petiole attachment to lamina basal, nonpeltate, petiole-lamina junction 

without lobules; wax pattern of abaxial lamina surface reticulate; venation camptodro-


ii 

' 

J~ ? 

FIG 40. Manihot jolyana. A, distribution; B, staminate flowers (Cruz 103, SP); C, inflorescence 

(Cruz 103, SP); D, foliaceous bracteoles (Kublmann s n, SP).


tion. 

FIG 41. Maniot androana. A, distribution; B, leaf (Kulmann s n, SP)M. po D distribu-


mous, veins on the abaxial lamina surface pubescent; lamina palmately 5 lobed; median 

lobes obovate-lanceolate, 5.0-10.0 cm long, 2.0-3.0 cm wide, margin entire, apex 

acuminate, base cuneate, base of lobes extremely constricted, less than 0.25 cm wide, 

width between base of sinus and petiole-lamina junction 0.25 cm; lowest lobes similar to 

median lobes in outline, slightly smaller. Inflorescence a monoecious, terminal, few 

flowered panicle, ca 4.0-6.0 cm long, rarely longer; bracteoles foliaceous, 1.0 cm long, 

0.5 cm wide, broadly elliptic, margins entire, occasionally serrate, pubescent; bractlets 

foliaceous. Pistillate flowers restricted to the upper half of the inflorescence; pedicels 1.0 

cm long, pubescent, tepal pubescent. Staminate buds more or less conical, flowers 

medium sized, tepal 1.0 cm long, pubescent, cleft 1/3 way down into 5 lobes, stamens 10, 

in 2 whorls of 5 each. Fruits and seeds non vidi. Fig 42A, B, C. 

TYPE. Peckolt 21: Brasil, Rio de Janeiro, Cantagallo, 1864 (syntypes, F, W). 

DISTRIBUTION. (Fig 41D). Brasil, in the states of Espirito Santo and Rio de Janeiro. 

BRASIL. Espirito Santo: Kublmann 06562 (PUL, RB) Rio Pancas, 30 Nov 1943; Luetzelburg 7209 

(NY) Urwald Am Rio Mutum, Norde Vom Rio Doce, Feb 1917. 

The geographic distribution of Manihot pohlii is sympatric with the two preceding 

species. Its close relationship with these may indicate that they represent a common gene 

pool. An alternative hypothesis is that these represent new isolates from a former 

common gene pool. The insufficient material does not allow a full evaluation of the 

parameters of variation. 

5. Manihot sect Anisophyllae Rogers & Appan, sect nov 

Manihot sect Parvibracteatae Pax subsect Guaraniticae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 74. 1910 pro parte. 

Manihot sect Heterophyllae Pax subsect Carthagineses Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 80. 1910 pro parte. 

Manihot sect Glaziovianae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 89. 1910 pro parte. 

Plantae caulinae, lignosae, infernae ad mediae frutices. Folia petiolatae, penitae 

lobatae. Inflorescentia monoecia racema. 

Plants caulescent, low to medium shrubs; leaves widely spaced on stem; petiolate, 

petiole attachment basal or peltate; lamina membranaceous, deeply lobed, lobes variously 

shaped but not linear. Inflorescence a monoecious raceme, single or a cluster of 2 or 3 

arising from a common base; bracts and bracteoles setaceous to semifoliaceous, margins 

entire or serrate. 

TYPE. Manihot anisophylla (Griseb.) Muell.-Arg. 


Key to the Species of Sect Anisophyllae 

1. Bracts and bracteoles setaceous, less than 0.2 cm wide, margin entire; stipule margin entire; 

flowers yellowish, without any purple pigmentation; petiole attachment basal. 34. M. anisophylla. 

1. Bracts and bracteoles semifoliaceous, more than 2.0 cm wide, margin serrate; stipule margin 

serrate; flowers mostly with considerable purple pigmentation; petiole attachment mostly 

peltate, rarely basal. 35. M. guaranitica. 

34. Manihot anisophylla (Grisebach) Mueller von Argau, Jour. Bot. 12: 230. 1874; Pax in 


Janipha anisophylla Grisebach, Abh. Konigl. Ges. Wiss. Gottingen 19: 47. 1874. 

Manihot carthaginensis var anisophylla 0. Kuntze, Rev. Gen. 3(2): 288. 1898. ex parte (see also 

35a). Type. Kuntze s n (syntypes, NY-2, US). 

Shrubs ca 3.0 m tall. Stems glabrous. Leaves alternate; stipules setaceous, glabrous, 

margin entire, caducous; petioles 6.0-12.0 cm long, terete, glabrous, petiole attachment


FIG 42. Manibot pohlii. A, leaf (Kuhlmann 06562, PUL); B, inflorescence (Peckolt 21, RB); 

C. 

foliaceous bracteoles (Kuhlmann 06562, RB).

,Si-.c^ 

;Y , _ -?' ' ~ r 

---- - 

t~~~~~ v. * / 

-? v 

r/~~~? /.. 

,,, ; 

s 

~~~~~~~~~~~~~~~~~~ 

I:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~C 

?C..4 

C 

~ ~t 

\,- * ^ 

r , 

^ -.. 

' ~ ~ ~ ~ 


to lamina basal, nonpeltate; abaxial lamina surface wax pattern smooth; venation 

camptodromous, veins on abaxial lamina surface glabrous; lamina palmately 5 lobed; 

median lobes obovate-elliptic, margin entire, ca 8.0 cm long, ca 3.0 cm wide, apex 

acuminate, base breve-angustatus, base of lobes 0.5 cm wide, width between sinus base 

and petiole-lamina junction 0.6 cm wide; lowest lobes similar to median lobes in outline, 

ca half in size. Inflorescence a monoecious, terminal raceme, sometimes a cluster of 2 or 3 

racemes arising from a basal point, ca 10.0 cm long, all parts glabrous; bracteoles 

setaceous, less than 0.5 cm long, bractlets setaceous, 0.3 cm long. Pistillate flowers 

restricted to the base of the inflorescence, pedicels 1.5 cm long, tepal 1.1 cm long, 

yellowish green without any purple pigmentation, cleft to base into 5 lobes, ovary 

subglobose. Staminate buds ovoid-ellipsoid, flowers medium sized, tepal 1.0 cm long, 

yellowish green without any purplish pigmentation, cleft 1/3 way down into 5 lobes, 

stamens 10, arranged in 2 whorls of 5 each. Capsules 1.25 cm long, surface smooth, apex 

rounded, dehiscence septicidal. Seeds 1.0 cm long, elliptic, with a well developed 

caruncle. Fig 44B, C. 

TYPE. Lorentz 297: Argentina, Cordoba: near Ascochinga, 1871 (syntypes, Photo F, 

G-4 Photo, NY). 

DISTRIBUTION. (Fig44A). ARGENTINA, in the provinces Salta,Catamarca,Tucuman, 

la Rioja, and San Luis. Alt to ca 1000 m. ARGENTINA. Salta: Peirano s n (LIL) Las Conchas 

Alemania a Cafayate, Department Guachipas, 27 Nov 1933. Catamarca: Castillon 14176 (LIL) 

Alrededores de La Capital, 12 Nov 1910; Castillon 1720 (LIL) El Rodea, Jan 1910; Castillon 1949 

(LIL) Pomancillo, Dec 1910; Hunziker, Cocucci & Subils 18466 (US) Sierra de Ambato, entre 

Rincon Y Saujil, Department Poman, 8 Dec 1965;Jorgensen 1225 (GH, LIL, MO, UC, US) Andalgala, 

Department Andalgala, 3 Feb 1916; Schreiter 6454 (LIL) La Chilca a Andalgal, Department 

Andalgala, 3 Nov 1930; Spegazzini s n (BAB-2) Department Poman, Poman, Dec 1909. Tucuman: 

Dinelli s n (BAB) Department Trancas, Vipos, 22 Dec 1907; Lillo 7278 (LIL) Vipos, 22 Dec 1907; 

Lillo 7918 (LIL) Vipos, 25 Mar 1908; Schreiter s n (LIL) Vipos, Department Trancas, 24 Jan 1926; 

Schreiter 1795 (LIL) Vipos, 8 Dec 1921; Venturi 1611 (A, LIL, US) Vipos, Department Trancas, 3 

Dec 1921; Venturi 7400 (US) Department Trancas, 15 Jan 1928. La Rioja: illegible 139 (S) La 

Aguadita, Department Capital, 30 Nov 1948; Venturi 8095 (GH, US-2) Nouogasta, Department 

Chilecito, 27 Dec 1928. Cordoba: Ariza 18565 (US) Inmediaciones de Carlos Paz, a Orillas Del Lago 

San Roque, Department Punilla, Dec 1965; Giardelli 828 (LIL, US) Ascochinga, 21 Nov 1936; 

Hunziker 10722 (US) Serranias Entre Villa de Maria Y San Miguel, Department Rio Seco, 11 Feb 

1955; Hunziker 9897 (US) Cerca de Cuchi Yaco, Al Este de Taninga, Department Pocho; 24 Feb 

1952; Kuntze s n (NY-2, US) Cordoba, Dec 1891; O'Donnell 4847 (F, S, W) Cruz Del Eje (Dique), 

Department Cruz Del Eje, 19 Jan 1947; Stuckert 12462 (LIL) Calera, Department Santamaria, 31 Dec 

1902; Stuckert 12756 (LIL) Department Santa Maria, Alta Gracia, 13 Feb 1903; Stuckert 18873 

(LIL) Sierra Chica de Cordoba, Mar 1908; Stuckert 22154 (LIL) Dique a Casa Bamba, Department 

Colon, 17 Feb 1911; Stuckert 4270 (LIL) Sierra Chica de Cordoba, 21 Jan 1898; Villafane 187 (S) 

Cruz Del Eje, Department Cruz Del Eje, 30 Dec 1946. San Luis: Galander s n (K) Quebrada Del Salada 

(Bebida de Las Vacas), 9 Mar 1882; Kurtz 8572 (NY) Sierra de San Luis, Bajo de Velis, El Totoral, 

Jan-Feb 1895. Province Unknown/Uncertain: Gerth s n (L) Quebrada de La Hurta, Bajo de Velis, 5 

Dec 1910. 

Manihot anisophylla, while distinct, shares many attributes with M. davisiae in 

section Parvibracteatae, in North America. 

35. Manihot guaranitica Chodat & Hassler. Bull. Herb. Boissier 2(5): 671. 1905. 

Shrubs, to 4.0 m tall, base swollen. Young stems glabrous, grayish brown or purplish 

brown. Leaves alternate; stipules ca 1.0 cm long, often persistent, glabrous, margin 

dentate, greenish yellow or purplish; petioles ca 14.0 cm long, terete, yellowish green or 

with purplish pigmentation, glabrous, petiole attachment to lamina mostly peltate, rarely 

basal, width between basal edge of lamina and petiole-lamina junction usually 1.0 cm or 

less; abaxial lamina surface wax pattern smooth; venation camptodromous, veins 

glabrous; lamina palmately 5 lobed rarely less than 5; median lobes obovate, obovate


: 

,- 

, 

-...- - - -,- -' 

i;~~~~~~~~~~~~~~~~~~~~~~" 

? 

C 

-?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~--?~~~~~~~~~~~~~ 

o \i~~~~~~~~~~~~~~~~~~ 

FG4.M.otispfl.A, di s i le btoB af(crir ,LL~C nlrsec V~ 

turi 8095, GH) M. guaranitic. D, distributon, dots repreent subsp guarnitica and theletter A repre 

~nts subsp~ ~~ ~ ~ 

oivin.. 

turi 8095, GH). M. guaranitica. D, distribution, dots represent subsp guaranitica and the letter A repre- 

sents subsp boliviana. 

'


pandurate or rhombic, ca 12.0 cm long, ca 7.0 cm wide, apex obtuse, acute or shortly 

angustate, base of lobes 1.0-2.0 cm wide, width between base of sinus and petiole-lamina 

junction ca 2.25 cm; lowest lobes ca 1/3 as long as median lobes, oblong with an obtuse 

apex. Inflorescence a monoecious, terminal, cluster of subspicate racemes arising from 

one basal point, ca 8.0 cm long, sometimes longer, usually with considerable amount of 

purplish pigmentation on peduncles, pedicels and tepals, sometimes without purplish 

pigmentation, usually with a bluish bloom over peduncles, pedicels and tepals, all parts 

glabrous; bracteoles setaceous to semifoliaceous, 1.0 cm long, 0.4 cm wide, margin 

smooth to dentate. Pistillate flowers restricted to the base of the upper half of the 

inflorescence, pedicels 1.25 cm long, tepal 1.0 cm long, cleft to base into 5 ovate lobes, 

ovary subglobose. Staminate buds more or less globular, flowers medium sized, tepal 1.0 

cm long, cleft 1/3 way down into 5 lobes, stamens 10, in 2 whorls of 5 each. Capsules 1.5 

cm long, subglobose, surface smooth, apex rounded. Seeds 1.25 cm long, caruncle poorly 

developed. 

DISTRIBUTION. (Fig 44D). Bolivia, Depts. of Cochabamba, Santa Cruz and Tarija; 

Paraguay, Depts. of Olimpo and Boqueron; Argentina, Prov. of Jujuy, Salta, Chaco, 

Catamarca and Tucuman. 

Key to the Subspecies of Manihot guaranitica 

1. Leaf lobes obovate, apex obtuse or acute; petioles and midribs usually purplish tinged. 

35a. M. guaranitica subsp guaranitica. 

1. Leaf lobes rhombic, apex acuminate; petioles and midribs without purplish tinge. 

35b. M. guaranitica subsp boliviana. 

35a. Manihot guaranitica Chodat & Hassler subsp guaranitica 

Janipha violacea Grisebach, Abh. Konighl. Ges. Wiss. Gottingen 24: 52. 1879; Manihot 

carthaginensis var anisophylla 0. Kuntze, Rev. Gen. 3(2): 288. 1898. ex parte (see also 34); 

Manihot recognita Pax in Engler, Pflanzenreich IV. 147(Heft 44): 91. 1910. Type. Lorentz 

& Hieronymus 330 (syntypes, F, NY-2, S). 

Manihot guaranitica Chodat & Hassler, Bull. Herb. Boissier 2(5): 671. 1905; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 75. 1910. Type. Hassler 3466 (syntype, W). 

Manihot fiebrigii Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 44): 75. 1910. Type. 

Fiebrig 10469 (syntypes, G-4). 

Manihot grandistipula Pax in Engler, Pflanzenreich IV. 147(Heft 44): 81. 1910. Type. Hassler 

2675 (syntypes, BM, G-2, K, P). 

Manihot anisitsii Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 197. 1924. 

Type. Anisits 1931 (syntypes, S). 

Petioles usually purplish tinged. Lamina widely peltate, rarely basal, width between 

basal edge of lamina and petiole-lamina junction ca 1.0 cm; median lobes obovate, apex 

obtuse or acute. Bracteoles semifoliaceous, margin dentate. Fig 45A, B. 

TYPE. Hassler 3466: Paraguay: Cordillera de Altos (syntype, W). 

DISTRIBUTION. (Fig 44D). BOLIVIA. Tarija: Pflanz 4035 (US) Villamontes. PARAGUAY. 

Olimpo: Fiebrig 10469 (G-4) Fuerte Olimpo, 1907; Rojas 13718 (S, W-2) Fuerte Olimpo, Chaco 

Paraguayo, 30 Oct 1946. Boqueron: Pedersen 4170 (P, UC, US) Puerto Casado et Vic., 24 Oct 1956. 

Province Unknown/Uncertain: Anisits 1931 (S) Ost Armonia, Auf Felsen, 17 Jan 1900;Balansa 1717 

(K, P-3) En Guarani, 31 Dec 1876; Hassler 1669 (BM, NY) Cordillera de Altos, Dec; Hassler 2675 

(BM, G-2, K, P) Gran Chaco, Santa Elisa, Dec 1903; Hassler 8497 (G-2) in Regione Cursus Superioris 

Fluminis Apa, Feb; Rojas 7206 (A-2) Puerto Casado, Chaco Paraguayo, Monte Ralo, Kilom. 160, Mar 

1935; Rojas 7221 (A) Picuiba, Chaco Paraguayo, Mar 1935. ARGENTINA. Jujuy: Jorgensen s n 

(BAB), Mar 1911; Lillo 5269C (LIL) Ledesma, 23 Dec 1906. Salta: Lorentz & Hieronymus 330 (F, 

NY-2, S) Am Rio Juramento, Feb 1873; Meyer 4805 (BM-2, F, LIL, UC) Campamento Y.P.F. Rio 

Pescado, Department Oran, 26 Feb 1943; Schreiter 10992 (LIL) Rio Tartagal, Department Oran, 4 

Jan 1940; Schreiter 3402 (LIL) Tartagal, Picada de Los Canas, 25 Oct 1924; Schreiter 4106 (LIL) 

Tartagal, Department Oran, Camino a La Picada de Las Canas, Feb 1925; Spegazzini s n (BAB), Mar 

1905; Venturi 8690 (US) Tartagal, Department Oran, 16 Dec 1926. Chaco: Meyer 2186 (GH, LIL) 

Zapallar, Mar 1937; Schultz 1550 (U) Las Brenas, Dec 1932; Schultz 1551 (U-3) Colonia Elisa, Nov 

1936. Catamarca: Schreiter 9068 (BM, F, LIL, UC) Cuesta de La Chilca, Department Andalgala, Dec


C 

~1 .. 

FIG 45. Manihot guaranitica subsp guaranitica. A, leaf (Schreiter 9068, LIL); B, inflorescence 

(Hassler 8497, G). M. guaranitica 

subsp subsp boliviana. C, leaf (Herzog (Herzog 1233, L).


1932. Tucuman: Bailetti 96 (LIL) Department Burroyaco, Jan 1918; Gonzales 21583 (LIL) 

Department Burroyaco, 20 Apr 1910; Lillo s n (LIL) Alurralde; Monetti 4897 (LIL) Department 

Burroyaco, 4 Apr 1914; Scbreiter s n (LIL) Department Trancas, 14 Nov 1929; Venturi 10006 (BM, 

GH, LIL, MO, S, US) Camino a Vipos, Department Trancas, 5 Nov 1929; Venturi 2685A (US) El 

Durazuito, Department Capital, 16 Feb 1924; Venturi 669 (LIL, US) Chanar Pozo, 27 Nov 1919. 

Province Unknown/Uncertain: Lillo 2531 (LIL), 13 May 1900;Schreiter 552 (LIL) Arenales Del Valle 

de Santa Maria, Dec 1918. Doubtfully assigned to this taxon: BRASIL. Minas Gerais: Saint-Hilaire s n 

(P-5) precise local unknown. 

LOCAL NAMES. Higuerilla (Gonzales 21583), Higuerita (Venturi 669). 

The characters given in the key distinguish this subspecies from boliviana (See Fig 

45A, B). Subspecies guaranitica occupies the southern part of the range of the species and 

occurs generally at lower altitudes than subspecies boliviana. 

35b. Manihot guaranitica Chodat & Hassler subsp boliviana (Pax & K. Hoffmann) Rogers 

& Appan, stat nov 

Manihot boliviana Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 63) Addit. V.: 402. 

1914. 

Petioles yellowish green. Lamina very narrowly peltate, rarely basal, width between 

basal edge of lamina and petiole-lamina junction less than 0.2 cm; median lobes rhombic, 

apex acuminate. Bracteoles setaceous, margin smooth. Fig 45C. 

TYPE. Herzog 1233: Bolivia, Santa Cruz: bei Charagua, Dec 1910 (syntypes, L, S). 

DISTRIBUTION. (Fig 44D). BOLIVIA. Cochabamba: Cardenas 3340 (F) Sailapata, Prov. 

Ayopaya, Nov. 1935. Santa Cruz: Cardenas 4729 (US) Camiri, Province Cordillera, Feb 1951. 

6. Manihot sect Carthaginenses Rogers & Appan, sect nov 

Manihot sect Heterophyllae Pax subsect Carthaginenses Pax in Engler, Pflanzenreich IV. 147(heft 

44): 80. 1910 pro parte. 

Plantae caulinae, lignosae, altae frutices. Folia petiolatae, penitae lobatae. Inflorescentia 

monoecia panicula. 

Plants caulescent; tall shrubs; leaves widely spaced on stem; petiolate, petiole 

attachment basal; lamina membranaceous, deeply lobed, lobes usually obovate and deeply 

pandurate. Inflorescence a monoecious panicle; bracts and bracteoles entire. 

setaceous, margins 

TYPE. Manihot carthaginensis 


(Jacquin) Muell.-Arg. 

Key to the Species of Sect Carthaginenses 

1. Erect shrubs; root system not significantly spreading laterally; filaments less than 1.0 cm 

long. 

36. M. carthaginensis. 

1. Decumbent shrubs; root system extensively spreading laterally, usually 3.0 m long or longer 

horizontally; filaments as long as 1.5 cm. 37. M. filamentosa. 

36. Manihot carthaginensis (Jacquin) Mueller von Argau in de Candolle, Prodr. 15(2): 

1073. 1866; in Martius, Fl. Bras. 11(2): 481. 1874; Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 81. 1910. 

Janipha frutescens Loefling, Iter Hispanicum 309. 1758 (fide Mueller 1866-Janipha frutescens 

Loefling, It. p 309 in quibusdam libris receptum omnio delendum est, frutescens enim non 

est nomen specificum, sed primum Verbum phraseos diagnosticae). 

Jatropha carthaginensis Jacquin, Select. Stirp. Am. Hist. 256 t. 162 f. 1. 1763; Jatropha 

cataginensis Jacquin ex Steudel, Nomencl. ed. 2. 1: 799. 1840 (orthographic variant). 

Jatropha frutescens Loefling, Iter Hispanicum (German edn.) 397. 1766.

0o 70 

ts~~~ OQ~ t . '|l_ _w_ 

FIG 46. Manihot sect Cartaginenses.. Geographical range.


Jatropha janipha L., Mant. 1: 126. 1771; Manihot janipha Pohl, P1. Bras. Ic. et Descr. 1: 55. 

1827. 

Janipha loeflingii H. B. K., Nov. Gen. et Spec. 2: 85. 1817. Type. Humboldt & Bonpland 1040 

(photo of syntype, P); Humboldt & Bonpland s n n v. 

Janipha yuquilla H. B. K., Nov. Gen et Spec. 2: 85. 1817. Type. Humboldt & Bonpland 149 

(photo of syntype, P); Janipha juquilla H. B. K. ex Steudel (orthographic variant);Janipha 

juquilla H. B. K. ex Muell.-Arg. in DC., Prodr. 15(2): 1073. 1866 (orthographic variant). 

Manihot pittierii Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 63): 401. 1914. 

Type. Pittier 1567 (syntypes, US-2). 

Manihot meridensis Pittier, Jour. Wash. Acad. 19: 352. 1929. Type. Jahn 1000 (syntypes, GH, 

US, VEN). 

Manibot remotiloba Pittier, Jour. Wash. Acad. 19: 352. 1929. Type. Pittier 11761 (syntypes, K, 

M, NY, P, US, VEN). 

Tall shrubs (1.5 m) to small trees (5.0 m); trunk at base 15.0-20.0 cm diam bark 

dark gray or reddish, smooth, almost glossy, peeling off in places. Roots tuberous, 

irregularly swollen; not significantly spreading laterally; epidermis smooth, dark brown, 

subepidermis white, with streaks of light pink; cortex very soft. Young stems 

obtuse-angled in cross section, glabrous, yellowish green; mature stems glabrous, light 

brownish gray, internally pale orange-yellow (7.5 YR 9/4). Leaves alternate; stipules 

setaceous, caducous, margin denticulate; petioles ca 20.0 cm long, terete, glabrous, 

greenish with maroon suffused or entirely dark red (2.5 R 3/7), petiole attachment to 

lamina basal, nonpeltate; lamina firmly membranaceous, dull blue-green above, silvery 

below, abaxial surface wax pattern smooth; venation camptodromous, veins glabrous; 

lamina palmately 5 lobed; median lobes obovate, usually deeply pandurate with a 

prominently dilated more or less triangular shaped apical region, rarely entire, ca 15.0 cm 

long, ca 6.0 cm wide, apex obtuse or acute, base breve angustatus, base of lobes ca 1.5 cm 

wide; lowest lobes ca 1/2 as long as median lobes, obovate, slightly nonsymmetric. 

Inflorescence a monoecious, terminal, sparsely branched panicle, ca 15.0 cm long, 

sometimes shorter, all parts glabrous; bracteoles and bractlets setaceous; pistillate flowers 

restricted to the base of the inflorescence, pedicels 2.0-3.0 cm long; tepal 1.1 cm long; 

disc yellow. Staminate buds, ovoid-ellipsoid, sometimes very slightly constricted in the 

middle, flowers medium sized, tepal 1.1 cm long, brilliant yellow (2.5 Y 9/9) at apex, 

deep maroon at base, cleft 1/3 way down into 5 lobes, stamens 10 arranged in 2 whorls of 

5 each, filaments of the superior whorl 0.9 cm long, of inferior whorl 0.7 cm long. 

Capsules subglobose, 1.25-1.5 cm long, 1.0 cm wide, pendent; surface smooth, without 

prominent wings; apex rounded; dehiscence septicidal; seeds 1.25 cm long, oblong, 

blackish brown. Fig 47B, C, D; 48A. 

TYPE. Jacquin s n: Colombia, Bolivar: Cartagena (holotype, P n v., isotypes, Photo 

at F, W). (Mueller 1866 erroneously cites the type local as Venezuela). 

DISTRIBUTION. (Fig 47A). West Indies, in Bonaire; Colombia, in LaGuajira Intendency 

and in the Depts. Atlantico, Magdalena, Bolivar and Cundinamarca; Venezuela in 

the states Zulia, Falcon, Nueva Esparta, Yaracuy, Aragua, Sucre, Merida, Bolivar and 

Distrito Federal; Trinidad and Tobago, Patos Island. 

Xerophytic scrub forest on limestone. Frequent in dry, open places and thickets by 

the sea coast. WEST INDIES. Bonaire: Boldingh 7430 (NY, U), 1909; Stoffers 534 (U), Jan 1954. 

COLOMBIA. La Guajira Intendency: Dawe 508 (US) Goajira, Oct; Dawe 552 (K-2, US) Goajira, 31 

Mar 1917. Atlantico: Dugand 4654 (NY, UC, US) alrededores de Barranquilla, 28 Jun 1953; Dugand 

5275 (W) costa del Caribe, Salgar, lomas arenosas cerca del mar, 14 Aug 1960; Dugand 5283 (W) costa 

del Caribe, Sabanilla, fruticetum espinoso circa del Mar, 14 Aug 1960; Dugand & Jaramillo 3231 (US) 

entre Puerto Columbia Y Salgar, 23 Jul 1943; Dugand & Jaramillo 3246 (US) Puerto Columbia, 

colinas aridas entre los Km 16 Y 17 de La Carretera, 24 Jul 1943; Elias 1546 (A, F, US) El Palmar, 

Region of Barranquilla, Jul 1937. Magdalena: Dugand 224/61 (F) Santa Rosa, near Barranquilla, Nov 

1932; Haught 4156 (A, F, US) near Papayal, 13 May 1944; Pittier 1567 (US-2) along roads around 

Barranquilla, Jun 1906; Pittier 1723 (US) lower hills around Rio Frio, Jun-Jul 1906; Smith 365 

(NY-3, S, U) Playa Brava, Santa Marta, Sep 1898. Bolivar: Curran 428 (GH, US) Norosi-Tiquisio Trail, 

Lands of Loba, Apr-May 1916; Killip & Smith 14032 (A, BM, F, GH, NY, US) vicinity of Cartagena,


(Rogers 494, NY); D, leaf with pandurate lobes (Dugand 4654, NY). 

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Nov 1926. Cundinamarca: Triana s n (BM-2, G, K, P) Tequendama Rio Seco, Valle du Magdalena, 

1853. Province Unknown/Uncertain: Dugand 46 (F), 6 Aug 1932; Espina 59 (F) Santa Marta Region, 

1931. VENEZUELA. Zulia: Plee 64 (P) Maracaybo. Falcon: Lasser & Aristequieta 3319 (VEN) 

Paraguana, Dec 1953. Nueva Esparta: Croizat s n (F) La Asuncion, Island of Margarita, Sep 1948. 

Yaracuy: Pittier 11761 (K, M, NY, P, US, VEN) between Yaritagua and Urachiche, Yaracuy, 10 Apr 

1925. Aragua: Aristiguieta & Saldivias 542 (VEN) La Cienega, Orumare de La Costa, Aragua, 19 Mar 

1951. Distrito Federal: Aristeguieta 4701 (VEN) near Arrecife, Jun 1961; Rogers 494 (F-2, G-2, K-2, 

MEXU, NY-3, US, W) 18 km W of Arrecife, on Caribbean Coast, 26 Feb 1962. Sucre: Funck 607 

(OXF) Cumana; Funcke 607 (K) Cumana; Steyermark & Agostini 91228 (VEN) Peninsula de Paria, 

Puerto de Hierro, colinas semixerofiticas a la base del Cerro, Jul 1962. Merida: Jahn 1000 (GH, US, 

VEN) Lagunillas, 13 Mar 1922. Bolivar: Aristeguieta 2234 (NY, VEN) Ciudad Piar, Apr 1954. 

Province Unknown/Uncertain: Curran & Haman 462 (GH, US) Vela de Coro, 2 Apr 1917. TRINIDAD 

TOBAGO. Patos Island: Broadway 7202 (F, NY, S, US) road to Flagstaff, 9 Jun 1929; Williams s n 

(NY) Trinidad, 25 Jun 1917; Williams 12516 (NY) Trinidad, 9 May 1931. 

LOCAL NAMES. Boracher (Curran & Haman 462); yuca antigua (Dugand & Jaramillo 

3231); yuca cimarrona (Dugand 46); yuca de monte (Dugand 4654); yuca escussonera 

(Espina 59); yuca sylvestre (Rogers 494). 

The history of the nomenclature of this species, first established by Jacquin in 1763, 

was confused by transfers from the genus Jatropha to Janipha by H. B. K., 1817, by 

recognition (by several authors incl. Linnaeus, 1771, and others) of Janipha frutescens as 

a separate, distinct species name given by Loefling, in 1758, and by the transfer of 

Jatropha janipha L. to Manihot janipha by Pohl. 

Mueller von Argau in de Candolle, Prodromus (1866), clarified the issue by 

recognizing that Loefling had not designated his plant with a separate species name, 

"frutescens," but that Loefling had merely used the term frutescens as a part of a 

description of a plant collected in Cumana (Venezuela). Pax (1910) first recognized that 

Janipha H. B. K. was wholly synonymous with Manihot although Pohl and other authors 

had earlier transferred species of Janipha to Manihot. 

Manihot carthaginensis grows on the drier coastal islands near Trinidad, westward 

along the north shore of S. America to Colombia. Its variable leaf form, common among 

many species of Manihot, caused much nomenclatural confusion and no doubt influenced 

the acceptance of Loefling's illegitimate name. Many plants of Manihot from Mexico and 

Central America have been misidentified by botanists as M. carthaginensis. 

Pohl (1827) redesignated Jatropha janipha Lour. (1793) as Manihot loureirii Pohl. 

Pax (1910) noted thatM. loureirii Pohl was probably a synonym of M. carthaginensis. We 

have placed M. loureirii Pohl (and Jatropha janipha Lour.) as synonyms of M. esculenta 

because of the extreme unlikelihood that M. carthaginensis would be found in southeast 

Asia, and that M. esculenta was probably already an accepted cultigen in that area by that 

time. 

The confused nomenclature is indicative of the variability within the species. It is 

probable that several ecotypic variations are discernable, but the herbarium material is 

sufficiently fragmentary to make decisions difficult. 

Manihot carthaginensis may play a role in plant breeding with M. esculenta, provided 

the breeder is seeking draught resistant cultivars, because the habitat of the species is the 

dry, xerophytic shrub zone immediately adjacent to the Caribbean Sea along the north 

coast of South America. Manihot carthaginensis is a very distinct species whose nearest 

relative is M. annisophylla. 

37. Manihot filamentosa Pittier, Jour. Wash. Acad. 20: 11. 1930. 

Shrubs 2.0- 3.0 m tall, with a tendency to be decumbent. Root system extensively 

spreading laterally, usually more than 3.0 m long horizontally. Stems glabrous. Leaves 

alternate; stipules setaceous, ca 1.0 cm long, margin denticulate, glabrous, caducous;


petioles ca 15.0 cm long, terete, glabrous, petiole attachment to lamina basal, nonpeltate; 

abaxial lamina surface wax pattern smooth, venation camptodromous, veins glabrous; 

lamina palmately 5 lobed; median lobes obovate, deeply pandurate with a prominently 

dilated more or less triangular apical region, ca 10.0 cm long, ca 4.0 cm wide, apex 

broadly acute, base breve angustatus, base of lobes ca 0.8 cm wide; lowest lobes ca z/ as 

long as median lobes, obovate, slightly nonsymmetric. Inflorescence a monoecious, 

terminal, sparsely branched panicle, ca 10.0 cm long, all parts glabrous, bracteoles and 

bractlets setaceous, pistillate flowers restricted to the base of the upper half of the 

inflorescence, pedicels 1.5 cm long, tepal non vidi; staminate buds ovoid-ellipsoid, flowers 

medium sized, tepal 0.9 cm long, cleft 1/3 way down into 5 lobes, stamens 10, in 2 

whorls of 5 each, filaments of fully developed androecium non vidi. (Fide original 

description, filaments very long, ca 1.5 cm in length). Fruits and seeds non vidi. 

Fig 48B, D and 49A. 

TYPE. Saer 366: Venezuela, Lara: Vicinity of Barquisimeto, Aug 1929 (syntypes, 

photo at F, US). 

DISTRIBUTION. (Fig 48C). Venezuela in the states of Lara and Guarico. Along 

margins of dry stream beds (fide Rogers 495. See Fig 48B). VENEZUELA. Lara: Saer 191 

(US, VEN) La Ruesga, Barquisimeto, 4 Apr 1925; Saer 367 (VEN) espinares arenosos de Las Vegas de 

La Quebrada la Ruesga, Vic. of Barquisimeto, Aug 1929. Guarico: Aristeguieta 4675 (VEN) circa de 

San Juan de Los Morros, Via Ciudad Bolivar, Jun 1961; Rogers 495 (G, NY, US)-stem and root 

specimens only-dry creek bed, near San Juan de los Morros, 27 Feb 1962. 

LOCAL NAME. Yuca sibidigua (Saer 191). 

Manihot filamentosa remains poorly defined because of the general paucity of good 

herbarium materials. Manihot carthagenensis is its closest relative from which it is 

distinguished mainly by its decumbent habit and its distribution, inland from the coast. 

7. Manihot sect Quinquelobae Pax emend Rogers & Appan 

Manihot sect Grandibracteatae Pax subsect Glabrescentes Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 27. 1910 pro parte. 

Manihot sect Grandibracteatae Pax subsect Coerulescentes Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 29. 1910 pro parte. 

Manihot sect Parvibracteatae Pax subsect Tristes Pax in Engler, Pflanzenreich IV. 147(Heft 44): 



44): 66. 1910 pro parte. 

Manihot sect Quinquelobae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 87. 1910. 

Plants caulescent; low to medium shrubs; leaves widely spaced on stem; petiolate, 

petiole attachment basal or narrowly peltate; lamina coriaceous, deeply lobed, lobes 

variously shaped but not linear. Inflorescence a monoecious raceme, simple or a cluster of 

several racemes arising from a common base; bracts and bracteoles setaceous to 

foliaceous, margins usually entire. 

TYPE. Manihot quinqueloba Pohl. 


Key to the Species of Sect Quinquelobae 

1. Bracts and bracteoles setaceous, less than 0.2 cm wide; leaf lobes not overlapping at sinus. 

2. Inflorescence subspicate. 

3. Leaves 3 lobed; median lobe midribs straight; bracteoles glabrous. 

4. Inflorescence usually less than 4.0 cm long; leaf lobes less than 2.0 cm wide; midrib 

of basal lobes curved down; leaves held in a vertical position on the plants. 

5. Inflorescence a cluster of racemes arising from a common base; leaf lobes gladiate; 

midrib of basal lobes slightly curved. 38. M. acuminatissima. 

5. Inflorescence a simple raceme; leaf lobes ovate-elliptic; midrib of basal lobes 

strongly curved. 39. M. sagittato-partita.


FIG 49. Manihot filamentosa. A, inflorescence (Saer 191, US). 

4. Inflorescence more than 4.0 cm long; leaf lobes more than 2.0 cm wide; midrib of 

basal lobes straight; leaves held horizontally. 

6. Base of leaf lobes constricted, less than 0.5 cm wide, lobes usually less than 8.0 cm 

long; midribs slender, less than 0.1 cm in diam; petioles less than 5.0 cm long. 

40. M. xavantinensis. 

6. Base of leaf lobes not constricted, more than 0.5 cm wide; lobes more than 8.0 cm 

long; midribs stout, more than 0.1 cm in diam near the petiole-lamina junction; 

petioles more than 5.0 cm long. 

41. M. sparsifolia. 

3. Leaves 7-11 lobed; median lobe midribs strongly curved, (reflexed); bracteoles 

pubescent. 

42. M. falcata. 

2. Inflorescence racemose, not subspicate. 

7. Stipules persistent; petiole attachment peltate; petioles less than 5.0 cm long; leaf lobes 

less than 7.0 cm long. 

43. M. pruinosa. 

7. Stipules caducous; petiole attachment basal; petioles more than 5.0 cm long; leaf lobes 

more than 7.0 cm long. 

8. Inflorescence less than 5.0 cm long; leaf lobes obovate, more than 2.5 cm wide, apices 

truncate. 44. M. quinqueloba. 

8. Inflorescence more than 5.0 cm long; leaf lobes digitiform, less than 2.5 cm wide; 

apices acute. 45. M. alutacea. 

1. Bracts and bracteoles foliaceous or semifoliaceous, usually more than 0.2 cm wide; leaf lobes 

overlapping at sinus. 

9. Inflorescence a simple raceme; leaf lobes less than 8.0 cm long. 

10. Inflorescence less than 5.0 cm long, few-flowered, flaccid; leaf lobes obovate or 

elliptic. 

46. M. violacea. 

10. Inflorescence more than 5.0 cm long, many flowered, erect; leaf lobes ovate to 

rotundate. 47. M. jacobinensis. 

9. Inflorescence a cluster of several racemes arising from a common base; leaf lobes greater 

than 8.0 cm long. 

11. All parts except interior surface of tepals glabrous. 

12. Inflorescence sparsely branched, with few flowers; leaf lobes less than 5.0 cm wide; 

apices acuminate. 48. M. divergens. 

12. Inflorescence profusely branched, with many flowers; leaf lobes more than 5.0 cm 

wide; apices obtuse, acute, or caudate.


1LI? Cf? 

,' Z 4 

r.. ,,~ 

FIG 50. Manihot sect Quinquelobae. Geograph~~~iclrne


13. Bracts and bracteoles semifoliaceous, less than 0.5 cm wide; leaves 3 lobed; tepal 

lobes usually purplish tinged. 

49. M. irwinii. 

13. Bracts and bracteoles foliaceous, more than 0.5 cm wide; leaves usually 5 lobed, 

rarely 3; tepal lobes without purplish pigmentation. 

50. M. cecropiaefolia. 

11. Leaf lobes, petioles, peduncles, bracts, bracteoles, tepal etc., densely pubescent. 

51. M. mossamedensis. 

38. Manihot acuminatissima Mueller von Argau in Martius, Fl. Bras. 11(2): 455. 1874; 

Pax in Engler, Pflanzenreich IV, 147(Heft 44): 66. 1910. 

Subshrubs. Stems glabrous. Leaves alternate; stipules caducous; petioles ca 7.0 cm 

long, terete, glabrous, petiole attachment to lamina basal, nonpeltate; lamina, adaxial 

surface dark green, abaxial surface glaucous, abaxial surface wax pattern smooth; 

venation camptodromous, veins glabrous, midribs prominent; lamina 3 lobed; median 

lobes gladiate, ca 8.0 cm long, ca 1.75 cm wide, rarely larger, apex acuminate, sinus of 

median lobes forming a wide angle of more than 90?; lowest lobes more or less same as 

median lobes, midribs slightly curved downwards. Inflorescence monoecious, terminal, a 

cluster of short subspicate racemes arising from one basal point, ca 3.5 cm long, all parts 

glabrous, peduncles, pedicels and tepals with a bluish white bloom; bracteoles and 

bractlets setaceous, less than 0.3 cm long, less than 0.1 cm wide. Pistillate flowers 

restricted to the base of the upper half of the inflorescence; pedicels 0.4 cm long; tepal 

0.6 cm long, cleft to base into 5 lobes; ovary subglobose. Staminate buds ovoid-ellipsoid; 

flowers small, tepal 0.5 cm long, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls 

of 5 each. Fruits and seeds non vidi. Fig 51B, C. 

TYPE. Gardner 3446: Brasil, Goias, 1841 (syntypes BM-2, photo at F, G, K-2, photo 

at NY, OXF, P, W-2). 

DISTRIBUTION. (Fig 51A). Brasil in states Goias and Bahia. BRASIL. Goias: Burchell 

7774 (K). Bahia: Luetzelburg 12253 (NY) Rio de Contas, Formosas, Jul 1913. 

Manihot acuminatissima is a close relative of M. sagittato-partita, joining with the 

latter at a similarity value of .90 (Fig 28). The main difference between the two is in the 

nature of the inflorescence, the major key character. 

39. Manihot sagittato-partita Pohl, PI, Bras. Ic. et Descr. 1. 22, t. 15. 1827; Muell.-Arg. in 

DC., Prodr. 15(2): 1060. 1866; in Martius, Fl. Bras. 11(2): 454. 1874; Pax in 


Jatropha sagittato-partita Steudel, Nomencl. ed. 2. 1: 800. 1840. 

Subshrubs, to 0.5 m tall. Stems glabrous. Leaves alternate; stipules caducous; petioles 

ca 5.0 cm long, terete, glabrous, petiole attachment to lamina basal, nonpeltate; lamina 

coriaceous, held vertically on the plant, adaxial surface dark green, abaxial surface wax 

pattern smooth; venation camptodromous, veins glabrous; lamina sagittately 3 lobed; 

median lobes ovate-elliptic, ca 8.0 cm long, ca 1.75 cm wide, apex acuminate, sinus of 

median lobes forming a wide angle of more than 90?; lowest lobes more or less same as 

median lobes, midribs strongly curved downwards. Inflorescence a monoecious, terminal, 

short raceme, ca 3.0 cm long, all parts glabrous; bracteoles and bractlets setaceous less 

than 0.2 cm long, less than 0.1 cm wide. Pistillate flowers restricted to the base of the 

inflorescence, several flowers arising from the base of the rachis more or less as a whorl; 

pedicels 1.0 cm long; tepal 1.0 cm long, cleft to base into 5 lobes, yellowish green; ovary 

subglobose. Staminate buds ovoid-ellipsoid, flowers small, tepal 1.1 cm long, yellowish 

green, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Fruits and 

seeds non vidi. Fig 52A, B.


i 

''~~~~~~~~~~~~~~~~~~~~~~~' 

' 

FIG 51. Manihot acuminatissima. A, distribution; B, leaf (Gardner 3446, K); C, inflorescence 

(Gard ner 3446, BM) . M. sagittato -partita. D, distribu tion.

Systematic 


i1 

i11.1 f. 

FIG 52. Manihot sagittato-partita. A, leaf (Pohl 1704, G); B, inflorescence (Irwin, Souza & dos 

Santos 9966, NY). M. xavantinensis. C, distribution; D, leaf (Argent, Ramos, Richards & Souza6675, 

K) 

i


TYPE. Pohl 734 (and on some specimens also 1704): Brasil: ca Villa Paracatu do 

Principe, Capitaniae Minas Geraes, et ad Serra do Cristaes, Capitaniae Goias. (Syntypes, F, 

G, K-2, W-2.) 

DISTRIBUTION. (Fig 5 1D). Brasil in the states Goias and Minas Gerais. Alt ca 

1000 m.BRASIL. Goias: Irwin, Souza & Dos Santos 9966 (F, G, K, MO, NY, RB, SP, UB, US) Circa 

20 km W of Cristalina, 5 Nov 1965; Pohl Icon Tab. 15 (W-2). 

40. Manihot xavantinensis Rogers & Appan, sp nov 

Frutices gracili, ad 1.0 m altus. Folia trilobus; lobi median ellipticis, ca 7.0 cm longa, 

ca 3.0 cm lata; basis loborum constrictus, 0.4 cm latae. Inflorescentia fasciculus 

subspicatorum racemi, ca 8.0 cm longa, omnies partes glabra; bracteolae et ramuli 

setiformis. Fructi parvi, 1.1 cm longae. Semina prominens caruncula. 

Slender shrubs, to 1.0 m tall, latex white. Young stems glabrous, brown with 

purplish tinge; mature stems glabrous, dark brown with purplish tinge. Leaves alternate; 

stipules caducous; petioles less than 5.0 cm long, glabrous, terete, purplish, petiole 

attachment to lamina basal, nonpeltate; lamina coriaceous, abaxial surface dark green, 

abaxial surface glaucous, abaxial surface wax pattern reticulate with discrete rings at 

magnifications of X 40; venation camptodromous, veins glabrous, midribs slender, less 

than 0.1 cm in diameter; lamina palmately 3 lobed; median lobes elliptic, ca 7.0 cm long, 

ca 3.0 cm wide, apex acute, base acute, base of lobes constricted, 0.4 cm wide, width 

between base of sinus and petiole-midrib juhction less than 0.3 cm, lobes slightly 

overlapping at sinus; lowest lobes more or less same as median lobes in outline and size, 

slightly nonsymmetric. Inflorescence monoecious, terminal, a cluster of subspicate 

racemes arising from one basal point, ca 8.0 cm long; all parts glabrous; bracteoles and 


restricted to the base of the upper 1/3 of the inflorescence, pedicels 1.5 cm long, tepal 

0.8 cm long, yellowish green internally, purplish tinged externally, cleft to base into 5 

strap-shaped lobes, ovary elongated. Staminate buds ovoid-ellipsoid, flowers small, tepal 

0.7 cm long, yellowish green internally, purplish tinged externally, cleft 1/3 way down 

into 5 lobes, stamens 10, in 2 whorles of 5 each, superior whorl 0.6 cm long, inferior 

whorl 0.5 cm long, anthers deep yellow. Capsules small, 1.1 cm long; surface smooth, 

without wings; apex rounded; dehiscence septicidal. Seeds small 0.8 cm long, 0.5 cm 

wide, oblong; caruncle very prominent, nearly 1/3 as long as the seeds. Fig 52D and 53A. 

TYPE. Harley, Souza & Fereira 10381: Brasil, Mato Grosso: Route 3, ca 2 km SE of 

expedition (Royal Society/Royal Geographical Society expedition 1967-1969) base 

camp at 12? 49' S, 51? 46' W, 1 Oct 1968 (holotype, K; isotype, NY). 

DISTRIBUTION. (Fig 52C). BRASIL. Mato Grosso: Argent, Ramos, Richards & Souza 6675 

(K-2) Circa 1 km W of Base Camp at 12.49? S-51.460 W secondary vegetation on airstrip, 6 Oct 1967; 

Philcox, Fereira & Bertoldo 3283 (K) Circa 1 km W of km 247 Xavantina-Cachimbo Road, 28 Nov 

1967; Pbilcox, Ramos & Sousa 3105 (K-2) km 264 Xavantina-Cachimbo Road, 17 Nov 1967. 

LOCAL NAME. Mandioca (Argent et al 6675); mandioca braba (Harley et al 10381) 

This distinct new species was represented by very adequate herbarium collections. Its 

closest relative is Manihot sparsifolia from which it is distinguished by the constricted leaf 

lobe bases, slender midribs and shorter petioles. 

41. Manihot sparsifolia Pohl, P1. Bras. Ic. et Descr. 1. 26, t. 20. 1827; Muell. Arg. in DC., 

Prodr. 15(2): 1060. 1866; in Martius, Fl. Bras. 11(2): 452. 1874; Pax in Engler, 


Jatropha sparsifolia Steudel, Nomencl. ed. 2. 1: 800. 1840. 

Manibot amaroleitensis Baillon, Adansonia 4: 281. 1863. Type. Weddell 2862 (syntypes, F, P).


Ascending subshrubs, to 1.0 m tall. Root not prominently tuberous; epidermis dark 

brown, rough. Stems glabrous, purplish tinged. Leaves alternate; stipules caducous; 

petioles ca 10.0 cm long, terete, glabrous, purplish tinged, petiole attachment to lamina 

basal, nonpeltate; lamina coriaceous, adaxial surface dark green, abaxial surface glaucous, 

abaxial surface wax pattern spongiose; venation camptodromous, veins glabrous, midribs 

prominently stout, 0.2 cm in diam near the petiole-lamina junction; hastately 3 lobed; 

median lobes ovate-lanceolate to elliptic, ca 13.0 cm long, ca 4.0 wide, entire, apex 

acuminate, base acute, base of lobes ca 2.0 cm wide, lobes at sinus not overlapping, width 

between base of sinus and petiole-lamina junction ca 1.5 cm; lowest lobes similar to 

median lobes in outline and size. Inflorescence monoecious, terminal, a cluster of long 

subspicate racemes arising from one basal point, ca 15.0 cm long; all parts glabrous; 

bracteoles and bractlets setaceous, less than 0.3 cm long, less than 0.1 cm wide. Pistillate 

flowers restricted to the base of the upper half of the inflorescence; pedicels 0.3 cm long; 

tepal 0.7 cm long, yellowish green with purplish tinge, cleft to base into 5 lobes. 

Staminate buds ovoid-ellipsoid; flowers small; tepal 0.6 cm long, yellowish green with 

purplish streaks along lobe edges, cleft 1/3 way down into 5 lobes; stamens 10, in 2 

whorls of 5 each. Capsules subglobose to slightly elongated; surface smooth, without ribs; 

apex rounded. Seeds non vidi. Fig 53C and D; 54A. 

TYPE. Pohl 2206: Brasil, Goias: ad Trahiras (syntypes, F, G, W-2). 

DISTRIBUTION. (Fig 5 3B). Brasil in the state Goias. Alt ca 750 m. BRASIL. Goias: 

Gardner 3974 (K), Mar 1840; Irwin, Maxwell, & Wasshausen 18926 (NY, UB) 75 km N of Corumba de 

Goias on Road to Niquelandia, Goias in Valley of Rio Maranhao, 21 Jan 1968; Irwin, Maxwell & 

Wasshausen 18997 (NY, UB) 75 km N of Corumba de Goias on Road to Niquelandia, Goias in Valley 

of Rio Maranhao, 22 Jan 1968; Irwin, Maxwell & Wasshausen 19082 (F, NY, UB) 60 km N of 

Corumba de Goias on Road to Niquelandia, 23 Jan 1968; Irwin, Souza & dos Santos 11757 (F, GH, K, 

MO, NY, RB, SP, UB, US) Campo and Cerrado Sandstone Summit of Serra Dourada, 18 Jan 1966; 

Pohl Icon Tab (W); Ule 50 (P) Serra de Balisa, Sep 1892; Weddell 2862 (F, P) Sertao d'Amaroleite, 

Sep-Oct 1844. 

42. Manihot falcata Rogers & Appan, sp nov9 

Subfrutices ad 0.5 m altus. Folia 7-11 lobatus; omniae lobi falcatus reflexus; lobi 

mediani ca 6.0 cm longae, oblongus-linearis. Inflorescentia fasciculus subspicatorum 

racemi, ca 5.0 cm longa, pistillatus et staminatus flores portati in separati pedunculi in 

eadam inflorescentia, pedunculi pistillati plerumque ferens 2 flores ad apicalis extremum; 

bracteolae pubescens, capsulae 1.25 cm longae, subglobosi. Semina non vidi. 

Ascending subshrubs, to 0.5 m tall. Roots not prominently tuberous; epidermis 

brown, rough. Young stems glabrous, grayish brown; mature stems glabrous, brownish 

with some purplish tinge. Leaves alternate; stipules setaceous, 0.5 cm long, glabrous, 

sometimes serrate; petioles ca 7.0 cm long, terete, glabrous, petiole attachment to lamina 

basal, nonpeltate; lamina coriaceous, adaxial surface dark green, abaxial surface wax 

pattern verrucate, edges of lamina purplish; venation camptodromous, veins glabrous, 

midribs purplish tinged towards the base; lamina palmately 7-11 lobed, all lobes falcately 

reflexed; median lobes oblong-linear, ca 6.0 cm long, apex acuminate, base acute, base of 

lobes 0.5 cm wide; lowest lobes similar to median lobes in outline, smaller in size. 

Inflorescence monoecious, terminal, a cluster of subspicate racemes, ca 5.0 cm long, 

pistillate and staminate flowers borne on separate peduncles in the same inflorescence; 

bracteoles setaceous, 0.4 cm long 0.2 cm wide, pubescent, bractlets setaceous, pubescent. 

Pistillate flowers borne at the apical end of peduncles, usually 2 flowers on each 

peduncle; pedicels curved down, 0.4 cm long, tepal 0.7 cm long, purplish externally, 

yellowish internally, interior surface pubescent, cleft to base into 5 lobes; ovary 

subglobose, glabrous. Staminate buds ovoid-ellipsoid; flowers small, tepal 0.6 cm long, 

9The leaf lobes are falcate, or distinctly curved.


FIG 53. Manihot xavantinensis. A, inflorescence (Harley, Souza & Fereira 10381, K). M. sparsifolia. 

B, distribution; C, leaf (Irwin, Maxwell & Wasshausen 19082, NY); D, inflorescence (Irwin, Maxw~ell 

well & Wasshausen Wassbausen 18997, NY).


purplish externally, yellowish internally, interior surface pubescent, cleft 1/3 way down 

into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules 1.25 cm long, subglobose, 

surface smooth, apex rounded. Seeds non vidi. Fig 54C, D. 

TYPE. Irwin, Souza & dos Santos 11039: Brasil, Goias: near Rio Descoberto, on the 

Goias-Distrito Federal Boundary between Anapolis and Brasilia. Alt 1010 m, 4 Dec 1965 

(holotype, NY; isotypes, F, G, K, UB, US). 

DISTRIBUTION. (Fig 54B). Brasil, Goias and Distrito Federal. Alt ca 1000 m. BRASIL. 

Goias: Glaziou 22137A (P) Guariroba, Au Morro Cubatio Entre Les Pierres, 10 Oct 1894. Distrito 

Federal: Irwin, Souza & dos Santos 10727 (F, S, UB, US) Campo near Aparecido Goias ca 50 km W of 

Brasilia, 29 Nov 1965. 

The specific epithet designates the most outstanding feature of this species. Its 

closest relative is Manihot xavantinensis. 

43. Manihot pruinosa Pohl, PI. Bras. Ic. et Descr. 1: 28. t. 22. 1827. 

Jatropba pruinosa Steudel, Nomencl. ed. 2. 1: 800. 1840; Manihot pruinosa Pohl emend. 

Muell.-Arg. var genuina Muell.-Arg. in DC., Prodr. 15(2): 1060. 1866. 

Manibot pruinosa Pohl emend. Muell.-Arg. var pumila Muell.-Arg. in DC., Prodr. 15(2): 1061. 

1866. Type. Riedel s n n v. 

Manihot burchellii Muell.-Arg. in Martius, Fl. Bras. 11(2): 457. 1874. Type. Burchell 7849 

(syntypes, F, G, K). 

Manibot pseudopruinosa Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 44): 61. 

1910 pro parte (see also 46b). Type. Glaziou 22131 pro parte (syntype, K). 

Shrubs, to 2 m tall, ascending, sometimes with a tendency to be prostrate, with 

copious white latex. Stems glabrous. Leaves alternate; stipules persistent, ca 1.0 cm long, 

ca 0.1 cm wide, laciniate, glabrous; petioles ca 4.0 cm long, rarely longer, terete, glabrous, 

petiole attachment to lamina narrowly peltate, width between the basal edge of lamina 

and petiole-lamina junction ca 0.2 cm; lamina coriaceous, adaxial surface dark bluish 

green, abaxial surface pruinose and bluish white, abaxial surface wax pattern pusticulate; 

venation camptodromous, veins glabrous; lamina palmately 3-5 lobed, leaves associated 

with inflorescence frequently nonlobed; median lobes obovate, ca 6.0 cm long, ca 2.5 cm 

wide, apex obtuse or acute, base acute, base of lobes 0.8 cm wide, lobes at sinus 

sometimes slightly overlapping; lowest lobes more or less similar to median lobes in 

outline, slightly smaller, nonsymmetric. Inflorescence a monoecious, terminal, raceme, ca 

12.0 cm long; bracteoles and bractlets setaceous, glabrous, ca 0.8 cm long, ca 0.1 cm 

wide. Pistillate flowers restricted to the base of upper 1/3 of the inflorescence; pedicels 

0.9 cm long, glabrous; tepal 0.7 cm long, interior surface pubescent, cleft to base into 5 

lobes; ovary subglobose, glabrous. Staminate buds ovoid-ellipsoid, slightly constricted in 

the middle; flowers small, tepal 0.7 cm long, cleft 1/3 way down into 5 lobes; stamens 10, 

in 2 whorls of 5 each. Capsules ovoid-ellipsoid, surface not winged, apex rounded. Seeds 

non vidi. Fig 55B, C. 

TYPE. Pohl 1705 (and on some specimens also 2469): Brasil, Goias: ad flumen Rio 

Claro (syntypes, F, G, K-2, P, W-2). 

DISTRIBUTION. (Fig 55A). Brasil, states of Mato Grosso and Goias. BRASIL. Mato Grosso: 

Gaudichaud 299 (P), 1833; Harley, Lima, Onashi & Souza 10583 (K) ca 8 km S of Base Camp at 

12.49?S-51.46?W, 10 Oct 1968. Goias: Burchell 7849 (F, G, K) Inter Goyaz et Cavalcante; Gardner 

3443 (BM, K, W), Jan 1840; Glaziou 22131 (K), 1898; Pohl Icon Tab (W); Ule 475 (P-2) Mossamedes, 

Jan 1893. Province Unknown/Uncertain: Tamberlik s n (W). 

Doubtfully assigned to this species: 

Brasil. Mato Grosso: Irwin & Soderstrom 6788 (NY, UB) vic. of Xavantina, 12 Oct 1964; 

Maguire, Pires, Maguire & Silva 56836 (NY) 350-440 km E of Vilhena, 25 Sept 1963. 

LOCAL NAME. Mandioca braba (Harley et al 10583).


223-AP. 1 

/ 

{'sI I 

i~~~~~~~~~~~~~~~~~~? 

W 

x~~~~r 

*I 4 aio prioi.A tmnt lwes(ri,Mxel&Wshue 89,N) 

M.flaa ,dsrbto;C ef(ri,Suaf o ats109 Y;,ifoecne(lzo 

FIG 54. Manihot sparsifolia. A, staminate flowers (Irwin, Maxwell & Wasshausen 18997, NY). 

M. falcata. B, distribution; C, leaf (Irwin, Souza & dos Santos 11039, NY); D, inflorescence (Glaziou 

22137-A, P). 

01 

fI


/4 7 9 

( I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

~~~~~~i~~~~~~~~~~~--~~~~~~~~i,' ?i-1 

o 

..i 

_ 

- C , f N Xt, 

;~~ 

FIG 55. Manihot pruinosa. A, distribution; B, leaf (Pobl 1705, K); C, inflorescence (Harley, 

Lima, Onasbi & Souza 10583, K). M. quinqueloba. D, distribution. 

