Observations on

Stengelîoid Species of Vernonia

by

Ç. i:arle, Smith, Jr.

Plant Science Research Division

Agriculture Handbook No. 396

Agricultural Research Service

UNITED STATES DEPARTMENT OF AGRICULTURE

shington, P.C. ^ I.g^^j September 1971

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

Washington, B.C. 20402 - Price $1
Acknowledgments

I wish to thank the curators of the following herbaria who have loaned

specimens for this study : Biology Department, Gordon College, Rawalpindi,

Pakistan (RAW) ; Botanical Survey of India, Calcutta (CAL) ; British

Museum (Natural History) (BM) ; Conservatoire et Jardin Botaniques,

Genève (G) ; East African Agricultural and Forestry Research Organization, Nairobi (EA) ; Federal Herbarium, Salisbury (SRGH) ; Gray Herbarium and Arnold Arboretum of Harvard University (GH and AAH) ;

Herbarium Universitatis Florentini, Firenze (FI) ; Jardin Botanique de

FEtat, Bruxelles (BR) ; Royal Botanic Gardens, Kew (K) ; Land Utilization

Department, Mbabane, Swaziland (MBA) ; Missouri Botanical Garden, St.

Louis (MO) ; New York Botanical Garden (NY) ; Republic of South Africa

Department of Agriculture, Botanical Gardens, Durban (NH) ; Republic of

South Africa Department of Agricultural Technical Services, Pretoria

(PRE). Except for the abbreviation for the Mbabane herbarium, abbreviations are those used in Lanjouw and Stafleu (1964).

I am very grateful for the photographs of type collections made for me

by the Botanischer Garten und Museum, Berlin-Dahlem ; British Museum

(Natural History); Jardim e Museu Agrícola do Ultramar, Lisbon; and

Dr. K. Walther, Staatsinstitut für Allgemeine Botanik und Botanischer

Garten, Hamburg, who provided an achene with photographs of specimens.

I had previously been permitted to study and photograph specimens at

Botanische Staatsammlung, München and at Muséum National d'Histoire

Naturelle, Laboratoire de Phanerogamic, Paris, and I thank the directors

and staffs for their cooperation.

Mrs. Regina Hughes prepared the line drawings. I am very grateful for

her contributions to my paper.

My exploration of eastern Africa could hardly have been accomplished

without the excellent help of many people and institutions along the way.

Everyone from my Ethiopian driver, Ato Sesai, to Dr. Brian Rycroft,

Director of the National Botanic Gardens of South Africa, ensured that

I saw all of the vernonias in their area. I am particularly indebted to Mrs.

Mell of Duiwelskloof, Transvaal; Mr. W. Duthie, Durban, Natal; Mrs.

H. M. Richards, Abercorn, Zambia ; and, especially, my excellent field companion, Dr. D. B. Fanshawe, Principal Scientific Officer, Forest Department, Kitwe, Zambia. For a year after my visit, Dr. Fanshawe continued

to collect and submit achenes and herbarium specimens of the stengelioid

vernonias which I had missed.

I have also received excellent assistance from the librarians of the

National Agricultural Library, the Library of the Harvard University

Herbaria, and the Library of the Philadelphia Academy of Natural Sciences.

Dr. S. B. Jones, University of Georgia, Athens, has very generously

provided me with a number of unpublished chromosome counts for species

of section Stengelia. I thank him very much.
Contents

PAGE

History of the section Stengelia 1

Section Stengelia today 4

Chemical constituents of stengelioid Vernonia species 6

Ecology of Vernonia species in eastern Africa 6

Biological units within section Stengelia 10

Preparation of the report 12

Selection of lectotypes 13

Systematic treatment 14

Literature cited 85

Appendix . 86

List of vernacular names for stengelioid Fernoma species 86
Observations on

Stengelioid Species of Vernonia

By C. Earle Smith, Jr.,1 botanist.