129


The specimens of Glaziou 22131 (the type of Pax's Manihot pseudopruinosa) 

represent a mixture between this species and M. violacea subsp recurvata. The specimen at 

Kew is M. pruinosa. The specimens of Glaziou 22131 at Arnold Arboretum, Paris and 

Stockholm are M. violacea subsp recurvata. 

44. Manihot quinqueloba Pohl, PI. Bras. Ic. et Descr. 1: 21. t. 14. 1827; Muell.-Arg. in 

DC., Prodr. 15(2): 1058. 1866; in Martius, Fl. Bras. 11(2): 446. 1874; Pax in Engler, 


Jatropha quinqueloba Steudel, Nomencl. ed. 2. 1: 800. 1840. 

Manihot subquinqueloba Muell. Arg. in Martius, Fl. Bras. 11(2): 446. 1874. Type. Manso s n 

(syntypes, F, G). 

Manihot polyantha Pax in Engler, Pflanzenreich IV. 147(Heft 44): 88. 1910. Type. Glaziou 

22133 (syntypes, F, G, K, NY, P, US). 

Shrubs. Young stems glabrous, with a bluish bloom; mature stems glabrous. Leaves 

alternate; stipules caducous; petioles ca 9.0 cm long, terete, glabrous, with a bluish 

bloom, petiole attachment to lamina basal, nonpeltate; lamina coriaceous, adaxial surface 

dark green, abaxial surface with thick bluish bloom, abaxial surface wax pattern 

pusticulate; venation camptodromous, veins glabrous, midribs prominently stout, ca 0.15 

cm in diam near the petiole midrib junction; lamina palmately 5 lobed; median lobes 

obovate, ca 9.0 cm long, ca 5.0 cm wide, apex obtuse or truncate, base of lobes ca 2.25 

cm wide, width between base of sinus and petiole-lamina junction ca 2.25 cm; lowest 

lobes nonsymmetric. Inflorescence monoecious, terminal, a cluster of racemes arising 

from one basal point, ca 4.0 cm long, all parts except the interior surface of tepal 

glabrous, peduncles, pedicels and tepals with a thick bluish bloom; bracteoles and 


restricted to the base of the upper 2/3 of the inflorescence; pedicels 1.0 cm long; tepal 

0.9 cm long, cleft to base into 5 lobes, interior surface of tepal pubescent; ovary 

subglobose. Staminate buds ovoid-ellipsoid, slightly constricted in the middle; flowers 

small; tepal 0.8 cm long, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 

each. Capsules 1.4 cm long, dehiscence septicidal. Seeds non vidi. Fig 56 A, B. 

TYPE. Pohl 1701 (and on some specimens also 1902) Brasil, Goias: ad Trahiras et 

Cocal (Syntypes, F, G, K-2, W-2). 

DISTRIBUTION. (Fig 55D). Brasil, states of Mato Grosso and Goias. BRASIL. Mato Grosso: 

Manso s n (F, G). Goias: Glaziou 22133 (F, G, K, NY, P, US), 22 Jan 1895; Pobl Icon Tab (W). 

Neither Manso nor Glaziou collections give the precise locality which accounts for 

the single dot in the distribution map of this species. 

45. Manihot alutacea Rogers & Appan, sp nov 10 

Frutices, ca 1.5 m altus. Lamina alutaceus in textura, petioli, vene et margines 

laminarum vividum rubrum; palmatus 5 lobatus; lobi mediani digitiformi, ca 9.0 cm 

longae, ca 2.0 cm latae. Inflorescentia fasciculus racemorum exoriens unum punctum 

basalis, ca 15.0 cm longa, tepala flavovirens purpureo pigmentum basi. Fructi et seminae 

non vidi. 

Shrubs, ca 1.5 m tall; with strong HCN smell. Young stems, glabrous, deep red; 

mature stems glabrous, with some purplish pigmentation. Leaves alternate; stipules 

caducous; petioles ca 12.0 cm long, obtuse-angled, glabrous, bright red, petiole 

attachment to lamina basal, nonpeltate; lamina alutaceus in texture, adaxial surface dark 

green, glossy, abaxial surface wax pattern smooth, venation camptodromous, veins 

glabrous, bright red, midribs stout, 0.1 cm in diameter near the petiole midrib junction; 

10 = tanned leather.


i2 & : '~ !iii!i' ? ~ 

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auae.C d 

6 

FI aiotuin A eoa la(Gaio223,P;Binlrsec(Gaou21,P) 

, 

isrbto;Dlef(alyBarseta.10,K)


lamina palmately 5 lobed; median lobes digitiform, ca 9.0 cm long, ca 2.0 cm wide, apex 

acute, base of lobes 1.0 cm wide, width between lobe sinus and petiole-lamina junction 

1.5 cm, edge of lamina bright reddish, lowest lobes more or less similar to median lobes in 

outline, slightly smaller. Inflorescence monoecious, terminal, a cluster of racemes arising 

from one basal point, long, ca 15.0 cm in length, all parts except the interior surface of 

tepals glabrous; bracteoles and bractlets setaceous, less than 0.3 cm long, less than 0.2 cm 

wide, glabrous, margin sometimes serrate. Pistillate flowers restricted to the base of the 

upper 1/2 of the inflorescence; pedicels 1.5 cm long; tepal 1.0 cm long, yellowish green 

with purplish pigmentation at the base, interior surface pubescent, cleft to base into 5 

lobes, ovary subglobose with reddish ribs. Staminate buds ovoid-ellipsoid; flowers 

medium sized, tepal 1.0 cm long, yellowish green with purplish pigmentation at the base, 

cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each, the superior whorl 

0.8 cm long, the inferior 0.6 cm long. Fruits immature with bright red ribs; mature fruit 

ca 1.3 cm long, globose, surface punctate. Seeds 0.9 cm long, caruncle well developed. 

Fig 56D, 57A. 

TYPE. Harley, Barroso et al 11404: Brasil, Goias: Chapada dos Veadeiros, ca 65 km 

due N of Brasilia, in rocky cerrado at summit of serra, 21 Dec 1968 (holotype, K; 

isotype, NY). 

DISTRIBUTION. 

(Fig 56C). Brasil, Goias, alt ca 1250 m.BRASIL. Goias: Irwin, Harley & 

Smith 32953 (NY) Campo and cerrado on outcrops, ca 22 km N of Alto do Paraiso, 22 Mar 1971. 

This quite distinct species has as its closest relative Manihot quinqueloba from which 

it is distinguished by its digitiform leaf lobes. The texture of the leaf is like tanned 

leather. 

46. Manihot violacea Pohl, PI. Bras. Ic. et Descr. 1: 43. t. 35. 1827. 

Subshrubs to 1.0 m tall; several ascending stems (3-8) from a woody base. Young 

stems glabrous, sometimes with a light bloom; mature stems glabrous. Leaves alternate; 

stipules caducous, filiform, less than 0.5 cm long, glabrous; petioles ca 5.0 cm long, 

terete, glabrous, purplish tinged, petiole attachment to lamina peltate; lamina coriaceous, 

adaxial surface dark green, abaxial surface with a thick bluish bloom or light bloom, 

sometimes without any bloom, abaxial surface wax pattern smooth, abaxial surface with 

short shiny bristles visible at magnifications of X 40; venation camptodromous, veins 

glabrous, purplish tinged towards the base; lamina 3 lobed; leaves associated with the 

inflorescence frequently nonlobed, lobes overlapping at sinus; median lobes obovate to 

elliptic, ca 6.0 cm long, 2.0-3.5 cm wide, apex obtuse or acute, base of lobes usually 

constricted, less than 0.5 cm wide, lowest lobes more or less similar to median lobes in 

size, sometimes recurved. Inflorescence a monoecious, terminal raceme, ca 4.0 cm long, 

flaccid, usually pendent, all parts glabrous, often with a bloom; bracteoles foliaceous, 

1.0-2.0 cm long, ca 1.25 cm wide, purplish tinged, margin undulate, rarely serrate; 

bracteoles foliaceous, ca 1.0 cm long, ca 0.75 cm wide. Pistillate flowers restricted to the 

base of the upper half of the inflorescence, tepal yellowish green, 1.0 cm long, cleft to 

base into 5 lobes, ovary subglobose. Staminate buds ovoid-ellipsoid; flowers medium 

sized; tepal 1.0 cm long, yellowish green, cleft 1/3 way down into 5 lobes; stamens 10, in 

2 whorls of 5 each. Capsules small, 1.0 cm long, subglobose, surface smooth, apex 

rounded, dehiscence septicidal. Seeds small, 0.8 cm long, oblong, caruncle prominent. 

DISTRIBUTION. (Fig 57B). Brasil, states of Mato Grosso, Goias, Minas Gerais and 

Distrito Federal. Alt ca 1000 m, common in burned-over cerrado. 

Key to the Subspecies of Manihot violacea 

1. Leaf lobes obovate, apex obtuse; basal lobes straight. 

46a. M. violacea subsp violacea. 

1. Leaf lobes elliptic, apex acute; basal lobes distinctly recurved. 46b. M. violacea subsp recurvata.


-' 

to 

? .~~~~~~~~~~~~~~~~~\ 

~~~~~~~~~~~~~~~~~~~~~~~~c~ 

FIG 57. Manbot alutace. A, infioescence (Haley, Barros et al. 1104, K). M. iolacea. B 

distributin, dots reresent subp violaceaand the leter O reprsents subs recumata.M. violace 

subp volcea C lef Pob 189,G) D,iniorscnce(Iwin Suza& os anos 101, ?) 

FIG 57. Manibot alutacea. A, inflorescence (Harley, Barroso et al. 11404, K). M. violacea. B, 

distribution, dots represent subsp violacea and the letter 0 represents subsp recurvata. M. violacea 

subsp violacea. C, leaf (Pohl 1189, G); D, inflorescence (Irwin, Souza & dos Santos 11051, NY).

134 

46a. Manihot violacea Pohl subsp violacea 


Manihot violacea Pohl, PI. Bras. Ic. et Descr. 1: 43. t. 35. 1827; Jatropha violacea Steudel, 

Nomencl. ed. 2. 1: 800. 1840; Manihot violacea Pohl emend. Muell. Arg. vargenuina Muell. 

Arg. in DC., Prodr. 15(2): 1070. 1866; in Martius, Fl. Bras. 11(2): 468. 1874; Pax in Engler, 


Plant parts usually with a thick bluish white bloom. Median lobes obovate, apex 

obtuse, midribs usually stout, lamina at sinus prominently overlapping; basal lobes more 

or less straight, not prominently recurved. Fig 57 C, D. 

TYPE. Pohl 1189: Brasil, Goias: circa Corumba, Corgo do Jaragua et Ourofino 

(syntypes, F, G, W-2). 

DISTRIBUTION. (Fig 57B). Brasil in states Mato Grosso, Goias, Minas Gerais, and 

Distrito Federal. BRASIL. Mato Grosso: Irwin & Soderstrom 6778 (NY, UB) ca 90 km N of 

Xavantina, 12 Oct 1964. Goias: Glaziou 22132 (P) Serra Dos Pyreneos, 18 Feb 1894;Heringer 7752 

(NY), 15 Oct 1960; Irwin, Souza & dos Santos 11051 (F, K, NY, SP, UB) Campo and Cerrado near 

Descoberto, 4 Dec 1965; Pohl Icon Tab (W). Distrito Federal: Irwin, Souza & dos Santos 7977 (F, 

NY, UB, US) Chapada da Contagem, 4 Sep 1965. Minas Gerais: Barreto 2676 (F) Serra Do Taquaril, 

Bello Horizonte, 7 Feb 1933; Glaziou 19856 (K, P) Entre Barreiros et Melancias, 19 Mar 1892. 

Province Unknown/Uncertain: Pohl s n (M). 

While this subspecies is quite variable in morphology, its distinctive bloom readily 

distinguishes it from subsp recurvata. 

46b. Manihot violacea Pohl subsp recurvata Rogers & Appan, subsp nov 

Manihot pseudopruinosa Pax in Engler, Pflanzenreich IV. 147(Heft 44): 61. 1910 Pro parte (See 

also 43). Type. Glaziou 22131 pro parte (syntypes A, P, S). 

Subfrutices, ad 1.0 m altus. Folia trilobus, lobi elliptici, apex acutus, lobi basali 

manifeste curvus sursum. Inflorescentia racemus, ca 4.0 cm longus, bracteolae et ramuli 

foliaceus, purpureus suffusus. Fructi parvi, 1.0 cm longi, subglobosi. Seminae 0.8 cm 

longae, oblongus. 

Plant parts with a light bloom or without any bloom. Median lobes elliptic, apex 

acuminate, midribs slender; lamina at sinus slightly overlapping, occasionally not 

overlapping; basal lobes prominently recurved. Fig 58A. 

TYPE. Irwin, Souza & dos Santos 11202: Brasil, Distrito Federal: Campo slope 

immediately east of Lagoa Paranoa, D. F. Elevation 975 m, 11 December, 1965. 

(holotype, NY, isotypes, F, G-2, H, MO, S, SP, UB, UC, US). 

DISTRIBUTION. (Fig 57B). Brasil, state of Goias and Distrito Federal. BRASIL. Goias: 

Glaziou s n (P) Rio do Gama, 4 Nov 1894; Glaziou 22131 (A, P, S) Vao do Parana, a Itiquira, 2 Feb 

1895; Irwin, Grear, Souza & dos Santos 14222 (F, GH, NY, UB) Cerrado Rio Parana circa 35 km N of 

Formosa on Road to Sao Gabriel, 29 Mar 1966. Distrito Federal: Irwin, Grear, Souza & dos Santos 

15392 (F, MO, NY, RB, UB, US) Cerrado near Corrego Taquari E of Lagoa Paranoa, 28 Apr 1966; 

Irwin & Soderstrom 5398 (NY, UB) circa 35 km E of Brasilia, 21 Aug 1964; Irwin, Souza & dos 

Santos 10092 (UB) 1 km N of Sobradinho, 8 Nov 1965; Irwin, Souza & dos Santos 10168 (F, G, GH, 

K, LE, MO, NY, RB, S, SP, UB, US) Cerrado adjacent to Parque, Municipality do Gama, 10 Nov 1965; 

Irwin, Souza & dos Santos 11243 (F, GH, IAN, LIL, MO, NY, SP, UB, US, W) Dam at Lagoa Paranoa, 

12 Dec 1965; Irwin, Souza & dos Santos 8257 (UB) Summit of Chapada da Contagem, 10 km E of 

Brasilia, 13 Sep 1965; Irwin, Souza & dos Santos 8297 (F, NY, UB) Corrego Urubu, slopes of Chapada 

da Contagem, 14 Sep 1965; Irwin, Souza & dos Santos 8563 (F, NY, UB) circa 15 km E of Brasilia on 

Road to Paranoa, 22 Sep 1965; Irwin, Souza & dos Santos 8627 (NY) 24 Sep 1965; Irwin, Souza & 

dos Santos 9113 (F, MO, NY, UB) 10 km E of Planaltina, 10 Oct 1965; Irwin, Souza & dos Santos 

9167 (F, GH, NY, SP, UB, US) burned over Cerrado between Brasilia and Sobradinho, 13 Oct 1965; 

Pereira 4765 (RB) Campo Cerrado, 16 Nov 1958. 

The distinctive quality of this subspecies is its recurved lateral lobes on the mature 

vegetative leaves.


~~~~~~~~~~~~~~~. 

. . ......| 

Es ,.~~~~~~~~ ,., 

r 

" ' 

; -+~~~~~~~ 

X 

b x o~~~~~~~~ 

,0-1 

=~~~~~~ N t 

FIG 58. Manibot violacea subsp recurvata. A, leaf (Irwin, Souza & dos Santos 11243, NY). M. 

jacobinensis. B, distribution; C, leaf and inflorescence (Blancbet 2553, G). M. divergens. D, distribu- 

tion. 

,


47. Manihot jacobinensis Mueller von Argau, Linnaea 34: 205. 1865; in DC., Prodr. 

15(2): 1070. 1866; in Martius, Fl. Bras. 11(2) 471. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 31. 1910. 

Manihot occidentalis Muell.-Arg. in Martius, Fl. Bras. 11(2): 468. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 31. 1910. Type. Tamberlik s n (syntypes, F-2, G, W). 

Manihot rigidifolia Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 194. 1924. 

Type. Luetzelburg 484a (syntypes F, M); Luetzelburg 484b (syntype, M). 

Shrubs. Stems glabrous. Leaves alternate; stipules caducous; petioles ca 6.0 cm long, 

terete, glabrous, petiole attachment to lamina basal, nonpeltate; lamina coriaceous, 

adaxial surface dark green, abaxial surface with a light bloom, abaxial surface wax pattern 

spongiose; venation camptodromous, veins glabrous; lamina 3 lobed, lobes prominently 

overlapping at sinus; median lobes ovate to almost rotundate, ca 5.0 cm long, ca 4.0 cm 

wide, apex broadly acute, base of lobes constricted ca 0.5 cm wide, lowest lobes more or 

less similar to median lobes in outline and size. Inflorescence a monoecious, terminal 

raceme, ca 12.0 cm long, held erect, all parts glabrous, with light bloom on peduncles, 

pedicels, tepals, etc; bracteoles foliaceous, 1.5 cm long, 0.8 cm wide; bractlets foliaceous, 

1.0 cm long, 0.4 cm wide. Pistillate flowers restricted to the base of the upper 2/3 of the 

inflorescence, tepal 1.0 cm long, cleft to base into 5 lobes, ovary subglobose, glabrous. 

Staminate buds slightly bifusiform; flowers medium sized, tepal 1.0 cm long, cleft 1/3 

way down into 5 lobes, stamens 10, in 2 whorls of 5 each. Fruits and seeds non vidi. 

Fig 58C. 

TYPE. Blanchet 2553: Brasil, Bahia: Serra da Jacobina, 1836 (syntypes, BM, F-3, 

G-3, K, NY, P-2, W-2). 

DISTRIBUTION. (Fig 58B). Brasil, states of Mato Grosso and Bahia. Alt ca 1000 m. 

BRASIL. Mato Grosso: Maguire, Pires, Maguire & Silva 56464 (NY) Gallery forest, Rio Juruena, on 

Brasilia-Acre Highway, 1 Sep 1963. Bahia: Blanchet 3553 (G) Province Jacobina, 1843; Froes 12630 

(NY) Central Portion of State, Oct 1942; Luetzelburg 484a (F, M) Lencoes, Pedra de Chapeo, 1914; 

Luetzelburg 484b (M) Lencoes, Pedra de Chapeo, 1914; Pereira 2023 (PUL-2, RB) Morro Do Chapeu, 

11 Sep 1956; Pereira 2023A (RB) Morro Do Chapeu, 11 Sep 1956. Province Unknown/Uncertain: 

Riedel s n (P). Province Unknown/Uncertain: Tamberlik s n (F-2, G, W) Brazilia Occid. 

The specimen collected by Maguire et al (56464, NY) fits the descriptions and type 

of Manihot jacobinensis in its major features, but differs in that the bracts are setaceous, 

while those of jacobinensis are foliaceous. However, species of section Quinquelobae to 

which jacobinensis clearly belongs are quite variable in this character, and the possibility 

of genetic recombination must not be ruled out. Quite possibly, the Maguire specimen 

represents a distinct taxon, but we feel that this single collection does not give sufficient 

basis for such designation. Therefore, we maintain Maguire et al 56464 as a variation 

within the M. jacobinensis complex. 

48. Manihot divergens Pohl, P1. Bras. Ic. et Descr. 1: 41. t. 33. 1827. 

Manihot arcuata Pohl, PI. Bras. Ic. et Descr. 1: 42. 1827; Manihot violacea Pohl emend. Muell. 

Arg. var arcuata (Pohl) Muell. Arg. in DC., Prodr. 15(2): 1069. 1866; Jatropha arcuata 

Steudel, Nomencl. ed. 2. 1: 799. 1840. Type. Pobl 1183 (syntypes, (F-2, G, W-2). 

Jatropha divergens Steudel, Nomencl. ed. 2. 1: 799. 1840; Manihot violacea Pohl emend. 

Muell.-Arg. var divergens (Pohl) Muell. Arg. in DC., Prodr. 15(2): 1069. 1866. 

Tall shrubs, to 2.5 m tall. Young stems, glabrous, reddish brown; mature stems 

glabrous. Leaves alternate; stipules caducous, glabrous, ca 1.0 cm long; petioles ca 10.0 

cm long, obtuse-angled, glabrous, petiole attachment to lamina basal, nonpeltate; lamina 

coriaceous, adaxial surface dark green, glossy, abaxial surface glaucous, abaxial surface 

wax pattern smooth; venation camptodromous, veins glabrous, midribs stout, 0.15 cm in 

diam near the petiole-lamina junction; lamina 3 lobed, leaves associated with the 

inflorescence frequently nonlobed; median lobes oblong-obovate, 10.0-15.0 cm long,


4.0-7.0 cm wide, lamina edges purplish, apex obtuse to broadly acute, base of lobes ca 

1.0 cm wide; lowest lobes similar to median lobes in size and outline. Inflorescence 

monoecious, terminal, a cluster of short racemes arising from one basal point ca 4.0 cm 

long, occasionally as long as 8.0 cm; all parts glabrous, sometimes with a bluish bloom; 

bracteoles foliaceous, 1.25 cm long, 0.8 cm wide; bractlets foliaceous, 0.75 cm long, 0.4 

cm wide. Pistillate flowers restricted to the base of the upper 1/2 of the inflorescence, 

pedicels 1.0 cm long, tepal 1.0 cm long, cleft to base into 5 lobes, ovary subglobose. 

Staminate buds ovoid-ellipsoid; tepal 1.0 cm long, cleft 1/3 way down into 5 lobes; 

stamens 10, in 2 whorls of 5 each. Capsules 1.25 cm long; surface smooth; apex rounded, 

dehiscence septicidal. Seeds 0.8 cm long, oblong, with a well-developed caruncle. 

Fig 59A, B. 

TYPE. Pohl 6040 (and on some specimens also 1657). Brasil, Goias: ad Rio Macaco 

et Areas (syntypes, F, G, K, W-2). 

DISTRIBUTION. (Fig 58D). Brasil, states of Goias, Minas Gerais and Distrito Federal. 

Alt ca 1000 m. BRASIL. Goias: Dawson 14149 (F) region of Chapada dos Veadeiros, 13 Apr 1956; 

Irwin, Grear, Souza & dos Santos 12443 (F, MICH, NY, SP, UB, US) Chapada dos Veadeiros, 9 Feb 

1966; Irwin, Grear, Souza & dos Santos 13694 (F, G, NY, RB, UB, UC) Cerrado, circa 20 km N df 

Cristalina, 7 Mar 1966; Irwin, Grear, Souza & dos Santos 13696 (F, GH, K, MO, NY, SP, UB, US) 

Cerrado, circa 20 km N of Cristalina, 7 Mar 1966; Irwin, Grear, Souza & dos Santos 15565 (F, NY, 

UB) Corrego Itaquira, circa 30 km N of Formosa, 2 May 1966; Irwin, Souza & dos Santos 10014 (F, 

MO, NY, RB, S, SP, UB) 75 km N of Cristalina on road to Brasilia, 6 Nov 1965; Irwin, Souza & dos 

Santos 9502 (F, GH, MICH, MO, NY, RB, SP, UB, US, W) near waterfall, 24 km NW of Veadeiros, 

road to Cavalcante, 22 Oct 1965;Macedo s n (PUL) Jacuba, Niquelandia, 24 Feb 1956;Pohl Icon Tab. 

33 (W); Pobl Icon Tab. 34 (W); Pohl 1183 (F-2, G, W-2) ad Rio Macacco et Areas; Prance & Silva 

58227 (NY-2) 5-10 km N of Veadeiros, Valley of Rio Parana, Cerrado, 19 Jul 1964. Distrito Federal: 

Irwin, Souza & dos Santos 10723 (F, NY, UB) Campo near Aparecido, circa 50 km W of Brasilia, 29 

Nov 1965. Minas Gerais: Macedo 4264 (PUL, RB) Araxa, 5 Feb 1956; Saint-Hilaire 488 (F, P). 

49. Manihot irwinii Rogers & Appan, sp nov n 

Frutices, ca 1.5 m altus, aliquot ascendens caules ab ligneum basim. Folia coriaceus, 

glaucissima in abaxialis pagina; venae rubelli. Inflorescentia aliquot racemi exoriens uno 

basalis punctum, bracteolae et ramuli semifoliaceus, ca 1.0 cm longi, ca 0.3 cm lati, 

purpurascens; tepalum purpurascens. Capsulae 1.6 cm longae. Semina 1.0 cm longae, 

oblongae. 

Shrubs, ca 1.5 m tall, with several ascending stems from woody base. Young stems 

obtuse-angled in cross section, glabrous, glaucous, purplish tinged; mature stems glabrous. 

Leaves alternate; stipules caducous, 0.5 cm long, 0.1 cm wide, glaucous; petioles ca 9.0 

cm long, terete, glabrous, reddish, petiole attachment to lamina basal, nonpeltate; lamina 

coriaceous, adaxial surface dark green, abaxial surface with a thick bloom, abaxial surface 

wax pattern smooth; venation camptodromous, veins glabrous, reddish, lamina 3 lobed, 

leaves associated with the inflorescence frequently nonlobed; median lobes ca 6.0 cm 

long, ca 1.0 cm wide, elliptic, apex acute, obtuse or caudate, base of lobes ca 1.0 cm 

wide, lamina prominently overlapping at sinus; lowest lobes similar in size to median 

lobes, nonsymmetric. Inflorencence monoecious, terminal, a cluster of several racemes 

arising from one basal point, ca 7.0 cm long, all parts except interior surface of tepals 

glabrous, with a thick bloom; bracteoles semifoliaceous, 1.0 cm long, 0.3 cm wide, 

purplish tinged, glabrous, margin entire; bracteoles semifoliaceous, 0.7 cm long, 0.2 cm 

wide, purplish tinged, margin entire. Pistillate flowers restricted to the base of the upper 

half of the inflorescence; pedicels 1.25 cm long; tepal 0.9 cm long, cleft to base into 5 

strap-shaped lobes, interior surface of lobes pubescent, yellowish green with considerable 

amount of purlish pigmentation; ovary subglobose, slightly elongated, the trifid stigma 

moderately lobed and lobulate. Staminate flowers medium sized, tepal 0.9 cm long, 

' This species is named in honor of Dr. Howard S. Irwin of The New York Botanical Garden. Dr. 

Irwin's studies of the flora of the central Brazilian plateau have contributed greatly to our knowledge 

and understanding of this little known region.

138 

I: 

= 


F~IG iegn.A, 

aio 9 

);B,ifoecne(wn,Sua&dsats 

lef(oh, 

10723~~~~~~~~~~~~~~~~~~~~~~,N?..irni.C, ta.13 ars ef(aly ) 

dsrbto;D 

FIG 59. Manihot divergens. A, leaf (Pohl 6040, W); B, inflorescence (Irwin, Souza & dos Santos 

10723, NY). M. irwinii. C, distribution; D, leaf (Harley, Barroso et al. 11325, K). 

...


yellowish green with considerable amount of purplish pigmentation, cleft 1/3 way down 

into 5 lobes; disc 10 lobed; stamens 10, in 2 whorls of 5 each, the superior whorl 0.7 cm 

long, the inferior whorl 0.5 cm long. Capsules 1.6 cm long, subglobose, slightly elongated, 

surface smooth, without prominent wings, apex rounded, dehiscence septicidal. Seeds 1.0 

cm long, oblong, with a prominent caruncle. Fig59D; 60A. 

TYPE. Irwin, Souza & dos Santos 10898: Brasil, Goias: Serra dos Pirineus (16? S, 

49? W), ca 11 km N of Corumba de Goias, Cerrado, rocky slopes, alt 1100 m, 2 Dec 1965 

(holotype, NY; isotypes, F, K, UB). 

DISTRIBUTION. (Fig 59C). Brasil, state of Goias. Alt ca 1000 m. On rocky slopes and 

sandstone summits. BRASIL. Goias: Harley, Barrosa et al 11325 (K) Serra Dourada, near Goias, 18 

Dec 1968; Irwin, Grear, Souza & dos Santos 12959 (B, F, G, GH, IAN, MO, NY, SP, UB, US, W) circa 

30 km NW of Veadeiros, 16 Feb 1966; Irwin, Maxwell & Wasshausen 18562 (COL, F, GH, MICH, MO, 

NY, RB, SP, UB, US) 25 km N of Corumba de Goias on road to Niquelandia, 13 Jan 1968; Irwin, 

Maxwell & Wassbausen 19359 (F, MG, MO, NY, S, SP, UB, UC, US, W) circa 20 km NW of Corumba 

de Goias, near road to Niquelandia 28 Jan 1968; Irwin, Souza & dos Santos 11752 (F, G, GH, K, MO, 

NY, RB, SP, UB, US) campo and cerrado sand stone summit of Serra Dourada, 18 Jan 1966;Macedo 

3476 (NY, S, US) Serra Dourada, 13 Dec 1951;Macedo 4347 (K, US) Corumba de Goias, 17 Feb 

1956. 

Manihot irwinii makes a simultaneous connection to M. cecropiaefolia and M. 

violacea subsp violacea at a similarity value of .86 (Fig. 28). This level of similarity 

indicates that this is quite a distinct species. Its distinctive features are the semifoliaceous 

bracteoles and the purple tepal lobes. 

50. Manihot cecropiaefolia Pohl, PI. Bras. Ic. et Descr. 1: 49. t. 42. 1827. 

Jatropha cecropiaefolia Steudel, Nomencl. ed. 2. 1: 799, 1840; Manihot violacea (Pohl) emend. 

Muell.-Arg. var cecropiaefolia (Pohl) Muell.-Arg. in DC., Prodr. 15(2): 1069. 1866; in 

Martius, Fl. Bras. 11(2): 467. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 30. 

1910. 

Shrubs, ca 1.5 m tall, with several divergent branches. Stems glabrous. Leaves 

alternate; stipules caducous; petioles ca 15.0 cm long, terete, glabrous, petiole attachment 

to lamina basal, nonpeltate; lamina coriaceous, adaxial surface dark green, abaxial surface 

with a bloom, abaxial lamina surface wax pattern pusticulate; venation camptodromous, 

veins glabrous, midribs stout, 0.1 cm in diam near petiole-midrib junction; lamina 

palmately 5 lobed, rarely 3; median lobes ovate-elliptic, 10.0-15.0 cm long, 5.0-8.0 cm 

wide, apex obtuse, base of lobes ca 2.0 cm wide, lamina at lobe sinus prominently 

overlapping; lowest lobes same as median lobes in length, prominently nonsymmetric and 

curved up. Inflorescence monoecious, terminal, a cluster of several racemes arising from 

one basal point, usually with several divergent principal peduncles, often pendent, ca 10.0 

cm long, occasionally as long as 20.0 cm, with a light bloom, all parts except interior 

surface of tepals glabrous; bracteoles foliaceous, ca 2.0 cm long, ca 1.0 cm wide, 

occasionally as long as 3.0 cm, and 1.5 cm wide, with a light purplish tinge, margin entire; 

bracteoles foliaceous, 1.0 cm long, 0.6 cm wide. Pistillate flowers restricted to the base of 

the upper half of the inflorescence; pedicels 1.0 cm long; tepal 1.0 cm long, cleft to base 

into 5 lobes; ovary subglobose. Staminate buds ovoid-ellipsoid; flowers medium sized; 

tepal 1.0 cm long, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. 

Capsules 1.25 cm long, subglobose, slightly elongated, surface smooth, apex rounded, ribs 

pinkish, dehiscence septicidal. Seeds 0.9 cm long, oblong, caruncle prominent. 

Fig 60C, and D. 

TYPE. Pohl 3919 (and on some specimens also 1649). Brasil, Goias: ad Villa Boa, 

Corgo do Jaragua (syntypes F, G, K, W-3). 

DISTRIBUTION. (Fig 60B). Brasil, state of Goias, and Distrito Federal. Alt ca 1000 

m. In gallery forest and in adjacent cerrado. BRASIL. Goias: Dawson 15083 (F) 18 km E of 

Formosa, Serra Dourado region, 23 May 1956; Glaziou 22134 (K, P, S) Entre Taboquinha et Lages, 20


'-. 

'd.4n 

!.4 ~~~~ ~ ~S' 

-1 4 

IIIIRA I,F (,)OA. MWlAL, 

FIG 60. Manibot irwinii. A, inflorescence (Macedo 3476, NY). M. cecropiaefolia. B, distribu- 

tion; C, leaf (Pohl 3919, W); D, inflorescence (Pohl 3919, W).


Feb 1895; Glaziou 22135 (F-3, G, K, P) Formosa, 4 Feb 1895; Irwin, Grear, Souza & dos 

Santosl 7841 (F, NY, UB) circa 50 km S of Caiaponia, road to Jatai, 27 Jun 1966; Irwin, Maxwell & 

Wasshausen 19083 (F, NY, UB) 60 km N of Corumba de Goias on road to Niquelandia, 23 Jan 1968; 

Pohl Icon Tab. 42 (W). Distrito Federal: Irwin, Grear, Souza & dos Santos 13019 (F, G, GH, K, MICH, 

MO, NY, RB, SP, UB, US) near waterfall, circa 25 km SW of Brasilia, 19 Feb 1966. Province 

Unknown/Uncertain: Weddell 2909 (P) Entre Goyas et Cujaba. 

Doubtfully assigned to this taxon: BRASIL. Goias: Irwin & Soderstrom 7364 (NY, UB) ca 50 km 

S. of Caiaponia, on road to Jatai, 25 Oct 1964. 

The profusely branched inflorescence is a distinctive character of both Manihot 

cecropiaefolia and M. irwinii. The 5 lobed leaf and yellowish green tepal lobes distinguish 

this species from M. irwinii. 

51. Manihot mossamedensis Taubert, Bot. Jahrb. Syst. 21: 442. 1896; Pax in Engler, 


Shrub, to 3.0 m tall, growing from a woody base. Young stems pubescent; mature 

stems glabrous. Leaves alternate; stipules caducous; petioles ca 20.0 cm long, obtuseangled, 

pubescent, petiole attachment to lamina basal, nonpeltate; lamina coriaceous, 

adaxial surface dark green, glossy, abaxial surface glaucous, abaxial surface wax pattern 

smooth; venation camptodromous, veins pubescent, midribs stout, 0.1 cm in diam near 

the petiole-midrib junction; lamina palmately 5-7 lobed, rarely 3, lamina at sinus 

overlapping; median lobes oblong, ca 15.0 cm long, ca 5 cm wide, occasionally as long as 

20.0 cm, 8.0 cm wide, apex acute to obtuse-acuminate, base of lobes ca 2.0 cm wide; 

lowest lobes similar in outline to median lobes but ca 1/2 as long. Inflorescence 

monoecious, terminal, a cluster of several racemes arising from one basal point, ca 8.0 cm 

long, pendent, bracts, bracteoles and flowers forming a dense cluster at the apical region 

of peduncles, peduncles and pedicels pubescent; bracteoles foliaceous, 2.5 cm long, 1.25 

cm wide, pubescent, margins entire; bractlets foliaceous, 1.5 cm long, 0.7 cm wide, 

pubescent. Pistillate flowers restricted to the base of the upper half of the inflorescence; 

pedicels 0.75 cm long; tepal 1.25 cm long, cleft to base into 5 lobes, pubescent; ovary 

nonglobose, pubescent. Staminate buds ovoid-ellipsoid; flowers medium sized, tepal 1.1 

cm long, cleft 1/3 way down into 5 lobes, pubescent; stamens 10, in 2 whorls of 5 each. 

Capsules nonglobose, 1.5 cm long, surface smooth, apex rounded, ribs purplish, 

dehiscence septicidal. Seeds 1.0 cm long, oblong, caruncle moderately prominent. 

Fig 61B, C. 

TYPE. Ule 3081: Brasil, Goias: bei Mossamedes (syntypes photo F, photo NY). 

DISTRIBUTION. (Fig 61A). Brasil, state of Goias. Alt ca 800 m, on disturbed forest 

slopes. BRASIL. Goias: Irwin, Grear, Souza & dos Santos 14989 (F, G, GH, K, MO, NY, RB, SP, UB, 

US) Corrego Estrema, 35 km NW of Formosa, 18 Apr 1966; Irwin, Souza & dos Santos 11850 (F, G, 

GH, MO, NY, RB, S, SP, UB, US) near Summit of Serra Dourada, circa 20 km SE of Goias Velho, 20 

Jan 1966; Irwin, Souza, Grear & dos Santos 15250 (F, NY, UB) Corrego Estrema, circa 35 km NE of 

Formosa, 22 Apr 1966; Ule 470 (P-2). 

The densely pubescent inflorescence is a characteristic feature of this taxon. 

8. Manihot sect Graciles Rogers & Appan, sect nov 

Manihot sect Grandibracteatae Pax subsect Rigidulae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 35. 1910 pro parte. 

Manihot sect Grandibracteatae Pax subsect Angustifoliae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 40. 1910 pro parte. 

Manihot sect Sinuatae Pax subsect Laciniosae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 45. 

1910 pro parte. 

Manihot sect Parvibracteatae Pax subsect Humiles Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

58. 1910 pro parte.


FIG . M a n i o' m n , d, le_ in So".,,, ,, 

,o , an,o 

' 1 i 

- - .........i 

14989, NY); C, inflorescence (Irwin, Souza & dos Santos 118 50, NY).* 

_,. 

/W~~~~~~~~~~~C 

_~~~----'-'I~ 

FIG 61. Manibot mossamedensis. A, distribution; B, leaf (Irwuin, Great, Souza & dos Santos 

14989, NY); C, infiorescence (Irwin, Souza & dos Santos 1 1850, NY).


Manihot sect Parvibracteatae Pax subsect Stenophyllae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 72. 1910. 

Manihot sect Parvibracteatae Pax subsect Graciles Pax in Engler, Pflanzenreich IV. 147(Heft 44): 


Plantae caulinae, lignosae, graciliae infernae ad mediae frutices. Folia petiolatae, 

penitae lobatae. Inflorescentia monoecia panicula aut racema. 

Plants caulescent; slender, low to medium, straggling shrubs; leaves widely spaced on 

stem; petiolate, petiole attachment basal; lamina usually membranaceous, deeply lobed, 

lobes linear or narrowly lanceolate. Inflorescence monoecious, paniculate or racemose; 

bracts and bracteoles setaceus to foliaceous, margins entire to laciniate. 

TYPE. Manihot gracilis Pohl emend Rogers & Appan. 


Key to the Species of Sect Graciles 

1. Inflorescence an erect subspicate raceme. 

2. Bracts and bracteoles foliaceous, more than 0.5 cm wide; leaf lobes, petioles, bracts, 

bracteoles, peduncles, tepals, etc. pubescent; plants more than 1.0 m tall. 52. M. flemingiana. 

2. Bracts and bracteoles setaceous, less than 0.25 cm wide; all parts glabrous; plants less than 

1.0 m tall. 

3. Leaves 5 lobed; venation frequently craspedodromous, sometimes camptodromous; 

lamina between base of median lobe sinus and petiole midrib junction less than 2.5 cm 

wide; inflorescence a cluster of racemes arising from a common base. 53. M. hunzikeriana. 

3. Leaves 3 lobed; venation camptodromous, never craspedodromous; lamina between 

base of median lobe sinus and petiole-midrib junction more than 2.5 cm; inflorescence 

a simple raceme. 54. M. hassleriana. 

1. Inflorescence a lax raceme (not subspicate), or a panicle. 

4. Inflorescence a panicle, bracts and bracteoles setaceous, less than 0.25 cm wide. 

5. Inflorescence usually less than 12.0 cm long, flowers clustered at terminal region of 

peduncles; leaf lobes held horizontally; plants more than 0.5 m tall. 55. M. triphylla. 

5. Inflorescence usually more than 12.0 cm long; flowers evenly positioned in inflorescence; 

leaf lobes drooping; plants usually less than 0.5 m tall. 56. M. fruticulosa. 

4. Inflorescence a lax raceme (not subspicate); bracts and bracteoles semifoliaceous or 

foliaceous, more than 0.25 cm wide. 

6. Leaf lobes linear, or narrowly lanceolate, less than 1.0 cm wide. 

7. Leaves 5 lobed. 

8. All parts glabrous; leaf lobes usually more than 6.0 cm long. 57. M. pentaphylla. 

8. Petioles, midribs, young stems, peduncles, pedicels, bracts, bracteoles, etc. 

pubescent; leaf lobes usually less than 6.0 cm long. 

58. M. tenella. 

7. Leaves 3 lobed. 59. M. gracilis. 

6. Leaf lobes lanceolate, usually 1.0-3.0 cm wide. 

9. Leaves 5 lobed; venation camptodromous, never craspedodromous; 

bracts and 

bracteoles foliaceous, more than 0.5 cm wide; inflorescence more than 3.0 cm long. 

60. M. paviaefolia. 

9. Leaves 3 lobed; venation frequently craspedodromous, sometimes camptodromous; 

bracts and bracteoles semifoliaceous, less than 0.25 cm wide; inflorescence usually 

less than 3.0 cm long. 

61. M maguireiana. 

52. Manihot flemingiana Rogers & Appan, sp nov 12 

Frutices effusi, ad 2.0 m altus. Juvenis caules parce pubescens. Folia 5 lobatus, 3 

major et 2 parvior; mediani lobi lineari, ca 10.0 cm longi, ca 0.5 cm lati, tenens ad dexter 

anguli 2 exterior lobi, inferimus lobi attenuatus, ca 2.0 cm longus, venae hirsutae. 

Inflorescentia brevis subspicatorum racemi, ca 4.0 cm longi, bracteolae et ramuli 

12 This species is named in honor of Henry Stanton Fleming, our valued colleague who, in 

addition to his great knowledge of the natural history of South America and the West Indies, is one of 

the greatest students of taximetrics.

o I 'L 1 ` \ : / 

FIG 62. Manibot sect. Graciles. Geographical Range. (The distribution of M. maquireiana 

Venezuela, was omitted from this sectional map). 

_..


foliaceus, hirsutae; pistillatus flores portati in longi pedicelli basi inflorescentiarum. 

Staminatae gemmae parvae, globosi, pubescens filamenta. Capsulae 1.1 cm longae, 

oblongae, sine marmoreae. 

Straggling shrubs, to 2.0 m tall. Young stems sparsely pubescent, grayish brown; 

mature stems glabrous, grayish brown. Leaves alternate; stipules 0.5 cm long, filiform, 

hirsute; petioles short, usually less than 4.0 cm long, terete, pubescent, petiole 

attachment to lamina basal, nonpeltate; lamina membranaceous, abaxial surface wax 

pattern reticulate; venation camptodromous, rarely craspedodromous, veins hirsute; 

lamina 5 lobed, 3 major lobes and 2 basal lobules; median lobes linear, ca 10.0 cm long, 

ca 0.5 cm wide, held almost perpendicular to the 2 outer lobes; apex acuminate, base of 

lobes 0.5 cm wide, width between base of sinus and petiole-midrib junction 0.4 cm; 

lowest lobes attenuate, ca 2.0 cm long, held parallel to the petiole. Inflorescence a 

monoecious, terminal, subspicate raceme, ca 4.0 cm long; peduncles and pedicels hirsute; 

bracteoles foliaceous, 0.75 cm long, 0.6 cm wide, hirsute, yellowish green with purplish 

tinge, margins entire; bractlets foliaceous 0.5 cm long, 0.3 cm wide, hirsute, margins 

entire. Pistillate flowers restricted to the base of the inflorescence, borne on ca 2.5 cm 

long pedicels; pedicels hirsute; tepal 0.8 cm long, cleft to base into 5 lobes, yellowish 

green with purplish pigmenation, hirsute; disc prominent; ovary subglobose, tomentose; 

the trifid stigma moderately lobed and lobulate. Staminate buds globular; flowers small, 

tepal 0.7 cm long, yellowish green with purplish pigmentation, hirsute, cleft 1/3 way 

down into 5 lobes, lobe apex obtuse; disc 10 lobed; stamens 10, in 2 whorls of 5 each, the 

superior whorl 0.6 cm long, the inferior whorl 0.5 cm long, filaments pubescent, anthers 

bright yellow. Capsules small, 1.1 cm long from base to apex, subglobose, slightly 

elongated, surface without prominent wings, hirsute, apex rounded, dehiscence septicidal. 

Seeds small, 0.8 cm long, oblong, grayish black without mottlings, caruncle trapeziform, 

not prominent. Fig 63B, C. 

TYPE. Philcox & Fereira 3868: Brasil, Mato Grosso: Cerrado E of km 242, 

Xavantina-Cachimbo road, 4 Jan 1968 (holotype, K; isotype NY). 

DISTRIBUTION. (Fig 63A). Brasil, state of Mato Grosso. BRASIL. Mato Grosso: Philcox 

& Fereira 4542 (K, NY) 1 km E of km 244, Xavantina-Cachimbo Road, 15 Mar 1968. 

Manihot flemingiana, which is well isolated (see Fig. 28, objects 64 and 65) from its 

nearest relative (M. hunzikeriana) is distinguished by its foliaceous bracteoles. Manihot 

hunzikeriana has setaceous bracteoles. The subspicate raceme of M. flemingiana is much 

more compact (ca 3-4 cm long) than is the inflorescence of M. hunzikeriana. 

53. Manihot hunzikeriana Martinez-Crovetto, Bonplandia 1(4): 273-277. 1964. 

Subshrubs ca 30 cm tall. Stems glabrous. Leaves alternate; stipules persistent, ca 0.5 

cm long, setaceous, glabrous; petioles short, usually less than 4.0 cm long, terete, 

glabrous, petiole attachment to lamina basal, nonpeltate; lamina membranaceous, abaxial 

surface wax pattern smooth; venation frequently craspedodromous, sometimes campto- 

dromous, veins glabrous; lamina palmately 5 lobed, rarely with 2 more short lobules; 

median lobes linear, frequently with falcate lobules near the base, ca 20.0 cm long, 0.6 

cm wide, apex acuminate, base of lobes 0.6 cm wide, width between base of sinus and 

petiole-lamina junction ca 2.0 cm; lowest lobes (in case of 5 lobed leaves) 1/2 as long as 

median lobes, in case of 7 lobed leaves attenuate and ca 1.0 cm long. Inflorescence 

monoecious, terminal, a cluster of long subspicate racemes arising from one basal point, 

ca 15.0 cm long, all parts glabrous; bracteoles and bractlets setaceous, less than 1.0 cm 

long, less than 0.2 cm wide. Pistillate flowers restricted to the base of the inflorescence, 

tepal 0.9 cm long, cleft to base into 5 lobes. Staminate buds ovoid-ellipsoid; flowers 

small; tepal 0.8 cm long, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5


9 L 

1 1.?1 !1 ..................... , 

FIG 63. Manihot flemingiana. A, distribution; B, leaf (Philcox & Fereira 3868, K); C, inflores- 

cence (Philcox & Fereira 3868, K). M. hunzikeriana. D, distribution.


each. Capsules 1.5 cm long, subglobose, slightly elongated, surface without prominent 

wings, apex rounded, dehiscence septicidal. Seeds 1.0 cm long, oblong, caruncle 

moderately prominent. Fig 64A, B. 

TYPE. Crovetto 9643: Argentina, Misiones: Santo Pipo, Nov 1962 (holotype, CORD 

n v; photo of holotype NY). 

DISTRIBUTION. (Fig 63D). Brasil, state of Rio Grande do Sul; Argentina, province 

of Misiones. In disturbed grasslands. BRASIL. Rio Grande do Sul: Anon s n (RB) Candelaria 

Rincao, Apr 1949. ARGENTINA. Misiones: Schulz 7053 (LIL) San Javier, Department San Javier, 7 

Feb 1948; Schulz 7178 (LIL) Santo Pipo, 19 Feb 1948;Schwarz 4981 (LIL) Santo Pipo, Department 

San Ignacio, 20 Oct 1947; Schwarz 5199 (TEX) Gobernador Roca, Department San Ignacio, 18 Nov 

1947; Schwarz 6369 (LIL, MO, TEX) Gobernador Roca, Department San Ignacio, 13 Oct 1948. 

LOCAL NAME. Timbo (anon. s n RB). 

54. Manihot hassleriana Chodat, Bull. Herb. Boissier 2(5): 672. 1905; Pax in Engler, 


Subshrubs. Stems glabrous. Leaves alternate; stipules persistent, ca 0.5 cm long, 

setaceous, glabrous; petioles short, usually less than 0.4 cm long, terete, glabrous, petiole 


pattern smooth; venation camptodromous, veins glabrous; lamina 3 lobed; median lobes 

linear, ca 25.0 cm long, ca 1.0 cm wide, apex acuminate, base of lobes 1.0 cm wide, width 

between base of sinus and petiole-lamina junction ca 3.0 cm; lowest lobes held at an angle 

of less than 45? to median lobes, slightly shorter than median lobes. Inflorescence a 

monoecious, terminal subspicate raceme, ca 7.0 cm long, all parts glabrous; bracteoles and 


restricted to the base of the upper half of the inflorescence, tepal 0.9 cm long, cleft to 

base into 5 lobes; ovary elongated. Staminate buds ovoid-ellipsoid; flowers small; tepal 

0.8 cm long, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Fruits 

and seeds non vidi. Fig 64D; 65A. 

TYPE. Hassler 4576: Paraguay: campo prope flumen Carimbatay, Sep 1898 

(syntypes, G-2). 

DISTRIBUTION. (Fig 64C). Paraguay. 

In the Graciles section Manihot hassleriana, M. hunzikeriana, M. flemingiana and M. 

maguireiana are the most distinct clements (See Fig 28, objects 66 & 67, 68 & 69, 64 & 

65 and 41 & 42). Of these 4 species, M. hassleriana is most closely related to M. 

hunzikeriana, both morphologically and geographically. 

55. Manihot triphylla Pohl, P1. Bras. Ic. et Descr. 1: 37. t. 28. 1827; Muell. Arg. in 

Martius, Fl. Bras. 11(2): 462. 1874. 

Jatropha triphylla Steudel, Nomencl. ed. 2. 1: 800. 1840; Manihot gracilis var triphylla Muell. 

Arg. in DC., Prodr. 15(2): 1066. 1866; Manihot triphylla Pohl var genuina Pax in Engler, 


Manihot angustifrons Muell. Arg. in Martius, Fl. Bras. 11(2): 461. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 72. 1910. Type. Riedel 1026 (syntypes, F, G). 

Manihot stenophylla Pax in Engler, Pflanzenreich IV. 147(Heft 44): 73. 1910. Type. Glaziou 

22129 (syntypes, P, F, G, K, NY). 

Shrubs, ca 1.0 m tall. Stems glabrous. Leaves alternate; stipules caducous; petioles ca 

8.0 cm long, terete, glabrous, petiole attachment to lamina basal, nonpeltate; lamina 

membranaceous, abaxial surface wax pattern reticulate; venation camptodromous, veins 

glabrous; lamina 3 lobed, median lobes linear, ca 15.0 cm long, ca 0.8 cm wide, apex 

acuminate, base of lobes extremely constricted and less than 0.1 cm wide, width between 

base of sinus and petiole-lamina junction less than 0.1 cm; lowest lobes form an angle of


4,- ~ ~ ~ ~ ~ ~ ~ ~ -, 

, , 

Ja~J 

FIG 64. Manibot hunzikeriana. A, leaf (Schulz 7178, LIL); B, inflorescence (Scbwarz 6369, 

MO). M. bassleriana. C, distribution; D, leaf (Hassler 4576, G).


less than 45" to the median lobes, slightly shorter than median lobes. Inflorescence a 

monoecious, terminal panicle, ca 10.0 cm long, few branched, flowers clustered at the 

terminal region of peduncles, with a bluish white bloom, all parts glabrous; bracteoles and 

bractlets setaceous. Pistillate flowers restricted to the base of the upper 1/3 of the 

inflorescence; pedicels 1.0 cm long; tepal 0.8 cm long, cleft to base into 5 lobes. 

Staminate buds bifusiform, flowers small, tepal 0.8 cm long, cleft 1/3 way down into 5 

lobes; stamens 10, in 2 whorls of 5 each. Capsules and seeds non vidi. Fig 65C, D. 

TYPE. Pohl 1184 (and on some specimens also 1708). Brasil, Goias: ad Serra de 

Cristaes (syntypes, F, K, M, W-3). 

DISTRIBUTION. (Fig 65B). Brasil, states of Goias, Minas Gerais and Distrito Federal. 

Alt ca 1000 m, among rocks. BRASIL. Goias: Glaziou 22129 (P, F, G, K, NY) Serra da Baliza, 4 

Jan 1895; Pobl Icon Tab. 28 (W). Distrito Federal: Irwin, Souza & dos Santos 12052 (F, G, K, MICH, 

NY, SP, UB, US) Corrego Landim, circa 25 km N of Brasilia, 27 Jan 1966;Maguire, Pires, Maguire & 

Silva 57088, (NY) Limit parque do Gama (Distrito Federal) and Goias, 15 Oct 1963. Minas Gerais: 

Assis 148 (GH) Fazenda Da Chicaca, Municipality Santa Luzia, 20 Nov 1945; Maguire & Maguire 

44732 (NY-3), 49 mi from Diamantina, Alt 3800', 22 Dec 1959; Occhioni s n Estrada de Nova Lima, 

29 Nov 1940. 

Formerly recognized as distinct species, Manihot angustifrons and M. stenophylla 

join at a C-value of 0.96 with M. triphylla, and thereafter remain as a distinct cluster until 

they join several other members of the Graciles section at C-value. 897. 

56. Manihot fruticulosa (Pax) Rogers & Appan, stat nov 

Manibot triphylla Pohl var fruticulosa Pax in Engler, Pflanzenreich. IV. 147(Heft 44): 74. 1910. 

Slender subshrubs ca 0.5 m tall, rarely to 1.0 m tall. Stems glabrous. Leaves 

alternate; stipules caducous; petioles ca 8.0 cm long, terete, glabrous, reddish, petiole 


pattern reticulate; venation camptodromous, veins glabrous; lamina 3 lobed, lobes 

drooping; median lobes linear, ca 10.0 cm long, ca 0.8 cm wide, base of lobes constricted, 

0.2 cm wide, width between base of sinus and petiole-lamina junction 0.2 cm; lowest 

lobes similar to median lobes in outline, slightly smaller. Inflorescence a monoecious, 

terminal panicle usually ca 18.0 cm long, occasionally shorter, all parts glabrous; 


flowers restricted to the base of the upper 1/2 of the inflorescence; pedicels 1.25 cm long; 

tepal 0.8 cm long, yellowish green with purplish streaks, cleft to base into 5 lobes; ovary 

subglobose. Staminate buds ovoid-ellipsoid, flowers small, tepal 0.8 cm long, yellowish 

green with purplish streaks, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 

each. Capsules 1.0 cm long, nonglobose, slightly elongated, surface smooth, apex 

rounded, dehiscence septicidal. Seeds small, 0.7 cm long, oblong, caruncle moderately 

prominent. Fig 66B, C. 

TYPE. Riedel s n (2820?): Brasil, Minas Gerais: bei Alegres (syntypes, K, NY, P). 

The Riedel specimens examined were without a collector's number. Pax (1910) cites 

Riedel 2820M, which we assume to be the same. 

DISTRIBUTION. (Fig 66A). Brasil, states of Goias, Minas Gerais and Distrito Federal. 

Alt ca 1000 m. Goias: Irwin, Souza & dos Santos 10013 (F, G, MO, NY, UB) 75 km N of 

Cristalina on road to Brasilia, 6 Nov 1965;Irwin, Souza & dos Santos 10493 (F, K, NY, UB, US) circa 

7 km E of Cabeceiras, 18 Nov 1965; Irwin, Souza & dos Santos 10938 (F, NY, UB) Cerrado ca 9 km S 

of Corumba de Goias, 2 Dec 1965; Irwin, Souza & dos Santos 11039A (NY, UB) Campo and Cerrado 

near Rio Descoberto, 4 Dec 1965; Pohl s n (G) Serra dos Christaes. Distrito Federal: Irwin, Souza & 

dos Santos 10164 (F, NY, SP, UB) Cerrado adjacent to Parque, Municipality do Gama, 10 Nov 1965. 

Minas Gerais: Claussen 756 (P), 1838. Province Unknown/Uncertain: Burchell 5861 (K), 19 Sep 1827. 

The new information added from the excellent specimens collected by Irwin et al 

makes it possible to more precisely define this taxon. Pax, with only the material of


1Cc 

B ~~~~~~~'' [ 

1 '''""'-''"-'>-- ''''I-.' I./",' 

, 

\ w > | --S 

~I 

Xa .Pob 118, W1 D inlrsec Pb 14 ) 

c~ i 

\~ ~ 

I . 

)~~~~~~~~~~~~~~~~~~~~~~ 

i .-_=_ .I 

FIG 65. Manihot hassleriana. A, inflorescence (Hassler 4576, G).M. triphylla. B, distribution; C, 

leaf (Pohl 1184,, inflorescence 

(Pobl 1184, W).


Riedel to base his judgment, could only recommend varietal status for this species. In 

actuality the computer analysis indicates that M. fruticulosa is more closely related to the 

following species, M. pentaphylla than to the preceding species M. triphylla. The drooping 

leaf lobes (Fig 66B) and the small stature of the plant are characteristic of this species. 

57. Manihot pentaphylla Pohl, P1. Bras. Ic. et Deser. 1: 53. t. 46. 1827. 

Subshrubs to shrubs, to 2.0 m tall. Stems glabrous. Leaves alternate; stipules 

caducous; petioles to 10.0 cm long, glabrous, terete, petiole attachment to lamina basal, 

nonpeltate; lamina membranaceous, abaxial surface wax pattern reticulate; venation 

camptodromous, veins glabrous; lamina palmately 5 lobed, occasionally 3 or 7; median 

lobes linear, 6.0-15.0 cm long, less than 1.0 cm wide, apex acuminate; lobe bases usually 

narrowly constricted, less than 0.3 cm wide, width between base of sinus and 

petiole-lamina junction less than 0.3 cm; lowest lobes more or less similar to median lobes 

in size and outline. Inflorescence monoecious, terminal, usually a simple raceme, 

occasionally a cluster of racemes arising from one basal point, to ca 10.0 cm long; all 

parts glabrous; bracteoles and bractlets semifoliaceous or foliaceous. Pistillate flowers 

restricted to the base of the inflorescence, or base of the upper half of the inflorescence; 

pedicels sometimes as long as 5.0 cm; tepal usually ca 1.0 cm long, cleft to base into 5 

lobes; ovary subglobose to slightly elongated; the trifid stigma moderately lobed and 

lobulate. Staminate buds ovoid-ellipsoid; flowers small, tepal ca 1.0 cm long, cleft 1/3 

way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules small, ca 1.0 cm long, 

surface smooth, apex rounded; dehiscence septicidal. Seeds small, ca 0.8 cm long, oblong; 

caruncle not prominent. 

DISTRIBUTION. (Fig 66D). Brasil, states of Para, Goias and Minas Gerais; Paraguay. 

Alt ca 1000 m on steep slopes and sandstone summits. 

This highly variable species has confused taxonomists as is evident by the number of 

misidentifications on herbarium labels. The computer analysis clearly indicates the fact 

that the subunits are geographical isolates of a single species. The subspecies have all 

clustered at the high C-value of .938 (Fig 28). 

All members of this species occur in degraded, dry, interior, upland regions. This 

species, in common with many other species, have adaptions to frequent burning. The 

particular adaptation of Manihot pentaphylla to this ecology is a fleshy storage root with 

adventitious buds at the root crown. 

Key to the Subspecies of Manihot pentaphylla 

1. Bracteoles more than 0.5 cm wide, margin entire; leaf lobes more than 0.5 cm wide. 

2. Leaf lobes lax; inflorescence usually a cluster of pendent racemes arising from one basal 

point, more than 6.0 cm long, many flowered; bracteoles more than 1.0 cm long. 

57a. M. pentaphylla subsp pentaphylla. 

2. Leaf lobes held rigidly erect; inflorescence usually a single raceme, less than 6.0 cm long, 

few flowered; bracteoles less than 1.0 cm long. 

57b. M. pentaphylla subsp rigidula. 

1. Bracteoles less than 0.5 cm wide, margin serrate; leaf lobes less than 0.5 cm wide. 

3. Petioles less than 7.0 cm long; leaf lobes extremely narrow, usually less than 0.25 cm 

wide; inflorescence less than 4.0 cm long; pistillate flowers borne on short, less than 2.0 

cm long pedicels. 

57c. M. pentaphylla subsp tenuifolia. 

3. Petioles more than 7.0 cm long; leaf lobes usually more than 0.25 cm wide; inflorescence 

more than 4.0 cm long; pistillate flowers borne on 3-5 cm long pedicels. 

57d. M. pentaphylla subsp graminifolia. 

57a. Manihot pentaphylla Pohl emend Rogers & Appan subsp pentaphylla 

Manihot pentaphylla Pohl, PI. Bras, Ic. et Descr. 1: 53. t. 46. 1827; Muell.-Arg. in Martius, Fl. 

Bras. 11(2): 473. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 36. 1910;Manihot


:/ 

,..~~~~~~~.~' 

w.... _.... 

, 

-..-?,,y/' 

* *- * v 

C / 

inflorescence (Irwin, Souza & dos Santos 10013, NY). M. pentaphylla. D, distribution, the dots repre- 

sent subsp rigidula, the letter O represents subsp pentaphylla, the letter N represents subsp tenuifolia, 

and the letter A represents subsp graminifolia.


pentaphylla var genuina Muell.-Arg. in DC., Prodr. 15(2): 1071. 1866;Jatropha pentaphylla 

Steudel, Nomencl. ed. 2. 1: 800. 1840. 

Manibot uleana Pax in Engler, Pflanzenreich IV. 147(Heft 44): 37. 1910. Type. Ule 3073 

(syntypes, F, NY). 

Petioles less than 7.0 cm long; leaf lobes 0.5-1.0 cm wide, lobes lax, not erect. 

Inflorescence a cluster of pendent racemes arising from one basal point, ca 10.0 cm long, 

usually with more than 15 flowers; bracteoles prominently foliaceous, ca 1.5 cm long, ca 

0.8 cm wide, margins entire. Pistillate flowers usually restricted to the base of the upper 

half of the inflorescence, occasionally closer to the base; pedicels ca 1.5 cm long. 

Fig 67A, B. 

TYPE. Pohl 1186 (and on some specimens also 1706). Brasil, Goias: ca Corumba et 

Corgo do Jaragua (syntypes, F, G, M, W-2). 

DISTRIBUTION. (Fig 66D). Brasil, state of Goias. BRASIL. Goias: Irwin, Maxwell & Was- 

shausen 18765 (B, COL, F, G, GH, NY, RB, UB, US) 20 km N of Corumba de Goias on road to 

Niquelandia, 18 Jan 1968; Irwin, Maxwell & Wasshausen 19263 (F, IAN, MICH, MO, NY, S, SP, UB, 

UC, W) circa 20 km NW of Corumba de Goias, near road to Niquelandia, 27 Jan 1968; Irwin, Souza & 

dos Santos 11742 (F, NY, SP, UB, US) Campo and Cerrado Sand Stone Summit of Serra Dourada, 18 

Jan 1966; Irwin, Souza & dos Santos 11755 (F, G, NY, UB) Campo and Cerrado Sand Stone Summit 

of Serra Dourada, 18 Jan 1966; Irwin, Souza & dos Santos 11887 (F, GH, K, MO, NY, RB, SP, UB, 

US) Cerrado Middle Slopes of Serra Dourada, circa 30 km SE of Goias Velho, 21 Jan 1966;Macedo 

3436 (MO, NY, US) Serra Dourada, 13 Dec 1951;Macedo 3461 (S) Serra Dourada, 13 Dec 1951;Pohl 

Icon Tab. 46 (W); Ule 3073 (F, NY) Serra Dos Pyreneos; Ule 374 (P) Serra dos Pyreneos Dec 1892; 

Ule 472 (P) Nos Campos Perto de Mossamedes, Jan 1893. 

57b. Manihot pentaphylla Pohl emend Rogers & Appan subsp rigidula (Mueller von 

Argau) Rogers & Appan, stat nov 

Manihot rigidula Muell.-Arg. in Martius, Fl. Bras. 11(2): 474 t. 67. 1874. 

Manihot conulifera Muell.-Arg. in Martius, Fl. Bras. 11(2): 474. 1874. Type. Riedel s n 


Petioles less than 7.0 cm long, leaf lobes 0.5-1.0 cm wide, lobes held erect on the 

plant. Inflorescence a simple raceme, ca 5.0 cm long or less, usually with less than 15 

flowers; bracteoles foliaceous, ca 1.0 cm long, ca 0.6 cm wide, margins entire. Pistillate 

flowers usually occurring nearer the base of the inflorescence; pedicels ca 1.0 cm long. 

TYPE. Riedel s n n v: Brasil, Minas Gerais; Warming s n (1508?): Brasil, Minas 

Gerais: near Lagoa Santa (syntypes, F, G, P). Both Muell. Arg. and Pax cite Warming s n 

One of the specimens examined listed the collector's number as 1508. We believe this 

specimen to be the same as the unnumbered collections. 

DISTRIBUTION. (Fig 66D). Brasil, states of Goias and Minas Gerais. BRASIL. Goias: 

Irwin & Soderstrom 7006 (NY, UB) circa 38 km S of Ciaponia on road to Jatai, 18 Oct 1964; Irwin & 

Soderstrom 7380 (F, GH, K, MO, NY, RB, SP, UB, US) circa 50 km S of Caiaponia on road to Jatai, 

26 Oct 1964; Riedel s n (F, G) Alegres. Minas Gerais: Claussen s n (W) Lagoa Santa, 1843, Glaziou 

19855 (P) Serra dos Cristaes, Pres de Diamantina, Apr 1892. 

57c. Manihot pentaphylla Pohl emend Rogers & Appan subsp tenuifolia (Pohl) Rogers & 

Appan, stat nov 

Manihot tenuifolia Pohl, PI. Bras. Ic. et Descr. 1: 38. 1827; Jatropha tenuifolia Steudel,- 

Nomencl. ed. 2. 1: 800. 1840. Manihot gracilis Pohl emend Muell.-Arg. var tenuifolia (Pohl) 

Muell.-Arg. in DC., Prodr. 15(2): 1066. 1866; Manihot reflexa Klotzsch ex Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 78. 1910 pro syn. 

Manibot tenerrima Pohl, PI. Bras. Ic. et Descr. 1: 39. 1827; Jatropha tenerrima Steudel, 

Nomencl. ed. 2. 1: 800. 1840. Manibot gracilis Pohl emend Muell.-Arg. var tenerrima (Pohl) 

Muell.-Arg. in DC., Prodr. 15(2): 1066. 1866; Manihot esculenta Crantz subsp 

flabellifolia (Pohl) Ciferri var tenerrima (Pohl) Ciferri, Type. Pohl 1182 (syntypes, F, G, W). 

Petioles less than 7.0 cm long, leaf lobes less than 0.25 cm wide, lobes lax, not erect. 

Inflorescence a short simple raceme, usually less than 3.0 cm long, usually with less than


C -A 

I/ /1 ? 

C | mL1 D 

FIG 67. Manihot pentaphylla subsp pentaphylla. A, leaf (Pobl 1186, W); B, inflorescence (Ir- 

win, Maxwell & Wasshausen 19263, NY). M. pentaphylla subsp rigidula. C, leaf (Warming 1508, F); D, 

inflorescence (Warming 1508, F).


10 flowers; bracteoles semifoliaceous, 0.8 cm long, ca 0.3 cm wide, margins serrate. 

Pistillate flowers restricted to the base of the inflorescence, pedicels ca 1.0 cm long. Fig 

68-A, B. 

TYPE. Pohl 2421: Brasil, Goias: ad Rio dos Indios Grande, in via ad fluvium Rio 

Claro (syntypes, F, G, W-2). 

DISTRIBUTION. (Fig 66D). Brasil, states of Para and Goias. BRASIL. Para: Pires, Black, 

Wurdack & Silva 6203 (NY) Serra do Cachimbo, 14 Dec 1956. Goias: Irwin & Soderstrom 7090 (UB) 

circa 33 km S of Caiaponia on road to Jatai, 20 Oct 1964; Irwin, Souza & dos Santos 11838 (UB) 

Campo and Cerrado Sandstone Summit of Serra Dourada, 20 Jan 1966; Macedo 3473 (US) Serra 

Dourada, 13 Dec 1951; Pohl Icon Tab. 29 (W); Pobl Icon Tab. 30 (W); Pohl 1182 (F, G, W), circa 

Corumba ad Villa Boa. Province Unknown/Uncertain: Weddell 2939 (P-2) Entre Goias et Cujaba, 

Nov-Dec 1844. 

57d. Manihot pentaphylla Pohl emend Rogers & Appan subsp graminifolia (Chodat & 

Hassler) Rogers & Appan, stat nov 

Manihot graminifolia Chodat & Hassler, Bull. Herb. Boissier 2(5); 671. 1905. 

Petioles ca 10.0 cm long, occasionally as long as 15.0 cm; leaf lobes 0.25-0.5 cm 

wide, lobes lax, not erect. Inflorescence a long, simple raceme, ca 12.0 cm, usually with 

about 10 flowers; bracteoles semifoliaceous, ca 1.5 cm long, 0.3 cm wide, margins serrate. 

Pistillate flowers restricted to the base of the inflorescence, borne on 3.0-5.0 cm long 

pedicels. Fig 68C, D. 

TYPE. Hassler 5172: Paraguay: Sierra de Maracayu, Oct 1900 (syntypes, BM, F-2, 

G-4, K, P, W). 

DISTRIBUTION. (Fig 66D). Paraguay. 

58. Manihot tenella Mueller von Argau in Martius, Fl. Bras. 11(2): 484. 1874; Pax in 


Shrubs, stems with a tendency to be decumbent, ca 0.5 m tall. Stems pubescent. 

Leaves alternate; stipules persistent, 1.0 cm long, filiform, pubescent; petioles ca 7.0 cm 

long, terete, pubescent, petiole attachment to lamina basal, nonpeltate; lamina membranaceous, 

abaxial surface wax pattern smooth; venation camptodromous, veins pubescent; 

lamina palmately 5 lobed; median lobes linear, ca 5.0 cm long, ca 0.5 cm wide, apex 

acuminate, base of lobes narrowly constricted, less than 0.2 cm wide; lowest lobes ca 1/2 

as long as median lobes. Inflorescence a monoecious, terminal ca 6.0 cm long raceme, 

peduncles and pedicels pubescent; bracteoles semifoliaceous, 1.2 cm long, 0.2 cm wide, 

pubescent, margins laciniate; bractlets semifoliaceous, 0.8 cm long, 0.1 cm wide. Pistillate 

flowers restricted to the base of the inflorescence; pedicels ca 1.0 cm long, pubescent; 

tepal 0.8 cm long, interior surface pubescent, cleft to base into 5 lobes; ovary pubescent. 

Staminate buds ovoid-ellipsoid, flowers small, tepal 0.7 cm long, cleft 1/3 way down into 

5 lobes, stamens 10, in 2 whorls of 5 each. Fruits and seeds non vidi. Fig 69B. 

TYPE. Riedel s n: Brasil, Sao Paulo: ad Rio Pardo (syntypes, photo at F, G). 

DISTRIBUTION. (Fig 69A). Brasil, state of Sao Paulo. PARAGUAY. Trinidad-Asuncion. 

Rojas 6214, 6326, 6326A, 6326C (A) Cabecera del Rio Aquidaban, Sierra de Amambay, Sep 1933. 

59. Manihot gracilis Pohl, P1. Bras. Ic. et Descr. 1: 23. t. 16. 1827. 

Subshrubs, several (5-15) divergent stems arising from a woody base, stems usually 

ascending, sometimes prostrate, to 1.0 m tall. Young stems obtuse angled in cross section, 

glabrous; mature stems glabrous. Leaves alternate; stipules sometimes persistent, filiform, 

0.5 cm long, glabrous; petioles short, ca 4.0 cm long, terete, glabrous, petiole attachment 

to lamina basal, nonpeltate; lamina membranaceous, abaxial surface wax pattern

156 

X - / . " 


FIG 68. Manihot pentapbylla subsp tenuifolia. A, leaf (Pobhl 2421, W); B, inflorescence (Pobl 

2421, W). M. pentaphylla subsp graminifolia. C, leaf (Hassler 5172, G); D, inflorescence (Hassler 5172, 

G).


I ` : r *- ^, 

l 

/,i_i 

I 

. 

FI 9 aio eel.A itiuin 

th osrpeetsbpgaii s a nd thelte 

th lat(Rgrs34,N) 

,la 

erset 

(oa 36A ) .gaii.C itiuin 

us ains .gaii ubpaii.D 

FIG 69. Manihot tenella. A, distribution; B, leaf (Rojas 6326-A, A). M. gracilis. C, distribution, 

the dots represent subsp gracilis and the letter A represents subsp varians. M. gracilis subsp gracilis. D, 

the plant (Rogers 364, NY).


reticulate; venation camptodromous, veins glabrous; lamina 3 lobed; median lobes linear 

or narrowly lanceolate, to 9.0 cm long, apex acuminate; lowest lobes more or less similar 

to median lobes in outline, slightly nonsymmetric. Inflorescence monoecious, terminal, 

short racemes, simple or a cluster of 2-4 racemes arising from a common base, ca 3.0 cm 

long, all parts glabrous; bracteoles semifoliaceous, ca 1.5 cm long, ca 0.3 cm wide, 

margins laciniate; bractlets semifoliaceous, ca 1.0 cm long, 0.2 cm wide, margins laciniate. 

Pistillate flowers restricted to the base of the inflorescence, pedicels less than 1.0 cm long, 

tepal 0.9 cm long, cleft to base into 5 lobes, ovary subglobose, the trifid stigma sparsely 

lobed and lobulate. Staminate buds conical; flowers small; tepal 0.9 cm long, cleft 1/3 

way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules small, subglobose, 1.0 

cm long, surface smooth, apex rounded, dehiscence septicidal. Seeds small, 0.8 cm long, 

oblong, caruncle not prominent. 

DISTRIBUTION. (Fig 69C). Brasil, states of Goias, Minas Gerais, Sao Paulo and 

Distrito Federal; Paraguay. Alt ca 1000 m in cerrado, "canga" soil. 

Key to the Subspecies of Manihot gracilis 

1. Leaf lobes narrowly lanceolate, usually less than 7.0 cm long, drooping. 59a.M. gracilis subsp gracilis. 

1. Leaf lobes digitate, usually more than 7.0 cm long, held horizontally. 59b. M. gracilis subsp varians. 

59a. Manihot gracilis Pohl emend Rogers & Appan subsp gracilis 

Manihot gracilis Pohl, PI. Bras. Ic. et Descr. 1: 23. t. 16. 1827; Jatropha gracilis Steudel, 

Nomencl. ed. 2. 1: 799. 1840. Manihot gracilis (Pohl) emend Muell.-Arg. var genuina 

Muell.-Arg. in DC., Prodr. 15(2): 1065. 1866. 

Manihot pronifolia Pohl, PI. Bras. Ic. et Descr. 1: 24. t. 17. 1827; Jatropha pronifolia Steudel, 

Nomencl. ed. 2. 1: 800. 1840. Manihot gracilis Pohl emend Muell. Arg. var pronifolia (Pohl) 

Muell.-Arg. in DC., Prodr. 15(2): 1065, 1866. Type. Pobl 1185 (syntypes, G, W). 

Manihot depauperata Pax in Engler, Pflanzenreich IV. 147(Heft 44): 41. 1910. Type. Hassler 

10222 (syntype, G). 

Shorter subshrubs, usually less than 0.75 m tall, stems ascending or prostrate. Leaf 

lobes shorter, usually less than 5.0 cm long, linear to narrowly lanceolate, lobes usually 

drooping, lamina at lobe sinus often slightly overlapping. Fig 69D; 70A, B. 

TYPE. Pobl 3920 (and on some specimens also 1659). Brasil, Goias: ad Arrayal 

Meyaponte (syntypes, F, K-2, W-2). 

DISTRIBUTION. (Fig 69C). Brasil, states of Goias, Minas Gerais, Sao Paulo and 

Distrito Federal; Paraguay. BRASIL. Goias: De Otero s n (RB) Posto de Criacao, Planaltina, 11 

Apr 1959; Glaziou s n (P) Rio da Gama, 31 Aug 1894; Glaziou 22129 (S) Serra da Baliza, Pres de 

Cachoeiras; Glaziou 22130 (F, G-2, K, P, US) Guariroba, 16 Aug 1894; Irwin & Soderstrom 7498 

(NY, UB) circa 35 km S of Caiaponia on road to Jatai, 28 Oct 1964; Irwin, Souza & dos Santos 10301 

(F, NY, UB) Cerrado at Summit of Chapada, circa 5 km E of Goias Boundary, 16 Nov 1965; Irwin, 

Souza & dos Santos 10557 (F, NY, SP, UB) Cerrado circa 3 km E of Cabeceiras, 19 Nov 1965; Irwin, 

Souza & dos Santos 10769 (NY, UB) circa 14 km S of Corumba de Goias, 30 Nov 1965; Irwin, Souza 

& dos Santos 10769A (UB) circa 14 km S of Corumba de Goias, 30 Nov 1965; Macedo 3473 (SP) 

Serra Dourada, Municipality Goias, 13 Dec 1951; Macedo & Smith 4651 (SP) Anapolis, 13 Oct 1956; 

Pohl Icon Tab. 16 (W); Pohl Icon Tab. 17 (W); Pohl 1185 (G, W) Meyaponte; Smith & Macedo 4651 

(US) Aeroporto, Anapolis. 13 Oct 1956. Distrito Federal: Cruz 125 (SP) Brasilia, Gama Cultivada No 

Jardim Botanico de Sao Paulo, 19 Feb 1964; Heringer 7810/4 (NY) Horto Do Guara, Brasilia, 4 Jan 

1961; Heringer 8760/954 (NY) Fundacao Zoobotanica, Brasilia, 5 Nov 1961; Heringer 8850/1044 

(NY) Granja Da Republica, Brasilia, 19 Jan 1962; Irwin, Grear, Souza & dos Santos 15516 (F, K, 

MICH, NY, S, UB) circa 3 km S of Sobradinho, 1 May 1966; Irwin, Souza & dos Santos 10263 (NY, 

UB) Cerrado, Brasilia, 13 Nov 1965; Irwin, Souza & dos Santos 10593 (F, G, MO, NY, RB, SP, UB, 

US, W) Campo, 12 km E of Braslandia on road to Brasilia, 22 Nov 1965; Irwin, Souza & dos Santos 

10676 (F, MO, NY, S, UB) Cerrado circa 12 km W of Taguatinga on road to Braslandia, 26 Nov 1965; 

Irwin, Souza & dos Santos, 10715 (F, G, GH, IAN, NY, S, SP, UB, US, W) Cerrado W of Setor 

Industrial Brasilia, 27 Nov 1965; Irwin, Souza & dos Santos 11113 (F, GH, MICH, MO, NY, RB, SP,


C LL`r' 

FIG 70. Manibot gracilis subsp gracilis. A, leaf (Pobi 1185, W); B, inflorescence (Irwin & Soder- 

strom 7498, NY). M. gracilis subsp varians. C, leaf (Pob 2208, W); D, inflorescence (Pobl 2208, W). 

/


UB, US-2) Cerrado S of Brasilia, 8 Dec 1965; Irwin, Souza & dos Santos 8074 (F, GH, MO, NY, RB, 

UB) Campo near Border of Cerrado, 8 Sep 1965; Irwin, Souza & dos Santos 8208 (F, NY, UB) 

Summit of Chapada Da Contagem, circa 10 km NE of Brasilia, 11 Sep 1965; Irwin, Souza & dos 

Santos 8269 (UB) Summit of Chapada Da Contagem, circa 10 km E of Brasilia, 13 Sep 1965; Irwin, 

Souza & dos Santos 8304 (F, NY, UB) Corrego Urubu, Wt Slopes of Chapada Da Contagem, 14 Sep 

1965; Irwin, Souza & dos Santos 8359 (UB) circa 10 km NE of Brasilia near Rio Torto, 16 Sep 1965; 

Irwin, Souza & dos Santos 8539 (F, K, NY, SP, UB) Brasilia, Asasul, 22 Sep 1965; Irwin, Souza & dos 

Santos 8671 (UB) circa 20 km S of Brasilia on road to Goiania near Rio Melchoir, 25 Sep 1965; Irwin, 

Souza & dos Santos 9053 (F, G, NY, S, UB, US) circa 25 km E of Brasilia near Planaltina, 8 Oct 1965; 

Irwin, Souza & dos Santos 9166 (F, GH, HB, MICH, MO, NY, SP, UB, UC, US, VEN, W) burned over 

Cerrado between Brasilia and Sobradinho, 13 Oct 1965; Irwin, Souza & dos Santos 9704 (F, G, GH, 

MO, NY, RB, SP, UB, UC, US) Cerrado near Setor Industrial, 30 Oct 1965;Labouriau & Valio 1182 

(SP) Gama, Cidade Satelite de Brasilia, 19 Feb 1964; Pereira 4621 (PUL, RB) Campo Cerrado, Brasilia, 

14 Nov 1958; Pires, Silva & Souza 9174 (NY) Brasilia, 18 Apr 1963. Minas Gerais: Claussen s n (K, 

NY); Claussen 757 (A); Regnell III 1073 (S-3, US) Cidade de Uberava, 12 Dec 1848; Riedel 789 (G, 

GH, NY, P); Rogers 362 (NY) Lagoa Santa, 17 Feb 1961; Rogers 364 (NY) Lagoa Santa, 17 Feb 

1961; Warming s n (P-2) Lagoa Santa; Weddell 1882 (P). Sao Paulo: Gehrt s n (SP) Pedregulho, 10 Apr 

1920. Province Unknown/Uncertain: Claussen 445 (G); Sellow s n (P). PARAGUAY. Province 

Unknown/Uncertain: Hassler 10222 (G) Sierra de Amambay, Feb; Rojas 6326B (A) Cabecera Rio 

Aquidaban, Sierra de Amambay, Campo Colina, Sep 1933. 

LOCAL NAME. Mandioquinha (De Otero sn) 

59b. Manihot gracilis Pohl emend Rogers & Appan subsp varians (Pohl) Rogers & Appan, 

stat nov 

Manibot varians Pohl, PI. Bras. Ic. et Descr. 1: 53. t. 47. 1827; Jatropha varians Steudel, 

Nomencl. ed. 2. 1: 800. 1840. Muell.-Arg. in DC., Prodr. 15(2): 1072. 1866; in Martius, Fl. 

Bras. 11(2): 475. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 40. 1910. 

Manihot pardina Muell.-Arg. in Martius, Fl. Bras. 11(2): 484. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 46. 1910. Type. Riedel s n (syntypes, F, G). 

Taller subshrubs ca 0.75-1.0 m tall, stems ascending. Leaf lobes longer, ca 8.0 cm in 

length, lanceolate to digitate, lobes held horizontally, lamina at lobe sinus not 

overlapping. Fig 70C, D. 

TYPE. Pohl 1 711 (and on some specimens also 2208) Brasil, Goias: ad Rio do Peixe 

(syntypes, F, G, K-2, M, W-2). 

DISTRIBUTION. (Fig 69C). Brasil, states of Goias, Minas Gerais and Sao Paulo. BRASIL. 

Goias: Pohl Icon. Tab. 47 (W); Minas Gerais: Barreto 3369 (F) Barreiro, Municipality Bello Horizonte, 

31 Jan 1933. Sao Paulo: Riedel s n (F, G) Rio Pardo. 

60. Manihot paviaefolia Pohl, PI. Bras. Ic. et Descr. 1: 52. t. 45. 1827; Muell.-Arg. in 

Martius, Fl. Bras. 11(2): 472. 1874; Peckolt, Ber. Deutsch. Pharm. Ges. 16: 29. 

1906; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 35. 1910. 

Jatropha paviaefolia Steudel, Nomencl. ed. 2. 1: 800. 1840; Manibot pentaphylla Pohl emend 

Muell.-Arg. var paviaefolia (Pohl) Muell.-Arg. in DC. Prodr. 15(2): 1071. 1866. 

.Subshrubs, to ca 1.0 m tall, stems ascending. Stems glabrous. Leaves alternate; 

stipules caducous, filiform, glabrous, 0.4 cm long; petioles ca 4.0 cm long, terete, 


surface wax pattern reticulate; venation camptodromous, veins glabrous; lamina palmately 

5 lobed, 3 major and 2 smaller; median lobes lanceolate, ca 7.0 cm long, ca 2.25 

cm wide, apex acuminate, base of lobes less than 0.2 cm wide, width between base of 

sinus and petiole-lamina junction less than 0.2 cm; lowest lobes ca 1/3 as long as median 

lobes. Inflorescence monoecious, terminal, pendent racemes, usually simple, occasionally 

a cluster of 2 racemes arising from a common base, ca 8.0 cm long, all parts glabrous; 

bracteoles prominently foliaceous, ca 1.25 cm long, ca 0.6 cm wide, yellowish green, 

margins entire; bracteoles foliaceous, 1.0 cm long, 0.5 cm wide, yellowish green. Pistillate


flowers restricted to the lower half of the inflorescence, pedicels to 3.5 cm long, tepal 1.0 

cm long, yellowish green without purple pigmentation, cleft to base into 5 lobes, ovary 

subglobose. Staminate buds ovoid-ellipsoid; slightly elongated; flowers medium sized, 

tepal 1.0 cm long, yellowish green without purple pigmentation, cleft 1/3 way down into 

5 lobes; stamens 10, in 2 whorls of 5 each. Capsules and seeds non vidi. Fig 71B, C. 

TYPE. Pohl 1190: Brasil, Goias: ad Corumba, Corgo do Jaragua et Ouro fino 

(syntypes, F, G, M, W-2). 

DISTRIBUTION. (Fig 71A). Brasil, state of Goias. Alt 1200 m, on steep sandstone 

slopes. BRASIL. Goias: Irwin, Maxwell & Wasshausen 19304 (F, GH, K, NY, SP, UB, US) circa 20 

km NW of Corumba de Goias, near road to Niquelandia, 27 Jan 1968; Irwin, Souza & dos Santos 

11707 (NY, UB) Summit of Serra Dourada, 20 km SE of Goias Velho, 18 Jan 1966; Macedo 4346 (K) 

Pirineus, Municipality Corumba, 17 Feb 1956; Pobl Icon. Tab. 45 (W); Ule 375 (P-2) Serra dos 

Pyreneos, Dec 1892. Province Unknown/Uncertain: Burchell 6289 (K). 

61. Manihot maguireiana Rogers & Appan, sp nov 13 

Frutices gracili, 1.0-1.5 m altus. Folia trilobus, lobi cernui; mediani lobi lanceolati 

inciso margine, raro intigra margine, basis racemi, ca 3.0 cm longi. Capsulae parvae, ca 0.9 

cm longae. 

Slender shrubs, 1.0-1.5 m tall, stems ascending. Young stems grayish brown, 

glabrous; mature stems dark purplish brown, glabrous. Leaves alternate; stipules 

caducous, filiform, less than 0.5 cm long, glabrous; petioles short, less than 4.0 cm long, 

terete, glabrous, dark purplish tinged, petiole attachment to lamina basal, nonpeltate; 

lamina membranaceous, adaxial surface dark green, abaxial surface silvery gray, wax 

pattern on the abaxial lamina surface shows discrete rings with irregular sides at 

magnifications of x40; venation frequently craspedodromous, sometimes camptodromous, 

veins glabrous; lamina 3 lobed, lobes usually deflexed; median lobes lanceolate 

with incised margin, rarely with entire margin, ca 7.0 cm long, ca 2.0 cm wide, the lobules 

along the lobe margin attenuate, ending in a fine point, the extension of a secondary vein, 

apex acuminate, base of lobes extremely constricted, less than 0.2 cm wide, width 

between base of sinus and petiole-lamina junction less than 0.2 cm; lowest lobes more or 

less similar to median lobes in size and outline, slightly nonsymmetric. Inflorescence 

monoecious, terminal, short racemes, simple or a cluster of 2 arising from a common 

base, ca 3.0 cm long, all parts glabrous; bracteoles semifoliaceous, ca 1.0 cm long, 0.2 cm 

wide, margin entire, rarely serrate; bractlets semifoliaceous, ca 0.75 cm long, 0.1 cm wide. 

Pistillate flowers restricted to the base of the inflorescence, pedicels 1.0-2.0 cm long; 

tepal 0.9 cm long, yellowish green with purplish streaks on the outside, pale pinkish 

inside, cleft to base into 5 lobes; ovary subglobose; the trifid stigma sparsely lobed and 

lobulate. Staminate buds ovoid-ellipsoid; flowers small, tepal 0.9 cm long, yellowish green 

with pinkish streaks externally, pale pinkish internally, cleft 1/3 way down into 5 lobes; 

stamens 10, in 2 whorls of 5 each, the superior whorl 0.7 cm long, the inferior 0.5 cm 

long, Capsules small, 0.9 cm long, subglobose, surface smooth, without wings, apex 

rounded. Seeds small, 0.7 cm long, oblong, caruncle not prominent. Fig. 72A, B. 

TYPE. Cardona 530: Venezuela, Bolivar: Rio Paragua, Isla el Casabe, alt ca 265 m, 

21 Apr 1943 (holotype, VEN; isotypes, A, US). 

DISTRIBUTION. (Fig 71D). Venezuela, state of Bolivar. Alt to 500 m. Frequent on 

rocky ridges and in savannas. VENEZUELA. Bolivar: Cardona 764 (US) Cerro Perro, Alto Rio 

Paragua, Jul 1943;Steyermark 75296 (VEN) vicinity of Uriman, 30 Apr 1953; Wurdack & Guppy 176 

(NY, US, VEN) 3 km E of Cano Azul, Hato La Vergarena, 23 Oct 1954. 

13 This species is named in honor of Dr. Bassett Maguire, outstanding botanist of The New York 

Botanical Garden, one of the foremost students of the vast and little known region of northeastern 

South America, the "Lost World" or Roraima Shield.


I 

0 0 

'..,.r, 

I 

a ....-- :: .. .. 6 ...- . / 

NY); C, inflorescence (Irwin, Maxwell & Wasshausen 19304, NY). M. maguireiana. D, distribution.


=SrL~ 75|' 

I'rA 

FIG ,72. Manihot maguireiana. A, leaf (Cardona 530, VEN); B, inflorescence (Cardona 530, 

VEN). 

Manihot maguireiana is the most distinct species of section Graciles, remaining as a 

separate unit in the computer analysis until the c-value .80. Its closest relatives are M. 

gracilis subsp gracilis, and M. triphylla. 

Its geographical distribution is distinct, northernmost in the section Graciles. 

9. Manihot sect Sinuatae Pax emend Rogers & Appan 

Manibot sect Grandibracteatae Pax subsect Tomentosae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 25. 1910 pro parte. 

Manibot sect Sinuatae Pax subsect Warmingianae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 


Manibot sect Parvibracteatae Pax subsect Langsdorffianae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 65. 1910 pro parte. 

Manibot sect Parvibracteatae Pax subsect Anomalae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 76. 1910. 

Manibot sect Heteropbyllae Pax subsect Cujabenses Pax in Engler, Pflanzenreich IV. 147(Heft 

44); 84. 1910. 

Manibot sect Heteropbyllae Pax subsect Variifoliae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 84. 1910 pro parte. 


petiole attachment basal; lamina membranaceous, deeply lobed, lobes variously shaped 

but not linear. Inflorescence monoecious, a cluster of subspicate racemes arising from a 

common base; bracts and bracteoles setaceous to foliaceous, margins usually entire. 

TYPE. Manihot warmingii Muell.-Arg. 


Key to the Species of Sect Sinuatae 

1. Leaf lobes entire or shallowly pandurate. 

1. Leaf lobes deeply pandurate. 

. 

62. M. anomala. 

63. M. warmingii.

? 

c--" 

2"- I- 

"" 

X' 

j 

*' 

-t- 

F '~~~~~~~~~~~~~~~~~~~~~~~~IGI~ 

73. Manihot sectSinuatae. Geographical range. 

FIG 73. Manibot sect Sinuatae. Geographical range.


62. Manihot anomala Pohl, P1. Bras. Ic. et Descr. 1:27. t. 21. 1827. 

Shrubs, to 3.0 m tall, frequently branched. Roots with intermittent tuberous 

enlargements, with milky latex and slight odor of HCN; epidermis rough, light yellowish 

brown to dark brown; subepidermis light tan, cortex with a watery consistency. Young 

stems glabrous or pubescent; mature stems woody with small pith, glabrous or pubescent, 

externally brown with green undercast, internally greenish white. Leaves alternate; young 

leaf and bud color strong yellow-green (5 GY 5/6) stipules caducous, filiform, glabrous or 

pubescent; petioles ca 10.0 cm long, terete, glabrous or pubescent, yellowish green or 

dark red (2.5 R 3/7), petiole attachment to lamina basal, nonpeltate; lamina 

membranaceous, abaxial surface wax pattern smooth; venation camptodromous, rarely 

craspedodromous, veins glabrous or pubescent; lamina 3-, rarely 5 lobed, leaves associated 

with the inflorescence frequently nonlobed; median lobes obovate, obovate pandurate, 

elliptic or narrowly elliptic with a repand or incised margin, usually 10.0-15.0 cm long, 

occasionally longer, 2.0-10.0 cm wide, apex acute to acuminate, base of lobes 0.75-3.0 

cm wide, width between base of sinus and petiole-lamina junction 0.75-3.0 cm; lowest 

lobes prominently nonsymmetric. Inflorescence monoecious, terminal, a cluster of 

subspicate racemes, to ca 15.0 cm long, totally glabrous or with various degrees of 

pubescence; bracteoles and bractlets foliaceous, semifoliaceous or setaceous, margins 

usually entire; rarely serrate, yellowish green or with some purplish pigmentation. 

Pistillate flowers restricted to the base of the upper 2/3 of the inflorescence, pedicels 

1.0-2.0 cm long, tepal to 2.0 cm long, yellowish green or with various degrees of purplish 

pigmentation, in the latter case usually the exterior surface is green and interior surface 

with dark maroon stripes, cleft to base into 5 lobes; disc light yellow to reddish orange; 

ovary subglobse, the trifid stigma well lobed and lobulate. Staminate buds globose or 

ovoid-ellipsoid to slightly constricted in the middle, flowers small (0.75 cm long) to large 

(2.0 cm long); color of tepal same as that of pistillate tepal, cleft 1/3 way down into 5 

lobes; disc light yellow to reddish orange; stamens 10, in 2 whorls of 5 each. Capsules 

0.75-1.5 cm long, subglobose to slightly elongated, surface smooth without prominent 

wings, apex rounded, dehiscence septicidal. Seeds 0.6-1.2 cm long, oblong, caruncle 

usually prominent. 

DISTRIBUTION. (Fig 74A). Brasil, states of Para, Mato Grosso, Goias, Minas Gerais, 

Sao Paulo and in Distrito Federal; Paraguay; Peru in Depts Cusco and Apurimac; Bolivia, 

Depts La Paz and Santa Cruz; Argentina, Provinces Jujuy, Salta, Catamarca and Tucuman. 

Alt ca 300-1000 m. In disturbed gallery forest, rocky slopes, creek margins, savanna, and 

in flat country. 

Manihot anomala has one of the widest distributions of the wild species of the genus. 

As a result of this distribution, and the great variability in leaf morphology, many of the 

phenotypic variations have been described as species. The senior author's collections of 

one of the subspecies (pavoniana) indicate the wide variation in one population (Figs 

76B, C, D; 77A, B, C). The unifying character of the species is the cluster of subspicate 

racemes. 

The variability in the phenotypes of this species indicates a highly heterozygous gene 

pool. The disturbed habitats in which the plants occur indicate that man has played a role 

in the expression of this phenotypic plasticity, much as in the case of the consciously 

cultivated species,Manihot esculenta. 

Key to the Subspecies of Manihot anomala 

1. Bracts and bracteoles foliaceous, more than 0.5 cm wide; flowers yellowish green; plant 

parts densely pubescent. 

62b. M. anomala subsp pubescens. 

1. Bracts and bracteoles semifoliaceous or setaceous, less than 0.2 cm wide; flowers mostly 

purplish tinged, sometimes yellowish green; plant parts moderately pubescent to glabrous. 

2. Bracts and bracteoles usually less than 1.0 cm long; flowers yellowish green or with some 

purple pigmentation; plant parts moderately pubescent.


-C Ii. 

. 

i 

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. 

X 11 

FIG74 Manho a 

nml. A isrbuin,tedosrersetsus bsen,th ete 

rereets 

us 

anoaa h etrMrpeet us uaess h etrNrpeet us lb 

raaadtelte ersnssbppvnaa N.aoaasbpaoaa ,la Ph 17 ) 

C,ifoecne(lusn.2,N) .aoml us uecn.D ef(l $ ) 

FIG 74. Manihot anomala. A, distribution, the dots represent subsp pubescens, the letter A 

represents subsp anomala, the letter M represents subsp cujabensis, the letter N represents subsp glab- 

rata and the letter D represents subsp pavoniana. M. anomala subsp anomala. B, leaf (Pobl 1197, W); 

C, inflorescence (Claussen 322, NY). M. anomala subsp pubescens. D, leaf (Ule 473, P).


3. Staminate flowers usually less than 1.0 cm long; staminate buds more or less globose. 

4. Leaf lobes more than 2.5 cm wide; margin entire. 62aM. anomala subsp anomala. 

4. Leaf lobes less than 2.5 cm wide; margin repand or incised. 

62c. M. anomala subsp cujabensis. 

3. Staminate flowers usually more than 1.0 cm in length; staminate buds ovoid-ellipsoid. 

62d. M. anomala subsp glabrata. 

2. Bracts and bracteoles usually more than 1.0 cm long, rarely shorter, flowers usually with 

considerable purple pigmentation; plant parts glabrous, rarely sparsely pubescent. 

62e. M. anomala subsp pavoniana. 

62a. Manihot anomala Pohl emend Rogers & Appan subsp anomala 

Manihot anomala Pohl, PI. Bras. Ic. et Descr. 1: 27. t. 21. 1827; Jatropha anomala Steudel, 

Nomencl. ed. 2. 1: 799. 1840; Muell.-Arg. in DC., Prodr. 15(2): 1059. 1866; in Martius Fl. 


Manihot heterophylla Pohl, P1. Bras. Ic. et Descr. 1: 39. t. 31. 1827; Jatropha heterophylla 

Steudel, Nomencl. ed. 2. 1: 799. 1840; Muell.-Arg. in DC., Prodr. 15(2): 1073. 1866; in 

Martius Fl. Bras. 11(2): 479, 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 85. 1910. 

Type. Pohl 1254 (syntypes, F-2, G, W-3). 

Manihot caricaefolia Pohl, PI. Bras. Ic. et Descr. 1: 40. t. 32. 1827;Jatropba caricaefolia Steudel, 

Nomencl. ed. 2. 1: 799. 1840; Muell.-Arg. in DC., Prodr. 15(2): 1072. 1866; in Martius, Fl. 

Bras. 11(2): 480. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 84. 1910. Type. Pohl 

1187 (syntypes, F, W-2). 

Plant parts sparsely to moderately pubescent. Leaf lobes entire, or pandurate, more 

than 2.5 cm wide, lobe bases narrow, usually less than 1.25 cm wide, width of lamina 

betweeen lobe bases and petiole-lamina junction usually less than 1.25 cm; venation 

camptodromous. Bracts and bracteoles setaceous, usually less than 1.0 cm long and less 

than 0.2 cm wide; staminate buds more or less globose; pistillate and staminate tepals 

yellowish green, rarely with little purplish pigmentation; staminate tepal less than 1.0 cm 

long. Fig 74B, C. 

TYPE. Pohl 1197 (and on some specimens also 1702): Brasil, Goias: ad Corgo do 

Jaragua, Corumba et Ouro fino (syntypes, F, K, M-2, W-2). 

DISTRIBUTION. (Fig 74A). Brasil, states of Mato Grosso, Goias, Minas Gerais, and 

Sao Paulo. BRASIL. Mato Grosso: Gomes 2205 (SP) Municipality De Tres Lagoas, Fazenda 

Floresta, Planta da Margem Do Rio Brioso, 20 Sept 1964; Harley & Souza 11023 (K) Xavantina, 

14.38? S.-52.14? W, 11 Nov 1968; Lindman A2505 (S-3) Palmeiras, 14 Dec 1893; Lindman 2505 (P) 

Palmeiras, 14 Dec 1893; Sick B265 (PUL, RB) Rio Dos Mortos, Xavantina, Feb 1947. Goias: Irwin, 

Souza & dos Santos 11896 (F, G, GH, K, MO, NY, RB, SP, UB, US, W) Serra Dourada ca 30 km SE 

of Goias Velho, 21 Jan 1966; Pohl Icon Tab. 21 (W);Pohl Icon. Tab. 31 (W);Pohl Icon. Tab. 32 (W); 

Pohl 1187 (F, W-3) in Silva Matogrosso; Pobl 1254 (F-2, G, W-3) Villa Boa. Minas Gerais: Claussen s n 

(K, P); Claussen 332 (G-3, NY), 1840; Claussen 755 (A, GH, NY, P, S), 1838; Claussen s n (K), 1840; 

Claussen s n (G-2), 1839; Macedo 412b (US) Ituiutaba, 10 Jan 1956. Sao Paulo; Caldas s n (SP) Rio 

Preto, Sao Paulo, 9 Mar 1942; Mendes s n (SP) Mato Proper. Joaquin Leonel, Pindorama, 14 Dec 

1938. Province Unknown/Uncertain: Burchell 8451 (K); Weddell s n (P) Nov-Dec 1844. 

62b. Manihot anomala Pohl emend Rogers & Appan subsp pubescens (Pohl) Rogers & 


Manihot pubescens Pohl, PI. Bras. Ic. et Descr 1: 47. t. 40. 1827; Jatropha pubescens Steudel, 

Nomencl. ed. 2. 1: 800. 1840. Muell.-Arg. in DC., Prodr. 15(2): 1067. 1866; in Martius, Fl. 


Plant parts pubescent to moderately tomentose. Leaf lobes entire, more than 2.5 cm 

wide, lobe bases wide, usually more than 2.0 cm wide, width of lamina between lobe 

bases and petiole-lamina junction usually more than 2.0 cm; venation camptodromous. 

Bracts and bracteoles foliaceous, more than 1.0 cm long, more than 0.5 cm wide; 

staminate buds ovoid-ellipsoid; pistillate and staminate tepals yellowish green, rarely with 

little purplish pigmentation; staminate tepal less than 1.0 cm long. Fig 74D; 75A.


TYPE. Pohl 1196 (and on some specimens also 1198, 1658 and 2047): Brasil, Goias: 

ad Engenho do S. Sebastiao non procul a S. Luzia, ad corumba, corgo do Jaragua, Ouro 

fino et Cidade do Gioyaz (syntypes, A, F, W-3). 

DISTRIBUTION. (Fig 74A). Brasil, states of Para, Goias, Minas Gerais and Distrito 

Federal. BRASIL. Para: Froes 23548 (PUL) Breu Branco, Rio Tocantins, 28 Sep 1948; Pires, Black, 

Wurdack & Silva 6159 (NY) Serra do Cachimbo, 12 Dec 1956. Goias: Gardner 3441 (BM) near Villa 

de Natividade, Dec 1839; Glaziou 22125 (P, S) Formosa a Itiquira, 2 Feb 1895; Irwin, Maxwell & 

Wasshausen 19200 (F, LIL, MICH, MO, NY, SP, UB, UC, US, W) 75 km N of Corumba de Goias on 

road to Niquelandia, 25 Jan 1968; Irwin, Souza & dos Santos 10344 (F, G, GH, NY, POM, RB, S, SP, 

UB, UC, US) circa 14 km E of Cabeceiras, 16 Nov 1965; Irwin, Souza & dos Santos 11862 (F, GH, K, 

LE, MO, NY, RB, SP, UB, US) Lower Slopes of Serra Dourada, circa 30 km E of Goias Velho, 20 Jan 

1966; Macedo 4417 (US) Jacuba, Niquelandia, 24 Feb 1956; Pobl Icon TAB. 40 (W); Ule 473 (P) 

Mossamedes, Jan 1893. Distrito Federal: Irwin, Grear, Souza & dos Santos 14004 (F, GH, MO, NY, 

RB, SP, UB, US, W) Corrego Landim circa 25 km N of Brasilia, 16 Mar 1966; Irwin, Maxwell & 

Wasshausen 19430 (B, F, G, GH, MO, NY, RB, S, SP, UB, US) circa 25 km NE of Brasilia, Limestone 

Outcrop near Corrego Landim, 2 Feb 1968; Irwin, Souza & dos Santos 11316 (F, G, MICH, MO, NY, 

S, SP, UB, US) Corrego Landim circa 20 km N of Brasilia, 15 Dec 1965. Minas Gerais: Claussen 754 

(P-2, S), 1838; Heringer 9710 (NY) Paraopeba, 5 Oct 1960; Warming s n (A, P-2, US) Lagoa Santa. 

Province Unknown/Uncertain: Weddell 2525 (P), 1844. 

62c. Manihot anomala Pohl emend Rogers & Appan subsp cujabensis (Mueller von 


Manibot cujabensis Muell.-Arg, Linnaea 34: 207. 1865-1866; in DC., Prodr. 15(2): 1074. 1866; 

Pax in Engler, Pflanzenreich IV. 147(Heft 44): 84. 1910;Manihot cuiabensis Muell.-Arg. in 

Martius, Fl. Bras. 11(2): 479. 1874 (orthographic variant). 

Manibot alcicornis Klotzsch ex Pax in Engler, Pflanzenreich IV. 147(Heft 44): 84. 1910. pro syn. 

Plant parts sparsely pubescent or glabrous. Leaf lobes narrowly elliptic with repand 

or incised margin, usually less than 2.0 cm wide, lobe bases narrow, usually less than 1.25 

cm wide, width of lamina between lobe bases and petiole-lamina junction usually less 

than 1.25 cm; venation frequently craspedodromous. Bracts and bracteoles setaceous, 

usually less than 1.0 cm long and less than 0.2 cm wide; staminate buds more or less 

globose; pistillate and staminate tepal yellowish green; staminate tepal less than 1.0 cm 

long. Fig 75B, C. 

TYPE. Manso 76B: Brasil, Mato Grosso: near Cuiaba (or Cujaba) (syntypes, 2 photos 

at F, photo at NY). 

DISTRIBUTION. BRASIL. Mato Grosso: Gaudichaud 280 (P), 1833. Province Unknown/ 

Uncertain: Burchell 8850 (K-2). 

62d. Manihot anomala Pohl emend Rogers & Appan subsp glabrata (Chodat & Hassler) 

Rogers & Appan, stat nov 

Manihot pubescens f glabrata Chodat & Hassler, Bull. Herb. Boissier 2(5): 672. 1905; Manihot 

glabrata (Chodat & Hassler) Pax in Engler, Pflanzenreich IV. 147(Heft 44): 43. 1910. 

Manihot langsdorffii var glabra Chodat & Hassler, Bull. Herb. Boissier 2(5): 673. 1905;Manihot 

klingensteinii Pax in Engler, Pflanzenreich IV. 147(Heft 44): 66. 1910. Type. Hassler 7976 

(syntypes, BM, F-3, MO, NY-2, P, S, UC); Hassler 8121 (syntypes, A, BM, MO, NY, P, S, 

UC, W). 

Manihot multiflora Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 44): 85. 1910. 

Type. Fiebrig 4360 (syntypes, BM, F, G-6, K, L, M). 

Manihot pseudoheterophylla Pax in Engler, Pflanzenreich IV. 147(Heft 44): 86. 1910. Type. 

Hassler 5649 (syntypes, BM, F-2, NY, P-2, W). 

Plants parts sparsely pubescent or glabrous. Leaf lobes entire or pandurate, more 

than 2.5 cm wide, lobe bases narrow, usually less than 1.25 cm wide, width of lamina 

between lobe bases and petiole-lamina junction usually less than 1.25 cm; venation 

camptodromous. Bracts and bracteoles setaceous, occasionally semifoliaceous, usually less 

than 1.0 cm long, usually less than 0.2 cm wide, when semifoliaceous, ca 0.3 cm wide;


C_, 

"'-.1' vI- 

Al 

FIG 75. Manihot anomala subsp pubescens. A, inflorescence (Pohl 1658, W). M. anomala subsp 

cujabensis. B, leaf (Gaudichaud 280, P); C, inflorescence (Gaudichaud 280, P). M. anomala subsp 

glabrata. D, leaf (Hassler 5730, F).


staminate buds ovoid-ellipsoid, slightly constricted in the middle; pistillate and staminate 

tepals yellowish green, occasionally with little purplish pigmentation; staminate tepal 

usually 1.0-2.0 cm long. Fig 75D; 76A. 

TYPE. Hassler 5730: Paraguay: ad ripam fluminis Jejui guazu (syntypes, BM, F-2, 

NY-2, P-2, S, UC, W). 

DISTRIBUTION. (Fig 74A). Paraguay. PARAGUAY. Province Unknown/Uncertain: Balansa 

1715 (P) Summit Du Cerro de Yaguaron, 29 Jan 1877; Fiebrig 4360 (BM, F, G-6, K, L, M) 

Zwischen Rio Apa Und Rio Aquidaban, Nov; Hassler 10741 (BM, MO, NY, S, UC, W) in Altaplanitie 

et Declivibus, Sierra de Amambay, Dec 1907; Hassler 5516 (BM, NY, P, W) in Regione Yerbalium de 

Maracayu, Paraguaria Euro-Austra, Nov; Hassler 5649 (BM, F-2, NY, P-2, W) in Regione Yerbalium de 

Maracayu, Paraguaria Euro-Austra, Dec; Hassler 7976 (BM, F-3, MO, NY-2, P, S, UC) in Regione 

Cursus Superioris Fluminis Apa, Nov; Hassler 8121 (A, BM, MO, NY, P, S, UC, W) in Regione Cursus 

Superioris Fluminis Apa, Dec; Rojas 3667 (A) Lima Alto Paraguay, Orillas de Bosque, Apr 1921; 

Rojas 6809 (A) Cerro Coro, Sierra de Amambay, Campo Orillas de Bosque, Mar 1934; Woolston 109 

(UC, US) Primavera, Alto Paraguay, 5 Mar 1954. 

62e. Manihot anomala Pohl emend Rogers & Appan subsp pavoniana (Mueller von 

Argau) Roger & Appan, stat nov 

Manihot pavoniana Muell.-Arg., Linnaea 34: 205. 1865; in DC., Prodr. 15(2): 1059. 1866; Pax in 

Engler, Pflanzenreich IV. 147(Heft 44): 65. 1910; Jatropha simayuca Ruiz & Pavon ex Pax 

in Engler, Pflanzenreich IV. 147(Heft 44): 65. 1910 pro syn. 

Manihot weberbaueri Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 194. 1924. 

Type. Weberbauer 6517 (syntype, F, G). 

Plant parts totally glabrous, rarely sparsely pubescent. Leaf lobes entire or 

pandurate, more than 2.5 cm wide, lobe bases narrow, usually less than 1.25 cm wide, 

width of lamina between lobe bases and petiole-lamina junction usually less than 1.25 cm; 

venation camptodromous. Bracts and bracteoles semifoliaceous, usually 1.0-2.0 cm long, 

ca 0.3 cm wide; staminate buds ovoid-ellipsoid, slightly constricted in the middle; 

pistillate and staminate tepals yellowish green with considerable amount of purplish 

pigmentation; staminate tepal usually less than 1.0 cm long, sometimes longer. 