Plant Science Research Division, Agricuhural Research Service

The present treatment of the species purported to belong to section Stengelia had its inception with the discovery of epoxyoleic acid

in the achene oil of Vernonia anthelmintica. In

its native Ceylon, and in India and Pakistan,

V, anthelmintica is semicultivated, i.e., it is allowed to persist in field borders because it is

favored as a home remedy. In spite of the industrial potential of the epoxyoleic oil fraction,

this species has proven, so far, to be agronomically unsuitable in the United States.

Consequently, a collection program was instituted to find related species. All of the relatives of V. anthelmintica are found in Africa.

None of these species of Vernonia are in cultivation although specimen labels indicate that

certain African species are encouraged by Africans who use them for medicinal or culinary

purposes. Until the discovery of the potential

value of the achene oil, no oil analyses had been

made for the species of section Stengelia. Preliminary study of borrowed herbarium specimens of species said to be members of section

Stengelia pointed up the impracticality of a

nomenclatural revision without field observations. This also afforded an opportunity to procure achene samples for oil analysis.

Exploration for species of section Stengelia

was begun in northern Ethiopia and continued

southward through most of the eastern African

countries, and through South Africa. When

mature achenes were found, samples were collected for analysis. Herbarium specimens were

prepared of all species encountered. Observations made in the field are used in interpreting

the variation within species in the species descriptions.

History of the section Stengelia

The first mention of Stengelia in botanical

literature was in an advertisement for centuries

of Ethiopian specimens that had been collected

by Wilhelm Schimper (Hochstetter, 1841).2 In

the list, on page 26, under b, Eupatorinae, are

listed 4 species of Vernonia. Following these is

the entry '*Stengelia, n. g., adoënsis C. H.

Schultz. B. (318)" indicating that Stengelia is

a new genus with the species adoensis. The

number in parentheses is Schimper's collection

number. Nowhere in the article are the new

1 Now Professor of Anthropoly and Biology, University of Alabama, Tuscaloosa.

2 References to Literature Cited (p. 85) are indicated

by the name of the author (authors) or title of the

publication followed by the year of publication.

taxa described, and the generic and specific

names are nomina nuda.

In August, 1841, (see Stafleu, 1967) SteudeFs

Nomenclátor listed Stengelia adoensis C. H.

Schultz and Stengelia kotschyana Hochstetter

(Steudel, 1841). Again, all names are nomina

nuda because the reference for S. adoensis is

the publication cited above and the reference

for S. kotschyana is an exsiccatae list of

Kotschy specimens.

Finally, W. G. Walpers (1843) published the

species as Vernonia adoensis and V, kotschyana,

citing Stengelia in synonymy. Under the International Code of Botanical Nomenclature (Lanjouw, 1966), publication in synonymy is ruled

invalid. The first validation by published description of the name Stengelia, so far as I am
AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

aware, is in a footnote in Peters (1862-4)

^^Reise nach Mossambique." The treatment, prepared by J. Steetz, unfortunately excludes

Schultz's original species, V, adoensis, but includes Vernonia abyssinica (as Stengelia controversa). Thus, the type of the genus Stengelia

is Stengelia eontroversa Sch.-Bip. ex Steetz =

Vernonia abyssinica Sch.-Bip. ex Walp. Because

the latter name was validly published in 1843,

Steetz was mistaken in setting up the first

species; he should have transferred the specific

name, abyssinica, to Stengelia. Be that as it

may, it has little bearing on the current use of

Stengelia. The species to which Schultz originally applied the generic name were placed by

Steetz in Ascaricida Cass.

The use of Stengelia as a sectional name was

not clearly stated until 1873 to the best of my

knowledge. However, to understand the rise of

the sectional name, it is necessary to go to A. P.

De Candolle (1836). In handling the nearly 300

species of Vernonia which he listed, De Candolle

divided the genus into sections, the ninth and

last of which was *1X Ascaricida Less." Within

his section, he grouped species in subsections

(unnamed) on the basis of the characteristics

of the phyllaries. In § 2. Involucri squamis

exterioribus angustioribus, he placed V. bojeri,

V. anthelmintica, and V. paucifiora, the second

being the one species growing outside of Africa

which is today considered to be a species of section Stengelia. The other two species are African and both have been associated with the

stengelioid species by several authors.