Fig 76B, C, D; Fig 77A, B, C. 

TYPE. Ruiz & Pavon s n: Peru (syntypes, BM, F-2, G, NY). 

DISTRIBUTION. (Fig 74A). Peru, Depts. of Cusco and Apurimac; Bolivia, Depts. of 

La Paz and Santa Cruz; Argentina, Provinces of Jujuy, Salta, Catamarca and Tucuman. In 

disturbed habitats, abandoned fields, and fence rows. PERU. Cusco: Vargas 14335 (NY) Limatombo, 

Sisal, Province Anta, 15 Mar 1963; Vargas 4884 (A, MO) Sisal Cunyac, Province Anta, 18 Jan 

1945. Apurimac: Ferreyra 2747A (NY-3, US-2) Entre Cuzco y Abancay, circa Del Puente Cunyac, 

Province Abancay; 20 Nov 1947; Vargas 10031 (GH, MO) Pachachaca, Province Abancay, 31 Dec 

1950; Vargas 26329 (UC) Hacienda Matara, Province Abancay, 25 Jan 1939; Vargas 5852 (NY) 

Kairanka, Province Grau, 9 Mar 1946. Province Unknown/Uncertain: Gay s n (P); Weberbauer 6517 

(F, G) NE of Pampus, Valle Del Mantaro. BOLIVIA. La Paz: Buchtien 4251 (GH, NY, US) Millihuaya, 

in Nord-Yungas, Dec 1917. Santa Cruz: Herzog 1419 (L, S) im Walde Bei Santa Cruz, Jan 1911; 

Rogers 430 (A, BM, COLO, G, K, M, MICH, MO, NY-2, P, S, SP, US-2, W) Experimental Station near 

General Saavedra, 80 km N of Santa Cruz, 11 Jan 1962; Rogers 431 (ARIZ, DAV, MG, UC, US) 

Experimental Station near General Saavedra, 80 km N of Santa Cruz, 11 Jan 1962; Rogers 435 (F-2, 

G, M, MICH, MO, NY, S, W) Province Warnes, Paichanetu, km 40 on road from Santa Cruz to 

Montero, 13 Jan 1962; Rogers 437-A (NY-2, W) Experimental Station near General Saavedra, 80 km 

N of Santa Cruz, 14 Jan 1962; Rogers 437-B (COLO) Experimental Station near General Saavedra, 80 

km N of Santa Cruz. 14 Jan 1962; Rogers 437-C (A, BM, F, G, K, NY-3, P, S) Experimental Station 

near General Saavedra, 14 Jan 1952; Rogers 437-D (G) Experimental Station near General Saavedra, 

80 km N of Santa Cruz, 14 Jan 1962; Steinbach 3143 (GH) Montecito de Delores, Cant. Buena Vista, 

Province del Sara, 4 Nov 1916; Steinbach 5202 (A, NY) Bosquerillos, Buena Vista, Province Sara; 

Steinbach 6709 (A, BM, F, MO, NY, S, U, UC) Bosques de Buena Vista, Province Sara, 29 Nov 1924. 

Province Unknown/Uncertain: Williams 233 (BM, NY-2) San Jose, 2 Feb 1902. ARGENTINA. Jujuy: 

Lillo 52696 (LIL) Ledesma, 23 Dec 1906; Spegazzini s n (BAB) Ledesma, 30 Jan 1906. Salta: 

Schreiter 3704 (LIL) Tartagal, Department Oran, 25 Feb 1925; Schreiter 5189 (LIL) Embarcacion 

F.C.C.N, 21 Dec 1925. Catamarca: Diaz s n (LIL) Aquadita, Department Capital, Dec 1931. 

Tucuman: Lillo 1822 (LIL) Tucuman, Apr 1895; Lillo 5235 (LIL) El Bosque, 7 Dec 1906; Venturi


ers 431, NY). 

ers 41 NY ) . 

FIG 76. Manihot anomala subsp glabrata; A, inflorescence (Fiebrig 4360, G). M. anomala subsp 

pavoniana. B, habit (Rogers 437, NY); C, roots (Rogers 435, NY); D, leaf with pandurate lobes (Rogers 

431, NY).


10191 (LIL, US) Department Burroyaco, Cerro Del Campo, 23 Feb 1930; Venturi 1690 (BAB, LIL, 

US) Durazuito, Department Capital; Venturi 1690B (US) El Cadillal, Department Tafi, 29 Dec 1923; 

Venturi 2865 (GH, LIL) El Durazuito, Department Capital, 16 Feb 1924. 

63. Manihot warmingii Mueller von Argau in Martius, Fl. Bras. 11(2): 481. 1874; Pax in 


Plant growth habit unknown. Stems non vidi. Leaf arrangement on the stem 

unknown; stipules filiform, ca 0.5 cm long, pubescent; petioles ca 4.0 cm long, terete, 

pubescent, petiole attachment to lamina basal, nonpeltate; lamina membranaceous, 

abaxial surface wax pattern smooth; venation camptodromous, veins tomentose; lamina 

palmately 5 lobed; median lobes obovate, deeply pandurate, ca 9.0 cm long, ca 3.5 cm 

wide, apex acuminate, base of lobes ca 1.0 cm wide; lowest lobes ca half as long as 

median lobes. Inflorescence structure unknown (only a fragment of the inflorescence 

seen); peduncles and pedicels tomentose; bracteoles foliaceous, ca 0.8 cm long, ca 0.5 cm 

wide, pubescent, margins entire; bractlets foliaceous. Pistillate flowers non vidi. Staminate 

buds ovoid-ellipsoid, flowers small, tepal ca 0.7 cm long, pubescent, cleft 1/3 way down 

into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules and seeds non vidi. 

TYPE. Warming s n (1507 ?): Brasil, Minas Gerais: near Lagoa Santa, 19 Dec 1865 

(syntypes, photo at F, G). 

DISTRIBUTION. (Fig 77D). Brasil, Minas Gerais. 

Manibot warmingii is represented either by photographic material or by fragments, 

and its status as a species is questionable. It may be a variant or synonym of M. anomala 

subsp pubescens. 

10. Manihot sect Variifoliae Rogers & Appan, sect nov 

Manihot sect Heteropbyllae Pax subsect Variifoliae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 84. 1910 pro parte. 


Plantae caulinae, lignosae, infernae ad mediae frutices. Folia petiolatae, penitae. 

Inflorescentia monoecia subspicata racema. 


petiole attachment peltate; lamina membranaceous, deeply lobed, lobes variously shaped, 

but not linear. Inflorescence monoecious, a cluster of subspicate racemes arising from a 

common base; bracts and bracteoles setaceous, margins entire. 

TYPE. Manihot variifolia Pax et K. Hoffmann. 


Key to the Species of Sect Variifoliae 

1. Leaf venation camptodromous; median lobes more than 5.0 cm wide, obovate. 64. M. mirabilis 

1. Leaf venation craspedodromous; median lobes less than 2.0 cm wide, hastate, with 2 

well-developed secondary lobes at base. 65. M. variifolia 

64. Manihot mirabilis Pax in Engler, Pflanzenreich IV. 147(Heft 44): 91. 1910. 

Shrubs. Young stems pubescent, obtuse-angled in cross section; mature stems 

sparsely pubescent or glabrous. Leaves alternate; stipules setaceous, pubescent, caducous; 

petioles ca 9.0 cm long, obtuse-angled, pubescent, petiole attachment to lamina peltate, 

width between basal edge of lamina and petiole-midrib junction ca 1.0 cm; lamina 

herbaceous to slightly coriaceous, abaxial surface wax pattern smooth; venation 

camptodromous, midribs stout, ca 0.2 cm in diam near base, primary, secondary, and 

tertiary veins prominent and bright yellowish, glabrous; lamina palmately 3 lobed, 

frequently nonlobed; median lobes obovate to almost rotundate, occasionally shallowly


~r/ 

FIG 77. Manibot anomala subsp pavoniana. A, leaf with entire lobes (Rogers 430, NY); B, non- 

lobed, entire-lobed and pandurate-lobed leaves on the same stem (Rogers 437-B, NY); C, inflorescence 

(Rogers 430, NY). M. warmingii. D, distribution.

}?\???)? **^^ ' 

/, 

i 

(, 

7 

I.M f range.. ... 

'O 

I ?^ 

FIG 78. Mani_ot sect Vatiifoliae. Geographical range. 

0 

I ..m. 

] 

. 

"- 

,.


pandurate, ca 12.0 cm long, 9.0-11.0 cm wide, apex obtuse, base of lobes narrow, ca 1.0 

cm wide, width between base of sinus and petiole-lamina junction ca 1.0 cm; lowest lobes 

prominently nonsymmetric. Inflorescence monoecious, terminal, a cluster of 2-6 

subspicate racemes arising from one common base, ca 12.0 cm long; peduncles and 

pedicels pubescent; bracteoles and bractlets setaceous, less than 0.5 cm long, pubescent. 

Pistillate flowers restricted to the base of the upper 2/3 of the inflorescence; pedicels 1.5 

cm long, pubescent; tepal 1.4 cm long, exterior and interior surfaces pubescent, cleft to 

base into 5 lobes; disc prominent; staminodes present; ovary subglobose, pubescent. 

Staminate buds ovoid-ellipsoid; flowers large, tepal 1.4 cm long, exterior and interior 

surface pubescent, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each, 

filaments pubescent. Capsules and seeds non vidi. Fig 79B, C. 

TYPE. Hassler 10711: Paraguay: Sierra de Amambay (syntypes, BM-2, F, NY, P, S, 

UC, W-2). 

DISTRIBUTION. (Fig 79A). Paraguay. 

65. Manihot variifolia Pax in Engler, Pflanzenreich IV. 147(Heft 44): 85. 1910. 

Manihot katharinae Pax in Engler, Pflanzenreich IV, 147(Heft 44): 87. 1910. Type. Hassler 2113 

(syntypes, F, NY-2, P, W). 

Shrubs. Young stems sparsely pubescent, obtuse-angled in cross section, purplish 

tinged; mature stems glabrous. Leaves alternate; stipules caducous, setaceous, less than 

0.5 cm long, sparsely pubescent; petioles ca 9.0 cm long, obtuse-angled, sparsely 

pubescent, petiole attachment to lamina peltate, width between base of lamina and 

petiole-midrib junction ca 0.5 cm; lamina membranaceous to slightly coriaceous, abaxial 

surface wax pattern smooth; venation craspedodromous, veins glabrous, primary, 

secondary, and tertiary veins prominent, bright yellowish; lamina palmately 3 or 5 lobed, 

rarely 7; median lobes hastate, ca 9.0 cm long, ca 1.5 cm wide, with 2 prominent lobules 

near the base, apex acute, base of lobes ca 1.0 cm wide, width between base of sinus and 

petiole-lamina junction ca 1.0 cm; lowest lobes ca 1/3 as long as median lobes. 

Inflorescence monoecious, terminal, a cluster of long subspicate racemes arising from one 

common base, ca 15.0 cm long; peduncles and pedicels purplish tinged, sparsely 

pubescent; bracteoles and bractlets setaceous, less than 0.5 cm long, less than 0.1 cm 

wide, pubescent, margins entire. Pistillate flowers non vidi. Staminate buds ovoid-ellipsoid; 

flowers medium sized; tepal 0.9 cm long, exterior and interior surfaces pubescent, 

cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules and seeds 

non vidi. Fig 80A, B. 

TYPE. Hassler 10897: Paraguay: Sierra de Amambay (syntype, G). 

DISTRIBUTION. (Fig 79D). Paraguay. PARAGUAY. Province Unknown/Uncertain: Hassler 

2113 (F, NY-2, P, W) Paraguaria Centralis, 1897. 

Doubtfully assigned to this species. Brasil. Goias: Gardner 3444 (K) upland Campo Duro, Oct 

1839. 

11. Manihot sect Glazioviannae Pax emend Rogers & Appan 

Manihot sect Grandibracteatae Pax subsect Glabrescentes Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 27. 1910 pro parte. 

Manibot sect Parvibracteatae Pax subsect Graciles Pax in Engler, Pflanzenreich IV. 147(Heft 44): 


Manihot sect Heterophyllae Pax subsect Carthaginenses Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 80. 1910 pro parte. 


Plants caulescent; tall shrubs to trees; leaves widely spaced on stem; petiolate, petiole 

attachment basal or peltate; lamina coriaceous to membranaceous, deeply lobed, lobes


, , ..1.. 

m " 

F ~~.. 1~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 

C is_ _A? 

FIG 79. Manibot mirabilis. A, distribution; B, leaf (Hassler 10711, P);C, inflorescence (Hassler 

10711, BM). M. variifolia. D, distribution. 

-I4 

f

Systematic 


A rB 

FIG 80. Manihot variifolia. A, leaf (Hassler 10897, G); B, inflorescence (Hassler 10897, G). 

variously shaped but not linear. Inflorescence a monoecious panicle or raceme; bracts and 

bracteoles setaceous to foliaceous, margins dentate or entire. 

TYPE. Manihot glaziovii Muell.-Arg. 


Key to the Species of Sect Glaziovianae 



3. Petiole attachment peltate. 

4. Inflorescence usually more than 10.0 cm long, many flowered; leaves usually 3 lobed; 

median lobes obovate, usually more than 3.0 cm wide; lobes usually overlapping at 

the sinus; basal lobes recurved. 66. M. glaziovii. 

4. Inflorescence less than 10.0 cm long, few flowered; leaves 5 lobed; median lobes 

oblong, usually less than 3.0 cm wide; lobes separated at sinus; basal lobes straight. 

67. M. pseudoglaziovii. 

3. Petiole attachment basal. 

5. Inflorescence totally glabrous (with the rare exception of the interior surface of 

tepal). 

68. M. epruinosa. 

5. Peduncles, pedicels, bracteoles, bractlets and filaments pubescent. 69. M. brachyandra. 

2. Bracts and bracteoles foliaceous, more than 0.5 cm wide. 70. M. maracasensis. 

1. Inflorescence a raceme. 

6. Staminate buds ovoid-ellipsoid; leaves usually 3 lobed; lobes never pandurate. 71. M. catingae. 

6. Staminate buds conical; leaves 5 lobed; lobes frequently pandurate. 72. M. dichotoma. 

66. Manihot glaziovii Mueller von Argau in Martius, Fl. Bras. 11(2): 446. 1874; Pax in 

Engler & Prantl, Pflzfam. 3(5): 79. 1890; Peckolt, Ber. Deutsch. Pharm. Ges. 16: 29. 

1906; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 89. 1910; Tobler, Ber. 

Deutsch. Bot. Ges. 38: 159. 1920. 

Tall shrubs to trees, to 10.0 m tall, trunks at base 20.0-30.0 cm in diam, with 

copious latex. Young stems and mature stems glabrous, the former often with a light

FI 8 

G ............................ , 0\ 

~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

i 

`? i....a 

I~ ~ ~ ~ ~ 

j_~~~~~~~~ 

~ ~~~~~ 

:r~~~~~~~~~~~~~~~~~~ 

FI 8. anbt Gorapia rne ec Gazoiaae 

O 

FIG 81. Manihot sect Glaziovianae. Geographical range.


bluish bloom. Leaves alternate; stipules caducous; petioles usually ca 20.0 cm long, 

occasionally as long as 45.0 cm, terete, glabrous, petiole attachment to lamina widely 

peltate, width between basal edge of lamina and petiole-midrib junction 1.0-3.0 cm; 

lamina membranaceous to coriaceous, abaxial surface wax pattern reticulate, abaxial surface 

with a bluish white bloom; venation camptodromous, veins glabrous; palmately 3 lobed, 

rarely 5; median lobes obovate, entire, never pandurate, ca 15.0 cm long, ca 7.0 cm wide, 

sometimes as long as 25.0 cm and as wide as 10.0 cm, apex obtuse to broadly acute, base 

of lobes 1.0-2.0 cm wide, width between base of sinus and petiole-midrib junction 

1.0-3.0 cm, lamina at sinus usually overlapping; lowest lobes prominently nonsymmetric 

and curved up. Inflorescence a monoecious, terminal, large, profusely branched, many 

flowered panicle, ca 30.0 cm long, all parts glabrous and with a light bluish bloom; 

bracteoles setaceous, ca 0.5 cm long, less than 0.25 cm wide, margins dentate; bracteoles 

setaceous, margins dentate. Pistillate flowers restricted to the base of the upper 2/3 of the 

inflorescence; pedicels ca 2.0 cm long; flowers large; tepal 1.5 cm long, greenish yellow 

with purplish tinge, cleft to base into 5 strap shaped lobes; disc prominent; ovary 

subglobose; the trifid stigma well lobed and lobulate. Staminate buds ovoid-ellipsoid to 

slightly conical; flowers large, tepal 1.5 cm long, greenish yellow with purplish 

pigmentation, cleft 1/3 way down into 5 lobes; disc prominent; stamens 10, in 2 whorls 

of 5 each. Capsules ca 2.0 cm long, surface without wings, apex rounded, dehiscence 

septicidal. Seeds 1.5 cm long, rounded, caruncle trapeziform. Fig 82B, C, D. 

TYPE. Glaziou 1022: Brasil, Rio de Janeiro: near Rio de Janeiro, cultivated 

(syntypes, photo at F, G). 

DISTRIBUTION. (Fig 82A). Native range. Brasil, states of Ceara, Paraiba, Pernambuco, 

and Bahia. Alt to ca 750 m. Cultivated in the following countries: USA, state of 

Hawaii; Mexico; Guatemala, Honduras; El Salvador; Costa Rica; West Indies; Venezuela; 

Trinidad and Tobago; Brasil; Bolivia; Belgian Congo (presently named Zaire); Uganda; 

Tanganyika; Ceylon; Laos; Malaysia; India; Philippines; Indonesia and Samoa. BRASIL. 

Ceara: Cruz 117 (SP) Belem do Machado, Dec 1965; Curran 32 (F, US) Ceara, 13 Apr 1918; Cutler 

8109 (MO, US) Pirapora near Maranguape, 24 Apr 1944; Cutler 8205 (MO, US) near Belem do 

Machado, 12 Aug 1944; Cutler 8293 (MO, US) 2 km W of Maranguape, 9 Mar 1945;Dusen 6488 (L), 

28 Apr 1899; Eugenio 782 (PUL, RB) Sitio B. Inacio de Azevedo, Serra de Baturite, 29 May 1939; 

Lofgren 190 (S), 10 Mar 1910; Lofgren 191 (S), 10 Mar 1910; Lofgren 225 (S), 15 Mar 1910; Rogers 

411 (G, NY-2, US) Sof Crato near Barbalha, 5 Mar 1961; Ule 9058 (L, NY, UC, US) Serra de 

Batirrite, Oct 1910. Paraiba: Coelho 972 (NY-2, U, UC, US) Alagoa Grande, 13 Jun 1953. 

Pernambuco: Pickel 1255 (MICH, US) Tapera, 14 Feb 1934; Rogers 386 (G, NY, US, W) along road 

N of Lagoa dos Gatos, circa 80 miles SW of Recife, 27 Feb 1961. Bahia: Luetzelburg 15400 (NY) 29 

Aug 1911. 

Collections representing introductions, growing outside the native range. 

USA. Hawaii: Fosberg 9907 (F, NY, UC) Hana Road, between Nahiku and Kaeleku, Maui, 20 

Aug 1933; Topping 9096 (MO, NY) between Hana and Hailsu, Maui. MEXICO. Jalisco: Palmer s n 

(MICH) Camela near Coast, 1943. GUATEMALA. Alta Verapaz: Tuerckheim 8752 (F, US) 

Cubilquitz, May 1904. Santa Rosa: Standley 60407 (F) near Los Verdes, 20 Dec 1938. HONDURAS. 

Francisco Morazan: Standley 28687 (F) La Leona, Tegucigalpa, 6 Sep 1951. EL SALVADOR. La 

Libertad: Carlson 215 (F-2, UC) Finca Germania near Comasagua, on Hillsides through a new Cafeta, 

22 Jan 1946. Santa Ana: Alien 7008 (F, NY) Area S of Santa Ana. San Salvador: Calderon 1815 (F), 

1923; Calderon 1817 (S), 1923. Province Unknown/Uncertain: Calderon 1499 (MO, NY) San 

Salvador, Feb 1923; Calderon 1780 (NY) Finca San Nicolas, 1923. COSTA RICA. Province 

Unknown/Uncertain: Pittier s n (NA). WEST INDIES. Cuba: Ekman 16905 (NY, S) Guayabal, 16 Jul 

1923; Jack 4651 (NY) Soledad, Cienfuegos, 9 Sep 1927. Haiti: Cook s n (NA) Saint Michel, 25 Aug 

1924; Cook s n (NA) Chapman Field, Apr 1931; Cook s n (NA-2) Bayeux, Jul 1927; Ekman 2691 (S) 

Bayeux, Department Du Nord, 29 Nov 1924. St. Thomas: Nelthropp 5 (NY), Dec 1930. Guadeloupe: 

Duss 4196 (NY), Dec 1899; Martinique: Duss 170 (NY) St. Pierre, 1882; Duss 4059 (NY) Saint Pierre, 

1899. VENEZUELA. Falcon: Tamayo 910 (US, VEN-2) Puebla Nuevo, Paraguana, Jan 1939. Aragua: 

Steyermark & Farinas 91440 (NY) in Valley 0.5 Km E of Tiara, 4 Nov 1962. Distrito Federal: 

Delgado 257 (F, US, VEN) La Quebradita, Caracas, 17 Jul 1939; Tamayo 831 (F, US, VEN) En 

Hondonades de Las Colinas de Barrancas, Jul 1939. TRINIDAD/TOBAGO. Togago: Broadway s n (F, 

NY, S-2) Highmoor, 11 Oct 1909; Broadway sn (F-2, NY, S) Scarborough, 12 Jun 1913. Trinidad:


0~~~~~~~~~~~~~~~ 

t ~ Ic"Zi~ 

4L 

'"~~~~~~~~~~~~~~~~~' 

pi Y 

iI ?~~~~~~~~~s~~~~i!~~~ 

"~-- t , ' 

_O~~~~~4 O I 

,, crr~~~~~~~~~~~~~~~~~ 

m~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~6 

r\ j? 4. 

'-t ~... ~ ~ ~ 

a!!lD 

;, ii l i nllll! 

i~~~~f 

i r~~~ 

FIG 82.Maibo lzoi.A itiuin(aierneol) ,hbt(oes36 Y;C 

leaf (Gaiu98,K;D nlrsec Rgr 8,N) 

FIG 82. Manihot glaziovii A, distribution (native range only); B, habit (Rogers 386, NY); C, 

leaf (Glaziou 9585, K); D, inflorescence (Rogers 386, NY).


Broadway sn (F, MO, S) Woodbrook, 1 Oct 1929. BRASIL. Para: Archer 8150 (F, US) Belem, 

Instituto Agronomico do Norte, South Woods, 21 Jan 1943; Archer 8151 (NY) Belem, Instituto 

Argonomico do Norte, South Woods, 21 Jan 1943; Huber 8173 (MG-2) Belem, Horto Botanico, 31 

Jan 1907; Nelson 7069 (MG) Turiassi, Nov 1905; Rogers 412 (F, W-2) Belem, Instituto Agronomico 

do Norte, 10 Mar 1961; Silva 391 (S, UC, US) Belem, Instituto Argonomico do Norte, 3 Jun 1946. 

Minas Gerais: Barreto 8122 (F-2) Estacao Experimental, Muncipality Belo Horizonte, 5 Mar 1937; 

Rogers 371 (NY, G) State Agriculture Experimental Station Belo Horizonte, 18 Feb 1961. Espirito 

Santo: Deslandes 189 (SP) Espirito Santo E Paraiba, Mar 1940. Sao Paulo: Lima 7147 (UC) Arvore 

Campo Experimental Institute Campinas, 2 Apr 1943; Mendes 6741 (UC) Campinas, Fazenda Santa 

Elisa, 31 Mar 1942; Rogers 352 (NY-4) Campinas, Grounds of Institute Agron. 7 Feb 1961; Usteri 32 

(F) Campinas; Viegas & Kiehl s n (SP) Fazenda Eduardo Prado, Campinas, 29 Mar 1939. Rio de 

Janeiro: Glaziou 9585 (G, K, S) Quinta da Boa Vista, 1878; Octavio s n (PUL, RB) Jardim Botanico, 

15 Mar 1945; Weddell 150 (P) Environs Rio de Janeiro, 1843. Parana: Dusen s n (NY, S) Paranagua, 4 

Mar 1911; Dusen 10104 (F, MO, S) Paranagua, 14 Aug 1910. BOLIVIA. Cochabamba: Eyerdam 

25317 (F, K, UC) at Quioma Silver Mines, Province Mizque, 3 Apr 1939. BELGIAN CONGO. 

Dacremont 152 (US), 1932. UGANDA. Mearns 2535 (US) Vicinity of Kabula Muliro, 26 Dec 1909. 

TANGANYIKA. Wigg 2288 (MO) Moragoro, 15 Apr 1948. CEYLON. Baker 112 (DS, MO, NY) 

Peradeniya, Royal Botanic Garden, Jan 1908; Horn s n (W) Peradeniya. LAOS. Poilane 2414 (UC) 

Xieng Kowany, Institute Scientifique de Saigon, 23 Nov 1920. INDO-CHINA. Pasquier 1859 (UC) Phu 

Ho, Jan 1925. MALAYSIA. Malaya: Collector Unknown/Uncertain 3005 (UC), 2 Dec 1917; Wycherly 

760 (F, G, NY, US) Number 5 Ridley Close, Kuala Lumpur, Jul 1962; Wycherly 761 (F, G-2, NY-2, 

US) Petaling Jaya, Jul 1962; Wycherly 762 (G, NY, US) Dr. Southorns Tree, Number 393 Road H, 

Ulu Klang, Jul 1962. PHILIPPINES. Cordero & Espiritu 91498 (L) Quezon, Palawan, 5 May 1964; 

Escritor 21457 (US) Bukidnon Sub. Province Mindanao, Jul-Aug 1913; Foxworthy 1619 (US) Lamas 

Forest Reserve, Bataan Province, Luzon, Oct 1906; Foxworthy 1886 (NY, US) Samao Forest Reserve, 

Bataan Province, Luzon, Dec 1906; Franco 22851 (US) Province Bataan, Luzon, Aug 1914; Merrill 

5010 (NY, US) Manila, Sep 1905; Nano 27434 (UC) Laguna Province, Luzon, 3 Feb 1910; Williams 

3055 (NY-2) Santa Cruz Province, Davao, SE Mindanao, 10 Jul 1905. INDONESIA. Koppel 4444 

(MO), Apr 1931. SAMOA. Rechinger 1707 (W), 20 May 1905. COUNTRY UNKNOWN/UNCER- 

TAIN. Drake s n (P). 

LOCAL NAMES. Caucho (Calderon 1780) caucho blanco (Calderon 1499), Ceara 

rubber (Pittier s n), Ceara India rubber (Baker 112), guamaro (Tamayo 910), manicoba 

(Rogers 386), maniva (Silva 391), mpira (Wigg 2288). 

USES. Commercial production of latex, and as an ornamental. In those parts of 

West Africa (i e Zaire) where the habit of eating many different species for vegetable 

greens is common, Manihot glaziovii is more frequently used for this purpose than for the 

extraction of latex (personal observation). Manihot glaziovii has been used in breeding 

work in Africa to provide resistance to mosaic virus disease in M. esculenta with some 

success (Jennings, 1957; Nichols, 1947; Storey and Nichols, 1938). 

67. Manihot pseudoglaziovii Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 

85): 196. 1924. 

Medium sized trees, ca 6.0 m tall. Stems glabrous. Leaves alternate; stipules 

caducous; petioles ca 15.0 cm long, terete, glabrous, petiole attachment to lamina peltate, 

width between basal edge of lamina and petiole-midrib junction ca 1.0 cm; lamina 

coriaceous, abaxial surface wax pattern reticulate, abaxial surface often with a light bluish 

bloom; venation camptodromous, veins glabrous; palmately 5 lobed; median lobes 

oblong, ca 10.0 cm long, ca 2.5 cm wide, apex acute, base of lobes ca 1.0 cm long, width 

between base of sinus and petiole-lamina junction 1.0-1.5 cm, lamina at sinus not 

overlapping; lowest lobes more or less similar to median lobes in size, slightly 

nonsymmetric; more or less straight, not prominently curved up. Inflorescence a 

monoecious, terminal, sparsely branched panicle, ca 8.0 cm long, all parts glabrous; 

bracteoles and bractlets setaceous, less than 0.5 cm long, less than 0.2 cm wide, margins 

dentate. Pistillate flowers restricted to the base of the upper 2/3 of the inflorescence, 

pedicels ca 2.0 cm long, occasionally as long as 3.0 cm; tepal 1.2 cm long, cleft to base


into 5 lobes; ovary subglobose. Staminate buds ovoid-ellipsoid, flowers medium sized, 

tepal 1.1 cm long, cleft 1/3 way down into 5 lobes, stamens 10, in 2 whorls of 5 each. 

Capsules ca 2.0 cm long, surface without wings, apex rounded, dehiscence septicidal. 

Seeds 1.5 cm long, rounded, caruncle small, not prominently developed. Fig 83B, C. 

TYPE. Luetzelburg 12888: Brasil, Rio Grande do Norte, Serra Parelhas, Mar 1920 

(syntype, M); Luetzelburg 12430: Brasil, Paraiba: Curraes Novos, Mar 1920 (syntype, M); 

Luetzelburg 12420 n v; Luetzelburg 12422 n v; Luetzelburg 12426 n v; Luetzelburg 

12429: Brasil, Paraiba: Gargalheira, Mar 1920 (syntypes, M, NY); Luetzelburg 12432 n v; 

Luetzelburg 12463: Brasil, Paraiba: Serra Borborema, Apr 1920 (syntype, M); Luetzelburg 

12414 n v; Luetzelburg 12415 n v; Luetzelburg 12421: Brasil, Paraiba: Acary, Mar 

1920 (syntypes, F-2, M); Luetzelburg 12431 n v; Luetzelburg 12479 n v. 

DISTRIBUTION. (Fig 83A). Brasil, states of Ceara, Rio Grande Norte, Paraiba. Alt to 

750 m. BRASIL. Ceara: Cutler 8113 (MO, NY, US) near Itapege, formerly Sao Francisco 24 Jun 

1944; Luetzelburg 12429 (M, NY) Gargalheira, Mar 1920. Province Unknown/Uncertain: Allemao 

1394 (P). 

CULTIVATED COLLECTION; BRASIL. Sao Paulo: Viegas s n (SP), cult. in Campo Experimental, 

Instituto, Campinas, 10 Dec 1941. 

USE. Minor latex supply (Cutler 8113). 

68. Manihot epruinosa Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 

196. 1924. 

Manihot floribunda Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 195. 1924. 

Type. Luetzelburg 12253 (syntypes, F, M). 

Tall shrubs to low trees, to 10.0 m tall. Young stems glabrous. Mature stems 

glabrous, grayish externally, bark smooth, brilliant yellow-green internally, with large 

pith. Roots tuberous, epidermis smooth, medium brown (5 R 3/13), subepidermis vivid 

red, cortex light yellowish pink (2.5 YR 9/3). Leaves alternate; stipules semifoliaceous, ca 

1.0 cm long, ca 0.3 cm wide, margins dentate, glabrous; petioles ca 15.0 cm long, terete, 

glabrous, greenish yellow with purplish pigmentation, petiole attachment to lamina basal, 

nonpeltate, rarely peltate; lamina coriaceous, abaxial surface wax pattern reticulate; 

venation camptodromous, veins glabrous; lamina palmately 5-7 lobed, rarely 3, lamina at 

sinus not overlapping; median lobes more or less elliptic to obovate, ca 10.0 cm long, ca 

3.0 cm wide, occasionally larger, apex obtuse, occasionally broadly acute, base of lobes 

1.0-1.5 cm wide, width between base of sinus and petiole-midrib junction ca 1.5 cm; 

lowest lobes slightly smaller than median lobes, slightly nonsymmetric. Inflorescence a 

monoecious, terminal panicle, with wide variation in size, small, less than 4.0 cm, sparsely 

branched, few flowered, to as long as 30.0 cm, profusely branched, and many flowered; 

all parts glabrous; bracteoles and bractlets setaceous, less than 0.5 cm long, less than 0.2 

cm wide, margins dentate. Pistillate flowers restricted to the base of the upper 2/3 of the 

inflorescence, tepal 1.1 cm long, yellowish green with considerable dark reddish (2.5 R 

3/7) pigmentation, sometimes almost blackish, cleft to base into 5 lobes, ovary 

subglobose. Staminate buds ovoid-ellipsoid; flowers medium sized; tepal 1.0 cm long, 

color same as pistillate tepal, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 

5 each. Capsules medium sized, ca 1.25 cm long, dark purplish red in color, surface 

perceptibly winged, apex slightly pointed. Seeds 0.9 cm long, rounded, caruncle small. 

Fig 84A, B, C. 

TYPE. Luetzelburg 12417: Brasil, Paraiba: Acary, Mar 1920 (syntypes, photo at F, 

M); Luetzelburg 12433: Brasil, Paraiba: Gargalheira, Mar 1920 (syntype, M); Luetzelburg 

15005: Brasil, Bahia: Serra de Sincora, Feb 1914 (syntype, M); Luetzelburg 15009 n v; 

Luetzelburg 15013: Brasil, Bahia: Serra das Almas, Mar 1914 (syntypes, M, NY). 

DISTRIBUTION. (Fig 83D). Brasil, states of Piaui, Ceara, Paraiba, Pernambuco and 

Bahia. In the lower altitudinal regions of the caatinga (dry scrub forest) with low rainfall.


. 

..' 

l^^feHB' ...i:? 

. ""' 

_HBB^v -:^MD" rv^- 

FIG 83. Manihot pseudoglaziovii. A, distribution; B, leaf (Luetzelburg 12421, M); C. inflores- 

cence (Luetzelburg 12429, M). M. epruinosa. D, distribution. 

_


a I. 

cence 

!q l 

t _X 

_1l~~~~~~~~~~~~~~~~~~~~~~~~il! !, it :. i , i ll 

_ ' 

.M.b. D u 

~~3 

Illllllli,, tll lllCflitfllt.. [l J 

_~~ ~~~~~~~~~ 

. 

S__ ~~~~~~~~~~~~~~~~~~~~~~~_ 

i~ _ 

0 

_ ~~~~~~~~~~~~~~~~~~~~~~~~ 

FIG 84. Manihot epruinosa. A, habit (Rogers 395, N?); B, leaf (Rogers 395, NY); C, inflores- 

cence (Rogers 395, N?). M. bracbyandra. 

D, distribution.


BRASIL. Piaui: Luetzelburg 15003 (M) Serra do Piauhy, Feb 1914. Ceara: Gardner 2437 (BM, K-2), 

Jul 1839; Rogers 409 (F, G, NY-2, US, W) near Crato, on Hillside on Road from Chapada, 5 Mar 

1961; Rogers 410 (G-2, NY) S of Crato near Barbalha, 5 Mar 1961. Paraiba: Coelho 2084 (NY, UC, 

US) Em Regioes Secas do Nordeste, 20 Apr 1959. Pernambuco: Cutler 8276 (MO) 8 km SE of 

Belmonte (New Name Manicobal) on Sitio Baixa da Extrema, 21 Jan 1945; Cutler 8277 (MO, US) 8 

km SE of Belmonte (Manicobal) on Sitio Baixa da Extrema, 21 Jan 1945; Cutler 8280 (MO, US) near 

Serra Talhada (formerly Villa Bella) 22 Jan 1945; Rogers 381 (G-2, NY, US) Gloria do Goita, 

Fazenda S Antonia, 25 Feb 1961; Rogers 385 (F, G, NY, US) Gloria do Goita, Fazenda S Antonia, 25 

Feb 1961; Rogers 394 (COLO, F, G, NY-2, W) near Caruaru, 2 Mar 1961;Rogers 395 (COLO, F, G, 

K, NY, W) near Pesqueira, 2 Mar 1961; Rogers 399 (F, G, NY-2, US) near San Jose do Belmonte 

(Manicobal), 3 Mar 1961; Rogers 403 (F, G, NY, US) Araripina Agricultural Experiment Station, 4 

Mar 1961; Sarmento s n (COLO, F, G, K, MO, NY, US, W) 2 miles W of Caruaru Aug 1961. Bahia: 

Labouriau 937 (RB) Caatinga, 4 Nov 1952; Luetzelburg 12253 (F, M) Serra Chuque. Sao Paulo: Lima 

s n (SP) Campo Experimental Instituto Campinas, cultivated, 2 Apr 1943; Rogers 353 (NY-4) 

Campinas, Grounds of Institute Agron., 7 Feb 1961; Rogers 354 (G-2) Campinas, Grounds of Institute 

Agron., 7 Feb 1961. 

69. Manihot brachyandra Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 

196. 1924. 

Plant growth habit unknown. Young stems pubescent; mature stems non vidi. Leaves 

alternate; stipules caducous; petioles 14.0 cm long (leaf description based on the only 

mature leaf of the type specimen), terete, pubescent, petiole attachment to lamina basal, 

nonpeltate; lamina coriaceous, abaxial surface wax pattern reticulate; venation camptodromous, 

veins pubescent; lamina palmately 7 lobed, lamina at sinus not overlapping; median 

lobes obovate, 8.0 cm long, 3.5 cm wide, apex obtuse, base of lobes 1.0 cm wide, width 

between base of sinus and petiole-midrib junction 1.0 cm; lowest lobes slightly smaller 

than median lobes, slightly nonsymmetric. Inflorescence a monoecious, terminal, panicle, 

4.0 cm long, peduncles and pedicels pubescent; bracteoles and bractlets setaceous, less 

than 0.5 cm long, less than 0.2 cm wide, pubescent. Pistillate tepal 0.7 cm long, cleft to 

base into 5 lobes, pubescent, ovary subglobose, pubescent. Staminate buds globose to 

ovoid-ellipsoid, tepal 0.6 cm long, cleft 1/3 way down into 5 lobes, pubescent; stamens 

10, in 2 whorls of 5 each, filaments pubescent. Capsules and seeds non vidi. Fig 85A. 

TYPE. Luetzelburg 12252 n v, ex parte (fide Pax): Brasil, Bahia: Rio de Contas, 

Itubira, Jul 1913. 

DISTRIBUTION. (Fig 84D). BRASIL. Bahia: Luetzelburg 12252A (photo at F, M) Rio de 

Contas, Itubira, Jul 1913. 

Pax stated that only a part of Luetzelburg 12252 was the type of his species. He did 

not state what the remaining portion of the specimen was referred to. We have examined 

only Luetzelburg 12252A, a fragmentary specimen, which fits the description of M. 

brachyandra, but the specimen may be the part of the specimen excluded by Pax. 

70. Manihot maracasensis Ule, Bot. Jahrb. Syst. 62: 221. 1908; Pax in Engler, 


Manihot maracasensis Ule var vestita Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 

85): 194. 1924. Type. Luetzelburg 12251 (syntype, M). 

Trees, 3.0-8.0 m tall. Stems pubescent. Leaves alternate; stipules caducous; petioles 

ca 6.0 cm long, terete, pubescent, petiole attachment to lamina basal, nonpeltate; lamina 

slightly coriaceous, abaxial surface wax pattern reticulate; venation camptodromous, 

veins on the abaxial lamina surface tomentose, on the adaxial surface pubescent; lamina 

palmately 3 lobed; median lobes elliptic to obovate, ca 5.0 cm long, ca 2.25 cm wide, 

apex acute, base of lobes constricted, less than 0.3 cm wide, width between base of sinus 

and petiole-lamina junction less than 0.3 cm; lowest lobes slightly smaller than median


p|- 

p 

I. S -. 

.r 

. 

r__ 

C~ D 

.i 

FIG 85. Manibot brachyandra. A, leaf (Luetzelburg 12252-A, M). M. maracasensis. B, distribu- 

tion; C, leaf (Luetzelburg 12251, M); D, inflorescence (Ule 7003, G). 

tion; C, leaf (Luetzelburg 12251, M); D, inflorescence (Ule 7003, G). 

co 

- 

'


lobes, slightly nonsymmetric. Inflorescence a monoecious, terminal, panicle, ca 8.0 cm 

long; peduncles and pedicels tomentose; bracteoles foliaceous, ca 1.0 cm long, ca 0.6 cm 

wide, tomentose, margins entire; bracteoles foliaceous, tomentose. Pistillate flowers 

restricted to the base of the upper 2/3 of the inflorescence; pedicels 1.0 cm long, tepal 

1.0 cm long, cleft to base into 5 lobes, exterior and interior surfaces of lobes tomentose; 

ovary subglobose, tomentose. Staminate buds ovoid-ellipsoid, tepal 1.0 cm long, 

tomentose, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules 

and seeds non vidi. Fig 85C, D. 

TYPE. Ule 7003: Brasil, Bahia: bei Maracas, Sep 1906 (syntypes, photo at F, G, L, 

photo at NY). 

DISTRIBUTION. (Fig 85B). Brasil, state of Bahia. BRASIL. Bahia: Luetzelburg 12251 

(M) Rio de Contas, Itubira. 

LOCAL NAME. Mani;oba (Luetzelburg 12251). 

71. Manihot catingae Ule, Bot. Jahrb. Syst. 62: 221. 1908; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 78. 1910. 

Trees, 2.0-5.0 m tall. Stems glabrous. Leaves alternate; stipules caducous, setaceous, 

glabrous, less than 1.0 cm long, margins dentate; petioles ca 5.0 cm long, terete, glabrous, 

petiole attachment to lamina basal, nonpeltate; lamina slightly coriaceous, abaxial surface 

wax pattern reticulate; venation camptodromous, veins glabrous; lamina palmately 3 

lobed; median lobes elliptic, ca 7.0 cm long, ca 2.0 cm wide, apex acute, base of lobes 

narrow, ca 0.75 cm wide, width between base of sinus and petiole-lamina junction ca 0.75 

cm; lowest lobes slightly smaller than median lobes, slightly nonsymmetric. Inflorescence 

a monoecious, terminal, short, few flowered raceme, ca 4.0 cm long, all parts except the 

interior surface of tepals glabrous; bracteoles and bractlets setaceous, less than 0.5 cm 

long, less than 0.2 cm wide, margins dentate. Pistillate flowers restricted to the base of 

the upper 2/3 of the inflorescence; pedicels 1.5 cm long; tepal 1.0 cm long, interior 

surface sparsely pubescent, cleft to base into 5 lobes; ovary subglobose. Staminate buds 

ovoid-ellipsoid; flowers medium sized, tepal 1.0 cm long, interior surface sparsely 

pubescent, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules 

1.25 cm long, subglobose, slightly elongated, surface smooth, apex rounded. Seeds non 


TYPE. Ule 7142: Brasil, Bahia: bei Remanso, Dec 1906 (syntypes, F, G, K, L, photo 

at NY). 

DISTRIBUTION. (Fig 86A). Brasil, state of Bahia. BRASIL. Bahia: Luetzelburg 365 (F, 

NY) Catinga Gebiet, Central Bahia, 1913. 

LOCAL NAME. Manicoba brava (Luetzelburg 365). 

72. Manihot dichotoma Ule, Notizbl. Bot. Gart. Berlin 5(41): 2. 1907; No. 41A: 16, 19, 

f. 1 A, B. 1908: Hook. Icon. pl. 29 t. 2876, 2877. 1909; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 83. 1910. 

Manibot dichotoma Ule var parvifolia Ule, Notizbl. Bot. Gart. Berlin 5(41a): 20. 1908; Pax in 

Engler, Pflanzenreich IV. 147(Heft 44): 83. 1910. Type. Ule 7362 (syntypes, K, L). 

Manihot dichotoma Ule var genuina Pax in Engler, Pflanzenreich IV. 147(Heft 44): 83. 1910. 

Trees 3.0-12.0 m tall. Stems glabrous. Leaves alternate; stipules caducous, setaceous, 

less than 1.0 cm long, margins dentate, glabrous; petioles ca 8.0 cm long, terete, glabrous, 

petiole attachment to lamina basal, nonpeltate; lamina slightly coriaceous to membranaceous, 

abaxial surface wax pattern reticulate; venation camptodromous, veins glabrous; 

lamina palmately 5 lobed, 3 major and 2 smaller; median lobes obovate, frequently


r, 

o _ 

t 

i_ ; t 

io~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ri 

`r~~~~~~~~~~ 

?? 

r 

?- 

t?270~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

CF~~~~~~~~~~~~~ 

..... L X. 

FIG 86. Manihot catingae. A, distribution; B, leaf (Ule 7142, G); C, inflorescence (Ule 7142, 

G). M. dichotoma. D, distribution.


pandurate, ca 8.0 cm long, ca 3.5 wide, apex acute, base of lobes narrow, ca 1.0 cm wide, 

width between base of sinus and petiole-midrib junction ca 1.0 cm; lowest lobes ca 1/2 as 

long as median lobes, nonsymmetric. Inflorescence a monoecious, terminal, short raceme, 

ca 4.0 cm long, all parts except the interior surface of tepals glabrous; bracteoles and 

bractlets setaceous, less than 0.5 cm long, less than 0.2 cm wide, glabrous, margins 

dentate. Pistillate flowers restricted to the base of the inflorescence, tepal 1.7 cm long, 

yellowish green externally, purplish internally, cleft to base into 5 strap-shaped lobes, 

ovary elongated with prominent wavy wings. Staminate buds conical, flowers large, tepal 

1.7 cm long, color same as pistillate tepal, cleft 1/3 way down into 5 lobes; stamens 10, in 

2 whorls of 5 each. Capsules and seeds non vidi. Fig 87A, B. 

TYPE. Ule 7045: Brasil, Bahia: Bei Calderao, Oct 1906 (syntypes, G, K, L, MG). 

DISTRIBUTION. (Fig 86D). Brasil, states of Pernambuco and Bahia. Plant introduced 

to several other tropical countries, but specimens of these either nonexistant or too 

fragmentary for positive identification. (See, for example, Cook s n (NA) and Ekman 

2690 (S), both from the West Indies, Haiti, included in this monograph as dubious 

specimens). Manihot dichotoma is also growing at the United States Department of 

Agriculture's Introduction Station at Miami, Florida. BRASIL. Pernambuco: Pickel 632 (SP). 

Bahia: Cnrz 109 (SP) Jequie, Cultivada na Fazenda Experimental Theodureto de Camargo, 7 Oct 

1965; Cruz 110 (SP) Jequie, Cultivada na Fazenda Experimental Theodureto de Camargo, 7 Oct 1965; 

Ule 7362 (K, L) Catinga Bei Tambury, Oct 1906. Province Unknown/Uncertain, Hooker Icon (W). 

12. Manihot sect Peruvianae Rogers & Appan, sect nov 

Manihot sect Grandibracteatae Pax subsect Peruvianae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 28. 1910. 

Manihot sect Parvibracteatae Pax subsect Elatae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 55. 1910 pro parte. 

FG8.1 MnodctmA,e1i70,)BinoseeVe05G 

~* N 

A - 

:!!~? 

-i--. 

FIG 87. Manibot dichotoma. A, leaf (Ule 7045, L); B, inflorescence (Ule 7045, G).


Manihot sect Parvibracteatae Pax subsect Guaraniticae Pax in Engler, Pflanzenreich IV. 

147(Heft 55): 74. 1910 pro parte. 

Plantae caulinae, lignosae, vita-simili frutices, super ceteris vegatatio scandens. Folio 

petiolatae, penitae lobatae, Inflorescentia monoecia panicula aut racema. 

Plants caulescent; vine-like shrubs clambering over other vegetation; leaves widely 

spaced on stem; petiolate, petiole attachment basal; lamina membranaceous, deeply 

lobed, lobes variously shaped but not linear. Inflorescence a monoecious panicle or a 

raceme; bracts and bracteoles setaceous or foliaceous, margins usually entire. 

TYPE. Manibot peruviana Muell.-Arg. 


Key to the Species of Sect Peruvianae 

1. Inflorescence a panicle; staminate flowers campanulate; staminate buds ovoid-ellipsoid; veins 

on lower surface of leaf glabrous or sparsely pubescent. 


3. Abaxial lamina surface wax pattern verrucate; all parts except interior surface of tepals 

usually glabrous, leaves 3 lobed. 73. M. brachyloba. 

3. Abaxial lamina surface wax pattern reticulate; petioles, midribs, peduncles, bracteoles, 

etc. sparsely pubescent; leaves frequenty 5 lobed, sometimes 3 lobed. 74. M. leptophylla. 

2. Bracts and bracteoles foliaceous, more than 0.5 cm wide. 75. M. quinquepartita. 

1. Inflorescence a raceme; staminate flowers gibbous; staminate buds bifusiform; veins on 

lower surface of leaf hirsute. 76. M. peruviana. 

73. Manihot brachyloba Mueller von Argau in Martius, Fl. Bras. 11(2) 451. 1874; Pax in 


Manihot rusbyi Britton, Bull. Torrey Club 28: 302. 1901; Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 98. 1910. Type. Rusby 888 (syntypes, F, NY, US). 

Manihot amazonica Ule, Verh. Bot. Vereins Prov. Brandenburg 50(1): 83. 1908 (1909); Pax in 

Engler, Pflanzenreich IV. 147(Heft 44): 57. 1910. Type. Ule 5264 (syntypes, F, G, K, L, 

MG, NY). 

Subscandent, woody, vine-like shrubs, clambering over other vegetation; branches to 

7.0 m long, slender, virgate; copious latex, odorless. Roots long, with no enlarged storage 

roots. Stems glabrous, glaucous. Leaves alternate; stipules caducous; petioles ca 10.0 cm 

long, terete, glabrous, yellowish green with some purple pigmentation near the lamina 

end, often with a light bloom, petiole attachment to lamina basal, nonpeltate; lamina 

membranaceous, abaxial surface wax pattern verrucate, abaxial surface often glaucous; 

venation camptodromous, veins yellowish green, never purplish tinged, glabrous; lamina 

palmately 3 lobed, never more than 3 lobed, leaves associated with the inflorescence 

frequently nonlobed; median lobes elliptic to lanceolate, ca 9.0 cm long, ca 3.5 cm wide, 

apex accuminate, base of lobes narrowly constricted, less than 0.3 cm wide, width 

between base of sinus and petiole-lamina junction less than 0.3 cm; lowest lobes more or 

less similar to median lobes in length, slightly nonsymmetric. Inflorescence a monoecious, 

terminal panicle, varying considerably in size, 10.0-30.0 cm long, all parts except the 

interior surface of tepals glabrous, often with a light bloom; bracteoles and bractlets 

setaceous, less than 0.5 cm long, less than 0.2 cm wide, margins entire. Pistillate flowers 

restricted to the base of the upper 2/3 of the inflorescence, tepal 1.2 cm long, yellowish 

green with considerable purple pigmentation, cleft to base into 5 lobes, ovary subglobose. 

Staminate buds ovoid-ellipsoid; flowers medium sized; tepal 1.2 cm long, color same as 

pistillate tepal, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. 

Capsules 1.5 cm long, subglobose, surface smooth, apex rounded, abruptly ending in a 

short pointed projection, dehiscence septicidal. Seeds 1.1 cm long, oblong, with well 

developed rib-like extensions along the lateral edges, dark brownish gray with few 

mottlings, caruncle very small and inconspicuous. Fig 89B, C.

SOUTH AMERICA 

.ONE COLLECTION COFLROlMCTI A flW 

fSOM 

IlB \fI_CA ''MIPICA 

FIG 88. Manibot sect Pevianae. Geographical range.


.' '.: :. :' "' T - 

-.. 

CI? / . 

u 19 A. / . 

""~ 

-'-'"-'"~"'~~~~~~~~~~~~~~~~~; 

"'~ ,,~i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~\ 

(Haugbt (Haught 1919, A). M. M. leptopbylla. D, distribution. 

distribution 

' 

...


TYPE. Martius s n: Brasil, Para: near Para: (syntypes, photo at F, G, M);Sieber s n n 

v. 

DISTRIBUTION. (Fig 89A). Costa Rica, Prov. Sta. Clara; West Indies, in Republica 

Dominicana; Colombia, Depts. Magdalena, Cordoba, Choco, Santander, Cundinamarca, 

and Meta; Venezuela, State of Zulia; Suriname; Guyane Francaise; Peru, Dept. Loreto; 

Brasil, Amapa Territory, states of Amazonas, Para, and in Rondonia Territory. Alt to ca 

500 m. Common in second growth, glades, and in shaded thickets on river banks. COSTA 

RICA. Santa Clara: Smith 6850 (F, GH, K, US) Siquirres, Apr 1896. WEST INDIES. Republica 

Dominicana: Abbott 2304 (NY, US) Vicinity of Laguna, Samana Peninsula, chiefly on the Pilon de 

Azucar, 9 May 1922; Abbott 2796 (GH, US) Loma de Perro, 2 miles S of Guarabo, SE of Jovero, 20 

Nov 1923; Ekman 14993 (F, NY, S) Peninsula de Samana, El Valle, in thickets along the river, 16 May 

1930; Ekman 15591 (S) Cordillera Central, Sabana de La Mar, at Rio Capitan, 7 Jul 1930; Ekman 

15736 (S) Cordillera Central, El Jovero, road to El Liar, 20 Jul 1930; Ekman 15854 (A, S) Peninsula 

Samana, Laguna, at Arroyo Seco, 5 Aug 1930;Marcano & Vega 4852A (F, G-2, NY-3, US, W) Samana 

Peninsula, Seccion de Arroyo Seco, Paraje la Loma de La Cruz; near San Juan River, 20 Dec 1964; 

Miller 1238 (US) Rio San Juan, 21 Mar 1928; Miller 1250 (US) Rio San Juan, 23 Mar 1928. 

COLOMBIA. Magdalena: Smith 365 (BM, F, G-2, GH, MO, P, TEX, UC, US) Santa Marta, Sep. 

Cordoba: Killip & Garcia 33395 (BM, F, US) Cordoba, 17 Feb 1939. Choco: Fernandez 324 (US) 

Bahia de Solano, Subiendo Por Quebrada Seca Hacia La Chorrera, 11 Jun 1950. Santander: Haught 

1664 (US) Vicinity of Puerto Barrio, between Carare and Magdalena Rivers, 29 Apr 1935; Haught 

1919 (A, F, S, UC, US) Vicinity of Barranca Bermeja, Magdalena Valley, between Sogamoso and 

Carare Rivers, 6 Sep 1936; Uribe 3077 (US) Bajo Magdalena, Dec 1957. Cundinamarca: Triana 3614 

(BM) Rio Seco, Jul 1853. Meta: Niceforo, M., Hno. 225 (DS) Villavicencio, 10 Dec 1939;Jaramillo, 

Mesa, Idrobo & Fernandez 413 (US) Intendencia Del Meta, Acacias, Aug 1946; Philipson, Idrobo & 

Fernandez 1463 (US) Rio Guejar, Sierra De La Macarena, 17 Nov 1949. VENEZUELA. Zulia: Bruijn 

1212 (NY) circa 62 km S of Machiques near Mission Los Angeles Del Tukuko, Perija District, 16 Oct 

1966; Steyernark 99980 (NY, VEN) Sierra de Perija, 31 Aug 1967. SURINAME. Pulle 213 (U) 

Voltzberg, 21 Aug 1920. GUYANE FRANCAISE. Sagot 1082 (G, W) La Mana. PERU. Loreto: 

Schunke 291 (A, F, NA, NY, UC, US) Gamitanacocha, Rio Mazan, 20 Feb 1935; Williams 2059 (A) 

Caballo-Cocha on the Amazon River, Aug 1929. BRASIL. Amapa Territory: Egler & Irwin 46075 

(NY) Rio Jari, 31 Jul 1961;Pires, Rodriques & Irvine 51202 (NY-2) Rio Araguari, 25 Sep 1961;Pires, 

Rodriques & Irvine 51499 (NY) Rio Araguari, 4 Oct 1961. Amazonas: Ducke, A. s n (MG, PUL), 

Coary (campo da Frequezia Velha), 14 Dec 1912; Froes 20947 (NY) Sao Paulo de Olivenca, Varsea, 

21 May 1945; Froes 23862 (U) Igarape Jandiatuba, 9 Jan 1949; Krukoff 6311 (BM, F, G, MICH, MO, 

NY, S, U, US) near Tres Casas, Municipality Humayata, Basin of Rio Madiera, 24 Sep 1934; Pires & 

Black 985 (VEN) Benjamin Constant, 24 Oct 1945; Ule 5264 (F, G, K, L, MG, NY) Rio Jurua, Nov 

1900. Para: Archer 7522 (US) Belem, Forest South of the I.A.N., 7 Dec 1942; Archer 8125 (NY, 

US-2) Belem, Instituto Agronomico Do Norte, South Woods, 2 Jan 1943; Archer 8291 (US-2) Belem, 

Instituto Agronomico Do Norte, North Woods, 12 Mar 1943; Ducke 14842 (MG, RB) Rio Cumina 

Mirim, Castanral Das Pedras, Beira De Matta, 27 Sept 1913; Goeldi 2834 (MG) Pedrevia Do Guama, 30 

Jul 1902; Guedes 2463 (MG), 31 Oct 1901; Huber 4694 (MG), 25 Apr 1904;Pires 17 (U) Belem, R. 