Walpers (1843) followed the arrangement of

the Prodromus in presenting the species of

Vernonia. In the supplement (p. 946) he specifically writes, ''Sectio IX, Ascaricida Less. § 2.

Involucri squamis exterioribus angustioribus.''

And he adds, *'A. Involucri squamis exterioribus oblongo-lanceolato-linearibus in appendicem

foliaceam variae magnitudinis expansis, quandoque foliis nonnullis, caulinis simplicibus circumdatis, achaeniis columnari-turbinatis, pappi

bi- V. pluriserialis radiis paleaceis, complanatis.

(=Stengelia C. H. Schultz Bipont. mss.).'' He

then describes Vernonia adoensis, V. kotschyana, and V. abyssinica. The establishment

of subsection Stengelia could not be much

clearer. I select Vernonia adoe^isis Sch.-Bip. ex

Walp. as the type species of subsection Stengelia (fig. 1). By inference, since Walpers did

not list all of the Prodromus species covered by

De Candolle but only species added after 1836,

the De Candollean species are included.

Although it is not significant for the purposes

of this report, Lessing (1831) defined his section Ascaricida to include those species which

De Candolle admitted. In a broad sense, Lessing-s section is the basal point from which

section Stengelia sprang.

With the publication of the Bentham and

Hooker (1873) ''Genera Plantarum,'' Stengelia

Sch.-Bip. was formally recognized as Section 3

of Vernonia, The diagnostic characters of

Genera Plantarum are essentially those of the

Walpers subsection, and the first cited literature

reference is the Walpers publication.

O. Hoffman in Engler and Prantl (1889) rearranged the sections of Vernonia, making

Stengelia section 2. The diagnostic characteristics are almost the same as those of prior publications except that they are brief and somewhat less specific. The only species mentioned

is Vernonia anthehnintica.

Through the latter half of the 19th century

and the early part of the 20th century many

collections went to Europe from Africa. Numerous botanists described new species of Vernonia

and often placed them to section at the time of

publication. Needless to say, Stengelia received

its share of new species. During this period, the

only publication broad enough to include a large

number of the species of section Stengelia was

the Oliver and Hiern's Compositae in the **Flora

of Tropical Africa'' (Oliver, 1877). The authors

used a revised grouping of the sections in which

Stengelia became section 6. Within the section,

they recognized 17 species which have since

been allowed to remain there by other authors.

Even then, the group was causing trouble because they had to key out F. purpurea^ V. nigritana^ and V. filigera in the key to sections,

rather than in that part of the species key covering section Stengelia.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA

PN-2021

Figure l.~Vernonia adoensis Sch.-Bip. ex Walp. is the lectotype species of Subsection Stengelia Sch.-Bip. ex Walp.

and Section Stengelia (Sch.-Bip. ex Walp.) Benth. & Hook. f. Drawn from Schimper 318 (Isotype, V. adoensis,

US) and Pappi 528 (FI),

No other treatment has covered this section

on so broad a geographic range. Most of the

species added after the ''Flora of Tropical

Africa'' was published have never been critically evaluated for inclusion in a flora and

they have never been monographed. At the time

that they were described, many of the species

were distinguishable, but the collections

amassed in the intervening years have erased

the boundaries between many of the species.

My study of the stengelioid group has necessarily been limited by interest in the achene oil,

and I cannot do the overall survey of African

Vernonia which is now required.
AGRICULTUEE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

Section Stengelia today

At the present time, section Stengelia may or

may not include as many as 120 taxa. I have

traced references to species and intraspecific

entities, and I have literature references to 120

names which have been placed in the section at

one time or another. Some were later removed

by one or another botanist, and I am sure that

I have missed many of these transfers in my

quest for prime references. In recent years,

very few new species have been described in the

section, but there has been some rearranging

of taxa within the section.