Guama, Capeoira, Terreno Alagavel, 8 Oct 1946; Pires & Black 1480 (U) Belem, Estrada Do Utinga 

Para Agua Preta Arbusto, 10 Apr 1947; Rogers 413 (COLO, F, G, K, NY, US) Belem, Instituto 

Agronomico Do Norte, 10 Mar 1961. Rondonia Territory: Baldwin 3145 (A, US) Guajara Mirim, 

Gaupore, 7 Dec 1943. Province Unknown/Uncertain: Ducke 7712 (MG) Baixo Rio Ica, 9 Sep 1906; 

Luetzelburg 20302 (M) Rio Gamopi (Pyapock), Jul 1927; Rusby 888 (F, NY, US) Falls of Madeira, 

Oct 1886. 

LOCAL NAMES. Manioc bicha (Sagot 1082), maniva brava (Froes 20947), maniva de 

veado (Rogers 413), sacharumo (Schunke 291), yuca cimarrona (Ekman 14993), yuca de 

indio (Haught 1919), yuca silvestre (Steyermark 99980). 

This is the only wild species known to us with a distribution in the West Indies 

(Dominica), Central America (Costa Rica) and the South American continent. In both 

Dominica and Costa Rica, Manihot brachyloba has a very limited distribution. There 

seems to be a distinctive geographical range for M. brachyloba (more to the east and 

north in South America) which differentiates this species from M. leptophylla (more to 

the west and south in South America). However, much more exploration is needed to 

precisely define the boundaries. 

The most outstanding unifying character of this species is the abaxial leaf surface


wax pattern, verrucate (Fig 1B). This character most readily distinguishes this species 

from its nearest relative, M. leptophylla, whose wax pattern is reticulate (Fig 1A). 

On the herbarium label of the specimen Ekman 14999 (F), Urban and Ekman list the 

name Manihot domingensis. This name was never published. 

74. Manihot leptophylla Pax in Engler, Pflanzenreich IV. 147(Heft 44): 57. 1910. 

Manihot palmata Muell.-Arg. var ferruginea Muell.-Arg. in DC., Prodr. 15(2): 1063. 1866; in 

Martius, Fl. Bras. 11(2): 458. 1874; Manihot dulcis (J. F. Gmel.) Pax var ferruginea 

(Muell.-Arg.) Pax in Engler, Pflanzenreich IV. 147(Heft 44): 71. 1910. Type. Poeppig 1523 

(syntypes, A, F, W-2): Poeppig 1410 (syntype, W). 

Subscandent, woody, vine-like shrubs, clambering over other vegetation; branches to 

7.0 m long, slender, virgate; copious latex, ordorless. Roots long, not tuberous; epidermis 

smooth, moderate brown (5 YR 3/3); subepidermis moderate orange-yellow (7.5 YR 8/8); 

cortex cream colored. Young stems pubescent; mature stems pubescent, moderate brown 

(5 YR 3/3) externally, cream colored internally. Young foliage at apex moderate 

yellow-green (2.5 GY 5/5); leaves alternate; stipules caducous, setaceous, pubescent; 

petioles ca 10.0 cm long, terete, pubescent, moderate reddish brown (7.5 R 3/6), petiole 


pattern reticulate; venation camptodromous, veins pubescent; lamina palmately 5 lobed, 

occasionally 3 lobed, leaves associated with the inflorescence frequently nonlobed; 

median lobes elliptic to lanceolate, ca 15.0 cm long, ca 4.0 cm wide, apex acuminate, base 

of lobes narrowly constricted, less than 0.3 cm wide, width between base of sinus and 

petiole-lamina junction less than 0.3 cm; lowest lobes more or less similar to median lobes 

in length, slightly nonsymmetric. Inflorescence a monoecious, terminal panicle, varying 

considerably in size, 10.0-30.0 cm long; peduncles and pedicels pubescent; bracteoles and 

bractlets setaceous, less than 0.5 cm long, less than 0.2 cm wide, pubescent, often 

purplish tinged, margins entire. Pistillate flowers restricted to the base of the upper 2/3 of 

the inflorescence, tepal 1.2 cm long, pale orange-yellow (7.5 YR 9/4) externally, dark red 

(5 R 3/7) internally, sparsely pubescent internally and externally, cleft to base into 5 

lobes, ovary subglobose, glabrous. Staminate buds ovoid-ellipsoid; flowers medium sized; 

tepal 1.2 cm long, color same as pistillate tepal, cleft 1/3 way down into 5 lobes; stamens 

10, in 2 whorls of 5 each. Capsules 1.5 cm long, subglobose, surface smooth, apex 

rounded, abruptly ending in a short pointed projection, dehiscence septicidal. Seeds 1.0 

cm long, oblong, dark brownish gray with few mottlings; caruncle very small and 

inconspicious. Fig 90A, B, C, D. 

TYPE. Eggers 15156: Ecuador, Manabi: bei El Recreo, 20 Mar 1897 (syntypes, 

photo at F, K, NY, S, US). 

DISTRIBUTION. (Fig 89D). Ecuador, Provinces Pichincha, Manabi, and Pastaza; Peru, 

Departments Loreto, San Martin, Huanuco, Junin, Cusco and Ayacucho; Brasil, States 

Amazonas, Para, Pernambuco and Acre. Alt to 2000 m. Among second growth low 

vegetation, forest openings, roadsides and close to river banks. ECUADOR. Pichincha: Cazalet 

& Pennington 5189 (NY, UC) 20 km W of Santo Domingo de Los Colorados, 29 Oct 1961. 

Pastaza: Skutch 4429 (F, US) Vicinity of Puyo, Eastern Foot Hills of the Andes, Aug 1939. PERU. 

Loreto: Carneiro & Dole s n (F, G, NY-4, US, W) Upper Innya River, Tributary of Urubamba, Mar 

1960; Ferreyra 4894 (US) Chambira, Yurimaguas, 11 Sep 1948; Huber 1460 (MG) Ucayali, 13 Nov 

1898; Huber 2600 (MG) Rio Ucayali, Jan 1902; Macbride 5074 (F, GH) Pampayacu, Hacienda at 

Mouth of Chinchao Rio, Jul 1923; Rogers 474 (F, G, K, NY-2, US, W) circa 24 km from Tingo Maria, 

on old logging road, 12 Feb 1962; Skutch 5009 (A, NA) Contamana, Rio Ucayali, 19 Oct 1940; 

Williams 5323 (A) Lower Rio Huallaga, Oct-Nov 1929; Woytkowski 34448 (F, MO, UC) Aguaitia, 28 

Aug 1946. San Martin: Klug 2662 (BM, F, MO, NY, US) Pongo de Cainarachi, Rio Cainarachi, 

Tributary of Rio Huallaga, Sept-Oct 1932; Klug 3861 (BM, F, GH, MO, NY, US) Juan Jui, Alto Rio 

Huallaga, Oct 1934. Huanuco: Ferreyra 1619 (GH, NY, US) Tulumayo, circa al puente, entre Tingo 

Maria Y La Divisoria, 26 Feb 1947; Ferreyra 6734 (US) Cayumba, Entre Huanuco Y Tingo Maria, 8 

Feb 1950; Landeman 3371 (K, OXF) Ganso Azul, Rio Pachitea, Oct 1942; Rogers 467 (COLO, F-2, 

G-2, K, M, MICH, MO, NY-3, S, US-2, W-2) Between Tingo Maria and Huanuco, 8 Feb 1962; Rogers


1_ ? , ., .t,* jil,:t./ ,, 

N 

FIG 0. aniotep pbyla.A,abi (Rog ers 47; la 

'' _- 

i.. ..-aD--e 

FIG 90. MFa nis o letphla 

(Roers (Roer 46,NY;D 4 ,i infoesec 

A,C' hai 

ege (Rges47 

Rgr 6,N) 

NY). 

,tepln Rgr 6,N) ,la


468 (F, G, K, NY, US, W) on main road from Tingo Maria to Huanuco, circa 8 km from Tingo Maria, 

8 Feb 1962. Junin: Killip & Smith 23791 (F, NY, US) La Merced, May-Jun 1929;Macbride 5472 (F, 

US) La Merced, in thicket on sandy valley floor, 10-24 Aug 1923. Cusco: Poeppig 1410 (W) Peruvia 

Subandina, Nov 1829; Poeppig 1523 (A, F, W-2) Cuchero, 1830; Vargas 11608 (NY) Pilocopata, 

Province Paucartambo, 30 May 1957. Ayacucho: Killip & Smith 22722 (A, F, NY, US) Aina, between 

Huanta and Rio Apurimac, May 1929; Killip & Smith 22964 (F, NY, US) Rio Apurimac Valley, near 

Kimpitiriki, 10 May 1929. BRASIL. Amazonas: Huber s n (PUL) Monte Verde, 28 Apr 1914; Huber 

4541 (MG, RB) Rio Purus, Monte Verde, 28 Apr 1904; Prance, Pena, Ramos & Videcki 2389 (NY-3) 

Boca do Acre, Rio Purus, Rio Acre, 19 Sep 1966. Para: Froes, R.L. 23548 (P, RB, U) Breu Branco, 

Rio Toncantins, 28 Sep 1948; Huber s n (MG) Belem, Jardim Botanico, Sep 1901; Huber 798 (MG) 

Rio Apim, 19 Jun 1897; Kuhlmann 1928 (PUL, RB) Boim, Tapajos, 7 Apr 1924;Monteiro Da Costa 

234 (F) Tapajos, Aramanahy, 8 Jan 1932; Silva 77 (US) Belem, Instituto Agronomico Do Norte, 2 

Feb 1944. Pernambuco: Leal 34 (PUL, RB) Restinga Do Rio Doce, Olinda, 12 Jan 1948. Acre: Ule 

9546 (L, MG, UC) Seringal San Francisco, Rio Acre, Mar-Apr 1911. Province Unknown/Uncertain: 

Bach 4162 (MG) Xingu, Providencia, 24 Dec 1903. Country unknown/uncertain. Pavon s n (BM). 

LOCAL NAMES. Maniva de veado (Monteiro da Costa 234), yuca del monte 

(Woytkowski 34448), yuca de raton (Eggers 15156), yuquilla (Rogers 467), yushiwa asti 

or spirits manioc (Carneiro & Dole s n). 

USES. According to Carneiro and Dole, this species is sometimes a volunteer in 

gardens of M. esculenta, and the roots are infrequently harvested for food. 

The epithet Manihot leptophylla is the first available, legitimate name for this 

species. We have reviewed the problems of M. palmata under species 28, M. leptopoda. 

The epithet, ferruginea is not available as a species name because Ule (Bot. Jahrb. Syst. 

1.(Beibl. 114): 2. 1914) preempted it for one of his own species with another type. 

Manihot ferruginea Ule = M. caerulescens Pohl emend Rogers & Appan, subsp caerules- 

cens Rogers & Appan. 

Refer to the remarks under the previous species (M. brachyloba) for geographical 

discussions and morphological distinctions. 

75. Manihot quinquepartita Huber ex Rogers & Appan, sp nov 

Frutex subscandens, super vegetationem scandens, ad 6.0 m alta. Caules pubescentes. 

Folia quinque-lobata, lobis-medianis 10.0-15.0 cm longis, venis pubescentibus, basibus 

lobarum valde constrictis. Inflorescentia paniculata monecia, 10.0-15.0 cm longa; 

bracteolis prominente foliaceis, pubescentibus, marginibus plerumque laciniatis. Capsulae 

grandes, circa 2.5 cm longae. Semina grandia, circa 2.0 cm longa. 

Subscandent, woody shrub, vine-like, clambering over other vegetation, to ca 6.0 m 

tall. Roots non vidi. Young stems pubescent; mature stems sparsely pubescent. Leaves 

alternate; stipules setaceous, less than 1.0 cm long, pubescent, caducous; petioles 

10.0-15.0 cm long, terete, sparsely pubescent, petiole attachment to lamina basal, 

nonpeltate; lamina membranaceous to almost coriaceous, abaxial surface wax pattern 

reticulate, often showing discrete irregular sided rings; venation camptodromous, veins 

pubescent; lamina palmately 5 lobed; median lobes elliptic-lanceolate, ca 11.0 cm long, ca 

3.5 cm wide, never pandurate, lamina edges slightly undulate, apex acuminate, base of 

lobes extremely constricted so that the leaves appear compound, lamina at lobe base less 

than 0.2 cm wide, width between base of sinus and petiole-lamina junction less than 0.2 

cm; lowest lobes slightly smaller than median lobes and slightly nonsymmetric. 

Inflorescence a moneocious, terminal panicle, 10.0-15.0 cm long; peduncles and pedicels 

pubescent; bracteoles prominently foliaceous, ca 1.5 cm long, ca 0.9 cm wide, pubescent, 

margins frequently laciniate, sometimes serrate or entire; bractlets foliaceous ca 1.0 cm 

long, ca 0.6 cm wide, margins laciniate, serrate, or entire. Pistillate flowers restricted to 

the base of the upper 2/3 of the inflorescence, pedicels ca 1.0 cm long, tepal 1.2 cm long, 

pubescent internally and externally, cleft to base,into 5 lobes; ovary subglobose, sparsely 

pubescent or glabrous. Staminate buds ovoid-ellipsoid, flowers medium sized, tepal 1.2


cm long, pubescent internally and externally, cleft 1/3 way down into 3 lobes; stamens 

10, in 2 whorls of 5 each, the superior whorl 0.9 cm long, the inferior 0.7 cm long, 

filaments pubescent. Capsules large, ca 2.5 cm long, surface smooth, apex rounded, 

dehiscence septicidal. Seeds large, 2.0 cm long, oblong, with well developed rib-like 

extensions along the lateral edges; caruncle very small and inconspicuous. Fig 91B, C. 

TYPE. Huber 915: Brasil, Para: Rio Capim, 2 Jul 1897 (holotype, MG, photo of 

type, F). 

DISTRIBUTION. (Fig 91A). Brasil, Amapa Territory and the states Para, Maranhao 

and Mato Grosso, on rocky slopes in clearings. BRASIL. Ampa Territory: Mattos & Mattos 

10208 (SP) Macapa, Serra do Navio. Para: Archer 8379 (US) 1 km N of Fazenda Urucurituba, oposite 

Fordlandia, Rio Tapajos, 15 Apr 1943; Cavalcante 139 (MG) Bacia do Trombetas, Jaramacaru, 29 May 

1957; Ducke 10542 (RB) Rio Tapajos, 23 Dec 1919; Ducke 11607 (MG) Parintins Capoeira Velha, 8 

May 1911; Ducke 15873 (MG, RB) Rio Tapajos, 11 Dec 1915;Guedes 215 (U) Belem, I.A.N., 25 Nov 

1949; Huber 3805 (MG), 24 Sep 1903; Pires 1857 (U-2, US) Belem, Provavelmente Trazida do 

R. Acara, 14 Jan 1950; Pires & Silva 4830 (NY) Braganna, Campo Alagado, 9 Apr 1955; Snethlage 

10103 (MG) Rio Jamauchim Tacumare, 1 Dec 1908. Maranhao: Ducke 530 (MG) Anil, Capoeira 

Velha; Froes 11545 (A, F, MO, NY, S, U) Island of Sao Luiz, Feb-Mar 1939. Mato Grosso: Krukoff 

1649 (BM, MICH, MO, NY, P, S, U) Source of Jatuarana River, Machado River Region. Province 

Unknown/Uncertain: Burchell 9942 (K). 

Huber had identified this species and provided the name on the type specimen, but 

did not publish the species. 

This is a well-defined species, whose nearest relative is Manihot brachyloba 

Muell.-Arg. The bracts and bracteoles of M. quinquepartita are foliaceous, while those of 

M. brachyloba are setaceous. The abaxial lamina surface wax pattern of this species is 

reticulate, that for M. brachyloba is verrucate. 

76. Manihot peruviana Mueller von Argau, Linnaea 34: 206. 1865; in DC., Prodr. 15(2): 


147(Heft 44): 29. 1910. 

Manihot beterandra Ule, Verh. Bot. Vereins Prov. Brandenburg 50(1): 84. 1908 (1909) nom 

rejic. (This name is rejected in accordance with article 63 of the International Code of 

Botanical Nomenclature. Manibot heterandra as circumscribed by Ule is based on the same 

type (Spruce 4287) as Manihot peruviana Muell.-Arg.). 

Subscandent, woody shrubs, clambering over other vegetation, ca 2.0 m tall. Young 

stems pubescent; mature stems sparsely pubescent. Leaves alternate; stipules caducous, 

setaceous ca 1.5 cm long, less than 0.2 cm wide, hirsute, margins entire; petioles ca 10.0 

cm long, terete, pubescent, petiole attachment to lamina basal, nonpeltate; lamina 

membranaceous, abaxial surface wax pattern verrucose; venation camptodromous, veins 

hirsute, lamina palmately 3 lobed, occasionally with 2 more smaller lobes; median lobes 

obovate to obovate-elliptic, ca 10.0 cm long, ca 3.5 cm wide, apex acuminate, base of 

lobes narrow, ca 0.75 cm wide, width between base of sinus and petiole-lamina junction 

ca 0.75 cm; lowest lobes slightly smaller than median lobes and slightly nonsymmetric. 

Inflorescence a monoecious, terminal, short raceme, simple or a cluster of 3 or 4 racemes 

arising from one common base, usually less than 7.0 cm long; peduncles and pedicels 

pubescent; bracteoles setaceous to nearly semifoliaceous, ca 1.5 cm long, ca 0.3 cm wide, 

pubescent, margins entire; bractlets setaceous, ca 1.0 cm long, ca 0.1 cm wide. Pistillate 

flowers restricted to the base of the inflorescence; tepal 1.0 cm long, interior surface 

pubescent, exterior surface sparsely pubescent, cleft to base into 5 lobes; ovary 

subglobose, glabrous. Staminate buds bifusiform, flowers small, tepal 1.0 cm long, 

exterior surface sparsely pubescent, interior surface pubescent, cleft 1/3 way down into 5 

lobes; stamens 10, in 2 whorls of 5 each. Capsules large ca 2.5 cm long, surface smooth, 

apex rounded, dehiscence septicidal. Seeds large, 1.8 cm long, 0.9 cm wide, narrowly 

oblong, caruncle very small, inconspicuous. Fig 92A, B.


157 U. 

i! AI . 

2 1 l ,% 5 , 7 . 

I 

G 9, d t l , ( , ; , n (ires 

... 

--v. D, dsiuo 

f 

185 7, U). M. peruviana. D, distribution 

a \ ''s-.. ; I ,'t 

4 L


TYPE. Spruce 4287: Peru, San Martin: ad Tarapoto, Oct 1855 (syntypes, BM, photo 

at F, G-2, K-2, NY, photo at NY, OXF, P, W). 

DISTRIBUTION. (Fig 91D). Peru, Departments Loreto and San Martin. PERU. Loreto: 

Ule 6635 (G, K, L, MG) Tarapoto, Dec 1902. San Martin: Ferreyra 5090 (NY, US) Arriba de 

Tarapoto, Cruzando El Rio Shilcayo, Province San Martin, 24 Sep 1948. 

LOCAL NAME. Yuquilla (Ferreyra 5090) 

13. Manihot sect Crotalariaeformes Rogers & Appan, sect nov 


147(Heft 44): 29. 1910 pro parte. 

Manihot sect Grandibracteatae Pax subsect Tripartitae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 37. 1910 pro parte. 




44): 66. 1910 pro parte. 

Plantae caulinae, semi-herbaceae, procumbens, infirmae acauli frutices. Folia 

petiolatae, penitae lobatae. Inflorescentia monoecia racema. 

Plants caulescent, semiherbaceous; procumbent, weak-stemmed shrubs; leaves widely 

spaced on stem; petiolate, petiole attachment basal or peltate; lamina membranaceous to 

slightly coriaceous, deeply lobed, lobes variously shaped but not linear. Inflorescence 

monoecious, a raceme; bracts and bracteoles setaceous or semifoliaceous, margins entire 

to laciniate. 

TYPE. Manihot crotalarieaformis Pohl. 


FIG 92. Maniot perin. A, lef (Sprce 4287, G);B, inflorescence (Spruce 4287, K). 

FIG 92. Manibot peruviana. A, leaf (Spruce 4287, G); B, inflorescence (Spruce 4287, K).

''* '> .." ' . 

.., /? 

I 

J*" ' 

- 

?? 

FIG 93. Manbot sect Crtalariaefores. Geograpical range 

~~~~~~~~~~~~~~~~~~~~~~ 

/: / . 

, 

?~~~~~~~~~~~~~~~- r ~ ~ ..~. ~ ~ ~ ~ 

i ~~~ ~~B~B~~B~~B~~L.... 

. ..... 

" 

?r.~~~~~~~~~~ 

~~~~~~~~TI~~~~~~~~~~~~~~~~~~~~~P 

FIG93 C rotalaria efr rnes 

r :: 

% "-~~


Key to the Species of Sect Crotalariaeformes 

1. Bract and bracteole margins usually serrate; rarely entire; petiole attachment narrowly 

peltate; abaxial leaf surface plain; petiole, midrib, young stem, etc., usually pubescent, rarely 

glabrous. 

2. Leaf lobes obovate or hastate. 77. M. procumbens. 

2. Leaf lobes pandurate. 

78. M. affinis. 

1. Bract and bracteole margins usually entire, rarely serrate; petiole attachment basal; abaxial 

leaf surface distinctly pruinose; all parts glabrous, never pubescent. 

3. Leaves usually 5 lobed, rarely 3. 79. M. reptans. 

3. Leaves 3 lobed. 80. M. crotalariaeformis. 

77. Manihot procumbens Mueller von Argau, Linnaea 34: 206. 1865; in DC., Prodr. 

15(2): 1072. 1866; in Martius, Fl. Bras. 11(2): 478. 1874; Pax in Engler, 


Manihot elegans Muell.-Arg. in Martius, Fl. Bras. 11(2): 485. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 47. 1910. Type. Riedel s n (476 ?) (syntypes, F, G, NY). 

Manihot procumbens Muell.-Arg. var grandifolia Chodat & Hassler, Bull. Herb. Boissier 2(5): 673. 

1905; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 40. 1910. Type. Hassler 4441 

(syntypes, F, G-4, K, NY, S, UC, W). 

Manihot procumbens Muell.-Arg. var genuina Pax in Engler, Pflanzenreich IV. 147(Heft 44): 40. 

1910. 

Manihot meeboldii Pax in Engler, Pflanzenreich IV. 147(Heft 44): 47. 1910. Type. Hassler 9525a 

(syntypes, G-2). 

Manihot sellowiana Klotzsch ex Pax in Engler, Pflanzenreich IV. 147(Heft 44): 40. 1910 pro syn. 

Procumbent, weak-stemmed, semiherbaceous shrubs, ca 0.4 m tall, numerous stems 

from woody base. Young stems obtuse-angled in cross section, glabrous or pubescent; 

mature stems glabrous or pubescent. Leaves alternate; stipules persistent, setaceous and 

less than 0.5 cm long to nearly semifoliaceous and 1.0 cm long, glabrous or pubescent, 

usually laciniate, rarely entire; petioles short, ca 2.0 cm long, terete, glabrous or 

pubescent, petiole attachment to lamina narrowly peltate, width between the petiolelamina 

junction and basal edge of lamina ca 0.2 cm; lamina membranaceous, abaxial 

surface wax pattern smooth; venation camptodromous or craspedodromous, veins 

glabrous or pubescent; lamina 5 lobed, occasionally 3 or 7; median lobes obovate or 

hastate, ca 3.5 cm long, ca 1.75 cm wide, rarely larger, apex acuminate or obtuse, base of 

lobes narrow, less than 0.3 cm wide, width between base of sinus and petiole-lamina 

junction ca 0.3 cm; lowest lobes slightly smaller than medium lobes, nonsymmetric. 

Inflorescence a monoecious, terminal, short, raceme, ca 4.0 cm long, most of the parts 

sparsely pubescent or pubescent, rarely glabrous; bracteoles semifoliaceous, ca 1.0 cm 

long, ca 0.3 cm wide, glabrous or pubescent, margins usually laciniate, rarely entire; 

bractlets semifoliaceous, ca 0.6 cm long, ca 0.2 cm wide. Pistillate flowers just above the 

base of the inflorescence, pedicels ca 1.5 cm long, rarely as long as 4.0 cm, tepal 1.1 cm 

long, yellowish green, pubescent or glabrous, cleft to base into 5 lobes, ovary subglobose, 

glabrous. Staminate buds ovoid-ellipsoid to conical, flowers medium sized, tepal 1.1 cm 

long, yellowish green, pubescent or glabrous, cleft 1/3 way down into 5 lobes; stamens 

10, in 2 whorls of 5 each. Capsules small, 1.2 cm long, surface smooth, apex rounded. 

Seeds non vidi. Fig 94B, C, D; 95A. 

TYPE. Sello s n "in Brasilia meridonali" (syntypes, G, US). 

DISTRIBUTION. (Fig 94A). Brasil, States Minas Gerais and Sao Paulo; Paraguay. Alt 

ca 1150 m. In cerrado, low woods, and on sandstone slopes. BRASIL. Minas Gerais: Macedo 

3142 (S, SP, US) Bareiro, Municipality Araxa, 10 Feb 1951; Riedel s n (476 ?) (F, G, NY) in Campis. 

Sao Paulo: Gaudichaud 4520 (P), 1833; Lofgren 2084 (SP) cerrado, 9 Jan 1893. Parana: Hatschbach 

7529 (L) Campo Mourao, 11 Dec 1961. PARAGUAY. Province Unknown/Uncertain: Hassler 4441 (F, 

G-4, K, NY, S, UC, W) In Regione Fluminis Capibary, Sep; Hassler 4984 (F, NY, P, UC, W) In 

Altoplanitie et Decliviis, Sierra de Maracayu, Oct; Hassler 9525 (BM, GH, NY, S, UC, W) In Regione 

Fluminis Yhu, Oct 1905; Hassler 9525A (G-2) in Regione Fluminis Yhu, Nov 1905.


4984, UC). 

T 

_ 

? 1 1.:2'%, 

--b 

-' ? 

-: 

. r c i 

!:1 1: -, 

FIG 94. Manibot procumbens. A, distribution; B, leaf with entire lobes (Hassler 9525-A, G); C, 

petiole attachment to lamina narrowly peltate (Hassler 4441, G); D, leaf with incised lobes (Hassler 

4984, UC). 

'


Manihot procumbens exhibits what appears on certain specimens as two distinct leaf 

lobe shapes (Fig 94B, D). For this reason, Muell.-Arg. erected M. elegans as distinct from 

M. procumbens, but more recent collections exhibit both leaf lobe types on the same 

plant. Variation in leaf lobe shape within the same species occurs in many other species of 

the genus Manihot, as has been demonstrated by collections of the authors in both 

Central and South America. 

78. Manihot affinis Pax in Engler, Pflanzenreich IV. 147(Heft 44): 48. 1910. 

Procumbent shrubs. Young stems obtuse angled in cross section, pubescent. Leaves 

alternate; stipules caducous; petioles 2.5 cm long, terete, pubescent, petiole attachment 

to lamina narrowly peltate, width between petiole-lamina junction and basal edge of 

lamina 0.2 cm (leaf description based on the single mature leaf of the type specimen), 

lamina membranaceous, abaxial surface wax pattern smooth; venation craspedodromous, 

veins pubescent; lamina 5 lobed; median lobes pandurate 4.0 cm long, 1.5 cm wide, apex 

obtuse, base of lobes narrow, 0.3 cm wide, width between base of sinus and 

petiole-lamina junction 0.3 cm; lowest lobes slightly smaller than median lobes. Capsules 

and seeds non vidi. Fig 95C. 

TYPE. Lofgren 5969: Brasil, Sao Paulo: Paranapanema, Nov 1899 (syntype, SP). 

DISTRIBUTION. (Fig 95B). There is insufficient information to make an accurate 

description or proper taxon designation of Manihot affinis. The computer analysis 

indicates that the species joins M. procumbens at a c-value of .94, and is, therefore, a 

possible synonym of M. procumbens. 

79. Manihot reptans Pax in Engler, Pflanzenreich IV. 147(Heft 44): 30. 1910. 

Procumbent, weak-stemmed, semiherbaceous shrubs, ca 0.4 m tall, numerous stems 

arising from a woody base; branches ca 1.0 m long; latex yellowish. Young stems glabrous, 

purplish tinged; mature stems glabrous. Leaves alternate; stipules persistent, filiform, less 

than 0.3 cm long, less than 0.1 cm wide, glabrous, margins entire; petioles short, ca 3.0 

cm long, terete, glabrous, purplish tinged, petiole attachment to lamina basal, nonpeltate; 

lamina membranaceous to slightly coriaceous, adaxial surface dark green, glossy, abaxial 

surface pruinose, abaxial surface wax pattern smooth; venation camptodromous, veins 

glabrous; lamina palmately 5 lobed, rarely 3; median lobes obovate-elliptic, ca 3.0 cm 

long, ca 1.25 cm wide, apex acute or obtuse, base of lobes narrow, less than 0.3 cm wide, 

width between base of sinus and petiole lamina junction ca 0.3 cm; lowest lobes slightly 

smaller than median lobes, prominently nonsymmetric. Inflorescence a monoecious, 

terminal, short raceme, ca 4.0 cm long, all parts except interior surface of tepals glabrous; 

bracteoles and bractlets setaceous, less than 0.5 cm long, less than 0.1 cm wide, purplish 

tinged, margins entire. Pistillate flowers restricted to the base of the inflorescence, 

pedicels ca 1.5 cm long, tepal 1.0 cm long, yellowish green with considerable purple 

pigmentation, interior surface pubescent, cleft to base into 5 lobes, ovary subglobose. 

Staminate buds ovoid-ellipsoid, flowers medium sized, tepal 1.0 cm long, color same as 

pistillate tepal, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. 

Capsules small, 1.1 cm long, surface smooth, apex rounded, dehiscence septicidal. Seeds 

small, 0.8 cm long, oblong, caruncle moderately prominent. Fig 96A, B. 

TYPE. Ule 3072: Brasil, Goias: Serra Dos Pyreneos (syntypes, photo at F, photo at 

NY). 

DISTRIBUTION. (Fig 95D). Brasil, States Goias and Minas Gerais. BRASIL. Goias: Irwin, 

Maxwell & Wasshausen 18615 (F, GH, IAN, MICH, MO, NY, RB, S, SP, UB, UC, US, W) 15 km 

N of Corumba de Goias on Road to Niquelandia, 15 Jan 1968; Irwin, Maxwell & Wasshausen 19236 

(F, G, GH, K, MO, NY, RB, SP, UB, US) 20 km NW of Corumba de Goias, near Pico Dos Pirineus, 26


~~~~~~~~~~~~~~~~~~I 

"\ 

9~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

~~~~~~~~~~~~~~~~~~~~~i 

~ ~ 

Id 

?~~~~~~~ 

C~~~~~~~~~~~~~~~~~~ 

?;\r 

1~ ~.. 

FIG 95. Manibot procumbens. A, inflorescence (Hassler 9525-A, G). M. affinis. B, distribution; 

C, leaf (Lofgren 5969, SP) . M. reptans. D, distribution. 

~. 

J 

O ~~~~/o 

FIG~~~~ 95 aio rcmes.A 

nlrsec Hsse 55A ) .afii.B itiuin 

C, leaf (LOfgren 5969, SP). M. reptans. D, d~~~~~~~~~~istiuin 

/?~~~~~~~~~~~~~~,/


A t 4 

-?' `'~?3: 

FIG 96. Manihot reptans. A, leaf (Macedo 3697 US); B, inflorescence (Macedo 4390, S). M. 

crotalariaeformis. 

crotalariaeformis. C, distribution; D, leaf (Poh1 (Pohl 494, W).


Jan 1968; Macedo 3697 (S, US) Pirineus, Municipality Corumba, 28 Jul 1952; Macedo 4390 (S, SP) 

Pirineus, Municipality Corumba, 17 Feb 1956; Ule 373 (P-2) Serra Dos Pyreneos, Dec 1892. Minas 

Gerais: Saint Hilaire 2328 (P-3). 

80. Manihot crotalariaeformis Pohl, P1. Bras. Ic. et Descr. 1: 24. t. 18. 1827; Muell.-Arg. 

in DC., Prodr. 15(2): 1061. 1866; in Martius, Fl. Bras. 11(2): 456. 1874; Pax in 


Jatropha crotalariaeformis Steudel, Nomencl. ed. 2. 1: 799. 1840. 

Procumbent, weak stemmed, semiherbaceous shrubs. Stems glabrous. Leaves 

alternate; stipules persistent, filiform, less than 0.3 cm long, less than 0.1 cm wide, 

glabrous, margins entire, petioles short, ca 3.0 cm long, terete, glabrous, petiole 

attachment to lamina basal, nonpeltate; lamina membranaceous to slightly coriaceous, 

abaxial surface wax pattern smooth; venation camptodromous, veins glabrous; lamina 

palmately 3 lobed, median lobes obovate-elliptic, ca 3.0 cm long, ca 1.25 cm wide, apex 

acute or obtuse, base of lobes narrow, less than 0.3 cm wide, width between base of sinus 

and petiole lamina junction ca 0.3 cm; lowest lobes slightly smaller than median lobes, 

nonsymmetric. Inflorescence a monoecious, terminal, short, raceme, ca 4.0 cm long 

(inflorescence description based on the fragmentary inflorescence of the type specimen); 

peduncles and pedicels glabrous; bracteoles and bractlets non vidi. Pistillate flowers non 

vidi. Staminate buds ovoid-ellipsoid, flowers medium sized, tepal 1.0 cm long, cleft 1/3 

way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules and seeds non vidi. 

Fig 96D. 

TYPE. Pohl 494: Brasil, Minas Gerais: ad Rio Parannahyba (syntypes, G, W). 

DISTRIBUTION. (Fig 96C). BRASIL. Minas Gerais: Pohl Icon Tab. 18 (W); Saint Hilaire 

255-0 (P). 

14. Manihot sect Stipulares Pax emend Rogers & Appan 

Manihot sect Stipulares Pax in Engler, Pflanzenreich IV. 147(Heft 44): 49. 1910. 

Manihot sect Parvibracteatae Pax subsect Nanae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

53. 1910. 

Plants nearly acaulescent, caespitose shrubs; leaves borne more or less in a rosette, 

internodes shortened; petiolate, petiole attachment basal; lamina membranaceous to 

slightly coriaceous, deeply lobed, lobes variously shaped but not linear. Inflorescence 

usually dioecious, sometimes monoecious; bracts and bracteoles setaceous to semifoliaceous, 

margins entire to laciniate. 

TYPE. Manihot stipularis Pax & K. Hoffmann. 


Key to the Species of Sect Stipulares 

1. Stipules foliaceous, usually more than 3.0 cm long; serrate or laciniate; bracteoles usually 

more than 1.0 cm long; petioles more than 15.0 cm long; leaves 7 lobed, occasionally 9 

lobed; lobes curved. 

2. Stipule margin laciniate. 81. M. stipularis. 

2. Stipule margin serrate. 82. M. pusilla. 

1. Stipules setaceous, less than 1.5 cm long; margin usually entire; bracteoles less than 1.0 cm 

long; petioles less than 15.0 cm long; leaves 3 to 5 lobed, rarely 7 lobed; lobes straight. 

3. Leaf lobes oblong or lanceolate, apex acute or acuminate. 

4. Plants usually less than 15.0 cm tall; petioles less than 8.0 cm long; leaf lobes less than 

1.5 cm wide. 83. M. oligantha. 

4. Plants usually more than 15.0 cm tall; petioles more than 10.0 cm long; leaf lobes more 

than 1.5 cm wide. 84. M. longepetiolata. 

3. Leaf lobes subrotundate, apex retuse or obtuse. 85. M. nana. 

81. Manihot stipularis Pax in Engler, Pflanzenreich IV. 147(Heft 44): 50. 1910.

, 

- 

.-; . / / ; 

^ , rI 

_' --\ ~ 

-> ., .* ' '~ . 

" ..- -.. ... . 

/*\V, :. '-'---*. . . ."'*' i 

.. ; , 

' 

?%I ?, ~ ~ 

..? .... ,~:- 

' ..... 

~: 

...,. 

FIG 97. Manihot sect Stipulares. Geographical range. 

...


Very short, nearly acaulescent, subshrubs, ca 20.0 cm tall; internodes very short, 

usually less than 0.5 cm long. Stems glabrous. Leaves borne more or less as a rosette; 

stipules persistent, foliaceous, ca 4.0 cm long, sometimes as long as 6.0 cm, ca 0.3-0.5 cm 

wide, glabrous, margins laciniate; petioles ca 20.0 cm long, terete, glabrous, purplish 

tinged, petiole attachment to lamina basal, nonpeltate; lamina membranaceous, abaxial 

surface wax pattern smooth; venation camptodromous, veins glabrous; lamina palmately 

7 lobed, occasionally 9, midribs of lobes usually reflexed; median lobes oblong, ca 6.0 cm 

long, ca 1.5 cm wide, apex acuminate, base of lobes narrowly constricted, less than 0.2 cm 

wide, width between base of sinus and petiole-lamina junction less than 0.2 cm; lowest 

lobes slightly smaller than median lobes. Inflorescence usually dioecious with pistillate 

and staminate inflorescences on separate plants, rarely on the same plant; all parts 

glabrous; bracteoles setaceous, ca 1.5 cm long, less than 0.1 cm wide, margin serrate; 

bractlets setaceous, ca 0.5 cm long. Pistillate inflorescence determinate, ca 7.0 cm long, 2 

flowers borne at the apex of a solitary peduncle; all parts glabrous; pedicels ca 1.5 cm 

long, curved down; tepal 1.2 cm long, purplish tinged, cleft to base into 5 lobes; ovary 

subglobose. Staminate inflorescence a panicle, ca 15.0 cm long; bracteoles and bractlets 

same as in pistillate inflorescence; pedicels 1.0 cm long, staminate buds ovoid-ellipsoid, 

flowers medium sized, tepal 1.2 cm long, with considerable purplish pigmentation, cleft 

1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules and seeds non 


TYPE. Glaziou 22128: Brasil, Goias: Guariroba 16 Oct 1894 (syntypes, A-2, F, G-2, 

K, MO, P, S). 

DISTRIBUTION. (Fig 98A). Brasil, state of Goias and Distrito Federal. Alt 1250 m. 

On rocky banks. BRASIL. Goias: Irwin, Souza & dos Santos 10012 (F, NY, UB, US) 75 km N of 

Cristalina on road to Brasilia, 6 Nov 1965; Irwin, Souza & dos Santos 11046 (F, G, GH, K, MICH, MO, 

NY, RB, SP, UB, US) campo and cerrado near Rio Descoberto, 4 Dec 1965;Maguire, Pires, Maguire & 

Silva 57096 (NY) Cerrado along Road to Rio Corumba, 30 km from Gama, Aug-Sep 1963. Distrito 

Federal: Heringer 9209 (NY) Rio Preto, Brasilia, 10 Aug 1963. 

This species (and others of the section Stipulares) are more consistently dioecious 

than monoecious, a condition reversed in all other species of the genus. 

Manihot stipularis is a handsome species, and with the species, M. oligantha, should 

make definite contributions to horticulture as a rock garden plant in warm climate. 

82. Manihot pusilla Pohl, P1. Bras. Ic. et Descr 1: 36. t. 26. 1827; Muell.-Arg. in Martius, 

Fl. Bras. 11(2): 460. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 50. 1910. 

Jatropha pusilla Steudel, Nomencl. ed. 2. 1: 800. 1840; Manihot palmata Muell.-Arg. var pusilla 

(Pohl) Muell. Arg. in DC., Prodr. 15(2): 1063. 1866. 

Very short acaulescent subshrubs; internodes very short, less than 0.5 cm long. Stems 

glabrous. Leaves borne more or less as a rosette (description based on the depauperate 

plant representing the type specimen); stipules persistent, foliaceous, ca 3.0 cm long, ca 

0.3 cm wide, glabrous, margins serrate; petioles 6.0 cm long (immature leaf), terete, 



7 lobed, midribs reflexed; median lobes oblong, 6.0 cm long, 1.5 cm wide, apex 

acuminate, base of lobes narrowly constricted, less than 0.2 cm wide, width between base 

of sinus and petiole lamina junction less than 0.2 cm; lowest lobes slightly smaller than 

median lobes. Inflorescence non vidi. Capsules and seeds non vidi. 

TYPE. Pohl 1181: Brasil, Goias: Serra de Cristaes (syntypes, photo at F, W). 

DISTRIBUTION. (Fig 98D). BRASIL. Goias: Pohl Icon Tab. 26 (W). 

83. Manihot oligantha Pax in Engler, Pflanzenreich IV. 147(Heft 44): 53. 1910.


P-, ,' e 

GI~ /E 

FIG 98. Manihot stipularis. A, distribution; B, pistillate plant (Irwin, Souza & dos Santos 

11046, NY); C, staminate plant (Heringer 9209, NY). M. pusilla. D, Distribution. 

i



usually less than 0.5 cm long. Stems glabrous. Leaves borne more or less as a rosette; 

stipules persistent, setaceous, ca 1.0 cm long, less than 0.1 cm wide, glabrous, margins 

entire, rarely serrate; petioles ca 5.0 cm long, terete, glabrous, purplish tinged towards the 

lamina end, petiole attachment to lamina basal, nonpeltate; lamina membranaceous to 

slightly coriaceous, adaxial surface dark green, abaxial surface lightly pruinose, abaxial 


3 or 5 lobed, occasionally with 2 more lobules, midribs of lobes straight; median lobes 

oblong to lanceolate, ca 3.0 cm long, ca 1.0 cm wide, apex acuminate, base of lobes 


petiole-lamina junction less than 0.2 cm, lamina at sinus usually overlapping; lowest lobes 

slightly smaller than median lobes. Inflorescence usually dioecious, rarely monoecious, all 

parts glabrous; bracteoles setaceous, ca 0.8 cm long, margins usually entire; bractlets 

setaceous, less than 0.5 cm long. Pistillate infloresence determinate, ca 7.0 cm long, 2 

flowers borne at the apex of a solitary peduncle; pedicels ca 1.5 cm long, curved down; 

tepal 1.1 cm long, purplish tinged, cleft to base into 5 lobes; ovary subglobose. Staminate 

inflorescence a panicle, ca 8.0 cm long; bracteoles and bractlets same as in pistillate 

inflorescence; pedicels 1.0 cm long, staminate buds ovoid-ellipsoid, flowers medium sized, 

tepal 1.2 cm long, with considerable purplish pigmentation, cleft 1/3 way down into 5 

lobes; stamens 10, in 2 whorls of 5 each. Capsules 1.0 cm long, subglobose, slightly 

elongated, surface smooth, glaucous, apex rounded, dehiscence septicidal. Seeds small, 

0.7 cm long, oblong, caruncle not prominent. Fig 99B. 

TYPE. Glaziou 22127: Brasil, Goias: entre la Fazenda de Boa Vista et Paraizo, 17 

Jan 1895 (syntypes, photo at F, G, K, photo at NY, P). 

DISTRIBUTION. (Fig 99A). Brasil, state of Goias. Alt ca 1250 m. Among outcrops 

and on rocky slopes. BRASIL. Goias: Irwin, Grear, Souza & dos Santos 13230 (F, GH, K, MO, NY, 

RB, SP, UB, UC, US, W) Campo and Cerrado circa 3 km N of Cristalina, 2 Mar 1966; Irwin, Grear, 

Souza & dos Santos 13230A (F, NY, UB) Campo and Cerrado circa 3 km N of Cristalina, 2 Mar 1966; 

Irwin, Souza & dos Santos 9906 (F, G, H, K, MO, NY, RB, SP, UB, US, W) circa 2 km N of Cristalina, 

5 Nov 1965. 

84. Manihot longepetiolata Pohl, P1. Bras. Ic. et Descr. 1: 25 t. 19. 1827; Muell.-Arg. in 

DC., Prodr. 15(2): 1061. 1866; in Martius, Fl. Bras. 11(2): 447. 1874. 

Manihot longipetiolata Pohl, Pax in Engler, Pflanzenreich IV. 147(Heft 44): 53. 1910. 

(orthographic variant). 

Jatropha longepetiolata Steudel, Nomencl. ed. 2. 1: 799. 1840. 

Short subshrubs, ca 20.0 cm tall; internodes short, usually ca 2.0 cm long. Young 

stems glabrous, glaucous; mature stems glabrous. Leaves borne more or less as a rosette; 


entire, rarely serrate; petioles ca 12.0 cm long, terete, glabrous, glaucous, petiole 

attachment to lamina basal, nonpeltate; lamina membranaceous to slightly coriaceous, 

adaxial surface dark green, abaxial surface pruinose, abaxial surface wax pattern smooth; 

venation camptodromous, veins glabrous; lamina palmately 3 lobed, midribs of lobes 

straight; median lobes oblong, ca 4.0 cm long, ca 2.0 cm wide, apex acute, base of lobes 


petiole-lamina junction less than 0.2 cm, lamina at sinus overlapping; lowest lobes slightly 

smaller than median lobes. Inflorescence usually dioecius, occasionally monoecius, ca 

10.0 cm long; all parts glabrous, most of the parts with a bluish white bloom; bracteoles 

setaceous, ca 0.8 cm long, margins usually entire; bractlets setaceous, Pistillate tepal 1.1 

cm long, cleft to base into 5 lobes; ovary subglobose. Staminate buds ovoid-ellipsoid, 

flowers medium sized, tepal 1.1 cm long, cleft 1/3 way down into 5 lobes; stamens 10, in 

2 whorls of 5 each. Capsules and seeds non vidi. Fig 99D. 

TYPE. Pohl 822: Brasil, Goias; ad serra de Cristaes (syntypes, F, G, W).


/ 

i 

i 

_i~~~~~~~~~~ -- 

/? I' 

C~~,I~ 

'j i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

J 

._//'~~ 

FIG 99. Manibot oligantba. A, distribution; B, the plant (Irwin, Great, Souza & dos Santos~~~~~~~~~~ 

133 \,N) .1neeilt.C disr i b tin th , lnt(o182 ) 

7~ 

r 

z


DISTRIBUTION. (Fig 99C). Brasil, states Goias and Minas Gerais. BRASIL. Goias: Ir- 

win, Grear, Souza & dos Santos 13636 (UB) Campo circa 15 km W of Cristalina, 6 Mar 1966; Irwin, 

Grear, Souza & dos Santos 13763 (NY, UB) Campo and Cerrado circa 15 km E of Cristalina, 8 Mar 

1966; Irwin, Souza & dos Santos 9783 (F, G, GH, K, LIL, MICH, MO, NY, RB, SP, UB, UC, US, W) 

Cerrado 5 km W of Cristalina, 2 Nov 1965;Pohl Icon Tab. 19 (W-2). Minas Gerais: Duarte 2785 (PUL, 

RB) Patos, Arredores, 18 Aug 1950. 

85. Manihot nana Mueller von Argau in Martius, Fl. Bras. 11(2): 448. 1874; Pax in 



usually less than 1.0 cm long. Stems glabrous. Leaves borne more or less as a rosette, 


entire, rarely serrate; petioles 3.0-11.0 cm long, terete, glabrous, petiole attachment to 

lamina basal, nonpeltate; lamina membranaceous to slightly coriaceous, adaxial surface 

dark green, abaxial surface lightly pruinose, abaxial surface wax pattern smooth; venation 

camptodromous, veins glabrous; lamina palmately 5 lobed, midribs straight or slightly 

curved; median lobes subrotundate, ca 4.0 cm long, ca 3.0 cm wide, apex retuse or 

obtuse, base of lobes narrow, less than 1.0 cm wide, width between base of sinus and 

petiole-lamina junction less than 1.0 cm, lamina at sinus prominently overlapping; lowest 

lobes slightly smaller than median lobes. Inflorescence non vidi. Capsules and seeds non 

vidi. (Description based on the fragmentary leaves in the type 

collection and the 

depauperate, sterile plant in Riedel s n) Fig 100B. 

TYPE. Riedel 2824: Brasil, Goias: ad Chapadao de S. Marcos (syntypes, Photo F, G). 

DISTRIBUTION. (Fig 100A). Brasil, states of Goias and Minas Gerais. BRASIL. Goias: 

Riedel s n (P). Minas Gerais: Heringer 7781 (NY), 31 Nov 1960. 

FIG 100. Maniot nana. A, distribution; B, the plant (Riedel s n, P). 

- _ . 

,--J "~~~~~~~~~/c-. .- 1 

!/'0.Mnhtnaa ,dsrbtinB h ln (idlsn )


The paucity of material of this species makes the decision difficult to either 

synonymize or maintain as a separate taxon. However, computer analysis indicates that 

the species is quite distinct, and that its closest relative is Manihot longepetiolata, which it 

joins at a C-value of .86. 

15. Manihot sect Grandibracteatae Pax emend Rogers & Appan 

Manihot sect Grandibracteatae Pax subsect Tomentosae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 25. 1910 pro parte. 


petiole attachment basal, lamina membranaceous, deeply lobed, lobes usually obovate, 

never linear. Inflorescence a monoecious cluster of several racemes arising from a 

common base; bracts and bracteoles foliaceous, margins entire. 

TYPE. Manihot tomentosa Pohl emend Rogers & Appan. 


86. Manihot tomentosa Pohl, PI. Bras. Ic. et Descr. 1: 50 t. 43. 1827. 

Subshrubs, ca 1.0 m tall, ascending stems arising from a woody root. Young stems 

tomentose, tomentum usually bright yellowish; mature stems tomentose. Leaves 

alternate; stipules caducous, setaceous, ca 1.5 cm long, ca 0.2 cm wide, tomentose; 

petioles ca 13.0 cm long, terete, tomentose, petiole attachment to lamina basal, 

nonpeltate; lamina coriaceous, abaxial surface wax pattern smooth; venation camptodro- 

mous, veins on adaxial and abaxial lamina surface tomentose; lamina palmately 5 or 7 

lobed, lamina at lobe sinus overlapping or separated; median lobes obovate, ca 13.0 cm 

long, ca 6.0 cm wide, apex acute or obtuse, base of lobes 1.0-2.0 cm wide, width 

between base of sinus and petiole-lamina junction 1.0-2.0 cm; lowest lobes slightly 

smaller than median lobes, nonsymmetric. Inflorescence monoecious, terminal, a cluster 

of ca 6 nearly subspicate racemes arising from one common base, 5.0-10.0 cm long; all 

parts tomentose; bracteoles foliaceous, ca 2.0 cm long, ca 0.6 cm wide, margins entire; 

bractlets foliaceous, ca 1.0 cm long, ca 0.4 cm wide, margins entire. Pistillate flowers 

restricted to the base of the upper half of the inflorescence, tepal 1.0 cm long, interior 

and exterior surface tomentose, cleft to base into 5 lobes; ovary subglobose, tomentose. 

Staminate buds ovoid-ellipsoid, flowers medium sized, tepal 1.0 cm long, exterior and 

interior surface tomentose, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 

each. Capsules 1.5 cm long, surface without wings, tomentose, apex rounded, dehiscence 

septicidal. Seeds 1.0 cm long, oblong, caruncle moderately prominent. 

DISTRIBUTION. (Fig 102A). Brasil, states of Goias and Minas Gerais. Alt ca 1000 m. 

Key to the Subspecies of Manihot tomentosa 

1. Leaves 5 lobed; lobes overlapping at sinus. 86a. M. tomentosa subsp tomentosa. 

1. Leaves 7 lobed, rarely 5; lobes usually separated at sinus. 86b. M. tomentosa subsp araliaefolia. 

86a. Manihot tomentosa Pohl emend Rogers & Appan subsp tomentosa 

Manihot tomentosa Poht, PI. Bras. Ic. et Descr. 50t.43. 1827; Muell.-Arg. in DC., Prodr. 15(2): 


147(Heft 44): 25. 1910;Jatropha tomentosa Steudel, Nomencl. ed. 2. 1: 800. 1840. 

Manihot araliaefolia Pax in Engler, Pflanzenreich IV. 147(Heft 44): 26. 1910 ex parte. Type. 

Glaziou 22136 (syntypes, US). 

Plant parts densely tomentose. Leaves 5 lobed, lamina at sinus overlapping. 

Inflorescence compact, peduncles never branched. Fig 102B, C. D.

!: '" . D' I 

, 

~:'~)L~~~~~~~~~~~~~~~~~~~~~~~~~ ,,?~~~? 

FI 

FIG 101. Manihot sect Grandibracteatae. Geographical range. 

, . 

t 

11 Mnio sc 

tfl 

' 

~ 

\ 1 CI` '? 

Grnibrceaa.


TYPE. Pohl 702 (and on some specimens also 767 & 1713) Brasil, Minas Gerais: ad 

Serra d'Urubu, ad Rio Paranahyba, S. Pedro d'Alcantara, Nossa Senhora, Paracatu 

(syntypes, F-2, W-2). 

DISTRIBUTION. (Fig 102A) Brasil, states of Goias and Minas Gerais. BRASIL. Goias: 

Glaziou 22136 (US) Entre Cocal et Lambary, 26 Feb 1895. Minas Gerais: Barreto 2617 (F) Lagoa 

Santa, Municipality Santa Luzia, 25 Mar 1933; Barreto 2618 (F) Lagoa Santa, Santa Luzia, 20 Nov 

1933; Burret & Brade 15967 (PUL, RB) Lagoa Santa, Dec 1937; Claussen 758 (NY-2, S), 1838; 

Claussen s n (OXF) Lagoa Santa, 1840; Claussen s n (F, W) Lagoa Santa. 1843. Glaziou 17753 (F, P) 

Serra do Senheiro, 20 Jan 1889; Pohl Icon Tab. 43 (W); Rogers 363 (NY-3) Lagoa Santa, 17 Feb, 

1961. 

LOCAL NAMES. Mandiocucu do campo (Barreto 2617) manioc (Claussen s n) 

86b. Manihot tomentosa Pohl emend Rogers & Appan subsp araliaefolia (Pax) Rogers & 


Manihot araliaefolia Pax in Engler, Pflanzenreich IV. 147(Heft 44): 26. 1910 ex parte. 

Manihot canastrana Glaziou, Bull. Soc. Bot. France 59(3): 628. 1912 (1913) nom rejic. 

Plant parts less densely tomentose. Leaves 7 lobed, rarely 5. Inflorescence loose, the 

central peduncle occasionally with lateral branches. Fig 103A. 

TYPE. Glaziou 22137: Brasil, Goias: Fazenda da Cava, pres du Morro da Canastra, 

23 Nov 1894 (syntypes, F-2, NY, P). 

DISTRIBUTION. (Fig 102A). BRASIL. Goias: Irwin, Grear, Souza & dos Santos 14195 (F, G, 

GH, K, MO, NY, RB, SP, UB, US, W) Cerrado Rio Parana circa 35 km N of Formosa on Road to Sao 

Gabriel, 28 Mar 1966; Irwin, Souza, Grear & dos Santos 15529 (F, NY, UB) Corrego Itaquera, circa 30 

km N of Formosa, 2 May 1966. 

16. Manihot sect Brevipetiolatae Pax emend Rogers & Appan 

Manihot sect Brevipetiolatae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 95. 1910. 

Manihot sect Weddellianae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 98. 1910. 

Plants caulescent; low to medium shrubs; leaves widely spaced on stem; subsessile, 

petiole attachment basal, lamina membranaceous to coriaceous, nonlobed. Inflorescence a 

monoecious subspicate raceme, single or 2 or 3 racemes arising from a common base; 

bracts and bracteoles setaceous to foliaceous, margins entire to laciniate. 

TYPE. Manibot orbicularis Pohl. 


Key to the Species of Sect Brevipetiolatae 

1. Leaves less than 10.0 cm wide, linear, lanceolate, gladiate, digitate, ovate or obovate, not 

orbicular. 

2. Leaves less than 1.25 cm wide; linear. 87. M. stricta. 

2. Leaves more than 1.25 cm wide; lanceolate, gladiate, digitate, ovate or obovate. 

3. Leaf venation camptodromous; leaves more than 2.0 cm wide, margins entire, no 

secondary lobes present; bract and bracteole margins entire, rarely serrate. 

4. Bracts and bracteoles setaceous, less than 0.25 cm wide; glabrous; leaf lobes less than 

15.0 cm long, more than 3.0 cm wide; lanceolate. 88. M. purpureo-costata. 

4. Bracts and bracteoles foliaceous, more than 0.5 cm wide; glabrous; leaf lobes less than 

15.0 cm long, more than 3.0 cm wide; ovate or obovate. 89. M. salicifolia. 

4. Bracts and bracteoles semifoliaceous, 0.25-0.5 cm wide, sparsely pubescent; leaf 

lobes more than 15.0 cm long, less than 3.0 cm wide; gladiate. 

90. M. attenuata. 

3. Leaf venation craspedodromous; lobes less than 2.0 cm wide, margins repand or incised, 

usually with attenuate secondary lobes at base; bract and bracteole margins laciniate. 

91. M. weddelliana. 

1. Leaves more than 10.0 cm wide, orbicular. 92. M. orbicularis.


27 

k 

" 

? .: .:~~~~~~~~~~~~~~~~~~~~~~* 

tlll~~~~~~~~~c 

(Rogers 363, NY); D, inflorescence (Barreto 2618, F). 

,i 

t 

~--~--~;~~`~--'o o 

5;e~~~~~~~~~~~~: 

FIG 102. Magihot tom ~ents.A ~ ~ itiuin osrpeetsbptmnoaadtelte 

FIG 102. Manibot tomentosa. A, distribution; dots represent subsp tomentosa and the letter A 

represents subsp araliaefolia. M. tomentosa subsp tomentosa. B, the plant (Rogers 363, NY); C, leaf


i: ; A 

FIG 103. Manihot tomentosa subsp araliaefolia. A, leaf (Glaziou 22137, F). 

87. Manihot stricta Baillon, Adansonia 4: 282. 1863-64; Muell.-Arg. in DC., Prodr. 

15(2): 1058. 1866; in Martius, Fl. Bras. 11(2): 444. 1874; Pax in Engler, 


Manihot linearifolia Muell.-Arg., Flora 55: 43. 1872; Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 96. 1910. Type. Haenke s n: Peru (syntypes, F-2, G, M). 

Slender subshrubs, ca 0.5 m tall, several ascending stems arising from a woody base. 

Stems glabrous. Leaves alternate; stipules caducous, filiform, glabrous; leaves subsessile, 

petioles less than 0.25 cm long, glabrous, petiole attachment to lamina basal, nonpeltate; 

lamina membranaceous, abaxial surface wax pattern smooth; venation camptodromous, 

veins glabrous; lamina nonlobed, linear, ca 15.0 cm long, ca 0.5 cm wide, sometimes as 

long as 25.0 cm long and 1.2 cm wide, apex acuminate, base attenuate. Inflorescence a 

monoecious, terminal, short, subspicate raceme, ca 3.0 cm long, all parts except the 

interior surface of tepals pubescent; bracteoles and bractlets setaceous, less than 0.5 cm 

long, less than 0.1 cm wide, purplish tinged, margins entire. Pistillate flowers restricted to 

the base of the inflorescence, tepal 1.0 cm long, yellowish green with purplish 

pigmentation, interior surface pubescent, cleft to base into 5 lobes, ovary subglobose. 

Staminate buds ovoid-ellipsoid to fusiform, flowers infundibuliform, tepal 1.0 cm long, 

interior surface pubescent, color same as that of pistillate tepal, cleft 1/3 way down into 

5 lobes; stamens 10, in 2 whorls of 5 each. Capsules very small, 0.8 cm long, subglobose, 

surface smooth, apex rounded, dehiscence septicidal. Seeds very small, 0.6 cm long, 0.3 

cm wide, oblong, caruncle not prominent. Fig 105B, C. 

TYPE. Gardner 3442: Brasil, Goias: near Villa de Natividade, 1841 (syntypes, BM-2, 

F, Photo F, G-2, K-2, P-2, W-2) 

DISTRIBUTION. (Fig 105A). Peru; Brasil, states of Mato Grosso and Goias. Alt to ca 

400 m. Among cerrado grasses. PERU. Province Unknown/Uncertain: Haenke s n (F-2, G, M).

-1~~~~~~ i , 

FIG 104. Manihot sect Brevipetiolatae. Geographical range.


' 

"., r _ _ 

\\~~~~~~~~i ' 

~~~~~~~~~~~~~~~~~~~"? 

i 

~~ 

~~~~~~~--- 

t


BRASIL. Mato Grosso: Argent, Ramos, Richards & Souza 6571 (K) Base Camp at 12.49" S-51.46" 

W, 30Sep 1967;Argent, Ramos, Richards & Souza 6685 (K) Base Camp at 12.49? S-51.460 W, 7 Oct 

1967; Harley & Souza 10341 (K) R 10, circa 12 km SW of Base Camp, near Lago de Leo, 28 Sep 

1968 Jrwin & Soderstrom 6761 (UB) circa 70 km N of Xavantina, 10 Oct 1964; Irwin & Soderstrom 

6774 (F, K, NY, UB) circa 90 km N of Xavantina, 14.40? S-52.200 W, 12 Oct 1964; Philcox & 

Fereira 3941 (K) Campo E of Base Camp at 12.49? S-51.460 W, 9 Jan 1968; Souza & Richards 6888 

(K) circa 66 km S along Road from Base Camp at 12.49? S-51.460 W, 9 Sep 1968. 

88. Manihot purpureo-costata Pohl, PI. Bras. Ic. et Descr. 1: 19. t. 11. 1827; Muell.-Arg. 

in DC., Prodr. 15(2): 1057. 1866; in Martius, Fl. Bras. 11(2): 442. 1874; Pax in 

Engler Pflanzenreich IV. 147(Heft 44): 96. 1910. 

Jatropha purpureo-costata Steudel, Nomencl. ed. 2. 1: 799. 1840. 

Subshrubs, ca 0.5 m tall. Stems glabrous. Leaves alternate; stipules caducous, 

filiform, glabrous; leaves subsessile, petioles less than 0.5 cm long, glabrous, glaucous, 

petiole attachment to lamina basal, nonpeltate; lamina slightly coriaceous, abaxial surface 

glaucous, abaxial surface wax pattern spongiose; venation camptodromous, veins 

glabrous; lamina nonlobed, lanceolate, ca 14.0 cm long, ca 4.0 cm wide, apex acute, base 

obtuse to acute. Inflorescence a monoecious, terminal, subspicate raceme, ca 15.0 cm 

long, all parts except interior surface of tepals glabrous, most of the parts glaucous; 

bracteoles and bractlets setaceous, less than 0.5 cm long, less than 0.1 cm wide, margins 

entire. Pistillate flowers restricted to the base of the upper half of the inflorescence, tepal 

1.0 cm long, dark purplish, interior surface pubescent, cleft to base into 5 lobes, ovary 

subglobose. Staminate buds ovoid ellipsoid to fusiform, flowers infundibuliform to nearly 

tubular, tepal 1.0 cm long, interior surface pubescent, color same as that of pistillate 

tepal, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules and 

seeds non vidi. Fig 106A, B. 

TYPE. Pohl 2130: Brasil, Goias: circa Cavalcante (syntypes, F, Photo F, M, W). 

DISTRIBUTION. (Fig 105D). 

Cavalcante. 

BRASIL. Goias: Pobl Icon Tab. 11 (W); Pobl 1703 (G, W) 

89. Manihot salicifolia Pohl, PI. Bras. Ic. et Descr. 1: 18. t. 10. 1827; Muell.-Arg. in DC., 



Jatropha salicifolia Steudel, Nomencl. ed. 2. 1: 800. 1840. Manihot riedeliana Muell.-Arg. in 

Martius, Fl. Bras. 11(2): 443. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 97. 

1910. Type. Riedel 613 (syntypes, F, G-2). 

Manibot mattogrossensis Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 197. 

1924. Type. Malme 2344 (syntypes, S-3). 

Subshrubs ca 0.5 m tall; several ascending stems arising from a woody base. Stems 

glabrous. Leaves alternate; stipules caducous, filiform, glabrous; leaves subsessile, petioles 

less than 0.5 cm long, glabrous, petiole attachment to lamina basal, nonpeltate; lamina 

membranaceous to slightly coriaceous, abaxial surface wax pattern spongiose; venation 

camptodromous, veins glabrous; lamina nonlobed, ovate or obovate, ca 12.0 cm long, ca 

5.5 cm wide, apex acute, base attenuate to breve-angustatus. Inflorescence a monoecious, 

terminal, subspicate raceme, 15.0-20.0 cm long, sometimes pendent, all parts except 

interior surface of tepals glabrous; bracteoles foliaceous, ca 1.2 cm long, ca 0.6 cm wide, 

margins entire; bractlets foliaceous, ca 1.0 cm long, ca 0.5 cm wide, margins entire. 

Pistillate flowers restricted to the base of the inflorescence, usually borne on long (ca 3.0 

cm) pedicels; tepal 1.3 cm long, yellowish green, interior surface pubescent, cleft to base 

into 5 lobes, ovary subglobose. Staminate buds ovoid-ellipsoid to fusiform, tepal 1.3 cm 

long, cleft 1/3 way down into 5 lobes, interior surface pubescent; stamens 10, in 2 whorls 

of 5 each. Capsules and seeds non vidi. Fig 106D; 107A, B.


~Z?' . 

I M 

FIG 106. Manihot purpureo-costata. A, leaf (Pobl 1703, W); B, inflorescence (Pohi 1703, W). 

M. salicifolia. C, distribution; D, leaf, ovate in outline (Malme 2344, S). 

M. salici~folia. C, distribution;D, leaf, ovate in outline (Malme 2344, S). 

r!


TYPE. Pohl 1194 (and on some specimens also 1660): Brasil, Goias: ca S. Luzia 

(syntypes, F, K, M, W-2). 

DISTRIBUTION. (Fig 106C). Brasil, states of Mato Grosso and Goias. Alt to ca 400 m, 

in cerrado. BRASIL. Mato Grosso: Irwin & Soderstrom 6675 (F, K, NY, UB) circa 75 km N of 

Xavantina, 9 Oct 1964; Malme 2344 (S-3) S Anna Da Chapada, 21 Sep 1902. Goias: Pohl Icon Tab. 10 

(W). Sao Paulo: Riedel 613 (F, G-2) Rio Pardo. 

90. Manihot attenuata Mueller von Argau in Martius, Fl. Bras. 11(2): 442. 1874; Pax in 

Engler, Pflanzenreich IV. 147(Heft 44); 95. 1910. 

Manihot brachystachys Pax in Engler, Pflanzenreich IV. 147(Heft 44): 97. 1910, Type. Glaziou 

22126 (syntypes, F, G, K, NY, P, S). 

Subshrubs, ca 0.5 m tall; several ascending stems arising from a woody base. Stems 

glabrous. Leaves alternate, held erect on plants; stipules caducous, filiform, glabrous; 

leaves subsessile, petioles less than 0.5 cm long, glabrous, petiole attachment to lamina 

basal, nonpeltate; lamina membranaceous to slightly coriaceous, abaxial surface wax 

pattern smooth; venation camptodromous, veins glabrous; lamina nonlobed, gladiate, ca 

20.0 cm long, ca 2.5 cm wide, apex acuminate, base acute. Inflorescence a monoecious, 

terminal, subspicate raceme, ca 7.0 cm long; bracteoles semifoliaceous, ca 0.75 cm long, 

ca 0.4 cm wide, sparsely pubescent, margins entire or serrate; bractlets semifoliaceous, ca 

0.5 cm long, ca 0.3 cm wide, sparsely pubescent. Pistillate flowers restricted to the base 

of the inflorescence, tepal 1.0 cm long, interior surface sparsely pubescent, cleft to base 

into 5 lobes, ovary subglobose. Staminate buds ovoid-ellipsoid to fusiform, tepal 1.0 cm 

long, interior surface pubescent, cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls 

of 5 each. Capsules small, 0.9 cm long, surface without wings, apex rounded, dehiscence 

septicidal. Seeds 0.7 cm long, oblong, caruncle not well developed. Fig 107D; 108A. 

TYPE. Burchell 7865: Brasil, Goias: inter Goyaz et Cavalcante (syntypes, Photo F, 

G, K). 

DISTRIBUTION. (Fig 107C). BRASIL. Goias: Glaziou 22126 (F, G, K, NY, P, S) Chapado 

Dos Veadeiros, 8 Jan 1895;Irwin, Grear, Souza & dos Santos 12654 (F, K, NY, SP, UB) Chapada Dos 

Veadeiros circa 15 km W of Veadeiros, 12 Feb 1966; Irwin, Souza & dos Santos 9421 (F, G, MO, NY, 

RB, UB, US) circa 12 km NW of Veadeiros, Road to Cavalcante, 21 Oct 1965. 

91. Manihot weddelliana Baillon, Adansonia 4: 281. 1863-1864; Pax in Engler, 


Subshrubs. Stems glabrous. Leaves alternate; stipules caducous, filiform, glabrous; 

leaves subsessile, petioles less than 0.5 cm long, glabrous, petiole attachment to lamina 

basal, nonpeltate; lamina membranaceous, abaxial surface wax pattern reticulate, showing 

discrete rings; venation craspedodromous, veins glabrous; lamina nonlobed, nearly 

digitiform, ca 10.0 cm long, ca 1.5 cm wide, often with 2 short attenuate lobules near the 

base, margin repand or incised, apex acuminate, base acute to obtuse. Inflorescence a 

monoecious, terminal, short, subspicate raceme, ca 3.0 cm long, all parts except the 

interior surface of tepals glabrous; bracteoles foliaceous, ca 1.5 cm long, ca 0.6 cm wide, 

margins laciniate; bractlets foliaceous, ca 1.5 cm long, ca 0.4 cm wide, margins laciniate. 

Pistilliate flowers non vidi. Staminate tepal 1.0 cm long, interior surface sparsely 

pubescent, cleft 1/3 way down into 5 lobes, stamens 10, in 2 whorls of 5 each. Capsules 

and seeds non vidi. Fig 108C, D. 

TYPE. Weddell 2908: Brasil, Goias: entre Goyaz et Cujaba, Nov-Dec 1844 (syntypes, 

F, P). 

DISTRIBUTION. (Fig 108B).


// < 

(Malme~~~~~~~~~~ 

234 S)._7 M. ~ 

ateut ,ditiuin D ef(lziu216 ) 

l___:!_ 211 3 .. 4. .. ..5 6 7 : I? .....! ... 

FIG 107. Manibot salicifolia. A, leaf, obovate in (Maie outline 

2344, B, S); inflorescence 

(Malie 2344, S). M attenuata. . C, distribution; D, leaf (Glaziou 22126 G). , 

?


_ _ 

?^^^^^... I 

sI I * 

_ i 

1 / 

FIG 108. Manihot attenuata. A, inflorescence (Glaziou 22126, S). M. weddelliana. B, distribu- 

tion; C, leaf (Weddell 2908, P); D, inflorescence (Weddell 2908, P).


92. Manihot orbicularis Pohl, PI. Bras. Ic. et Descr. 1: 20. t. 12. 1827; Muell.-Arg. in DC., 



Jatropha orbicularis Steudel, Nomencl. ed. 2. 1: 799. 1840. 

Plant growth habit unknown. Young stems glabrous. Leaves alternate; stipules 

caducous; leaves subsessile, petioles less than 0.5 cm long, glabrous, petiole attachment to 

lamina basal, nonpeltate; lamina coriaceous, abaxial surface wax pattern smooth; venation 

camptodromous, veins glabrous; lamina nonlobed, orbicular, ca 15.0 cm long, ca 15.0 cm 

wide, apex obtuse cum acumine base obtuse. Inflorescence non vidi. Capsules and seeds 

non vidi. Fig 109B. 

TYPE. Pohl 1992: Brasil, Goias; ad Corgo Coral, non procul loco ubi flumen 

Maranhao trajicitur ad Agua-quente (syntypes, F, G, W-2). 

DISTRIBUTION. (Fig 109A). BRASIL. Goias: Pohl Icon Tab. 12 (W-2). 

17. Manihot sect Peltatae Pax emend Rogers & Appan 

Manihot sect Peltatae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 91. 1910. 

Manihot sect Indwisae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 93. 1910. 


petiole attachment basal or peltate, lamina membranaceous to coriaceous, nonlobed or 

very shallowly lobed. Inflorescence a monoecious subspicate raceme, single or 2 or 3 

racemes arising from a common base; bracts and bracteoles setaceous to foliaceous, 

margins usually entire. 

TYPE. Manihbot peltata Pohl. 


FIG 109. Manibot orbicularis. A, distribution; B, leaf (Pohl 1992, W).

eE- 

~........ 

' 

t-0 i ',K 

6?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

i~~~~~~~~~~~~~ 

d 

I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

i~~~ 

i~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

ii~~ 

FIG 110. Manihot sect Peltatae. Geographical range.


Key to the Species of Sect Peltatae 

1. Bracts and bracteoles setaceous, less than 0.25 cm wide; petiole attachment peltate; leaf 

shallowly stellate, cordate or triangular. 

2. Leaf shallowly stellate, more than 10.0 cm wide; petiole attachment very widely peltate, 

lamina between petiole-midrib junction and basal edge of leaf more than 3.0 cm wide. 

93. M. peltata. 

2. Leaf cordate to triangular, less than 10.0 cm wide; petiole attachment narrowly peltate, 

lamina between petiole-midrib junction and basal edge of leaf less than 1.0 cm wide. 

94. M. populifolia. 

1. Bracts and bracteoles foliaceous, more than 0.5 cm wide; petiole attachment basal; leaf 

reniform. 95. M. reniform is. 

93. Manihot peltata Pohl, P1. Bras. Ic. et Descr. 1: 20. t. 13. 1827; Muell.-Arg. in DC., 



Jatropha peltata Steudel, Nomencl. ed. 2. 1: 800. 1840. 

Subshrubs, ca 1.0 m tall; several ascending stems arising from a woody base. Stems 

glabrous. Leaves alternate; stipules caducous, filiform, glabrous; petioles 3.0-11.0 cm 

long, terete, glabrous, petiole attachment to lamina widely peltate, width between 

petiole-lamina junction and basal edge of lamina ca 6.0 cm; lamina coriaceous, abaxial 

surface wax pattern smooth; venation camptodromous, veins glabrous, major veins 

purplish tinged; lamina shallowly 6 lobed, stellate in outline, lobe sinus less than 1/2 the 

length of the lobes, lamina ca 18.0 cm long, ca 18.0 cm wide, apex of the shallow lobes 

obtuse inflorescence a monoecious, terminal, short, subspicate raceme, ca 6.0 cm long, all 

parts glabrous; bracteoles and bractlets setaceous, less than 0.3 cm long, less than 0.1 cm 

wide. Pistillate flowers restricted to the base of the upper half of the inflorescence, 

pedicels ca 0.75 cm long, tepal 0.8 cm long, cleft to base into 5 lobes, ovary subglobose. 

Staminate buds nearly tubular, flowers small, tepal 0.8 cm long, cleft 1/3 way down into 

5 lobes; stamens 10, in 2 whorls of 5 each. Capsules 1.1 cm long, dehiscence septicidal. 

Seeds 0.8 cm long, oblong, caruncle very small. Fig 111B, C. 

TYPE. Pohl 1661 (and some specimens also 1901): Brasil, Goias: ca Trahiras 

(syntypes, F, G, K-2, W-2). 

DISTRIBUTION. (Fig 111A). Brasil, state of Goias, Alt ca 1000 m, in cerrado. BRASIL 

Goias: Irwin, Souza & dos Santos 10937 (F, G, GH, K, MO, NY, RB, SP, UB, US) Cerrado circa 9 km 

S of Corumba de Goias, 2 Dec 1965; Pobl Icon Tab. 13 (W). Province Unknown/Uncertain: Burchell 

7663 (K); Burchell 7683 (K, P). 

94. Manihot populifolia Pax in Engler, Pflanzenreich IV. 147(Heft 44): 93. 1910. 

Manihot cordifolia Pax in Engler, Pflanzenreich IV. 147(Heft 44): 94. 1910. Type. Fiebrig 5297 

(syntype, G). 


petioles 10.0-15.0 cm long, terete, glabrous, petiole attachment to lamina narrowly 

peltate, width between petiole-lamina junction and basal edge of lamina ca 0.5 cm; lamina 

membranaceous to slightly coriaceous, abaxial surface wax pattern smooth; venation 

camptodromous, veins glabrous; lamina nonlobed, cordate to triangular, ca 8.0 cm long, 

ca 8.0 cm wide, apex acuminate to acute, base cordate to truncate. Inflorescence a 

monoecious, terminal, subspicate raceme, simple or a cluster of 2 or 3 racemes arising 

from a common base, ca 10.0 cm long, sometimes as long as 20.0 cm, all parts glabrous; 


flowers restricted to the base of the upper half of the inflorescence, pedicels ca 1.0 cm 

long, tepal 1.0 cm long, cleft to base into 5 lobes, ovary subglobose. Staminate buds 

ovoid-ellipsoid, flowers medium sized, tepal 1.0 cm long, cleft 1/2 way down into 5 lobes; 

stamens 10, in 2 whorls of 5 each. Capsules and seeds non vidi. Fig 112A, B.


,i 

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'S 

-f'i' 

CY)~~~~~~~~~~~~~~~~~~~~~~~~~~~~~C 

\? ~ ~~~~~~~ I 

_- , * 

A .. 

:,: 

........~ i 

::~ 9 

:: ? 

FIG 1 1 aibo 1 . ? 61); Pb M ,lea istiuin ett.A C, inlrsec (Iri,Su 

za &dos ants 1037, Y).M. ppulfoli. D,disribuion 

FIG 111. Manibot peltata. A, distribution; B, leaf (Pobi 1661, G); C, inflorescence (Irwin, Souza 

& dos Santos 10937, NY). M. populifolia. D, distribution.


' 

:: 

_ 

ODC 

". . 

1 ^ -4 ^ 

^ 

)\ /'itiliiuliiiniiiiiiisliil 'iii 

1) 

i iii iiiiiiiiiill 

FIG 112. Manihot populifolia. A, leaf, cordate in outline (Fiebrig 5297, G); B, leaf deltoid in 

outline (Hassler 10896, G). M. reniformis. C, distribution; D, leaf (Martius 1935, K). 


DISTRIBUTION. Hassler 10896: Paraguay: Sierra de Amambay (syntype, G-2). 

DISTRIBUTION. (Fig 111D). PARAGUAY. Province Unknown/Uncertain: Fiebrig 4669 

(BM, K, L) Zwischen Rio Apa Und Rio Aquidaban, Dec; Fiebrig 5297 (G) Zwischen Rio Apa Und 

Aquidaban. 

95. Manihot reniformis Pohl, PI. Bras. Ic. et Descr. 1: 56. 1827; Muell.-Arg. in DC., 



Jatropha reniformis Steudel, Nomencl. ed. 2. 1: 800. 1840. 


petioles ca 5.0 cm long, terete, glabrous, petiole attachment to lamina basal, nonpeltate; 

lamina coriaceous, abaxial surface wax pattern spongiose; venation camptodromous, veins 

glabrous; lamina nonlobed, rarely shallowly 3 lobed, reniform, ca 7.0 cm long, ca 7.0 cm 

wide, apex rounded, base cordate. Inflorescence a monoecious, terminal raceme, ca 7.0 

cm long, all parts glabrous; braceteoles foliaceous, ca 1.2 cm long, ca 0.6 cm wide, 

margins entire; bractlets foliaceous, ca 1.0 cm long, ca 0.5 cm wide, margins entire. 

Pistillate flowers restricted to the base of the inflorescence, pedicels ca 1.0 cm long, tepal 

1.2 cm long, cleft to base into 5 lobes, ovary subglobose. Staminate buds ovoid-ellipsoid, 

slightly constricted in the middle, flowers medium sized, tepal 1.2 cm long, cleft 1/3 way 

down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules 1.0 cm long, surface 

smooth, apex rounded, dehiscence septicidal. Seeds 8.0 cm long, oblong, carunculate and 

slightly tapering, caruncle not prominent. Fig 112D; 113A. 

TYPE. Martius 1935; Brasil, Bahia; Serra de Sincora, 1818 (syntypes, G, K, L-2, M-4, 

MO). 

DISTRIBUTION. (Fig 112C). BRASIL. Bahia: Froes 20121 (NY) Entre Brejao E Iracema, 

Campos Gerais, 20 Feb 1943; Froes 20167 (NY) Entre Brejao E Iracema, Campos Gerais Regiao Serra 

Sincora, 19 Feb 1943; Froes 20170 (NY) Entre Brejao E Iracema, Campos Gerais Regiao Serra 

Sincora; Froes 20174 (NY) Entre Brejao E Jiquy, Campos Gerais Regiao Serra Sincora, 18 Feb 

1943;Froes 20177 (NY, US) Entre Jiquy E Iracema, Campos Gerais, 18 Feb 1943; Ule 7085 (L) Serra 

de Sincora, Nov 1906. Province Unknown/Uncertain;: Collector Unknown/Uncertain, Icon. (W). 

18. Manihot sect Tripartitae Rogers & Appan, sect nov 

Manihot sect Grandibracteatae Pax subsect Tripartitae Pax in Engler, Pflanzenreich IV. 147(Heft 

44): 37. 1910 pro parte. 

Manihot sect Grandibracteatae Pax subsect Augustifoliae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 40. 1910 pro parte. 



Manihot sect Parvibracteatae Pax subsect Humiles Pax in Engler, Pflanzenreich IV. 147(Heft 44): 


Plantae caulinae, lignosae, infernae ad mediae frutices. Folia petiolatae, penitae 

lobatae. Inflorescentia monoecia racema. 


petiole attachment basal, lamina membranaceous, deeply lobed, lobes variously shaped 

but not linear. Inflorescence a monoecious raceme, single or a cluster of racemes arising 

from one common base; bracts and bracteoles foliaceous, margins laciniate. 

TYPE. Manihot tripartita (Sprengel) Muell.-Arg. emend Rogers & Appan. 


96. Manihot tripartita (Sprengel) Mueller von Argau in de Candolle, Prodr. 15(2): 1068. 

1866. 

Subshrubs ca 1.0 cm tall, several ascending stems from a single base. Young stems 

obtuse-angled in cross section, tomentose to glabrous, grayish brown or with purplish


cm long, less than 0.2 cm wide, tomentose, glabrous, margins laciniate; petioles ca 6.0 cm 

long, terete, glabrous to tomentose, petiole attachment basal, nonpeltate, 2 short (less 

than 0.3 cm long) filiform and tomentose lobules present at the base of the petiole-midrib 

junction; lamina membranaceous, abaxial surface wax pattern reticulate; venation 

camptodromous or craspedodromous, veins tomentose to glabrous; lamina 3 or 5 lobed; 

median lobes obovate, obovate-pandurate, elliptic, or narrowly lanceolate with incised 

margins, 5.0-10.0 cm long, 1.0-4.0 cm wide, margins undulate, entire or laciniate, apex 

acute to acuminate, base of lobes narrowly constricted, less than 0.3 cm wide, width 

between base of sinus and petiole-lamina junction less than 0.3 cm; lowest lobe slightly 

smaller than median lobes, slightly nonsymmetric. Inflorescence a monoecious, terminal, 

pendent raceme 5.0-10.0 cm long; peduncles and pedicels glabrous to tomentose; 

bracteoles prominently foliaceous 1.0-2.0 cm long, 0.5-0.9 cm wide, occasionally larger, 

yellowish green or with purple pigmentation, tomentose to glabrous, margins laciniate; 

bractlets foliaceous, ca 1.0 cm long, ca 0.5 cm wide, tomentose to glabrous, margins 

laciniate. Pistillate flowers restricted to the base of the inflorescence, pedicels ca 1.0 cm 

long, sometimes as long as 3.0 cm, glabrous to tomentose; tepal 1.0-2.0 cm long, 

yellowish green or with varying degrees of purple pigmentation, interior and exterior 

surface glabrous to tomentose, cleft to base into 5 lobes; ovary subglobose, glabrous to 

tomentose. Staminate buds ovoid-ellipsoid, flowers campanulate, tepal 1.0-2.0 cm long, 

glabrous to tomentose, yellowish green or with purple pigmentation, cleft 1/3 way down 

into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules ca 1.25 cm long, surface without 

wings, glabrous or pubescent, apex rounded, dehiscence septicidal. Seeds ca 1.0 cm long, 

oblong, caruncle moderately prominent. 

DISTRIBUTION. (Fig 115A). Brasil, Parar, states of Mato Grosso, Goias, Distrito 

Federal, Bahia, Minas Gerais, Sao Paulo and Rio de Janeiro; Paraguay. Alt up to 1000 m. 

In cut over gallery forest, in margins of forest, in cerrado, and on rocky slopes, etc.

f ' * S s. - .. -- 

X~~~~~~~~~~~~~~~~~~~~~~~~i 

"~ ' o-' t . 

j 

'-. 

_ 

?c, w! ... E .. I 

\ '-. ,':-'t. , 

1-_ ' ' i' X 

\~~~~~~~~~~~~~ 

FIG 114. Manio.. .. 

FI .14 *aio _ec Trprtte .egapia 

i * _ -_ 

_ _~~~~~~~~ I i ~ ~ 

< 

_ * | wfi I~~~~~~~~ 

\


The lengthy synonymy of Manihot tripartita and its subspecies indicates the 

complexity of this species. Most attention in the past had been given to the vegetative 

characters, particularly the pubescence and leaf form, and relatively little attention to the 

reproductive characters, or to geographic considerations. 

The unifying characters of the species are the inflorescence type, a pendant raceme, 

with foliaceous bracteoles whose margins are serrate or laciniate. 

Examination of the "Skyline," Fig 28, indicates that all the subspecies of this species 

have united at a level of 0.90 similarity. The subspecies Manihot tripartita subsp vestita 

and M. tripartita subsp laciniosa are the last members to join the larger species cluster. An 

uncritical application of the computer analysis would indicate that these two could be 

designated as separate species. However, the characters which play a role in differentiation 

within this complex are largely those characters describing pubescence, and there is 

considerable variation in these characters. The characters of the reproductive structures 

are essentially the same for all of the variants, and therefore, are considered as the better 

determining force for designation of the taxa. 

Key to the Subspecies of Manibot tripartita 

1. Leaf lobes usually more than 2.0 cm wide; margin entire or undulate, rarely pandurate; 

venation camptodromous. 

2. Plants glabrous or pubescent; population growing in the central region or southeastern 

periphery of the range of this species (Fig 115A). 

3. Plants pubescent; population growing in the central region of the geographical range of 

this species. 

96a. M. tripartita subsp tripartita. 

3. Plants glabrous; population growing in the southeastern periphery of the range of this 

species. 

96b. M. tripartita subsp humilis. 

2. Plants tomentose; population growing in the western periphery of the range of this 

species. 

96c. M. tripartita subsp vestita. 

1. Leaf lobes usually less than 2.0 cm wide; margin laciniate; venation craspedodromous. 

96d. M. tripartita subsp laciniosa. 

96a. Manihot tripartita (Sprengel) Mueller von Argau emend Rogers & Appan subsp 

tripartita 

Jatropha tripartita Sprengel, Syst. Veg. 3: 76. 1826; Adenoropium tripartitum Pohl, PI. Bras. Ic. 

et Descr. 1: 15. 1827. Manibot tripartita (Sprengel) Muell. Arg. var genuina Muell.-Arg. in 

DC., Prodr. 15(2): 1068; in Martius, Fl. Bras. 11(2): 476. 1874; Type. Sprengel s n n v. 

Manihot dalecbanpiaeformis Pohl, PI. Bras. Ic. et Descr. 1: 44. t. 36. 1827; Jatropha 

dalechampiaefornis Steudel, Nomencl. ed. 2. 1: 799. 1840. Manihot tripartita (Sprengel) 

Muell.-Arg. var dalechampiaeformis (Pohl) Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

38. 1910. Type. Pohl 1192 (syntype, W);Pohl 1193 (syntype, W). 

Manihot tomientella Pohl, PI. Bras. Ic. et Descr. 1: 45 t. 37. 1827;Jatropha tomentella Steudel, 

Nomencl. ed. 2. 1: 800. 1840. Type. Pohl 477 (syntype, W). 

Manihot cajanifornis Pohl, PI. Bras. Ic. et Descr. 1: 45. t. 38. 1827; Jatropha cajaniformis 

Steudel, Nomencl. ed. 2. 1: 799. 1840; Manihot tripartita (Sprengel) Muell.-Arg. var 

cajaniformis (Pohl) Muell.-Arg. in DC., Prodr. 15:(2): 1068. 1866; in Martius, Fl. Bras. 

11(2): 477. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 39. 1910. Type. Pobl 

3921 (syntypes, G, W). 

Manihot porrecta Pohl, PI. Bras. Ic. et Descr. 1: 46. t. 39. 1827; Jatropha porrecta Steudel, 

Nomencl. ed. 2. 1: 800. 1840; Manihot tripartita (Sprengel) Muell.-Arg. var porrecta (Pohl) 

Muell.-Arg. in DC., Prodr. 15(2): 1068. 1866; in Martius, Fl. Bras. 11(2): 477. 1874; Pax in 

Engler, Pflanzenreich IV. 147(Heft 44): 38. 1910. Type. Pohl 1191 (syntypes, F, G, M, 

W-2);Pohl 1656 (syntype, K). 

Manihot sinuata Pohl, PI. Bras. Ic. et Descr. 1: 48. t. 41. 1827; Muell.-Arg. in Martius, Fl. Bras. 

11(2): 482. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 45. 1910;Jatropha sinuata 

Steudel, Nomencl. ed. 2. 1: 800. 1840;Manihot sinuata Pohl vargenuina Muell.-Arg. in DC., 

Prodr. 15(2): 1074. 1866. Type. Pohl 1655 (syntypes, F, G, K, M, W-2). 

Manihot cleomaefolia Pohl, PI. Bras. Ic. et Descr. 1: 51. t. 44 1827; Jatropha cleomaefolia 

Steudel, Nomencl. ed. 2. 1: 799. 1840. Type. Pobl 1707 (syntypes, K-2, W); Pohl 3598 

(syntypes, M, W); Pobl 3722 (syntype, F).


Manihot tripartita (Sprengel) Muell.-Arg. var lanceolata Muell.-Arg. in DC., Prodr. 15(2): 1068. 

1866; in Martius, Fl. Bras. 11(2): 478. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 

38. 1910. Type. Sello s n (B2104 C2066?) (syntypes, BM, GH). 

Manihot tripartita (Sprengel) Muell.-Arg. var glauca Muell.-Arg. in DC., Prodr. 15(2): 1069. 1866; 

in Martius, Fl. Bras. 11(2): 477. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 39. 

1910. Type. Blanchet 3324 pro parte (syntypes, BM, F, K, NY, P, W). 

Manihot tripartita (Sprengel) Muell.-Arg. var seminuda Muell.-Arg. in Martius, Fl. Bras. 11(2): 

477. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 38. 1910. Type. Gardner 3445 

(syntypes, G, K). 

Manihot tripartita (Sprengel) Muell.-Arg. var apaensis Chodat & Hassler, Bull. Herb. Boissier 2(5): 

651. 1905; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 38. 1910. Type. Hassler 7955 

(syntypes, BM, F, NY, P, S, UC). 

Plant parts generally pubescent. Leaf lobes more than 2.0 cm wide, obovate, 

obovate-pandurate or elliptic, margins entire or undulate; venation camptodromous. 

Fig 115B, C. 

TYPE. Pohl 1192 (1193 appears on same specimens): Brasil, Goias: ad Corumba, 

Corgo do Jaragua, Ouro fino (neotype, W-2). (Neotype selected by Rogers & Appan. All 

the original material in the Berlin Herbarium studied by Sprengel which circumscribed 

Jatropha tripartita Sprengel are believed to have been destroyed.) 

BRASIL. Para: Ducke 15585 (MG, PUL) Alcobaco, margem da linha na campina da Estrada Breu 

Branco, 1 Jan 1915. Mato Grosso: Argent, Ramos, Richards & Souza 6674 (K) circa 1 km W of Base 

Camp at 12.49? S-51.460 W, 6 Oct 1967; Argent, Ramos, Richards & Souza 6721 (K) circa 6 km S of 

Xavantina (R/A Transect), 11 Oct 1967; Collenette 188 (K) Burity, NE of Coyuba, on open campo, 

Oct 1927; Harley & Souza 10287 (K) near Base Camp at 12.49? S-51.460 W, 27 Sep 1968;Harley, 

Souza & Fereira 11000 (K) Vale de Sonhas, S of Xavantina, 10 Nov 1968; Lima s n (SP) Campo 

Grande, 10 Oct 1944; Pereira & Egler 383 (PUL) Municipality De Corumba, Fazenda Aguassuzinho, 

17 Oct 1953; Philcox & Fereira 3898 (K) km 241 Xavantina-Cachimbo Road, 5 Jan 1968;Philcox & 

Fereira 3899 (K) km 241 Xavantina-Cachimbo Road, 5 Jan 1968;Philcox, Fereira and Bertoldo 3283 

(K, NY) circa 1 km W of km 247 Xavantina-Cachimbo Road, 28 Nov 1967;Pbilcox, Ramos & Souza 

3005 (K) 1 km E of km 265 Xavantina-Cachimbo Road, 14 Nov 1967; Philcox, Ramos & Sousa 3019 

(K) circa km 274 Xavantina-Cachimbo Road, 14 Nov 1967; Robert 420 (BM) Santa Anna da Chapada, 

20 Jul 1902; Robert 558B (K) Santa Anna da Chapada, 20 Sep 1902; Scbwacke 4520 (PUL, RB) 

Cuiaba. Goias: Brade 15403 (PUL) Goiania, Dec 1936; Gardner 3445 (G, K) near Almas, dry hill, Oct 

1829; Glaziou 22138 (G, K) Tiborno, entre Ciganos et as Brancas; Glaziou 22139 (F, G, K, S) 

Tiborno, entre Ciganos et as Brancas; Irwin, Maxwell & Wasshausen 18560 (F, IAN, MICH, MO, NY, 

S, SP, UB, US, W) 25 km N of Corumba De Goias on road to Niquelandia, 13 Jan 1968; Irwin, 

Maxwell & Wasshausen 18671 (F, G, GH, MO, NY, RB, SP, UB, UC, US) 15 km N of Corumba De 

Goias on road to Niquelandia, 16 Jan 1968; Irwin, Maxwell & Wasshausen 18924 (UB) 75 km N of 

Corumba De Goias on road to Niquelandia, Goias in Valley of Rio Maranhao, 21 Jan 1968; Irwin, 

Maxwell & Wasshausen 18983 (NY, UB) 75 km N of Corumba De Goias on road to Niquelandia, Goias 

in Valley of Maranhao, 22 Jan 1968; Irwin, Maxwell & Wasshausen 19036 (F, G, GH, MO, NY, RB, 

SP, UB, US) 75 km N of Corumba De Goias on Road to Niquelandia, 22 Jan 1968; Irwin, Maxwell & 

Wasshausen 21033 (F, G, GH, MICH, MO, NY, RB, SP, UB, US) Rio Das Lontras, Araguaina, 12 Mar 

1968; Irwin, Souza & dos Santos 10015 (F, NY, UB) 75 km N of Cristalina on road to Brasilia, 6 Nov 

1965; Irwin, Souza & dos Santos 10480 (B, F, G, GH, IAN, LE, MICH, MO, NY, RB, SP, UB, UC, US, 

W) Cerrado circa 5 km E of Cabeceiras, 18 Nov 1965; Irwin, Souza & dos Santos 10897 (F, IAN, 

MICH, NY, S, SP, UB, UC, US, W) Cerrado circa 11 km N of Corumba De Goias, 2 Dec 1965; 

Labouriau & Valio 1172 (SP) Municipality Goiatuba, 15 Feb 1964; Pohl Icon Tab. 36 (W); Pohl Icon 

Tab. 38 (W); Pohl Icon Tab. 41 (W); Pohl 1191 (F, G, M, W-2) Corgo do Jaragua, Ouro Fino; Pohl 

1655 (F, G, K, M, W-2) Corgo do Jaragua, Ouro Fino; Pobl 1656 (K); Pohl 3921 (G, W) Burriti 

Pequeno; Weddell 2960 (A, P-2) entre Goias et Cujaba, Nov-Dec 1844; Distrito Federal: Heringer 

8895/1089 (NY) Barragem do Paranoa, Brasilia, 17 Feb 1962; Irwin, Souza & dos Santos 11195 (F, G, 

GH, K, MO, NY, RB, SP, UB, US) campo slope E of Lagoa Paranoa, 11 Dec 1965. Bahia: Blanchet 

3324 (BM, F, K, NY, P, W) Igreja Velha, 1841; Blanchet 3324B (G-3, MG, P, W) Igreja Vehla, 1841. 

Minas Gerais: Assis 244 (GH, SP, UC, US) Fazenda da Chicaca, Municipality Santa Luzia, 13 Dec 

1945; Assis 52 (MO) Fazenda da Chicaca, Municipality Santa Luzia, 20 Nov 1945; Barreto 14.104 

(NY) Lagoa Santa, 20 Nov 1933;Barreto 2619 (F) Villa Cruzeiro de Sul, Municipality Belo Horizonte, 

10 Dec 1932; Barreto 2625 (F) Marzago, Municipality Belo Horizonte, 6 Jan 1934;Barreto 2626 (F) 

Alto do Cercado, Municipality Belo Horizonte, 21 Jan 1934; Barreto 5055 (F) Lagoa Santa, 

Municipality Santa Luzia, 20 Nov 1933: Barreto 8126 (F) Lagoa Santa Luzia, 26 Feb 1933;Barreto 

8127 (F) Cardoso, Municipality Belo Horizonte, 30 Nov 1935; Barreto & Brade 1598 (PUL) Lagoa


, ' (3 

A B? 

.U--t 

I,- IIIjIIjIIIIjI NII I IH lit iiiI 

FIG 115. Manibot tripartita. A, distribution; dots represent subsp tripartita, the letter A repre- 

sents subsp humilis, the letter O represents subsp vestita and the letter N represents subsp laciniosa. M. 

tripartita subsp tripartita. B, leaf (Argent, Ramos, Richards & Souza 6674, K); C, inflorescence (Irwin, 

Maxwell & Wasshausen 18560, NY). M. tripartita subsp bumilis. D, leaf (Warming 1520, G).


Santa, Dec 1957; Claussen s n (W) Lagoa Santa; Claussen s n (NY); Claussen 752 (GH, NY-2, P-6, S), 

1838; Claussen 753 (P), 1838; Claussen 953 (P), 1843; Claussen s n (G-2, P), 1839; Claussen s n (K), 

1840; Claussen s n (F); Cruz 120 (SP) Varzea da Palma, Fazenda Belgominas, Feb 1964; Cruz 126 (SP) 

Paraopeba, Nativa no Horto Florestal, 10 Feb 1966; Gehrt s n (SP) Belo !orizonte, 15 Dec 1918; 

Glaziou 18476A (P); Glaziou 20484 (P) entre Santa Luzia and Fiadade, 18 Nor 1893; Hoebne s n (SP) 

Areiao, Sta. Barbara, 21 Jan 1921; Irwin, Maxwell & Wasshausen 20422 (F, NY, SP, UB) Serra Do 

Cipo, circa 134 km N of Belo Horizonte, 18 Feb 1968;Joly 1139 (SP) Belo Horizonte, Pampulha, 15 

Jan 1951; Kublmann s n (PUL) Lagoa Santa, Nov 1951; Lutz 926 (US) Capella Nova do Betim, Mar 

1916; Macedo 4177 (S) S Vicente, Municipality Ituiutaba, 16 Jan 1956; Markgraf 3509 (F) Serra de 

Grao Mogol, Municipality Grao Mogol, 12 Nov 1938; Occhionissn (PUL) Arredores do Horto Florestal, 

Cerrado Sujo, 27 Nov 1940; Pires & Black 3048 (NY), Jan 1951; Pohl Icon Tab. 44 (W); Pohl Icon 

Tab. 57 (W); Pohl 1707 (K-2, W) Inter Coccaes et Saint Cruz, Alto do Morro Ourobranco; Pobl 3598 

(M, W) Inter Coccaes et Saint Cruz, Alto do Morro Ourobranco; Pohl 3722 (F) Inter Coccaes et Saint 

Cruz, Alto do Morro Ourobranco; Pohl 477 (W) Comorea Rio das Mortes in Serra Urubu; Regnell III 

1071 A (S) Uberava, 18 Jan 1849; Regnel III 1071 B (S-2) Uberava, 29 Nov 1849; Saint-Hilaire s n 

(P-6); Sello B2104 C2066 (BM, GH) Meridionali, Pompeo, 1836; Warming s n (P-2, US) Lagoa Santa; 

Williams & Assis 5689 (GH) Serra da Matuca, Municipality Nova Lima, Feb 1945; Williams & Assis 

6061 (GH, US) near Lagoa Pampulha, Municipality Belo Horizonte, 8 Mar 1945. Sao Paulo: Gebrt s n 

(SP) Emas, 16 Dec 1919;Lofgren 1030 (SP, P) Araraquara, 21 Nov 1888;Lofgren 1452 (SP) Cerrado. 

Rio De Janeiro: Glaziou 13201 (A, BM, G, K) 1882; Glaziou 13202 (K, P, US) environs of Rio 

Janeiro, Feb 1882; Saint-Hilaire 700 (P). Province Unknown/Uncertain: Riedel s n (K, NY, P-2, W). 

PARAGUAY. Province Unknown/Uncertain: Hassler 10704 (BM, F, NY, P, UC, W) in Altaplanitie et 

Declivibus, Sierra De Amambay, Nov; Hassler 7955 (BM, F, NY, P, S, UC) In Regione Cursus 

Superioris Fluminis Apa, Nov. 

LOCAL NAMES. Mandioca (Philcox & Fereira 3898); mandioca braba (Harley, Souza 

& Fereira 11000) manioca (Claussen s n). 

96b. Manihot tripartita (Sprengel) Mueller von Argau emend Rogers & Appan subsp 

humilis (Mueller von Argau) Rogers & Appan, stat nov 

Manihot bumilis Muell.-Arg. in Martius, Fl. Bras. 11(2): 448. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 58. 1910. 

Manihot tripartita (Sprengel) Muell.-Arg. var glabra Muell. Arg. in Martius, Fl. Bras. 11(2): 478. 

1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 39. 1910. Type. Warming s n (1520) 

pro parte (see also M. tripartita subsp laciniosa, 96d) (syntype, G); Riedel s n n v. 

Plant parts glabrous. Leaf lobes more than 2.0 cm wide, obovate or elliptic, margins 

entire or undulate; venation camptodromous. Fig 115D; 116A. 

TYPE. Riedel s n: Brasil, Goias: Chapadao de S. Marcos (Syntype, G). 

DISTRIBUTION. (Fig 115A). Brasil, states of Goias, Minas Gerais and Sio Paulo. BRA- 

SIL. Minas Gerais: Warming 1520 (G) Lagoa Santa. Sao Paulo: Brade s n (SP) Itirapina, 28 Sep 

1921; Cruz 123 (SP) Itirapina, 26 Jan 1966; Kublmann 2889 (SP) Sao Carlos, 27 Oct 1953;Lofgren 

991 (SP) Cerrado 1 Oct 1888; Toledo & Gebrt s n (SP) Itirapina, 25 Sep 1940. Province Unknown/ 

Uncertain: Freyrey s n (S). 

Insertae Sedis: Irwin, Maxwell & Wasshausen 21651 (NY) Goias, ca 27 km S of Paraiso, 23 Mar 

1968. Irwin & Soderstorm 6319 (NY) Mato Grosso, ca 2 km S of Xavantina, 25 Sep 1964. 

96c. Manihot tripartita (Sprengel) Mueller von Argau emend Rogers & Appan subsp 

vestita (S. Moore) Rogers & Appan, stat nov 

Manihot tripartita (Sprengel) Muell.-Arg. var vestita S. Moore, Trans. Linn. Soc. London Bot. 4: 

466. 1895; Pax in Engler Pflanzenreich IV. 147(Heft 44): 38. 1910. 

Manibot tricbandra Pax in Engler, Pflanzenreich IV. 147(Heft 44): 39. 1910. Type. Robert 558 

pro parte (syntypes, F, NY). 

Most plant parts tomentose. Leaf lobes more than 2.0 cm wide, obovate or elliptic, 

margins entire or undulate; venation camptodromous. Fig 116B, C. 

TYPE. Moore 186: Brasil, Mato Grosso: Serra de Chapada (syntype, NY). 

DISTRIBUTION. (Fig 115A). Brasil, state of Mato Grosso. BRASIL. Mato Grosso: Mal- 

me s n (S) Santa Anna da Chapada, 13 Aug 1902; Malme 2331 (S) Santa Anna da Chapada; Robert 

558 (F, NY) Santa Anna da Chapada, 20 Sep 1902.


ubs esia B la (am 231 ),i 

D, leaf (Sin--H---- . 

_........ . 

FIG 116. Manihot tripartita subsp humilis. A, inflorescence (L6fgren 991, SP). M. tripartita 

subsp vestita. B, leaf (Malme 2331, S);C, inflorescence (Malme 2331, S). M. tripartita subsp laciniosa. 

D, leaf (Saint-Hilaire s n, P).


Pax lists Robert 558 pro parte as the type of Manihot trichandra. There is no 

mention in Pax (1910) of the remaining part of Robert 558, but he may have referred to 

Robert 558B, which is identified in this monograph as M. tripartita, subsp tripartita (q v). 

96d. Manihot tripartita (Sprengel) Mueller von Argau emend. Rogers & Appan subsp 

laciniosa (Pohl) Rogers & Appan, stat nov. 

Manihot laciniosa Pohl, PI. Bras. Ic. et Descr. 1: 54. t. 48. 1827. Jatropha laciniosa Steudel, 

Nomencl. ed. 2. 2: 99. 1841. Jatropha lanciniosa Steudel, Nomencl. ed. 2. 1: 799. 1840 

(orthographic variant). Manihot sinuata (Pohl) emend. Muell.-Arg. var laciniosa (Pohl) Muell. 

Arg. in DC., Prodr. 15(2) 1075. 1866. Manihot laciniosa (Pohl) emend. Muell.-Arg. var 

genuina Muell. Arg. in Martius, Fl. Bras. 11(2): 483. 1874; Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 46. 1910. Type. Pohl 2207 (syntypes, F, G, W). 

Manihot lagoensis Muell.-Arg. in Martius, Fl. Bras, 11(2): 475. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 42. 1910. Type. Warming s n (1509) (syntypes, F, G). 

Manihot tripartita (Sprengel) Muell.-Arg. var. subintegra Muell.-Arg. in Martius, Fl. Bras. 11(2): 

478. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 39. 1910. Type. Warming s n 

(1520 pro parte (see also M. tripartita subsp humilis, 96b). 1520/2) (syntypes, F, G). 

Manihot laciniosa (Pohl) emend Muell.-Arg. var lanata Muell.-Arg. in Martius, Fl. Bras. 11(2): 

483. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 46. 1910. Type. Riedel s n 


Manihot intercedens Muell.-Arg. in Martius Fl. Bras. 11(2): 483. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 46. 1910. Type. Blanchet 3324 pro parte (syntypes, BM, G-3, OXF, 

P); Warming s n (1520/1) (syntype, F). 

Most plant parts pubescent, occasionally tomentose. Leaf lobes less than 2.0 cm 

wide, narrowly lanceolate, margins laciniate with several attenuate lobules along the 

lateral edges of lobes; venation craspedodromous. Fig 116D; 117A. 

TYPE. Pohl 2207: Brasil, Goias: ad Rio do Peixe (syntypes, F, G, W). 

DISTRIBUTION. (Fig 115A). Brasil, states of Goias, Bahia, Minas Gerais and Rio de 

Janeiro. BRASIL. Goias: Pobl Icon Tab. 48 (W). Bahia: Blanchet 3324 (BM, G-3, OXF, P) Igreja 

Vehla, 1841. Minas Gerais: Burrett 10.107 (F) Cocal, Municipality Pirapora, 18 Dec 1937; Riedel s n 

(F, G) in campis Curvellos; Warming 1509 (F, G) Lagoa Santa; Warming 1520 (G) Lagoa Santa; 

Warming 1520/1 (F) Lagoa Santa; Warming 1520/2 (F) Lagoa Santa. Rio de Janeiro: Saint-Hilaire s n 

(P). 

19. Manihot sect Caerulescentes Rogers & Appan, sect nov 


147(Heft 44): 29. 1910 pro parte. 

Manihot sect Grandibracteatae Pax subsect Grandiflorae Pax in Engler, Pflanzenreich IV. 

147(Heft 44): 32. 1910. 

Manihot sect Sinuatae Pax subsect Warmingianae Pax in Engler, Pflanzenreich IV. 147(Heft 44): 


Plantae caulinae, lignosae, altae frutices ad arbores. Folia petiolatae, penitae lobatae. 

Inflorescentia monoecia racema. 

Plants caulescent; tall shrubs to trees; leaves widely spaced on stem; petiolate, petiole 

attachment basal, lamina coriaceous, deeply lobed, lobes variously shaped but not linear. 

Inflorescence a monoecious raceme, single or a cluster of racemes arising from one 

common base; bracts and bracteoles foliaceous, margins entire. 

TYPE. Manihot caerulescens Pohl emend Rogers & Appan. 


Key to the Species of Sect Caerulescentes 

1. Leaves 3-5 lobed; median lobes cuneate, obovate, or elliptic; lobe apices obtuse, acute or 

cuspidate. 

97. M. caerulescens. 

1. Leaves usually 7 lobed, rarely 5, median lobes oblong-pandurate, lobe apices acuminate. 

98. M. beptaphylla.


9 . .. . ...... ..1827} 

__ a 

lA f-5 

FIG 117. Manihot tripartita subsp laciniosa. A, inflorescence (Blanchet 3324, G). 

97. Manihot caerulescens Pohl, PI. Bras. Ic. et Descr. 1: 56. 1827. 

Tall shrubs to trees, usually more than 3.0 m tall, with copious latex. Young stems 

glabrous, often purplish tinged; mature stems glabrous. Leaves alternate; stipules 

caducous, setaceous, less than 0.5 cm long, glabrous, margins entire; petioles 10.0-20.0 

cm long, terete, glabrous, often purplish tinged, petiole attachment to lamina basal, 

nonpeltate; lamina coriaceous, adaxial surface dark green, glossy, abaxial surface wax 

pattern showing discrete rings (at magnifications of x 40); venation camptodromous, 

veins glabrous, sometimes purplish tinged; lamina palmately 5 lobed, occasionally 3; 

median lobes cuneate, obovate or elliptic, ca 10.0 cm long, ca 5.0 cm wide, occasionally 

larger, apex obtuse, acute, or cuspidate, base of lobes narrowly constricted, less than 0.3 

cm wide, width between base of sinus and petiole-lamina junction 

X ....- 

less than 0.3 cm; 

lowest lobes slightly smaller than median lobes, slightly nonsymmetric. Inflorescence a 

monoecious, terminal raceme, 5.0-20.0 cm long, occasionally a cluster of 2 or 3 racemes 

arising from a common base, all parts glabrous; bracteoles prominently foliaceous, 

1.0-2.5 cm long, 0.5-0.9 cm wide, margins entire, bractlets foliaceous, ca 1.0 cm long, 

0.6 cm wide, margins entire. Pistillate flowers restricted to the base of the inflorescence, 

pedicels ca 1.5 cm long; tepal 1.0-2.0 cm long, yellowish green, rarely with purplish 

tinge, cleft to base into 5 lobes; disc prominent; ovary subglobose. Staminate buds 

ovoid-ellipsoid to conical, flowers campanulate; tepals 1.0-2.0 cm long, yellowish green, 

rarely purplish tinged, cleft 1/3-1/2 way down into 5 lobes; stamens 10, in 2 whorls of 5 

each. Capsules 1.5-2.5 cm long, prominently winged, apex prominently pointed, 

dehiscence septicidal. Seeds 1.0-1.5cm long, oblong, caruncle prominent. 

DISTRIBUTION. (Fig 119A). Brasil, Amapa Territory, and in the states Para, Bahia, 

Maranhao, Piaui, Ceara, Pernambuco, Mato Grosso, Goias, Minas Gerais and Rio de 

Janeiro; Paraguay. Cult. in Florida (Cook s n (NA-2)). In dry, interior uplands to altitudes 

ca 1000 m, in sertao and agreste (subdivisions of the caatinga). Soils generally sandy.

_.- 

\ ~ 

[' 

i ~ ~ .j 

?~.:.:. --/ _:. .,..........l||s}E 

I. 

:. 

_ - N X 

i 

~~~~~~~~~~~~~~~~~~~~~~~~~~~~.,.,,6,, 

- -~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

x 

>. 

\ * a 

FIG 118. Manio t Caelescens. _ Geographical _ range. 

i ' I 

_ _ ,, . 

l 

l I 

l 

, ...... _.. r. 

' ' l ' 

FIG .18 Maio sec Gegap a .ae 

s 

X 

Canlses 

\ 

\ < ____ 

_ _ 

l . ' 

i i , 

' ' ' 

l 

I R


. A 

. 

* ' 

_, ' .... 

k', V B 

C,.. 

FIG 119. Manihot caerulescens. A, distribution, dots represent subsp caerulescens, the letter A 

represents subsp paraensis and the letter O represents subsp macrantha. M. caerulescens subsp caerules- 

cens. B, habit (Rogers 404, NY); C, leaf (Rogers 398, NY); D, leaf of a M. esculenta cultivar. This 

appears to be a hybrid between M. esculenta and M. caerulescens subsp caerulescens (Rogers 405, NY).


Key to the Subspecies of Manihot caerulescens 

1. Inflorescence short to medium sized, usually less than 10.0 cm long; leaf lobes cuneate or 

obovate; population growing in the central or northern periphery of the range of this species 

(Fig 119A). 

2. Inflorescence medium sized, usually 5.0-10.0 cm long; leaf lobes cuneate; population 

growing in the central region of the range of this species. 

97a. M. caerulescens subsp caerulescens. 

2. Inflorescence short, usually less than 5.0 cm long; leaf lobes obovate; population 

growing in the northern periphery of the range of this species. 

97b. M. caerulescens subsp paraensis. 

1. Inflorescence long, vigorous, usually more than 10.0 cm long; leaf lobes elliptic; population 

growing in the southern periphery of the range of this species. 

97d. M. caerulescens subsp macrantha. 

97a. Manihot caerulescens Pohl emend Rogers & Appan subsp caerulescens 

Manibot caerulescens Pohl, PI. Bras. Ic. et Descr. 1: 56. 1827;Manihot coerulescens Pohl emend 

Muell.-Arg. var genuina Muell.-Arg. in DC., Prodr. 15(2): 1070. 1866; in Martius, Fl. Bras. 

11(2): 469. 1874; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 32. 1910. 

Jatropha coerulea Steudel, Nomencl. ed. 2. 2: 99. 1841. Jatropha coerulea Ind. Kew. 2: 1251. 

1895. 

Jatropha coerulescens Steudel in Muell.-Arg. in DC., Prodr. 15(2): 1070. 1866. (fide Muell.-Arg. 

"lapsu sub J. coerulea"). 

Manihot coerulescens Pohl emend Muell.-Arg. var pubescens Muell.-Arg. in DC., Prodr. 15(2): 


147(Heft 44): 32. 1910. Type. Gardner 1837 (syntypes, BM, NY, P-2, US, W-2). 

Manihot speciosa Muell.-Arg. in Martius, Fl. Bras. 11(2): 470. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 34 1910. Type. Riedel 2822 (syntypes, F, G, K, NY, P); Manihot 

riedeliana Klotzsch ex Pax in Engler, Pflanzenfiech IV. 147(Heft 44): 34. 1910 pro syn. 

Manihot grandiflora Muell.-Arg. in Martius, Fl. Bras. 11(2): 471. t. 66. 1874; Pax in Engler, 

Pflanzenreich IV. 147(Heft 44): 34. 1910. Type. Warming s n (1502) (syntypes, G, P-2); 

Riedel 791 (syntypes, G-2, K). 

Manihot piauhyensis Ule, Notizbl. Bot. Gart. Berlin 5(41): 2. 1907; 5(41a): 18. 1908. Type. Ule 

7141a (syntype, UC); Ule 7141b n v; Ule 7141 (7141c) (syntypes, G-2, K, L, MG-2, UC, 

US). 

Manihot toledi Labroy ex Ule, Bot. Jahrb. Syst. l(Beibl. 114): 8. 1914. Type. Zehntner 695 

* (syntypes, F, NY); Zehntner 694 n v; Zehntner 692 n v. 

Manihot cuneata Ule, Bot. Jahrb. Syst. l(Beibl. 114): 1. 1914. Type. Zehntner 394 (syntypes, F, 

NY). 

Manibot ferruginea Ule, Bot. Jahrb. Syst. l(Beibl. 114): 2. 1914. Type. Zehntner 611 (syntypes, 

F, NY). See remarks for 74. M. leptophylla Pax & Hoffm. 

Manihot trifoliata Ule, Bot. Jahrb. Syst. 1(Beibl. 114): 3. 1914. Type. Zebntner 533 (syntypes, 

F-2, NY): Zehntner 532 n v; Zehntner 530 n v. 

Manibot rotundata Ule, Bot. Jahrb. Syst. 1(Beibl. 114): 4. 1914. Type. Zehntner 390 (syntypes, 

F, NY); Zehntner 382 n v. 

Manihot bahiensis Ule, Bot. Jahrb. Syst. 1(Beibl. 114): 4. 1914. Type. Zehntner 365 (syntypes, 

F, NY); Zehntner 371 n v. 

Manihot microdendron Ule, Bot. Jahrb. Syst. 1(Beibl. 114): 5. 1914. Type. Zebntner 689 

(syntypes, F, M, NY). 

Manihot labroyana Ule, Bot. Jahrb. Syst. 1(Beibl. 114): 6. 1914. Type. Zebntner 238 (syntypes, 

F, NY); Zebntner 275 n v; Zehntner 290 n v. 

Manihot discolor Ule, Bot. Jahrb. Syst. l(Beibl. 114): 7. 1914. Type, Zehntner 690 (syntypes F, 

NY). 

Manihot harmsiana Ule, Bot. Jahrb. Syst. 1 (Beibl. 114). 7. 1914. Type. Zehntner 691 (syntype, 

F. NY). 

Manihot lyrata Ule, Bot. Jahrb. Syst. 1(Beibl. 114): 9. 1914. Type. Zehntner 299 (syntypes, F, 

NY); Zebntner 282 n v. 

Manihot cearensis Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 85): 194. 1924. 

Type. Lofgren 327 (syntype, S). 

Leaf lobes cuneate. Inflorescence medium sized, usually 5.0-10.0 cm long. 

Fig 119B, C, D; 120A, B.


C FIG 12 ot caerulescens subsp caerulescen A, inflo 

't 

FIG 120. Manibot caerulescens subsp caerulescens A, inflorescence (Chase 7803, F); B, promi- 

nently winged fruits (Rogers 397, NY). M. caerulescens subsp paraensis. C, leaf (Ducke s n, PUL); D, 

inflorescence (Pires, Rodrigues & Irvine 51058, NY).


TYPE. Martius s n: Brasil, Bahia: Malhada ad Sincora (syntypes, photo at F, G, L, 

M-3). 

DISTRIBUTION. (Fig 119A). Brasil, states of Para, Maranhao, Piaui, Ceara, Pernambuco, 

Mato Grosso, Goias, Bahia, Minas Gerais and Rio de Janeiro. Cult. in Florida (Cook 

s n (NA)). BRASIL. Para: Pires, Black, Wurdack & Silva 6118 (NY) Serro do Cachimbo, 12 Dec 

1956; Pires, Black, Wurdack & Silva 6284 (NY) Serra do Cachimbo, 15 Dec 1956. Maranhao: Lisboa 

2343 (MG) Mirador, 18 Aug 1909; Lisboa 2363 (MG), 2 Sep 1909. Piaui: Zehntner 689 (F, M, NY) 

Caatingazone, Serra Branca. Ceara: Ducke s n (PUL) M. Calmon, 1909;Gardner 1837 (BM, NY, P-2, US, 

W-2) Serra de Araripe, Oct 1838; Lisboa 2431 (MG) Serra Tbiapaba, 16 Oct 1909; Lofgren 327 (S) S. 

Felix, Canascal, 18 Mar 1910; Luetzelburg 26414 (NY, US) Serra de Araripe, Araras, Carasco, 11 

Feb 1935. Pernambuco: Chase 7803 (F, US) Garanhuns, Nov 1924; Rogers 397 (COLO, F, G, K, 

MICH, NY, US, W) San Jose do Belmonte (formerly Manicobal) 3 Mar 1961; Rogers 404 (NY) 

Araripina Agriculture Experimental Station in chapada, 4 Mar 1961; Rogers 408 (G, NY, US) circa 50 

km SW of Crato on chapada, 5 Mar 1961. Mato Grosso: Harley, Lima, Souza & de Castro 10533 (K) 

near Base Camp at 12.49? S-51.460 W, 7 Oct 1968;Irwin, Grear, Souza & dos Santos 16210 (F, NY, 

SP, UB) Cerrado Rio Turvo circa 210 km N of Xavantina, 28 May 1966; Irwin, Grear, Souza & dos 

Santos 16461 (F, NY, UB) wet campo circa 84 km N of Xavantina, 2 June 1966; Philcox & Fereira 

4609 (K) 2 km W of km 264 Xavantina-Cachimbo Road, 22 Mar 1968; Philcox, Fereira & Bertoldo 

3340 (K) between main Xavantina road, Base Camp at 12.49? S-51.460 W, 2 Dec 1967; Philcox, 

Ramos & Souza 3109 (K) km 264 Xavantina-Cachimbo Road, 17 Nov 1967; Ratter, de Castro, dos 

Santos & Souza 834 (K) near Base Camp at 12.54? S-51.520 W, close to Xavantina-Sao Felix Road, 6 

Apr 1968; Ratter, de Castro, dos Santos & Souza 835 (K) near Base Camp at 12.54? S-51.520 W, 

close to Xavantina-Sao Felix Road, 6 Apr 1968. Goias: Irwin, Grear, Souza & dos Santos 14450 (F, G, 

GH, MO, NY, RB, SP, UB, US) cerrado Rio da Prata circa 6 km S of Posse, 6 Apr 1966. Bahia: Irwin, 

Grear, Souza & dos Santos 14609 (F, MICH, MG, MO, NY, S, SP, UB, UC, US, W) cerrado Rio Piau 

circa 225 km SYW of Barreiras on road to Posse, 12 April 1966; Irwin, Grear, Souza, & dos Santos 

14821 (F, GH, K, MO, NY, RB, SP, UB, UC, US) cerrado near Rio Piau circa 150 km SW of Barreiras, 

14 Apr 1966; Luetzelburg 12674 (NY) Tamandina, Catinga-Gebiet, 1914; Luetzelburg 1348 (NY) 

Serra do Rio Preto, 1913; Pereira 2055 (RB), 15 Sep 1956; Ule 7141 (G-2, K, L, MG-2, UC, US) bei 

Remanso cultivated, Dec 1906; Ule 7141A (UC), 1906; Zehntner s n (M-2) Chapada Velha; Zehntner 

238 (F, NY); Zehntner 299 (F, NY) Licury Und Gruna, Aug 1912; Zebntner 365 (F, NY) Santa Rita, 

16 Oct 1912; Zehntner 390 (F, NY) bei Santa Rita, Oct 1912; Zehntner 394 (F, NY): Zehntner 533 

(F-2, NY); Zehntner 611 (F, NY); Zehntner 690 (F, NY), bei Villa Nova; Zehntner 695 (F, NY) bei 

Villa Nova, June 1911. Minas Gerais: Barreto 9449 (F-2) Municipality Diamantina, 4 Nov 1937; 

Claussen s n (NY); Magalhaes 14887 (PUL, RB) Virgem da Lapa, Apr 1959; Riedel 791 (G-2, K); 

Warming 1502 (G, P-2) Lagoa Santa. Rio de Janeiro: Kuhlmann s.n. (PUL, RB) Jardim Botanico, 10 

Sep 1946; Saint-Hilaire s n (P-3). Province Unknown/Uncertain: Claussen s.n. (NY); Pohl 56 (W); 

Riedel 2822 (F, G, K, NY, P) inter Rio da Padre et Rio Sao Francisco. 

Cult: USA, Florida, Cook s n (NA) Homestead, Dade County, 20 Aug 1926; Cook s.n. (NA) 

Johnson's Place, Homestead, Dade County, May 1929. 

LOCAL NAMES. Manidoca de viado (Philcox, Fereira & Bertoldo 3340), mandioca 

braba (Harley, Lima, Souza & deCastro 10533) manicoba (Rogers 397), manicoba brava 

(Zehntner 690), manicoba de veado (Lisboa 2343), manicoba von gruna (Zehntner 299). 

USES. Source of latex (see remarks). 

The large synonymy of Manihot caerulescens subspecies cearulescens is due, in part, 

to the fact that the plants have been selected, cultivated and modified from wild plants in 

eastern Brasil. Variants, clones (or cultivars) have been selected for their production of 

latex, and it seems that this practise had its origins long before the great rubber boom at 

the end of the 19th and beginning of the 20th century. We base this hypothesis in large 

part on the fact that the natives of the region made a number of artifacts from the latex, 

such as toy rubber balls, shoes, and occasional other garments, in a region where Hevea 

brasiliensis Muell.-Arg. does not grow well. 

Manihot caerulescens has a "weedy" aspect. There are numbers of marginally 

developed (or poorly cared for) plantations, and nearby, in the bush, will be found 

various individual volunteer plants in a wide variety of forms. It is possible, but by no 

means certain, that these forms are the basis for the many names applied, particularly by 

Ule.


The color of the latex and the color of the seeds vary from white to cream to yellow, 

and the manner of collection of the latex varies from location to location. Each of these 

variations is given a local name, and these local names have also, undoubtedly, influenced 

the various botanists who have described the variants as species or varieties. Mario Barroso 

Ramas has written an interesting paper (title-"Borracha", Sao Paulo, 1951?) describing 

the latex producing species of Manihot, and some of their properties. 

The type materials of Manihot trifoliata Ule var platyphylla Ule and M. bahiensis Ule 

var microsperma Ule were not available for this study, and hence the taxonomic status 

could not be evaluated. However, considering the fact that most of the Ule (1914) species 

are only phenotypic forms of M. caerulescens subsp caerulescens, it is very likely that 

these taxa are also a synonym of M. caerulescens subsp caerulescens. 

97b. Manihot caerulescens Pohl emend Rogers & Appan subsp paraensis (Mueller von 


Manihot paraensis Muell.-Arg. in Martius, Fl. Bras. 11(2): 470. 1874; Pax in Engler, Pflanzenreich 

IV. 147(Heft 44): 32. 1910. 

Leaf lobes obovate. Inflorescence short, usually less than 5.0 cm long. Figs 120C, D, 

TYPE Spruce 186: Brasil, Para: vicinibus Santarem, Jan 1850 (syntypes, BM, F, G-3, 

K-2, M, NY-2, OXF, W). 

DISTRIBUTION. (Fig 119A). Brasil, Amapa Territory and in the state Para. BRASIL. 

Amapa Territory: Pires, Rodriques & Irvine 51058 (NY) Rio Araguari, 19 Sep 1961. Para: Ducke s n 

(MG, PUL, RB) Monte Alegre, Serra do Paitum, 5 Mar 1916;Pires, Black, Wurdack & Silva 6477 (NY) 

Santarem, 25 Dec 1956; Spruce 622 (P) Santarem, Jan 1850. 

97c. Manihot caerulescens Pohl emend Rogers & Appan subsp macrantha (Pax & K. 

Hoffmann) Rogers & Appan, stat nov 

Manihot macrantha Pax & K. Hoffmann in Engler, Pflanzenreich IV. 147(Heft 44): 32. 

1910; Manihot speciosa Chodat & Hassler, Bull. Herb. Boissier 2(5): 673. 1905 nom 

rejic. (The epithet speciosa has been used by Muell.-Arg. in Martius, Fl. Bras. 11(2): 

470. 1874.) 

Leaf lobes elliptic. Inflorescence long, usually ca 15.0 cm Fig 121A, B. 

TYPE. Hassler 5132: Paraguay: Sierra de Maracayu, Oct 1900 (syntypes, BM, F, 

photo at F, G-7, NY, photo at NY, P, S, UC, W). 

DISTRIBUTION. (Fig 119A). PARAGUAY. Province Unknown/Uncertain: Rojas 6792 (A) 

Cerro Cora, Sierra De Amambay, Mar 1934. 

98. Manihot heptaphylla Ule, Notizbl. Bot.Gart. Berlin 5(41): 2. 1908; 5(41a): 17, 21,f. 

2A-B. 1908; Pax in Engler, Pflanzenreich IV. 147(Heft 44): 43. 1910. 

Manihot beptaphylla X piauhyensis Ule, Bot. Jahrb. Syst. l(Beibl. 114): 6. 1914. Type. Zehntner 

381 (syntypes, M); Zehntner 385n v. 

Trees, 2.0-8.0 m tall. Stems glabrous. Leaves alternate; stipules caducous; petioles ca 

12.0 cm long, terete, glabrous, petiole attachment to lamina basal, nonpeltate; lamina 

coriaceous, adaxial surface dark green, abaxial surface wax pattern showing discrete rings 

(at magnifications of X 40); venation camptodromous, veins glabrous; lamina palmately 7 

lobed, rarely 5; median lobes oblong-pandurate, ca 10.0 cm long, ca 3.0 cm wide, apex 

acuminate, base of lobes narrowly constricted, less than 0.3 cm wide, width between base 

of sinus and petiole-lamina junction less than 0.3 cm; lowest lobes ca 1/2-2/3 as long as 

median lobes. Inflorescence a monoecious, terminal raceme, simple or a cluster of 2 

racemes arising from a common base, ca 6.0 cm long, all parts glabrous; bracteoles 

foliaceous, ca 1.5 cm long, ca 0.8 cm wide, margins entire; bractlets foliaceous, ca 1.0 cm 

long, ca 0.5 cm wide, margins entire. Pistillate flowers restricted to the base of the


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FIG 121. Manibot caerulescens subsp macrantba. A, leaf (Hassler 5132, UC); B, inflorescence 

(Hassler 5132, UC). M. beptapbylla. C, distribution; D, leaf (Ule 7206, G). 

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inflorescence, pedicels ca 1.0 cm long; tepal 1.2 cm long, cleft to base into 5 lobes; ovary 

subglobose. Staminate buds ovoid-ellipsoid, flowers medium sized, tepals 1.2 cm long, 

cleft 1/3 way down into 5 lobes; stamens 10, in 2 whorls of 5 each. Capsules and seeds 

non vidi. Fig 121D; 122A. 

TYPE. Ule 7206: Brasil, Bahia: Serra do Sao Ignacio am S. Francisco, Feb 1907 

(syntypes, G-2, K, L, MG, UC, US). 

DISTRIBUTION. (Fig 121C). BRASIL. Bahia: Zehntner 381 (M) Santa Rita, Caatingazone; 

Irwin, Harley & Smith 30966 (NY) ca 24 km N of Seabra, road to Aqua de Rega, elev ca 1000 m, 24 

Feb 1971. 

2. Manihotoides D. J. Rogers & S. G. Appan, gen nov 

TYPE SPECIES. Manihotoides pauciflora (T. S. Brandegee) Rogers & Appan. 

Arbores; cortex fusco-castaneus; folia trisecta lobis medianis ca 2.0 cm longis, ca 

2.25 cm latis; inflorescentia uniflora; flos staminatus 2.0-2.5 cm longus; capsulae 

globosae diametro ca 1.5 cm;semina ca 1.25 cm longa. 

Low trees ca 2.5 m tall, profusely branched from base onwards, branching not 

regularly dichotomous or trichotomous but very irregufar; outer and basal branches often 

lax, foliage characteristically borne on short, lateral stems with telescoped internodes. 

Stems glabrous, smooth, shiny, succulent, dark grayish brown. Leaves alternate; stipules 

persistent, pubescent, 0.2-0.3 cm long; petioles ca 3.0 cm long, occasionally longer, 

slender, greenish, glabrous, ventral surface shallowly canaliculate; lamina nonpeltate, 

surfaces green to pale green, glabrous, abaxial surface wax pattern smooth; venation 

camptodromous, lamina deeply lobed, appearing ternately compound, the lobes obovate 

FIG 122. Manihot heptaphylla. A, inflorescence (Ule 7206, G).


to nearly obcordate with a truncate apex, ca 2.0 cm long, ca 2.25 cm wide; base of lobes 

very narrowly constricted, the lobes frequently pendent. Inflorescence uniflorous, rarely 

2-flowered, monoecious, usually borne at the apex of the characteristic short stems, 

pedicels ca 2.0 cm long, greenish, glabrous, bearing 2 setaceous, pubescent bractlets, ca 

0.2 cm long. Pistillate tepals yellowish green, glabrous, ca 1.8 cm long, cleft into 5 

oblong, lanceolate, caducous lobes; disc fleshy, entire, pistil about 0.8 cm long; ovary 

subglobose, glabrous, the trifid stigma moderately lobed and lobulate. Staminate flowers 

nearly tubular; tepals usually 2.0-2.5 cm long, rarely shorter, yellowish green, glabrous, 

deeply cleft nearly to the base into 5 oblong-lanceolate lobes, slightly reflexed at anthesis; 

disc fleshy, shallowly 10 lobed; stamens 10, in 2 whorls, superior whorl ca 1.6 cm long, 

inferior whorl ca 1.3 cm long, anthers dehiscing longitudinally, cream colored, ca 0.3 cm 

long, dorsifixed, the point of attachment of the cream colored filament below center of 

anther. Fruit borne singly on pedicels 1.0-2.0 cm long; capsules more or less subglobose 

ca 1.5 cm long from base to apex, surface slightly verruculose, ribs not prominent, apex 

rounded; fruit dehisence septicidal. Seeds ca 1.25 cm long, oblong, carunculate end 

tapering and acutely pointed, surface shiny, grayish with dark mottlings, caruncle not 

prominent, trapeziform. Fig 123B, C, D; 124A, B. 

TYPE. Purpus 3418 (holotype UC; isotypes F, MO, NY, UC, US) 

DISTRIBUTION. (Fig 123A). Mexico: Puebla and Oaxaca. Narrowly localized in the 

upper and interior regions of the steep canyons formed by the tributaries of Rio Santo 

Domingo, southeast of Tehuacan City, at altitudes of 1000-1800 m. Habitat xerophytic, 

with thorn scrub cactus cover on igneous and sedimentary rocks, with friable light red 

soil. 

This taxon was described and designated as a species of Manihot (Manihot pauciflora 

T. S. Brandegee) in 1910. In the computer aided similarity-graph-clustering analysis, the 

13 specimens representing this taxon form a homogeneous cluster at a high "C-value" of 

.918 (Fig 5). This cluster then remains distinct, with an enormous "moat", till a very low 

"C-value" of .541, thus manifesting a remarkable phenotypic distinctness. 

The genus has several unique morphological attributes, which have not developed in 

any species of Manibot. The characteristic uniflorous inflorescence (Fig 123D) of 

Manihotoides (all the species of Manihot have many flowered inflorescences) is usually 

considered as a primitive character. The leaves are uniformly 3 lobed (Fig 123C) and are 

morphologically distinct. The foliage is borne characteristically on short, condensed stems 

arising from branchlets, a characteristic not occurring in any species of Manihot. Standley 

(1923) indicated that this taxon is very unlike other Manihot species, and the leaves 

resemble some species of Oxalis. Based on these indications this taxon has been raised to 

the status of a monotypic genus. 

1. Manihotoides pauciflora (T. S. Brandegee) D. J. Rogers & S. G. Appan, comb nov. 

Manihot pauciflora T. S. Brandegee, Univ. Calif. Publ. Bot. 4: 89. 1910. 

See generic description. 

TYPE Purpus 3418: Mexico, Puebla: vicinity of San Luis Tultitlanapa, near Oaxaca, 

Jun 1908 (holotype, UC; isotypes, F, MO, NY, UC, US). 

DISTRIBUTION. (Fig 123A). Mexico, Puebla, Oaxaca. MEXICO. Puebla: Purpus 5841 

(DS, UC), Tehuacan, June 1912; Rose, Painter & Rose 10114 (US) near Tehuacan, Aug.-Sep 1905; 

Smith, Peterson & Tejeda 3563 (F, TEX) Tehuacan area, near Coxcatlan on Cerro Ajuereado, July 

1961. Oaxaca: Conzatti 4130 (US) Cuesta de San Bernardina, 25 May 1921; Kimnach & Moran 161 

(UC) 4.8 miles beyond Teotitlan to Tecomavaca, 17 Nov 1959.


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FIG 123. Manihotoides pauciflora. A, distribution; B, a stem with fruits; C, leaf (Purpus 3418, 

UC); D, uniflorus inflorescence (Purpus 3418, UC). 

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FIG 124. Manibotoides pauciflora. A, habit (photo by C. E. Smith); B, branching habit (photo 

by C. E. Smith). 

-


Species Dubia 

Manihot bahiensis Ule var microsperma Ule, Bot. Jahrb. Syst. 1(Beibl. 114): 5. 1914: 

TYPE. Zehntner 384, 387, and 455. West Bahia Bei Santa Rita wild wachsend and 

angepflanzk, Oct 1912, non vidi. See remarks under 97a, M. caerulescens subsp 

caerulescens, 

Manihot hilariana Baillon, Adansonia 4: 282. 1864; Muell.-Arg. in DC., Prodr. 15(2): 

1065. 1866; in Fl. Bras. 11(2). 464. 1874; Pax in Engler Pflanzenr. IV. 147(Heft 44: 

79. 1910. 

TYPE. Hilaire cat B2, n. 2328. Brasil Minas Gerais, non vidi. 

Manibot johannis Pax in Engler Pflanzenreich IV. 147(Heft 44): 78. 1910. 

TYPE. Martius s n (M, G), Brasil, Bahia, in sylvis catingae inter Malhada et Sincora. 

Pax correctly renamed this species because the epithet, Manihot pohliana 

Muell.-Arg. had been preempted by M. pohlii Wawra. 

The fragmentary material available to us, and the very general description 

provided did not permit proper classification in the main body of this study. 

Manihot membranaceae Pax & Hoffmann in Engler, Pflanzenreich IV. 147(Heft 47): 111. 

1911. 

TYPE. Hoehne 1380, Brasil Matto Grosso, non vidi. 

Manihot pruinosa var pumila Muell.-Arg. in DC. Prodr. 15(2): 1061. 1866; in Fl. Bras. 

11(2) 453. 1874; Pax in Engler Pflanzenr. IV. 147(Heft 44): 62. 1910. 

TYPE. Reidel 2824 Brasil, Goias, non vidi. 

Manihot teissonieri A. Chevalier, Jour. Agr. Trop. 7: 357. 1907. 

TYPE. No type designated, no specimens cited. 

Chevalier described this species from plants growing in West Africa. In the 

description, Chevalier indicated that he thought the plant was similar to M. 

piauhyensis Ule. The description of the fruit ("une baie indehiscente et non une 

capsule") is not admissable for Manihot. Furthermore, the description of the seed 

is confusing..... "une graine elliptique lenticulaire, entierement recouverte d'une 

mince pulpe jaunatre (caroncle) ....." There is indeed a caruncle on the seeds of 

Manihot, but covering only the micropyle, not the whole seed. 

Manihot trifoliata Ule var platyphylla Ule, Bot. Jahrb. Syst. l(Beibl. 114): 3. 1914. 

TYPE. Zehntner 456a Bahia. Chapada do Jatoba, Oct 1912, non vidi. 

See remarks under 97a, M. caerulescens subsp caerulescens. 

Dubious or Excluded Taxa and Incorrect Author Names 

Jatropha angustifolia Steudel, Nomencl. ed. 2. 99. 1841. Steudel attributed this epithet 

to Pohl incorrectly. Pohl never published such a name in the genus Manihot. nomen 

nudum. 

Jatropha palmata Sesse & Mociino apud Cervantes. nomen dubium. See p 3. 

Jatropha silvestris. nomen dubium. See p 3, under Vellozo. 

Jatropha sylvestris. Steudel. orthographic variant of preceding. 

Jatropha triloba. nomen dubium. See p 3, under Sesse & Mocifio. 

Manihot aipi Rusby. incorrect author citation. Pax (Pflanzenr. IV. 147(Heft 44): 67. 

1910) incorrectly cites Rusby as author of epithet, while Rusby (Mem. Torrey Bot. 

6: 120. 1896) cites Pohl as the author. 

Manihot angustifolia. nomen nudum. See Jatropha angustifolia, above. 

Manibot berroana Beaverd, Bull. Soc. Bot. Geneve II. 4: 69. 1912, nomen nudum. 

Manihot cassava Cook & Collins, Contr. U. S. Nat. Herb. 8: 184. 1903. nomen nudum. 

Manihot coerulea Pohl (fide Steudel). nomen nudum. Pohl did not publish this name 

under Manihot. 

Manihot consanguinea Klotzsch in Sched. (Pax, Pflanzenr. IV. 147(Heft 44): 38. 1910). 

nomen nudum.


Manihot curcas Crantz, Inst. Rei Herb. 1: 167. 1766. = Jatropba curcas. 

Manihot digitata Sweet, Hort. Brit. ed. 2. 458. 1830. = Jatropha fischeri Steudel, fide 

Pax, Pflanzenr. IV. 147(Heft 44): 99. 1910. 

Manihot diversifolia Sweet, Hort. Brit. ed. 2. 458. 1830. = Jatropha diversifolia Steudel, 

fide Pax, Pflanzenr. IV. 147(Heft 44): 99. 1910. 

Manihot gossypifolia Crantz, Inst. Rei Herb. 1: 167. 1766. = Jatropha gossypifolia. 

Manihot guyanensis Klotzsch in Sched. (Pax, Pflanzenr. IV. 147(Heft 44): 84. 1910). 

nomen nudum. 

Manihot berbacea Crantz, Inst. Rei Herb. 1: 167. 1766. = Jatropha urens. 

Manihot heterophylla Chodat & Hassler. Species epithet wrongly attributed to these 

authors by Pax, Pflanzenr. IV. 147(Heft 44): 86. 1910. 

Manihot japonica Semler, Trop. Agricult. 2: 614. 1887. nomen nudum. 

Manihot moluccana Crantz, Inst. Rei Herb. 1: 167. 1766. = Aleurites triloba? 

Manihot multifida Crantz, Inst. Rei Herb. 1: 167. 1766. = Jatropha multifida. 

Manihot neoglaziovii Pax & K. Hoffmn. ex Luefzelb., Estud. Bot. Nordeste Braz. 3: 141. 

1923. 

Manihot pohliana Muell. Arg. Name preempted by M. pohlii Wawra. See p 251. 

Manihot preciosa Hort. (M. preciosa Schindler, fide Gray Card Index). Kew Bull. p 196. 

1910. nomen nudum. 

Manihot spinosissima Mill. ex Steudel, Nomencl. ed. 2. 1: 99. 1841. nomen nudum. 

Manihot urens Crantz, Inst. Rei Herb. 1: 167. 1766. = Jatropha urens. 

ACKNOWLEDGMENTS 

The senior author is particularly indebted to Dr. William C. Steere, Dr. Bassett 

Maguire, and Dr. Howard Irwin at The New York Botanical Garden, where this work was 

initiated. We wish to thank the librarians in The New York Botanical Garden, the Field 

Museum of Natural History, and the Hunt Botanical Library for their kind assistance on 

many occasions. Dr. William Weber of the University of Colorado provided much aid and 

assistance in the development of this monograph. 

We are deeply indebted to our wives, Constance and Rajam, who not only provided 

comfort and encouragement, but also contributed to the development of many parts of 

the monograph. 

We acknowledge, with thanks, the following agencies and institutions for support for 

field studies and allied activities, and computer program development: The National 

Science Foundation, grants number G-14129, GB-25244, and 656-OSIS; the Office of 

Naval Research, contract Non-R 3640 (00); The National Institutes of Health, grants 

number GM 11208 and GM 13974; The International Business Machines Corporation, 

Project Number 2112, and The Rocky Mountain Forestry Experiment Station, Project 

Number 1629.


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Rogers, D. J. 1963. Taximetrics-new name, old concept. Brittonia 15: 285-290. 

. 1967. A computer aided morphological classification of Manihot esculenta Crantz. Proc. 

Intern. Symp. Trop. Root Crops (Trinidad). 1(Sect. 1): 57-80. 

. 1970. Theoretical and practical considerations on data structuring for a computerized 

information retrieval system. In:Archeologie et Calcaulateurs. Ed. Cent. Nat. Rech. Scientif. Paris 

VII. pp. 145-154. 

, and Appan, S. G. 1969. Taximetric methods for delimiting biological species. Taxon 18: 

609-624. 

, and Appan, S. G. (in press). The North American Species of Manihot. Colo. Assoc. Univ. 

Press. Boulder. 

,and Fleming, H. S. 1973. Monograph of Manibot esculenta Crantz. Econ. Bot. 27: 1-114. 

-__, Fleming, H. S. and Estabrook, G. F. 1967. Use of computers in studies of taxonomy and 

evolution. In: Dobzhansky, Hecht and Steere (Eds.) Evolutionary Biology 1: 169-196. Appleton- 

Century Press. 

Sauer, J. 1951. Crop plants of ancient Peru modelled in pottery. Bull. Missouri Bot. Gard. 37: 

187-194. 

Sloane, H. 1696. Catalogus plantarum quae in insula Jamaica sponte proveniunt. 1: 130, t. 85. 

Stafleu, F. A. 1967. Taxonomic Literature. Int. Bur. PI. Tax. and Nomencl. Utrecht. 556 pp. 

Standley, P. C. 1923. Trees and shrubs of Mexico. Contr. U. S. Nat. Herb. 23: 571-848. 

Steudel, E. T. 1840, 1841. Nomenclator Botanicus. Part I, 1840, p. 779-800; Part II, 1841, p. 99. 

Storey, H. H. and Nichols, R. F. W. 1938. Studies of the mosaic of cassava. Ann. Appl. Biol. 25(4): 

790-806. 

Tournefort, J. P. 1700 Institutiones rei Herbariae. Parisii. p. 658, t. 438. 

Wirth, M., Estabrook, G. F. and Rogers, D. J. 1966. A graph theory model for systematic biology, 

with an example for the Oncidiinae (Orchidaceae). Syst. Zool. 15: 59-69.

1. Manihot 

sect 1. Manihot 

1. M. esculenta Crantz 

sect 2. Parvibracteatae Pax emend Rogers 

& Appan 

2. M. pringlei Watson 

3. M. auriculata McVaugh 

4. M. aesculifolia'(H.B.K.) Pohl 

5. M. rubricaulis I. M. Johnston 

a. subsp rubricaulis 

b. subsp isoloba (Standley) Rogers 

& Appan 

6. M. oaxacana Rogers & Appan 

7. M. chlorosticta Standley & Goldman 

8. M. davisiae Croizat 

9. M. angustiloba (Torrey) Muell.-Arg. 

emend Rogers & Appan 

10. M. rhomboidea Muell.-Arg. 

a. subsp rhomboidea 

b. subsp microcarpa (Muell.-Arg.) 

Rogers & Appan 

11. M. subspicata Rogers & Appan 

12. M. walkerae Croizat 

sect 3. Foetidae Rogers & Appan 

13. M. caudata Greenman 

14. M. michaelis McVaugh 

15. M. tomatophylla Standley 

16. M. websterae Rogers & Appan 

17. M. foetida (H.B.K.) Pohl 

18. M. crassisepala Pax & K. Hoffmann 

sect 4. Heterophyllae Pax emend Rogers & 

Appan 

19. M. zehntneri Ule 

20. M. marajoara Chermonte de Miranda 

apud Huber 

21. M. surinamensis Rogers & Appan 

22. M. tristis Muell.-Arg. 

a. subsp tristis 

b. subsp saxicola (Lanjouw) Rogers 

& Appan 

c. subsp surumuensis (Ule) Rogers 

& Appan 

23. M. janiphoides Muell.-Arg. 

24. M. grahami Hooker 

25. M. inflata Muell.-Arg. 

26. M. pilosa Pohl 

27. M. corymbiflora Pax & K. Hoffmann 

28. M. leptopoda (Muell.-Arg.) Rogers 

& Appan 

29. M. quinquefolia Pohl 

30. M. condensata Rogers & Appan 

31. M. jolyana N. D. Cruz 

32. M. handroana N. D. Cruz 

NUMERICAL LIST OF TAXA 

255 

33. M. pohlii Wawra 

sect 5. Anisophyllae Rogers & Appan 

34. M. anisophylla (Grisebach) Muell.- 

Arg. 

35. M. guaranitica Chodat & Hassler 

a. subsp guaranitica 

b. subsp boliviana (Pax & K. Hoff- 

mann) Rogers & Appan 

sect 6. Carthaginenses Rogers & Appan 

36. M. carthaginensis (Jacquin) Muell.- 

Arg. 

37. M. filamentosa Pittier 

sect 7. Quinquelobae Pax emend Rogers & 

Appan 

38. M. acuminatissima Muell.-Arg. 

39. M. sagittato-partita Pohl 

40. M. xavantinensis Rogers & Appan 

41. M. sparsifolia Pohl 

42. M. falcata Rogers & Appan 

43. M. pruinosa Pohl 

44. M. quinqueloba Pohl 

45. M. alutacea Rogers & Appan 

46. M. violacea Pohl 

a. subsp violacea 

b. subsp recurvata Rogers & Appan 

47. M. jacobinensis Muell.-Arg. 

48. M. divergens Pohl 

49. M. irwinii Rogers & Appan 

50. M. cecropiaefolia Pohl 

51. M. mossamedensis Taubert 

sect 8. Graciles Rogers & Appan 

52. M. flemingiana Rogers & Appan 

53. M. hunzikeriana Martinez-Crovetto 

54. M. hassleriana Chodat 

55. M. triphylla Pohl 

56. M. fruticulosa (Pax) Rogers & Ap- 

pan 

57. M. pentaphylla Pohl 

a. subsp pentaphylla 

b. subsp rigidula (Muell.-Arg.) Rog- 

ers & Appan 

c. subsp tenuifolia (Pohl) Rogers & 

Appan 

d. subsp graminifolia (Chodat & 

Hassler) Rogers & Appan 

58. M. tenella Muell-Arg. 

59. M. gracilis Pohl emend Rogers & 

Appan 

a. subsp gracilis 

b. subsp varians (Pohl) Rogers & 

Appan 

60. M. paviaefolia Pohl 

61. M. maguireiana Rogers & Appan 

sect 9. Sinuatae Pax emend Rogers & Ap- 

pan


62. M. anomala Pohl emend Rogers & 

Appan 

a. subsp anomala 

b. subsp pubescens (Pohl) Rogers 

& Appan 

c. subsp cujabensis (Muell.-Arg.) 


d. subsp glabrata (Chodat & Hassler) 


e. subsp pavoniana (Muell.-Arg.) 


63. M. warmingii Muell.-Arg. 

sect 10. Variifoliae Rogers & Appan 

64. M. mirabilis Pax 

65. M. variifolia Pax & K. Hoffmann 

sect 11. Glaziovianae Pax emend Rogers & 

Appan 

66. M. glaziovii Muell.-Arg. 

67. M. pseudoglaziovii Pax & K. Hoffmann 

68. M. epruinosa Pax & K. Hoffmann 

69. M. brachyandra Pax & K. Hoffmann 

70. M. maracasensis Ule 

71. M. catingae Ule 

72. M. dichotoma Ule 

sect 12. Peruvianae Rogers & Appan 

73. M. brachyloba Muell.-Arg. 

74. M. leptopbylla Pax & K. Hoffmann 

75. M. quinquepartita Huber ex Rogers 

and Appan 

76. M. peruviana Muell.-Arg. 

sect 13. Crotalariaeformes Rogers & Appan 

77. M. procumbens Muell.-Arg. 

78. M. affinis Pax & K. Hoffmann 

79. M. reptans Pax 

80. M. crotalariaeformis Pohl 

sect 14. Stipulares Pax emend Rogers & 

Appan 

81. M. stipularis Pax & K. Hoffmann 

82. M. pusilla Pohl 

83. M. oligantha Pax & K. Hoffmann 

84. M. longepetiolata Pohl 

85. M. nana Muell.-Arg. 

sect 15. Grandibracteatae Pax emend Rog 

ers & Appan 

86. M. tomentosa Pohl emend Rogers 

& Appan 

a. subsp tomentosa 

b. subsp araliaefolia (Pax) Rogers 

& Appan 

sect. 16. Brevipetiolatae Pax emend Rogers 

& Appan 

87. M. stricta Baillon 

88. M. purpureo-costata Pohl 

89. M. salicifolia Pohl 

90. M. attenuata Muell.-Arg. 

91. M. weddelliana Baillon 

92. M. orbicularis Pohl 

sect 17. Peltatae Pax emend Rogers & Ap- 

pan 

93. M. peltata Pohl 

94. M. populifolia Pax in Engler 

95. M. reniformis Pohl 

sect 18. Tripartitae Rogers & Appan 

96. M. tripartita (Sprengel) Muell.-Arg. 

emend Rogers & Appan 

a. subsp tripartita 

b. subsp humilis (Muell.-Arg.) Rog- 

ers & Appan 

c. subsp vestita (S. Moore) Rogers 

& Appan 

d. subsp laciniosa (Pohl) Rogers & 

Appan 

sect 19. Caerulescentes Rogers & Appan 

97. M. caerulescens (Pohl) emend Rog- 

ers & Appan 

a. subsp caerulescens 

b. subsp paraensis (Muell.-Arg.) 


c. subsp macrantha (Pax & K. 

Hoffmann) Rogers & Appan 

98. M. heptaphylla Ule 

2. Manihotoides 

1. M. pauciflora (T. S. Brandegee) 

Rogers & Appan

List of Exsiccate 257 

LIST OF EXSICCATE 

Note: in this list, 1-99 indicates a specimen dubium. The figures in parentheses refer 

to genus number (first), and species number (second). See numerical list of taxa 

Abbott, W. L. 

2304 (1-73); 2796 (1-734). 

Aguilar, J. I. 

1373 (1-4); 1546 (1-lOb). 

Allemao, F. 

1394 (1-67). 

Allen, P. H. & Armour, R. 

7008 (1-66). 

Anisits, J. D. 

1931 (1-35a). 

Annunzio, D. D. 

55748 (1-24). 

Archer, W. A. 

4184 (1-99); 7522 (1-73); 8125 (1-73); 

8150 (1-66); 8151 (1-66); 8291 (1-73); 

8379 (1-75). 

Argauaraz, J. L. 

451 (1-24). 

Argent, G., Ramos, J., Richards, P. W. & Souza, 

R. 

6571 (1-87); 6674 (1-96a); 6675 (1-40); 

6685 (1-87); 6721 (1-96a). 

Aristeguieta, L. 

2234 (1-36); 4675 (1-37); 4701 (1-36). 

Aristiguieta, L. & Saldivias, M. 

542 (1-36). 

Ariza, L. 

18565 (1-34). 

Arsene, Bro. G. 

s n (1-10a); 5272 (1-lOa); 12263 (1-24). 

Assis, V. 

52 (1-96a); 148 (1-55); 244 (1-96a). 

Bach, J. 

4162 (1-74). 

Badillo, V. 

1475 (1-1). 

Bailetti, E. 

96 (1-35a). 

Baker, C. F. 

112 (1-66). 

Balansa, B. 

1713 (1-24); 1713A (1-24); 1714 (1-24); 

1715 (1-62d); 1717 (1-35a). 

Baldwin, J. T., Jr. 

3025 (1-99); 3145 (1-73). 

Bangham, W. N. 

300 (1-4). 

Barreto, M. 

14.104 (1-96a); 2617 (1-86a); 2618 (1- 

86a); 2619 (1-96a); 2625 (1-96a); 2626 

(1-96a); 2676 (1-46a); 3369 (1-59b); 

5055 (1-96a); 5067 (1-26); 8122 (1-66); 

8126 (1-96a); 8127 (1-96a); 9449 (1- 

97a); 9624 (1-26). 

Barreto, M. & Brade, A. C. 

1598 (1-96a). 

Bartlett, H. H. 

10613 (1-11); 13695 (1-11). 

Bejarano, G. 

115 (1-1). 

Bertoni, M. S. 

511 (1-24); 2263 (1-24); 2299 (1-24); 

2537 (1-24); 3499 (1-24); 4638 (1-24); 

47971 (1-24). 

Blake, S. F. 

7688 (1-4). 

Blanchet, M. 

2553 (1-47); 3324 (1-96a & 1-96d); 

3324B (1-96a); 3553 (1-47). 

Boldingh, I. 

7430 (1-36). 

Brade, A. C. 

s n (1-96b); 15403 (1-96a). 

Brandegee, T. S. 

s n (1-4); s n (1-5a); s n (1-7); 550 (1-7). 

Brenes, A. M. 

3993 (1-4). 

Broadway, W. E. 

s n (1-66); s n (1-66); s n (1-66); 7202 

(1-36). 

Bruijn, J. de 

1212 (1-73). 

Buchtien, O. 

4251 (1-62e). 

Burchell 

5861 (1-56); 6289 (1-60); 7015 (1-99); 

7663 (1-93); 7682 (1-93); 7774 (1-38); 

7849 (1-43); 7865 (1-90); 8451 (1-62a); 

8850 (1-62c); 9942 (1-75). 

Burret, M. & Brade, A. C. 

15967 (1-86a). 

Burret, M., Brade, A. C. & Barreto, M. 

10.107 (1-96d). 

B. W. 

2778 (1-1); 2785 (1-1); 2841 (1-1); 2845 

(1-1)C, 2873 (1-1); 6331 (1-22b). 

Cabrera, A. L. 

5712 (1-24). 

Caldas, J. P. 

s n (1-62a). 

Calderon, S. 

776 (1-4); 1023 (1-lOb); 1499 (1-66);


1780 (1-66); 1815 (1-66); 1817 (1-66). 

Cardenas, M. 

3340 (1-35b); 4729 (1-35b). 

Cardona, F. 

530 (1-61); 764 (1-61); 1158 (1-99- 

Doubtfully assigned to 1-21). 

Carlson, M. C. 

215 (1-66). 

Carneiro, R. & Dole, G. 

sn (1-74). 

Carter, A. 

4982 (1-9); 5126 (1-9). 

Castillon, L. 

1720 (1-34); 1949 (1-34); 14176 (1-34). 

Castro, J. B. & Kiehl, J. 

s n (1-24); sn (1-26). 

Cavalcante, P. 

139 (1-75). 

Cazalet, P. C. D. & Pennington, T. D. 

5189 (1-74). 

Chase, A. 

7803 (1-97a). 

Choussy, F. 

73 (1-4). 

Christopher 

sn (1-1). 

Claussen 

s n (1-57b); s n (1-59a); s n (1-62a); s n 

(1-96a); s n (1-97a); 757 (1-59a). 

Claussen, M. 

s n (1-96a); s n (1-97a); 322 (1-62a); 445 

(1-59a); 752 (1-96a); 753 (1-96a); 754 

(1-62b); 755 (1-62a); 756 (1-56); 758 

(1-86a); 953 (1-96a). 

Claussen, P. 

s n (1-86a); s n (1-96a); s n (1-62a); s n 

(1-62a). 

Coelho, J. de M. 

972 (1-66); 2084 (1-68). 

Collenette, C. L. 

188 (1-96a). 

Conde, E. 

117 (1-24). 

Conzatti, C. 

2413 (1-6); 4130 (2-1). 

Cook, 0. F. 

s n (1-66); s n (1-66); s n (1-97a); s n 

(1-97a); s n (1-66); s n (1-99). 

Cook, O. F. & Doyle, C. B. 

311 (1-4); 648 (1-4). 

Cook, O. F. & Jenkins, W. H. 

s n (1-24). 

Cordero, P. & Espiritu, R. 

91498 (1-66). 

Craig, M. 

s n (1-24). 

Crisman, G. E. & Willis, W. D. 

206 (1-4). 

Croizat, L. 

s n (1-36). 

Crovetto, R. M. 

9643 (1-53). 

Crutchfield, J. & Johnston, M. C. 

s n (1-2); 5460A (1-11); 5523 (1-11); 

5572B (1-12); 5784E (1-11). 

Cruz, N. D. 

103 (1-31); 104 (1-26); 105 (1-26); 109 

(1-72); 110 (1-72); 113 (1-1); 114 (1- 

26); 116 (1-23); 117 (1-66); 119 (1-24); 

120 (1-96a); 123 (1-96b); 125 (1-59a); 

126 (1-96a). 

Curran, H. M. 

32 (1-66); 428 (1-36); 721 (1-24). 

Curran, H. M. & Haman, M. 

462 (1-36). 

Cutler, H. C. 

8109 (1-66); 8113 (1-67); 8205 (1-66); 

8276 (1-68); 8277 (1-68); 8280 (1-68); 

8293 (1-66). 

Dacremont 

152 (1-66). 

Darrow, R. A., Gould, F. W., Phillips, W. S. & 

Pultz, L. M. 

1937 (1-9). 

Davis, L. A. 

sn (1-8). 

Dawe, M. T. 

508 (1-36); 552 (1-36). 

Dawson, E. Y. 

14149 (1-48); 15083 (1-50). 

Deam, C. C. 

6097 (1-lOb). 

Delgado, E. 

257 (1-66). 

Deslandes, J. 

189 (1-66). 

De Otero, J. R. 

s n (-59a). 

Diaz, R. 

s n (1-62E). 

Dinelli, E. 

s n (1-34). 

Drake, E. 

s n (1-66). 

Dressier, R. L. 

1913 (1-2). 

Duarte, A. P. 

2785 (1-84). 

Ducke, A. 

s n (1-28); s n (1-97b); s n (1-73); sn 

(1-97a); 530 (1-75); 4808 (1-20); 7712 

(1-73); 10542 (1-75); 11607 (1-75); 

14842 (1-73); 15585 (1-96a); 15873 (1- 

75); 35706 (1-99). 

Dugand, A. 

46 (1-36); 224/61 (1-36); 4654 (1-36); 

5275 (1-36); 5283 (1-36). 

Dugand, A. & Jaramillo, R. 

3231 (1-36); 3246 (1-36). 

Duges, A. 

8 (l-10a) 203 (1-lOa). 

Dusen, P. 

s n (1-66); 5130 (1-26); 6488 (1-66);

List of Exsiccate259 

7812 (1-24); 10104 (1-66); 11158 (1- 

24); 16487 (1-99); 16822 (1-24); 17597 

(1-24). 

Duss, P. 

170 (1-66); 4059 (1-66); 4196 (1-66). 

Dutra, J. 

366 (1-24). 

Echeverria, J. A. 

4158 (1-4). 

Edwards, M. T. 

411 (1-11). 

Eggers, H. F. A. 

15156 (1-74). 

Egler, W. A. & Irwin, H. S. 

46075 (1-73). 

Eifrig, G. 

s n (1-24). 

Ekman, E. L. 

495 (1-24); 496 (1-24); 2690 (1-99); 

2691 (1-66); 14993 (1-73); 15591 (1- 

73); 15736 (1-73); 15854 (1-73); 16905 

(1-66). 

Elias, Bro. 

1546 (1-36). 

Emrick, G. M. 

140 (1-7); 156 (1-7). 

Escritor, L. 

21457 (1-66). 

Esc. Nac de Agricultura 

16203 (1-24). 

Espina 

59 (1-36). 

Eugenio, J. 

782 (1-66). 

Eyerdam, W. J. 

25317 (1-66). 

Faberge, A. C. 

sn (1-lOa). 

Falcao, J. I., Egler, W. A. & Pereira, E. 

329 (1-1). 

Feddema, C. 

1007 (1-4); 909 (1-lOb). 

Fernandez, A. 

324 (1-73). 

Ferreyra, R. 

1619 (1-74); 2747A (1-62e); 4894 (1- 

74); 5090 (1-76); 6734 (1-74). 

Ferris, R. S. 

6140 (1-7). 

Ferris, R. S. & Mexia, Y. 

5218(1-7). 

Fiebrig, K. 

4360 (1-62d); 4669 (1-94); 5297 (1-94); 

10469 (1-35a). 

Field, H. 

s n (1-99-doubtfully assigned to 22b). 

Fosberg, F. R. 

9907 (1-66). 

Foxworthy, F. W. 

1619 (1-66); 1886 (1-66). 

Franco, F. 

22851 (1-66). 

Freitas, O. 

sn (1-99). 

Freyreis 

sn (1-28). 

Freyrey 

s n (1-96b). 

Froes, R. L. 

1087 (1-99); 11545 (1-75); 12630 (1- 

47); 12662/28 (1-99); 12663/29 (1-99); 

12669/35 (1-99); 12703/69 (1-99); 

19996 (1-99); 20066 (1-99); 20084 (1- 

99); 20085 (1-99); 20121 (1-95); 20167 

(1-95); 20170 (1-95); 20174 (1-95); 

20177 (1-95); 20234 (1-28); 20947 (1- 

73); 23548 (1-62b & 1-99); 23862 (1- 

73). 

Fruchard, M. 

s n (1-24). 

Funck (or Funcke) 

607 (1-36). 

Gaillard, M. 

124 (1-22a). 

Galander, C. 

sn (1-34). 

Galeotti, M. 

3734 (1-lOa); 3735 (1-4); 3794 (1-lOb). 

Gardner 

1837 (1-97a); 2437 (1-68); 3441 (1-62b); 

3442 (1-87); 3443 (1-43); 3444 (1-65); 

3445 (1-96a); 3446 (1-38); 3974 (1-41). 

Garnier, H. A. 

sn (1-4). 

Gaudichaud, C. 

280 (1-62c); 299 (1-43); 4520 (1-77). 

Gaumer, G. F. 

1142 (1-4). 

Gaumer, G. F. & Sons 

23394 (1-4). 

Gay, M. Cl. 

sn (1-62e). 

Gehrt, A. 

s n (1-96a); s n (1-96a). 

Gehrt, G. 

sn (1-59a). 

Gentle, P. H. 

2539 (1-4). 

Gentry, H. S. 

1468 (1-5b); 2371 (1-9); 2372 (1-5b); 

2450 (1-8); 5026 (1-7); 6101 (1-4); 6583 

(1-5b); 7085 (1-7); 14282 (1-9); 14293 

(1-8); 17802 (1-9). 

Gentry, H. S., Correl & Arguelles 

18083 (1-5b). 

Gerth, H. 

sn (1-34). 

Ghiesbreght, M. 

s n (1-lOb). 

Giardelli, M. L. 

828 (1-34). 

Glaziou, A. 

s n (1-27); s n (1-46b); s n (1-59a); 1022 

(1-66); 8321 (1-24); 8322 (1-26); 9585 

(1-66); 13201 (1-96a); 13202 (1-96a);


13203 (1-27); 14242 (1-27); 14243 (1- 

27); 14244 (1-24); 17753 (1-86a); 18476 

(1-26); 18476A (1-96a); 19855 (1-57b); 

19856 (1-46a); 20484 (1-96a); 22125 

(1-62b); 22126 (1-90); 22127 (1-83); 

22128 (1-81); 22129 (1-55 & 1-59a); 

22130 (1-59a); 22131 (1-43 & 1-46b); 

22132 (1-46a); 22133 (1-44); 22134 (1- 

50); 22135 (1-50); 22136 (1-86a); 22137 

(1-86b); 22137A (1-42); 22138 (1-96a); 

22139 (1-96a). 

Godfrey, R. K. 

54844 (1-24); 60461 (1-24). 

Goeldi, A. 

2834 (1-73). 

Goes, O. C. & Constantino, D. 

817 (1-28). 

Goes, O. C. & Dionisio 

560 (1-24). 

Goldsmith, P. 

120 (1-10a). 

Gomes, A. I. 

s n (1-28). 

Gomes, J. C., Jr. 

2205 (1-62a). 

Gonzalez, S. 

21583 (1-35a). 

Goodding, L. N. 

35-57 (1-9); 5370 (1-9); 6548 (1-9); 

6549 (1-9). 

Gossweileri 

13553 (1-99); 13927 (1-99). 

Gould, F. W. & Haskell, H. S. 

3248 (1-9). 

Grabendorfer, C. 

550 (1-7). 

Graham, J. & Johnston, M. C. 

4721B (1-11). 

Gregg, J. 

198 (1-11). 

Gregory, D. P. & Eiten, G. 

321 (1-7). 

Guedes, M. 

2463 (1-73). 

Guedes, T. 

215 (1-75). 

Guillemin, M. 

s n (1-28); 132 (1-28). 

Haenke 

s n (1-87). 

Harley, R. M. & Barroso, G. M. et al. 

11325 (1-49); 11404 (1-45). 

Harley, R. M., Lima, A. M., Onashi, E. & Souza, 

R. 

10583 (1-43). 

Harley, R. M., Lima, A. M. Souza, R. & De 

Castro, R. 

10533 (1-97a). 

Harley, R. M. & Souza, R. 

10287 (1-96a); 10341 (1-87); 11023 (1- 

62a). 

Harley, R. M., Souza, R. & Fereira, A. 

10381 (1-40); 11000 (1-96a). 

Harrison, G. H. & Kearney, T. H. 

7983 (1-9). 

Harrison, G. J., Kearney, T. H. & Hastings, W. 

6031 (1-9). 

Harrison, G. J., Kearney, T. H. & Hope, C. 

8904 (1-9). 

Harris, J. A. 

16475 (1-8). 

Hassler, E. 

1669 (1-35a); 2113 (1-65); 2675 (1-35a); 

3466 (1-35a); 4441 (1-77); 4576 (1-54); 

4984 (1-77); 5132 (1-97c); 5172 (1-57d); 

5407 (1-24); 5413 (1-24); 5413A (1-24); 

5413B (1-24); 5516 (1-62d); 5517 (1- 

62d); 5649 (1-62d); 5730 (1-62d); 6756 

(1-24); 7955 (1-96a); 7976 (1-62d); 8121 

(1-62d); 8497 (1-35a); 9525 (1-77); 

9525A (1-77); 10222 (1-59a); 10680 (1- 

24); 10704 (1-96a); 10711 (1-64); 10741 

(1-62d); 10896 (1-94); 10897 (1-65). 

Hastings, J. R. & Turner, R. M. 

64-134 (1-9); 64-355 (1-7). 

Hatschbach, G. 

6404 (1-25); 7529 (1-77). 

Haught, O. 

1664 (1-73); 1919 (1-73); 4156 (1-36). 

Hayes, S. 

717 (1-4). 

Herald, D. F. & Clark, E. E. 

349 (1-lOb). 

Heringer, E. P. 

7752 (1-46a); 7781 (1-85); 7810/4 (1- 

59a); 8760/954 (1-59a); 8850/1044 (1- 

59a); 8895/1089 (1-96a); 9209 (1-81); 

9710 (1-62b). 

Herter, W. 

9673 (1-24). 

Herzog, Th. 

1233 (1-35b); 1419 (1-62e). 

Heyde et Lux 

4306 (1-1). 

Hinton, G. B. 

940 (1-lOa); 1207 (1-4); 3562 (1-17); 

3564 (1-18); 4149 (1-17); 4220 (1-lOa); 

4349 (1-lOa); 4374 (1-4); 4437 (1-10a); 

4467 (1-lOb); 4555 (1-4); 6188 (1-lOa); 

10376 (1-4). 

Hinton, G. B. et al. 

6264 (1-4); 6468 (1-4); 8029 (1-lOb); 

9090 (1-lOb); 10345 (1-7); 10488 (1- 

10b); 10939 (1-7); 13972 (1-4); 15072 

(1-lOb). 

Hoehne, F. C. 

sn (1-26). 

Holm, R. W. & Iltis, H. H. 

250 (1-4). 

Holway, E. W. D. 

513 (1-lOa). 

Hooker 

Icon (1-72). 

Horn, A. 

s n (1-66).

List of Exsiccate261 

Huber, J. 

s n (1-1); s n (1-74); sn (1-74); 798 

(1-74); 915 (1-75); 1460 (1-74); 2600 

(1-74); 2792 (1-20); 3805 (1-75); 4541 

(1-74); 4694 (1-73); 8173 (1-66); 9346 

(1-1 & 1-20). 

Hunziker, A. T. 

9897 (1-34); 10535 (1-24); 10722 (1- 

34). 

Hunziker, A. T., Cocucci, A. E. & Subils, R. 

18466 (1-34). 

Ibarrola, T. S. 

1009 (1-24); 1766 (1-24). 

Illegible 

Illeg. (1-2); s n (1-99); s n (1-24); sn 

(1-24); s n (1-24); s n (1-24); s n (1-24); 

s n (1-24); 139 (1-34); 164 (1-24); 3005 

(1-66); 32581 (1-24). 

Irwin, H. S., Grear, J. W., Souza, R. & dos 

Santos, R. R. 

12443 (1-48); 12654 (1-90); 12959 (1- 

49); 13019 (1-50); 13230 (1-83); 13230A 

(1-83); 13636 (1-84); 13694 (1-48); 

13696 (1-48); 13763 (1-84); 14004 (1- 

62b); 14195 (1-86b); 14222 (1-46b); 

14447 (1-99); 14450 (1-97a); 14609 (1- 

97a); 14698 (1-99); 14821 (1-97a); 

14989 (1-51); 15392 (1-46b); 15516 (1- 

59a); 15565 (1-48); 16210 (1-97a); 

16461 (1-97a); 17841 (1-50). 

Irwin, H. S., Harley, R. M. & Smith, G. L. 

30966 (1-98); 32953 (1-45). 

Irwin, H. S., Maxwell, H. & Wasshausen, D. C. 

18560 (1-96a); 18562 (1-49); 18615 (1- 

79); 18671 (1-96a); 18765 (1-57a); 

18924 (1-96a); 18926 (1-41); 18983 (1- 

96a); 18997 (1-41); 19036 (1-96a); 

19082 (1-41); 19083 (1-50); 19200 (1- 

62b); 19236 (1-79); 19263 (1-57a); 

19304 (1-60); 19359 (1-49); 19430 (1- 

62b); 19774 (1-26); 20422 (1-96a); 

20822 (1-26); 20862 (1-26); 21033 (1- 

96a); 21119 (1-99); 21651 (1-96b?); 

21762 (1-99). 

Irwin, H. S., Prance, G. T., Soderstrom, T. R. & 

Holmgren, N. 

54957 (1-99). 

Irwin, H. S. & Soderstrom, T. R. 

5398 (1-46b); 6319 (1-96b?); 6675 (1- 

89); 6761 (1-87); 6774 (1-87); 6778 

(1-46a); 6788 (1-43); 7006 (1-57b); 7090 

(1-57c); 7364 (1-99); 7380 (1-57b); 7498 

(1-59a). 

Irwin, H. S., Souza, R. & dos Santos, R. R. 

7977 (1-46a); 8074 (1-59a); 8208 (1- 

59a); 8257 (1-46b); 8269 (1-59a); 8297 

(1-46b); 8304 (1-59a); 8359 (1-59a); 

8539 (1-59a); 8563 (1-46b); 8627 (1- 

46b); 8671 (1-59a); 9053 (1-59a); 9113 

(1-46b); 9166 (1-59a); 9167 (1-46b); 

9421 (1-90); 9502 (1-48); 9704 (1-59a); 

9783 (1-84); 9906 (1-83); 9966 (1-39); 

10012 (1-81); 10013 (1-56); 10014 (1- 

48); 10015 (1-96a); 10092 (1-46b); 

10164 (1-56); 10168 (1-46b); 10263 (1- 

59a); 10301 (1-59a); 10344 (1-62b); 

10480 (1-96a); 10493 (1-56); 10498 (1- 

99); 10557 (1-59a); 10593 (1-59a); 

10676 (1-59a); 10715 (1-59a); 10723 

(1-48); 10727 (1-42); 10769 (1-59a); 

10769A (1-59a); 10897 (1-96a); 10898 

(1-49); 10937 (1-93); 10938 (1-56); 

11039 (1-42); 11039A (1-56); 11046 

(1-81); 11051 (1-46a); 11113 (1-59a); 

11195 (1-96a); 11202 (1-46b); 11243 

(1-46b); 11316 (1-62b); 11707 (1-60); 

11742 (1-57a); 11752 (1-49); 11755 (1- 

57a); 11757 (1-41); 11838 (1-57c); 

11850 (1-51); 11862 (1-62b); 11887 (1- 

57a); 11896 (1-62a); 12052 (1-55). 

Irwin, H. S., Souza, R., Grear, J. W. & dos 

Santos, R. R. 

15250 (1-51); 15529 (1-86b). 

Jack, J. G. 

4651 (1-66). 

Jacquin 

sn (1-36). 

Jahn, A. 

1000 (1-36). 

Jancey, R. C. 

320 (1-4); 321 (1-4); 322 (1-4); 323 

(1-4); 324 (1-4); 325 (1-4); 326 (1-4); 

327 (1-4); 328 (1-4); 329 (1-4); 330 

(1-14); 331 (1-14); 332 (1-14); 333 

(1-14); 334 (1-4); 335 (1-4). 

Jaramillo, Mesa, D., Idrobo, J. M. & Fernandez, 

A. 

413 (1-73). 

Johansen, H. 

38 (1-4). 

Johnston, M. C. 

5363B (1-12). 

Joly, A. B. 

1139 (1-96a). 

Jones, M. E. 

sn (1-5b); 127 (1-14). 

Jorgensen, P. 

s n (1-35a); 1225 (1-34); 3082 (1-24); 

4002 (1-24); 31062 (1-24). 

Jorgensen, P. & Hansen 

617 (1-24). 

Jurgensen, C. 

582 (1-10a). 

Karwinski 

s n (l-lOb). 

Kearney, T. H. & Peebles, R. H. 

14816 (1-9); 14928 (1-9). 

Kellogg, J. H. 

sn (1-24). 

Kenoyer, L. A. 

s n (1-lOa); s n (1-lOb); C142 (1-11); 

A-198 (1-2). 

Kerber 

185 (1-18).


Killip, E. P. & Garcia, H. 

33395 (1-73). 

Killip, E. P. & Smith, A. C. 

14032 (1-36); 22722 (1-74); 22964 (1- 

74); 23791 (1-74). 

Kimnach, M. & Moran, R. 

161 (2-1). 

King, R. M. 

366 (1-6); 737 (1-6); 1213 (1-6). 

Klug, G. 

2662 (1-74); 3861 (1-74). 

Knechtel 

507 (1-lOb). 

Knobloch, I. W. 

1335 (1-5b). 

Koppel, van de 

4444 (1-66). 

Krug, C. A. 

sn(1-26). 

Krukoff, B. A. 

1649 (1-75); 6311 (1-73). 

Kuhlmann, J. G. 

s n (1-28); s n (1-96a); s n (1-97a); 1928 

(1-74); 06022 (1-28); 06242 (1-25); 

06562 (1-33). 

Kuhlmann, M. 

s n (1-25); s n (1-31); sn (1-32); 2751 

(1-26); 2889 (1-96b). 

Kuhlmann, M. & Gehrt, A. 

sn (1-26). 

Kuntze, O. 

sn (1-34). 

Kurtz, F. 

8572 (1-34). 

Labouriau 

937 (1-68). 

Labouriau, L. & Valio 

1172 (1-96a); 1182 (1-59a). 

Labouriau, M. S. 

186 (1-1); 187 (1-24). 

Lamorton, O. H. de 

s n (1-24); s n (1-24); s n (1-24). 

Landb. 

922 (1-22b). 

Landeman, C. 

3371 (1-74). 

Lanfranchi, A. E. 

327 (1-24). 

Lanjouw, J. 

955 (1-22b). 

Lanjouw, J. & Donselaar, W. A. E. 

789 (1-21). 

Lasser, T. & Aristeguieta, L. 

3319 (1-36). 

Laughlin, R. M. 

1125(1-4). 

Lauro, G. 

sn (1-24). 

Leal, C. G. 

34 (1-74). 

Leavenworth, Wm. C. 451 (1-7). 

Leavenworth, Wm. C. & Hoogstraal, H. 

1402 (1-15); 1532 (1-7); 1548 (1-lOa); 

1556 (1-7). 

Lemmon, J. G. 

s n (1-8); 3054 (1-9); 70 3055 (1-8). 

Lesueur, H. 

246 (1-11); 1249 (1-5b). 

Levy, P. 

41 (1-4). 

Lillo, M. 

s n (1-35a); 1822 (1-62e); 2243 (1-24); 

2365 (1-24); 2531 (1-35a); 5235 (1-62e); 

5269C (1-35a); 7278 (1-34); 7287 (1- 

24); 7918 (1-34); 52696 (1-62e). 

Lima, A. S. 

s n (1-68); s n (1-96a); 7147 (1-66). 

Lindman, C. A. M. 

2505 (1-62a); A2505 (1-62a). 

Lisboa, A. 

2343 (1-97a); 2363 (1-97a); 2431 (1- 

97a); 2432 (1-99). 

Livingston & Thornber 

s n (1-8). 

Llamas, A. de 

sn (1-24). 

Lofgren, A. 

190 (1-66); 191 (1-66); 225 (1-66); 327 

(1-97a); 991 (1-96b) 1030 (1-96a); 1452 

(1-96a); 2084 (1-77); 5969 (1-78). 

Lorentz, P. G. 

297 (1-34). 

Lorentz, P. G. & Hieronymus, G. 

330 (1-35a). 

Luetzelburg, V. 

9A (1-1); 9B (1-1); 10 (1-1); 365 (1-71); 

484A (1-47); 484B (1-47); 1348 (97a); 

7209 (1-33); 12251 (1-70); 12252A (1- 

69); 12253 (1-38 & 1-68); 12417 (1-68); 

12421 (1-67); 12429 (1-67); 12430 (1- 

67); 12433 (1-68); 12463 (1-67); 12674 

(1-97a); 12888 (1-67); 15003 (1-68); 

15005 (1-68); 15013 (1-68) 15399 (1- 

28); 15400 (1-66); 20302 (1-73); 26414 

(1-97a). 

Lundell, C. L. & Lundell, A. A. 

7473 (1-4); 7737 (1-4); 7848 (1-4). 

Lutz, A. 

926 (1-96a); 976 (1-99). 

Lyonnet, E. 

301 (1-lOb). 

MacBride, J. F. 

5074 (1-74); 5472 (1-74). 

MacDaniels, L. H. 

212 (1-7). 

Macedo, A. 

s n (1-48); 2915 (1-26); 3142 (1-77); 

3436 (1-57a); 3461 (1-57a); 3473 (1-57c 

& 1-59a); 3476 (1-49); 3697 (1-79); 

4126 (1-62a); 4177 (1-96a); 4264 (1-48); 

4346 (1-60); 4347 (1-49); 4390 (1-79); 

4417 (1-62b). 

Macedo, A. & Smith, L. B. 

4651 (1-59a). 

Machado, O. 

sn (1-28).


Magalhaes, M. 

6080 (1-26); 14887 (1-97a). 

Maguire, B. & Maguire, C. K. 

44732 (1-55). 

Maguire, B., Pires, J. M., Maguire, C. K. & Silva, 

N. T. 

56464 (1-47); 56496 (1-99); 56836 (1- 

43); 57088 (1-55); 57096 (1-81). 

Malme 

s n (l-96c); 1272 (1-24); 2331 (1-96c); 

2344 (1-89). 

Manso 

sn (1-44); 76B (1-62c). 

Marcano & Vega, E. 

4852A (1-73). 

Markgraf, F. 

3509 (1-96a). 

Markgraf, F. & Brade 

3746 (1-24). 

Marsh, E. G. 

1163 (1-11). 

Martinez, M. 

15088 (1-lOb). 

Martius 

s n (1-29); s n (1-73); sn (1-97a); sn 

(Manihot johannis Pax. Species dubium); 

sn (1-1); sn (1-1); 942 (1-26); 1935 (1- 

95); 2427 (1-1). 

Mattos, J. & Mattos, N. 

10208 (1-75). 

Matuda, E. 

1665 (1-lOb); 4392 (1-lOb); 4813 (1-4); 

16375 (1-4). 

Maxon, W. R., Harvey, A. D. & Valentine, A. T. 

7107 (1-4); 7192 (1-4). 

McVaugh, R. 

13298 (1-10a); 13386 (1-lOb); 15037 

(1-13); 15283 (1-3); 15355 (1-4); 15486 

(1-lOb); 15502 (1-14); 15531 (1-lOa & 

1-lOb); 15571 (1-lOa); 15638 (1-7); 

15875 (1-7); 15954 (1-lOb); 16580 (1- 

lob); 17615 (1-lOa); 17951 (1-15); 

18128 (1-lOb). 

McVaugh, R. & Koelz, W. N. 

1553 (1-7); 1695 (1-7). 

Mearns, E. A. 

2535 (1-66). 

Medina, B. R. 

158 (1-24). 

Melhus, I. E. & Goodman, G. J. 

3652 (1-10a). 

Mendes, J. E. T. 

6741 (1-66). 

Mendes, O. T. 

sn (1-62a). 

Merrill, E. D. 

5010 (1-66). 

Mexia, Y. 

4109 (1-26); 4153 (1-26); 8778 (1-4). 

Meyer, T. 

2186 (1-35a); 4805 (1-35a); 5421 (1-24); 

5463 (1-24); 5850 (1-24); 11773 (1-24); 

11945 (1-24); 12043 (1-24). 

Miers, J. 

1240 (1-24). 

Milano, V. 

sn (1-24). 

Miller, G. S., Jr. 

1238 (1-73); 1250 (1-73). 

Millspaugh, Herb. 

3055 (1-8). 

Miranda, F. 

2092 (1-10a). 

Monetti, L. 

4897 (1-35a). 

Monteiro da Costa, R. C. 

234 (1-74). 

Montes, J. E. 

360 (1-24); 584 (1-24); 1443 (1-24); 

1448 (1-24); 1896 (1-24); 9524 (1-24). 

Moore, S. 

186 (1-96c). 

Morley, T. 

745 (1-lOb). 

Mosen, H. 

3279 (1-25); 4386 (1-26). 

Muller, C. H. 

3696 (1-5b). 

Nano, J. F. 

27434 (1-66). 

Nelson, E. W. 

2852 (1-4); 2899 (1-lOb). 

Nelson, E. W. & Goldman, E. A. 

7401 (1-7). 

Nelson, G. 

7069 (1-66). 

Nelthropp, A. 

5 (1-66). 

Nicefora 

225 (1-73). 

Normanha, E. 

sn (1-26). 

Novais, C. 

926 (1-26). 

Novelo, N. S. 

114 (1-4). 

Occhioni, P. 

sn (1-55); sn (1-96a). 

Octavio 

s n (1-66). 

Orbigny, A. d' 

1040 (1-99). 

Ortega, J. G. 

6345 (1-7); 6488 (1-7); 7024 (1-7); 7273 

(1-7). 

O'Donnell, C. A. 

205 (1-24); 4847 (1-34). 

Palacios, M. A. 

23 (1-24). 

Palmer, E. 

142=156 (1-lOa & 1-lOb); 201 (1-13); 

222 (1-9); 224 (1-5a); 233 (1-9); 266 (1- 

2); 272 (1-7); 1027 (1-7); 1027A (1-7); 

1536 (1-4).


Palmer, L. H. 

sn (1-66). 

Parks, H. B. 

s n (1-12). 

Pasquier, du 

1859 (1-66). 

Pavon 

s n (1-74). 

Pavon, Herb. 

34163 (1-lOa). 

Peckolt, Th. 

21 (1-33); 95 (1-26); 201B (1-1). 

Pedersen, T. M. 

7 (1-24); 4170 (1-35a). 

Peebles, R. H. 

8796 (1-8). 

Peebles, R. H., Harrison, G. J. & Kearney, T. H. 

4584 (1-9); 5588 (1-9). 

Peebles, R. H. & Kearney, T. H. 

8742 (1-9). 

Peirano 

s n (1-34). 

Pereira, E. 

2023 (1-47); 2023A (1-47); 2055 (1- 

97a); 4621 (1-59a); 4765 (1-46b). 

Pereira, E. & Egler, W. A. 

383 (1-96a). 

Perkins, A. E. & Hall, J. M. 

3565 (1-11). 

Pflanz, K. 

4035 (1-35a). 

Phares, D. L. 

11(1-24). 

Philcox, D. & Fereira, A. 

3868 (1-52); 3898 (1-96a); 3899 (1-96a); 

3941 (1-87); 4542 (1-52); 4609 (1-97a). 

Philcox, D., Fereira, A. & Bertoldo, J. 

3283 (1-40 & 1-96a); 3340 (1-97a). 

Philcox, D., Ramos, J. & Sousa, R. 

3005 (1-96a); 3019 (1-96a); 3105 (1-40); 

3109 (1-97a). 

Philipson, W. R., Idrobo, J. M. & Fernandez, A. 

1463 (1-73). 

Pickel, D. B. 

632 (1-99); 1255 (1-66). 

Pires, J. M. 

17 (1-73); 1857 (1-75); 3603 (1-99). 

Pires, J. M. & Black, G. A. 

985 (1-73); 1480 (1-73); 3048 (1-96a). 

Pires, J. M., Black, G. A., Wurdack, J. J. & Sil- 

va, N. T. 

6118 (1-97a); 6159 (1-62b); 6203 (1- 

57c); 6284 (1-97a); 6477 (1-97b). 

Pires, J. M., Rodrigues, Wm. & Irvine, G. C. 

5100 (1-1); 50815 (1-1); 51058 (1-97b); 

51202 (1-73); 51499 (1-73). 

Pires, J. M. & Silva, N. T. 

4830 (1-75). 

Pires, J. M., Silva, N. T. & Souza, R. 

9174 (1-59a). 

Pires, J. M. & Westra, L. Y. Th. 

48837 (1-22b). 

Pittier, H. 

s n (1-66); 132 (1-lOb); 1567 (1-36); 

1723 (1-36); 11761 (1-36). 

Pittier, H. & Durand, Th. 

6579 (1-4). 

Plee 

64 (1-36). 

Poeppig 

1410 (1-74); 1523 (1-74). 

Pohl 

s n (1-46a); s n (1-56); Icon (1-99); Icon 

tab. 10 (1-89); Icon tab. 11 (1-88); Icon 



tab. 16 (1-59a); Icon tab. 17 (1-59a); Icon 


tab. 20 (1-41); Icon tab. 21 (1-62a); Icon 




tab. 29 (1-57c); Icon tab. 30 (1-57c); Icon 





tab. 40 (1-62b); Icon tab. 41 (1-96a); 

Icon tab. 42 (1-50); Icon tab. 43 (1-86a); 

Icon tab. 44 (1-96a); Icon tab. 45 (1-60); 

Icon tab. 46 (1-57a); Icon tab. 47 (1- 

59b); Icon tab. 48 (1-96d); Icon tab. 57 

(1-96a); 56 (1-97a); 477 (1-96a) 494 (1- 

80); 702 (1-86a); 734 (1-39); 767 (1- 

86a); 822 (1-84). 1181 (1-82); 1182 (1- 

57c); 1183 (1-48); 1184 (1-55); 1185 

(1-59a); 1186 (1-57a); 1187 (1-62a & 1- 

99); 1188 (1-1); 1189 (1-46a); 1190 (1- 

60); 1191 (1-96a); 1192 (1-96a); 1193 

(1-96a); 1194 (1-89); 1196 (1-62b); 1197 

(1-62a); 1198 (1-62b); 1254 (1-62a); 

1371 (1-1); 1649 (1-50); 1655 (1-96a); 

1656 (1-96a); 1657 (1-48); 1658 (1-62b); 

1659 (1-59a); 1660 (1-89); 1661 (1-93); 

1701 (1-44); 1702 (1-62a); 1703 (1-88); 

1704 (1-39); 1705 (1-43); 1706 (1-57a); 

1707 (1-96a); 1708 (1-55); 1709 (1-1); 

1711 (1-59b); 1713 (1-86a); 1901 (1-93); 

1902 (1-44); 1992 (1-92); 2047 (1-62b); 

2130 (1-88); 2206 (1-41); 2207 (1-96d); 

2208 (1-59b); 2421 (1-57c); 2469 (1-43); 

3598 (1-96a); 3722 (1-96a); 3775 (1-1): 

3776 (1-1); 3777 (1-1); 3778 (1-1); 3780 

(1-1); 3781 (1-1); 3919 (1-50); 3920 

(1-59a); 3921 (1-96a). 

Porto, P. C. 

sn (1-99). 

Prance, G. T., Pena, B. S., Ramos, J. F. & Vi- 

decki, E. R., Jr. 

2389 (1-74). 

Poilane, M. 

2414 (1-66). 

Prance, G. T. & Silva, N. T. 

58227 (1-48). 

Pringle, C. G. 

76 (1-9); 127 (1-9); 2243 (1-12); 3558 

(1-2); 3826 (1-2); 5159 (1-lOa); 8687


(1-13); 11318 (1-lOa); 11339 (1-18); 

13938 (1-4). 

Pulle, A. 

213 (1-73). 

Purpus, C. A. 

517 (1-7); 3418 (2-1); 5465 (1-2); 5841 

(2-1); 6112 (1-4); 7353 (1-6); 8448 (1- 

4); 8484 (1-4); 9322 (1-lOb); 10213 (1lob). 

Raizada, M. B. 

s n (1-24) 

Rambo, B. 

31287 (1-24); 45076 (1-24). 

Ratter, J. A., Castro, R. A. de, Santos, R. R. 

dos & Souza, R. 

834 (1-97a); 835 (1-97a). 

Rechinger, K. U. L. 

1707 (1-66). 

Regnell, A. F. 

186 (1-28); 408 (1-24); III 1070 (1-26); 

III 1070A (1-26); III 1071A (1-96a); III 

1071B (1-96a); III 1073 (1-59a). 

Reiche, K. 

1072 (1-2). 

Richard, L. C. 

sn (1-22b). 

Reidel 

s n (1-47); s n 2820? (1-56); s n (1-57b); 

s n (1-58); s n (1-59b); s n 476? (1-77); s 

n (1-85); s n (1-96a) s n (1-96b); s n (1- 

96d); 190 (1-25); 191 (1-28); 613 (1-89); 

789 (1-59a); 791 (1-97a); 1026 (1-55); 

1530 (1-26); 1848 (1-26); 2822 (1-97a); 

2824 (1-85). 

Robert, A. 

420 (1-96a); 558 (1-96c); 558B (1-96a). 

Rodriguez 

164 (1-24); 4922 (1-24). 

Rodriguez, J. V. 

1528 (1-lOb). 

Rogers, D. J. 

77 (1-1); 84 (1-1); 100 (1-1); 101 (1-1); 

51 (1-1); 106 (1-1); 119 (1-1); 120 (1-1); 

123 (1-1); 128 (1-1); 131 (1-1); 236 (1- 

1); 237 (1-1); 243 (1-1); 249 (1-1); 252 

(1-1); 257 (1-1); 261 (1-1); 263 (1-1); 

264 (1-1); 267 (1-1); 268 (1-1); 270 (1- 

1); 271 (1-1); 277 (1-1); 278 (1-1); 286 

); 3(1-1); 293 (1-1); 29 (1-1); 296 (1-1); 

300 (1-1); 303 (1-1); 312 (1-1); 313 (1- 

1); 321 (1-1); 323 (1-1); 335 (1-28); 352 

(1-66); 353 (1-68); 354 (1-68); 356 (1- 

99); 357 (1-23); 362 (1-59a); 363 (1- 

86a); 364 (1-59a); 365 (1-1); 370 (1-1); 

371 (1-66); 372 (1-26); 373 (1-26); 381 

(1-68); 385 (1-68); 386 (1-66); 389 (1- 

1); 390 (1-1); 394 (1-68); 395 (1-68); 

397 (1-97a); 398 (1-97a); 399 (1-68); 

403 (1-68); 404 (1-97a); 405 (1-1); 408 

(1-97a); 409 (1-68); 410 (1-68); 411 (1- 

66); 412 (1-66); 413 (1-73); 417 (1-1); 

419 (1-1); 422 (1-1); 423 (1-1); 430 (1- 

62e); 431 (1-62e); 432 (1-1); 435 

(1-62e); 437-A (1-62e); 437-B (1-62e); 

437-C (1-62e); 437-D (1-62e); 442 (1- 

1); 448 (1-1); 453 (1-1); 467 (1-74); 468 

(1-74); 474 (1-74); 490 (1-1); 494 (1- 

36); 495 (1-37); 504 (1-4); 505 (1-6); 

506 (1-6); 507 (1-6); 508 (1-lOa); 508A 

(1-10a); 508B (1-lOa); 508C (1-lOa); 

508D (1-10a); 508E (1-10a); 508F (1- 

10a); 508G (1-lOa); 508H (1-10a); 5081 

(1-lOa); 509 (1-13); 510 (1-4); 511 (1- 

7); 512 (1-7); 513 (1-14); 514 (1-14); 

515 (1-4); 516 (1-15); 517 (1-15); 519 

(1-9); 520 (1-4); 521 (1-4); 522 (1-12). 

Rogers, D. J., Appan, S. G. & Rogers, R. 

525 (1-11); 526 (1-11); 528 (1-2); 529 

(1-11); 530 (1-1); 533 (1-11). 

Rogers, D. J. & Godfrey, R. K. 

499 (1-24). 

Rogers, D. J., Thieret, J. W. & Reese, Wm. 

496 (1-24); 497 (1-24); 498 (1-24). 

Rojas, T. 

s n (1-24); s n (1-24); 196A (1-24); 3667 

(1-62d); 4252 (1-24); 4753 (1-24); 6214 

(1-58); 6326 (1-58); 6326A (1-58); 

6326B (1-59a); 6326C (1-58); 6469 (1- 

24); 6792 (1-97c); 6809 (1-62d); 7206 

(1-35a); 7221 (1-35a); 13718 (1-35a). 

Rombouts, H. E. 

409 (1-21); 464 (1-21). 

Rosengurtt 

B-4975 (1-24). 

Rose, J. N. 

1608 (1-lOa); 1611 (1-4); 2499 (1-13); 

3204 (1-4); 3266 (1-7). 

Rose, J. N. & Hay, R. 

6201 (1-lOa). 

Rose, J. N. & Hough, W. 

4751 (1-lOb). 

Rose, J. N., Painter, J. H. & Rose, J. S. 

10114 (2-1). 

Rose, J. N. & Rose, J. S. 

11190 (1-lOa). 

Rose, J. N. & Russell, P. G. 

19910 (1-99). 

Ruiz & Pavon 

s n (1-62e). 

Rusby, H. H. 

2 (1-4); 178 (1-13); 886 (1-1); 888 (1- 

73); 1297A (1-30). 

Ryan, C. L. & Floyed, H. A. 

11 (1-lOb). 

Saer, J. 

191 (1-37); 366 (1-37); 367 (1-37). 

Sagot 

1082 (1-73). 

Saint-Hilaire 

s n (1-96a); s n (1-96d); s n (1-97a); s n 

(1-99); s n (1-35a); 255-0 (1-80); 488 

(1-48); 700 (1-96a); 2328 (1-79). 

Sarmento, A. 

sn (1-68).


Schomburgk, R. H. 

s n (1-1); 426 (1-1); 426/694 (1-1); 694 

(1-1). 

Schott 

518 (1-4). 

Schott, A. 

s n (1-24); III 8 (1-9); 52 (1-12). 

Schreiter, R. 

s n (1-34); s n (1-35a); 552 (1-35a); 1795 

(1-34); 3402 (1-35a); 3704 (1-62e); 4106 

(1-35a); 5189 (1-62e); 6454 (1-34); 9068 

(1-35a); 10992 (1-35a). 

Schulz, A. G. 

1550 (1-35a); 1551 (1-35a); 7053 (1-53); 

7178 (1-53). 

Schulz, J. P. & Donselaar, J. van 

10593 (1-22b). 

Schunke, J. M. 

291 (1-73). 

Schwacke 

4520 (1-96a). 

Schwarz, G. J. 

4981 (1-53); 5199 (1-53); 6369 (1-53); 

6629 (1-24); 6738 (1-24); 7143 (1-24); 

8790 (1-24). 

Schwindt, E. 

487 (1-24); 1019 (1-24); 2759 (1-24); 

3091 (1-24); 3326 (1-24); 4839 (1-24). 

Seler, C. & Seler, E. 

2814 (l-10a). 

Sello 

s n B2104 C2066? (1-96a); s n (1-77). 

Sellow 

sn (1-59a). 

Sesse & Mocino 

30856 (1-10a). 

Sesse, Mociiio, Castillo & Maldonado 

4218 (1-lOa); 4222 (1-lOa); 4224 

(1-lOb); 4229 (1-17); 4573 (1-lOa). 

Shreve, F. 

6371 (1-9); 6753A (1-Sb). 

Sick, H. 

B265 (1-62a). 

Silva, A. 

391 (1-66); 77 (1-74). 

Skehan, J. 

sn (1-24). 

Skutch, A. F. 

4429 (1-74); 5009 (1-74). 

Smith, C. E. 

s n (2-1). 

Smith, C. E., Peterson, F. A. & Tejeda, N. 

3563 (2-1). 

Smith, H. H. 

365 (1-36 & 1-73). 

Smith, J. D. 

6850 (1-73). 

Smith, L. B. & Klein, R. M. 

11024 (1-24); 11711 (1-24); 13076 (1- 

24). 

Smith, L. B. & Macedo, A. 

4651 (1-59a). 

Smith, L. B. & Reitz, Pe. R. 

9083 (1-24); 9647 (1-24); 9702 (1-24); 

9896 (1-24). 

Snethlage, D. E. 

10103 (1-75). 

Sota, A. V. de la 

3341 (1-24). 

Souza, R. & Richards, P. W. 

6888 (1-87). 

Spegazzini, C. 

s n (1-35a); s n (1-62e); 2351 (1-24); 

18385 (1-24). 

Spegazzini, L. 

16407 (1-24). 

Spegazzini, P. L. 

sn (1-34). 

Spruce, R. 

186 (1-97b); 622 (1-97b); 825 (1-1); 

3604 (1-22a); 4287 (1-76). 

Stahel 

107 (1-22b). 

Standley, P. C. 

9657 (1-4); 11491 (1-4); 19836 (1-4); 

28687 (1-66) 53767 (1-1); 60407 (1-66); 

73669 (1-4); 73683 (1-4); 73736 (1-4); 

73783 (1-4); 74305 (1-4); 79452 (1-4). 

Steere, W. C. 

1673 (1-4); 1947 (1-4). 

Steinbach, J. 

3143 (1-62e); 5202 (1-62e); 6709 (1- 

62e). 

Steyermark, J. A. 

47778 (1-4); 47779 (1-4); 50614 (1-10a 

& 1-lOb); 50866 (1-lOb); 51316 (1-4); 

51474 (1-lOa); 75228 (1-99); 75296 (1- 

61); 99980 (1-73). 

Steyermark, J. A. & Agostini, G. 

91228 (1-36). 

Steyermark, J. A. & Farinas, M. 

91440 (1-66). 

Stoffers, A. L. 

534 (1-36). 

Stuckert, T. 

14270 (1-34); 12462 (1-34); 12756 

(1-34); 18873 (1-34); 22154 (1-34). 

Swingle, W. T. 

sn (1-9). 

Tamayo, F. 

775 (1-99); 831 (1-66); 910(1-66). 

Tamberlik 

s n (1-43); sn (1-47). 

Tessmann, G. 

6012 (1-24). 

Thackery, F. A. 

487 (1-9). 

Thieme, C. 

5473 (1-4). 

Thornber 

sn (1-8). 

Tisher 

sn (1-8). 

Toledo & Gehrt, A. 

sn (1-96b). 

Tonduz, Ad. 

13977 (1-4).

Index 267 

Topping, D. L. 

9096 (1-66). 

Townsend, C. H. T. & Barber, C. M. 

404 (1-5b). 

Triana, J. 

s n (1-36); 3614 (1-73); 3620 (E) (1-1). 

Tuerckheim, H. von 

8752 (1-66). 

Tweedie, J. 

s n (1-24); 103 (1-1); 104 (1-1). 

Tyson, E. L. 

4108 (1-4). 

Ule, E. 

50 (1-41); 373 (1-79); 374 (1-57a); 375 

(1-60); 470 (1-51); 471 (1-1); 472 (1- 

57a); 473 (1-62b); 475 (1-43); 3072 (1- 

79); 3073 (1-57a); 3081 (1-51); 4605 (1- 

26); 5264 (1-73); 6635 (1-76); 7003 (1- 

70); 7045 (1-72); 7085 (1-95); 7141 (1- 

97a); 7141A (1-97a); 7142 (1-71); 7206 

(1-98); 7362 (1-72); 7645 (1-1); 7944 

(1-22c); 9058 (1-66); 9546 (1-74). 

Uribe, L. U. 

3077 (1-73). 

Usteri, A. 

32 (1-66). 

Vargas, C. 

10031 (1-62e); 11608 (1-74); 14335 (1- 

62e); 26329 (1-62e); 4884 (1-62e); 5852 

(1-62e). 

Velez, I. 

2465 (1-1). 

Venturi, S. 

63 (1-24); 63E (1-24); 134 (1-24); 669 

(1-35a); 1611 (1-34); 1690 (1-62e); 

1690B (1-62e); 2865 (1-62e); 2685A (1- 

35a); 7241 (1-24); 7400 (1-34); 8095 (1- 

34); 8690 (1-35a); 9880 (1-24); 10006 

(1-35a); 10191 (1-62e). 

Versteeg 

419 (1-22b); 801 (1-22b). 

Victoria, J. 

8 (1-24). 

Viegas, A. P. 

s n (1-23); s n (1-24); s n (1-67); s n (1- 

26); sn (1-26); sn (1-26). 

Viegas, A. P. & Kiehl, J. 

sn (1-66). 

Villafane, M. 

187 (1-34). 

Walker, E. J. 

1003 (1-12). 

Warming, M. 

s n (1-23); s n (1-59a); s n (1-62b); s n 

(1-96a); 1502 (1-97a); 1503 (1-23); 1507 

or s n? (1-63); 1508 or s n? (1-57b); 1509 

(1-96d); 1519 (1-26); 1520 (1-96b & 1- 

96d); 1520/1 (1-96d); 1520/2 (1-96d). 

Waterfall, U. T. 

12808 (1-8). 

Weberbauer, A. 

6517 (1-62e). 

Webster, G. L. 

11229 (1-2). 

Webster, G. L., Miller, K. & Miller, L. 

11641 (1-6); 12955 (1-6); 13007 (1-6); 

13074 (1-16); 13080 (1-1Oa); 13080A 

(1-10a); 13080B (1-lOa); 13080C (1- 

10a). 

Weddell, M. A. 

s n (1-62a); 150 (1-66); 1882 (1-59a); 

2525 (1-62b); 2862 (1-41); 2908 (1-91); 

2909 (1-50); 2939 (1-57c); 2960 (1-96a). 

Weintraub, F. C. & Roller, J. 

122 (1-lOa). 

West, E. 

s n (1-24). 

White, O. E. 

1019 (1-30 & 1-99). 

White, S. S. 

542 (1-8 & 1-9); 2758 (1-5b); 2930 (1- 

5b); 3013 (1-9); 3103 (1-5b); 3647 (1- 

Sb). 

Wiggins, I. L. 

7056 (1-9); 7155 (1-9). 

Wiggins, I. L. & Rollins, R. C. 

221 (1-8 & 1-9). 

Wigg, T. L. 

2288 (1-66). 

Wilbur, R. L. & Wilbur, C. R. 

1743 (1-lOa); 2088 (1-lOa). 

Williams, L. 

2059 (1-73); 5323 (1-74); 9867 (1-4); 

13132 (1-22a); 15893 (1-22a); 16095 

(1-22a). 

Williams, L. E. 

sn (1-24). 

Williams, L. O. & Assis, V. 

5689 (1-96a); 6061 (1-96a); 6876 (1-26). 

Williams, L. O. & Molina, A. R. 

10116 (1-lOb). 

Williams, R. O. 

s n (1-36); 12516 (1-36). 

Williams, R. S. 

233 (1-62e); 361 (1-99); 3055 (1-66). 

Woolston, A. 

109 (1-62d). 

Wornock, B. H. & Barkley, F. A. 

147 (1-12). 

Woytkowski, F. 

34448 (1-74). 

Wright, C. 

1811 (1-9). 

Wright, D. 

sn (1-24). 

Wurdack, J. J. & Guppy, N. G. L. 

176 (1-61). 

Wycherly, P. R. 

760 (1-66); 761 (1-66); 762 (1-66). 

Zabala, S. 

15 (1-24). 

Zehntner 

s n (1-97a); 238 (1-97a); 299 (1-97a); 

365 (1-97a); 381 (1-98); 390 (1-97a); 

394 (1-97a); 533 (1-97a); 598 (1-19); 

611 (1-97a); 689 (1-97a); 690 (1-97a); 

691 (1-97a); 695 (1-97a). 

Zingg, R. M. 

10 (1-lOb).


Adenoropium tripartitum 233 

Janipha 19 

aesculifolia 41 

anisophylla 105 

foetida 74 

frutescens 112 

juquilla 114 

loeflingii 114 

var multifida 88 

manihot 25 

var angustiloba 53, 62 

violacea 110 

yuquilla H.B.K. 114 

Jatropha 19 


angustifolia 251 

anomala 167 

arcuata 136 

cajaniformis 233 

caricaefolia 167 

cataginensis 112 

carthaginensis 112 

cecropiaefolia 139 

cleomaefolia 233 

coerulea Ind. Kew. 242 

coerulea Steudel 242 

coerulescens 242 

crotalariaeformis 206 

dalechampiaeformis 233 

diffusa 27 

digitiformis 27 

divergens 136 

dulcis 25 

flabellifolia 27 

foetida 74 

frutescens 112 

gracilis 158 

heterophylla 167 

janipha Linnaeus 114 

janipha Loureiro non Linnaeus 25 

laciniosa 238 

lanciniosa 238 

longepetiolata 210 

loureirii 27 

manihot Linnaeus 25 

mitis 25 

orbicularis 225 

palmata Sesse & Mocinio apud Cervantes 3 

palmata Arrabida in Steudel 96 

palmata Vellozo 96 

paniculata 27 

paviaefolia 160 

peltata Steudel 227 

pentaphylla 151 

pilosa 92 

porrecta 233 

INDEX OF SCIENTIFIC NAMES 

pronifolia 158 

pruinosa 127 

pubescens 167 

purpureo-costata 220 

pusilla 208 

quinquefolia 98 

quinqueformis 98 

quinqueloba 130 

reniformis 230 

sagittato-partita 121 

salicifolia 220 

silvestris 3 

simayuca 170 

sinuata 233 

sparsifolia 124 

stipulata Arrabida in Steudel 25 

stipulata Vellozo 25 

sylvestris 251 

tenerrima 153 

tenuifolia 153 

tomentella 233 

tomentosa 213 

triloba 3 

tripartita 233 

triphylla 147 

varians 160 

violacea 134 

Mandioca 19 

dulcis 27 

utilissima 27 

Manihot 19 

Sect Anisophyllae 105 

Sect Brevipetiolatae Pax emend Rogers & 

Appan 215 

Sect Caerulescentes 238 

Sect Carthaginenses 112 

Sect Crotalariaeformes 199 

Sect Foetidae 65 

Sect Glaziovianae Pax 105, 172, 175 

Sect Glaziovianae Pax emend Rogers & 

Appan 175 

Sect Graciles 141 

Sect Grandibracteatae Pax 

Subsect Angustifoliae 141, 230 

Subsect Coerulescentes 77, 118, 199, 

238 

Subsect Glabrescentes 118 

Subsect Grandiflorae 238 

Subsect Papillosae 65 

Subsect Peruvianae 189 

Subsect Rigidulae 77, 141 

Subsect Tomentosae 163, 213 

Subsect Tripartitae 199, 230 

Sect Grandibracteatae Pax emend Rogers & 

Appan 213 

Sect Heterophyllae Pax

Index269 

Subsect Carthaginenses 25, 36, 77, 

105, 112, 175 

Subsect Cujabenses 163 

Subsect Variifoliae 163, 172 

Sect Heterophyllae Pax emend Rogers & 

Appan 77 

Sect Indivisae Pax 225 

Sect Manihot 25 

Sect Parvibracteatae Pax 

Subsect Anomalae 163 

Subsect Elatae 34, 77, 189 

Subsect Graciles 34, 143, 175 

Subsect Guaraniticae 105, 190 

Subsect Humiles 141, 230 

Subsect Langsdorffianae 77, 163 

Subsect Nanae 206 

Subsect Stenophyllae 143 

Subsect Tristes 77, 118 

Subsect Utilissimae 25, 118, 199 

Sect Parvibracteatae Pax emend Rogers & 

Appan 34 

Sect Peltatae Pax emend Rogers & Appan 

225 

Sect Peruvianae 189 

Sect Quinquelobae Pax emend Rogers & 

Appan 118 

Sect Sinuatae Pax 

Subsect Laciniosae 141, 199, 230 

Subsect Warmingianae 34, 163, 238 

Sect Sinuatae Pax emend Rogers & Appan 

163 

Sect Stipulares Pax emend Rogers & Appan 

206 

Sect Tripartitae 230 

Sect Variifoliae 172 

Sect Weddellianae Pax 215 

acuminatissima 121 

acutiloba 53 


affinis 203 

aipi Pohl 27 

var lanceolata 27 

var latifolia 27 

var lutescens 27 

aipi Rusby 251 

alcicornis 168 

alutacea 130 

amaroleitensis 124 

amazonica 190 

angustifolia 251 

angustifrons 147 

angustiloba (Torrey) Muell.-Arg. emend 

Rogers & Appan 53 

anisitsii 110 

anisophylla 105 

anomala Pohl 165, 167 

anomala Pohl emend Rogers & Appan 167 

subsp anomala 167 

subsp cujabensis 168 

subsp glabrata 168 

subsp pavoniana 170 

subsp pubescens 167 

araliaefolia 213, 215 

arcuata 136 

attenuata 222 

aurictilata 39 

bahiensis 242 

var microsperma 251 

berroana 251 

boliviana 112 

brachyandra 185 

brachyloba 190 

brachystachys 222 

brasiliensis 92 

brevipedicellata 92 

burchellii 127 

caerulescens Pohl 239, 242 

caerulescens Pohl emend Rogers & Appan 

239 

subsp caerulescens 242 

subsp macrantha 245 

subsp paraensis 245 

cajaniformis 233 

canastrana 215 

caricaefolia 167 

carthaginensis 112 

var anisophylla 105, 110 

cassava 251 

catingae 187 

caudata 68 

cearensis 242 

cecropiaefolia 139 

chlorosticta 50 

cleomaefolia 233 

coerulea 251 

coerulescens Pohl emend Muell.-Arg. 242 

var genuina 242 

var pubescens 242 

colimensis 50 

condensata 100 

consanguinea 251 

conulifera 153 

cordifolia 227 

corymbiflora 96 

crassisepala 74 

crotalariaeformis 206 

cuiabensis 168 

cujabensis 168 

cuneata 242 

curcas 252 

dalechampiaeformis 233 

davisiae 51 

depauperata 158 

dichotoma 187 


var parvifolia 187 

diffusa 27 

digitata 252 

digitiformis 27 

discolor 242 

divergens 136 

diversifolia 252 

domingensis 194 

dulcis (J.F. Gmelin) Pax 25 

var aipi 27 

var diffusa 27 

var ferruginea 194 

var flabellifolia 27 

var leptopoda 96 


edule 27 

elegans 201


enneaphylla 88 

epruinosa 182 

esculenta 25 

var sprucei 27 

subsp flabellifolia 28 

var argentea 28 

var coalescens 28 

var debilis 28 

var digitifolia 28 

var flavicaulis 28 

var fuscescens 28 

var nodosa 28 

var tenerrima 153 

subsp grandifolia 28 

sect alboerecta 28 

var domingensis 28 

var hispaniolensis 28 

var luteola 28 

var mutabilis 28 

var ramosissima 28 

var rufescens 28 

sect diffusa 28 

var communis 28 

var fertilis 28 

var jamaicensis 28 

var pohlii 28 

var zimmermannii 28 

falcata 125 

ferruginea 242 

fiebrigii 110 

filamentosa 117 

flabellifolia 27 

flemingiana 143 

flexuosa 27 

floribunda 182 

foetida 74 

fruticulosa 149 

glabrata 168 

glaziovii 177 

gossypifolia 252 

gracilis Pohl 155 

gracilis Pohl emend Muell.-Arg. 153, 158 


var pronifolia 158 

var tenerrima 153 

var tenuifolia 153 

var triphylla 147 

gracilis Pohl emend Rogers & Appan 155 

subsp gracilis 158 

subsp varians 160 

grahami 88 

graminifolia 155 

grandiflora 242 

grandistipula 110 

gualanensis 41 

guaranitica Chodat & Hassler 108, 110 

subsp boliviana 112 

subsp guaranitica 110 

guyanensis 252 

handroana 102 

harmsiana 242 

hassleriana 147 

hemigynandra 92 

hemitrichandra 92 

heptaphylla 245 

heptaphylla x piauhyensis 245 

herbacea 252 

heterandra 197 

heterophylla Chodat & Hassler 252 

heterophylla Pohl 167 

hilariana 251 

humilis 236 

hunzikeriana 145 

inflata 91 

intercedens 238 

intermedia 41 

irwinii 137 

isoloba 46 

jacobinensis 136 

janipha 114 

janiphoides 86 

japonica 252 

johannis 251 

jolyana 100 

katharinae 175 

klingensteinii 168 

labroyana 242 

laciniosa 238 


var lanata 238 

lagoensis 238 

langsdorffii 92 

var glabra 168 

leptophylla 194 

leptopoda 96 

linearifolia 217 

lobata 88 

loeflingii 88 


longepetiolata 210 

longipetiolata 210 

loureirii 27 

ludibunda 59 

lyrata 242 

macrantha 245 

maguireiana 161 

manihot 27 

maracasensis 185 

var vestita 185 

marajoara 80 

mattogrossensis 220 

meeboldii 201 

melanobasis 27 

membranacea 251 

meridensis 114 

mexicana 59 

meyeriana 92 

michaelis 70 

microcarpa 60 

microdendron 242 

mirabilis 172 

mobilis 50 

moluccana 252 

mossamedensis 141 

multifida 252 

multiflora 168 

nana 212 

neoglaziovii 252 

oaxacana 46 

occidentalis 136

Index 

olfersiana 41 

oligantha 208 

orbicularis 225 

orinocensis 84 

palmata (Vellozo) Muell.-Arg. 

var aipi 27 

var diffusa 27 

var digitiformis 27 

var ferruginea 194 

var flabellifolia 27 


var leptopoda 96 


var pusilla 208 

palmata (Vellozo) Pax 96 

paraensis 245 

pardina 160 

parvicocca 60 

pauciflora 248 

paviaefolia 160 

pavoniana 170 

pedicellaris 92 

peltata 227 

pentaphylla Pohl 151, 160 


var paviaefolia 160 

pentaphylla Pohl emend Rogers & Appan 

subsp graminifolia 155 

subsp pentaphylla 151 

subsp rigidula 153 

subsp tenuifolia 153 

peruviana 197 

piauhyensis 242 

pilosa 92 

pittieri 114 

pohliana 252 

pohlii 102 

polyantha 130 

populifolia 227 

porrecta 233 

preciosa 252 

pringlei 37 

procumbens 201 


var grandifolia 201 

pronifolia 158 

pruinosa 127 


var pumila 127, 251 

pseudoglaziovii 181 

pseudoheterophylla 168 

pseudopruinosa 127, 134 

pubescens 167 

forma glabrata 168 

purpureo-costata 220 

pusilla 208 

quenqueformis 98 

quinquefolia 98 

quinqueloba 130 

quinquepartita 196 

recognita 110 

reflexa 153 

remotiloba 114 

reniformis 230 

reptans 203 

271 

rhomboidea Muell.-Arg. 59 

rhomboidea Muell.-Arg. emend Rogers & 

Appan 57 

subsp microcarpa 60 

subsp rhomboidea 59 

riedeliana Klotzsch ex Pax 242 

riedeliana Muell.-Arg. 220 

rigidifolia 136 

rigidula 153 

rotundata 242 

rubricaulis I.M. Johnston emend Rogers & 

Appan 45 

subsp isoloba 46 

subsp rubricaulis 46 

rusbyi 190 

sagittato-partita 121 

salicifolia 220 

saxicola 84 

sellowiana 201 

sinuata 233 


var laciniosa 238 

sparsifolia 124 

speciosa Chodat & Hassler 245 

speciosa Muell.-Arg. 242 

spinosissima 252 

sprucei 27 

stenophylla 147 

stipularis 206 

stricta 217 

subquinqueloba 130 

subspicata 62 

surinamensis 80 

surumuensis 86 

teissonnieri 251 

tenella 155 

tenerrima 153 

tenuifolia 153 

theveti 2 

toledi 242 

tomatophylla 70 

tomentella 233 

tomentosa Pohl 213 

tomentosa Pohl emend Rogers & Appan 

213 

subsp araliaefolia 215 

subsp tomentosa 213 

trichandra 236 

trifoliata 242 

var platyphylla 251 

tripartita (Sprengel) Muell.-Arg. 230 

var apaensis 234 

var cajaniformis 233 

var dalechampiaeformis 233 


var glabra 236 

var glauca 234 

var lanceolata 234 

var porrecta 233 

var quinqueloba 102 

var seminuda 234 

var subintegra 238 

var vestita 236 

tripartita (Sprengel) Muell.-Arg. emend


Rogers & Appan 230 

subsp humilis 236 

subsp laciniosa 238 

subsp tripartita 233 

subsp vestita 236 

triphylla 147 

var fruticulosa 149 


tristis Muell.-Arg. 82 

subsp saxicola 84 

subsp surumuensis 86 

subsp tristis 84 

tubuliflora 92 

tweedieana 88 

var lobata 88 

forma nana 88 

uleana 153 

urens 252 

utilissima Pohl 27 

var castellana 27 

var sutinga 27 

varians 160 

variifolia 175 

violacea Pohl 132 

violacea Pohl emend Muell.-Arg. 

var arcuata 136 

var cecropiaefolia 139 

var divergens 136 


violacea Pohl emend Rogers & Appan 132 

subsp recurvata 134 

subsp violacea 134 

walkerae 63 

warmingii 172 

weberbaueri 170 

websterae 72 

weddelliana 222 

xavantinensis 124 

zehntneri 78 

Manihotoides 247 

pauciflora 248

- -7 Lo 

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FIG 28. "Skyline"-computer produced graph of relations between species of Manibot in South 

America. The numbers on the horizontal axis above and below represent the specimens included in the 

computer analysis. On the vertical axis are shown the similarity values. In the Skyline, horizontal lines' 

indicate that all the specimens above the line are connected. By examination of the graph, one may 

observe that most species are delimited at, or near, the 0.90 value, although so.ne very distinct species 

show their first connections at a much lower similarity value. The horizontal lines in the graph also 

give one indication of the sectional divisions, with the average similarity value of 0.75 for differentiation 

of the sections. 

In the graph, the specimen assignment to species is as follows: SECTION 18: 1, 2, 3, 6, 4, 7, 5, 8, 

11 = 96a; 9, 10 = 96b; 14, 14 = 96c; 19, 331, 23, 24, 25, 26 = 96d. SECTION 8: 33, 34, 35, 36 = 55; 

37, 38 = 56; 51, 52, 53, = 57b; 58, 59, 60 = 57a; 61, 62, 63 = 57c; 56, 57 = 57d; 54, 55 = 58; 39, 40 = 

60; 66, 67 = 54; 43, 44, 45, 49 = 59a; 46, 47, 48 = 59b; 68, 69, = 53; 64, 65 = 52; 41, 42 = 61. 

SECTION 4: 70, 71, 72, 322, 332, 333 = 24; 320, 338 = 25; 73, 93, 94 = 23; 85, 88, 89, 90, 91 = I*; 

92 = 19* 328, 340, 96, 97, 98, 99, 95, 324 = 1*; 325 = 20; 326, 327 = 21; 86, 87 = 22b; 323, 339 = 

22c; 101, 102, 103, 104, 108, 106 = 26; 111, 112 = 28; 321 = 29; 109, 110 = 27; 78, 79 = 30; 82, 83,

O 

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51 2,12 8 27 2 3;19 30=4;13,12 33=1 34 3 4.....ON11 

280 281, 28 ,6,. . 1 8,2% 7 8,26 87 8,20 9,29 8 9,26 3 

....9,39=72 = 70 !ETOJ 9, 9=6. 

2 0,32 0,34 0 3 0,37 0 

= 74 [ 0,3 1 5 1,33=7.SCIN9:20 3 2;22=6;24 3,26 3 

=~~~~~~~~~~i 

62;2829=6cI4,4,4,4,4,4 =6d 4,4,4 2.SCIN1:29 

25 4;2125,23 5 ECIN13 5,35_79 5 8;37,3836,5,3331 

36 =!7 6 8 ETO 4 9,9 1 9,13=8;14 9 3 9,17=8;18 

CD(D 

7 

Ln (O CD D ((0 

r ooco r-^rcoooooo 

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O Nc) (O) N) 7-) (f) 19 =l5 ETO 5 5,5 - - - - 6;3233=86.SCIN1:1011=9;121314 

- - - - - 

;t No. 

= 4 661718 101118i 5 SCIN16 6,7011= 7 7, 7 8;19 

9 

17,7,16=90 7,78=9; 8,84=9 ECIN19 0,222320,0,2726 

84 = 22a; 75, 76 = 31; 77, 107 = 32; 80, 81, 336 = 33. SECTION 6: 270, 271, 272, 274, 275 = 36; 

273, 366 = 37. SECTION 5: 260, 261 = 34; 262, 263 = 35b; 264, 265, 266, 267, 268, 269 = 35a. 

SECTION 7: 115, 116, 118, 117, = 44; 119, 120 = 45; 136, 137 = 46a; 138, 139 = 46b; 144, 145 = 

49; 148, 149, 150 = 48; 140, 142, 154, 143 = 47; 146, 147 = 50; 121, 122; 123, 124 = 43; 151, 152 = 

51; 125, 126 = 38; 127, 128 = 39; 129, 130 = 40; 131, 132, 133 = 41; 134, 135 =42. SECTION 11: 

280, 281, 282 = 66; 297 = 71; 283, 284 = 67; 285, 286, 287, 288, 290, 291, 289 = 68; 295, 296, 334 

= 70; 298, 299, 329 = 72; 293, 294 = 69. SECTION 12: 301, 302, 303, 304, 305 = 73; 306, 307, 308 

= 74; 309, 310, 311 = 75; 312, 313 = 76. SECTION 9: 230, 231 = 62b; 232 = 63; 234, 235, 236, 237 

= 2,36=$7SETO5:2026=$426,23=$b26,2,26,2726,298=35. 

62a; 238, 239 = 62c; 240, 241, 242, 244, 245, 243 = 62d; 246, 247, 248 = 62e. SECTION 10: 249, 

250 = 64; 251, 252, 253 = 65. SECTION 13: 354, 355 = 79; 356 = 80; 357, 358, 362, 359, 363, 361, 

360 = 77; 364 = 78. SECTION 14: 190, 191 = 81; 192, 193 = 82; 194, 195 = 83; 196, 197 = 84; 198, 

199 = 85. SECTION 15: 350, 351 = 86a; 352, 353 = 86b. SECTION 17: 160, 161 = 93; 162, 163, 164 

= 94; 166, 167, 168 = 95. SECTION 16: 169, 170, 171 = 87; 172, 173 = 88; 179, 180, 181, 182 = 89; 

174, 175, 176 = 90; 177, 178 = 91; 183, 184 = 92. SECTION 19: 201, 202, 203, 204, 205, 207, 206, 

208, 209, 210, 211, 212, 214, 215, 216, 213, 217 = 97a; 218, 219 = 97b; 220, 221 = 97c; 222, 223 = 

98. 

- NN 

N

Manihot Manihotoides (Euphorbiaceae) - CNCFlora

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