The boundaries of the section have not been

redefined since the publication of subsection

Stengelia by Walpers in 1843. Therefore, the

characters given by Hoffman in Engler and

Prantl (1889) still stand—"Inner involucral

bracts with membranous, colored appendages

(fig. 2) ; . . .; outer pappus bristles or scales

short, inner sometimes scattered." These are

Walpers' criteria (see above) in a slightly abbreviated statement. The section is now a heterogeneous melange of species groups which share

the sectional characters. In 1915, De Wildeman

reported the following in discussing the characters of V. bukamaensis an,_ its sectional affinity, "Les bractées involúcrales sont relativement peu modifiées dans leur partie supérieure.

Ce caractère est d'ailleurs un plus ou moins ;

fréquement, les plantes que l'on doit rapporter à

ce sous-genre, ne présentent pas d'une façon

nette la transformation foliacée de l'estrémité

de la bractée, de sorte que l'on pourrait les rapporter au sous-genre Decaneuron qui possède

parmi ses représentants des passages indiscutables vers les Stengelia."

Unfortunately, the sectional characters have

led to a strange mixture of plants which are

morphologically different and have many different growth habits. The lectotype species, V.

adoensis, is an herbaceous perennial that grows

to 2 m. tall from a large root crown. V. guineensis and its allies produce shorter aboveground shoots yearly from a root crown from

which radiate tuberous roots. Several other

species are woody shrubs that grow to 4 or 5

m. tall. Two species are trees which reach 10

m. ultimately. Another complex of stengelioid

species produces a root crown bearing a rosette

of leaves above which are borne nearly naked

scapes which support the heads. Still others in

this group produce the leaves at one season and

the inflorescences separately at another season.

The stengelioid species of Vernonia can be

separated into 2 large groups on morphological

characters. The herbaceous perennials, shrubs,

and trees all have florets with a narrow tube

which is as long as the limb or longer ; the limb

widens abruptly at the top of the tube, is urceolate and has relatively short deltoid or short

«'

/'

/

PN-2022

Figure 2.—Two extremes of phyllary diflPerentiation are

illustrated by (A) the large, often white phyllary tips

of V. adoensis and (B) the phyllaries of V. abyssinica

with small herbaceous tips and large, exposed coriaceous bases.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA

among- them; introgression may be occurring

due to disturbance in habitats in recent times ;

or they may be truly valid species complexes,

sharing a common ancestry and might be considered subsections.

My study of the stengelioid species of Vernonia is too limited to establish effective secPN-2023

Figure 3.—The upright, herbaceous perennial and woodyspecies of section Stengelia usually share (A) florets

(V. abyssinica) with a distinct narrow tube flaring

into the limb with deltoid or short lanceolate lobes and

B, pollen grains (V. guinecnsis) with a heavy, often

regular, reticulura.

lanceolate lobes, which are firm in texture. Pollen grains of this group usually have a regular,

heavy reticulum (fig. 3).

The rosette group of the stengelioid species

of Vernonia generally have flowers with little

or no tube which widens gradually into the

conical limb with long, lanceolate, thin lobes.

The pollen grains of many of these species are

decorated with a thin, irregular reticulum

(Smith, 1969) (fig. 4).

Within these 2 major groupings, other less

well-defined groups appear. For example, V.

guineensis and the species which I have placed

immediately thereafter all seem to have tuberous roots, lanate under surfaces on the leaves,

and phyllaries so arranged that the coriaceous

basal portion is prominent at maturity. Such

groups may indicate only an incomplete isolation of species and the continuing gene flow

PN-2024

Figure 4.—The species of section Stengelia with a basal

rosette of leaves often have (A) florets (V. acrocephala) in which the tube gradually enlarges into the

limb with thin, long, lanceolate lobes and (B) pollen

grains (V. praemorsa) with a thin, irregular recticulum.
6 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

tion limits. This would entail the study of many

species from other sections, some of whose species approach the stengelioid species closely. In

fact, Vernonia afromontana and its relatives

are so similar to the stengelioid species that I

made herbarium specimens and achene collections whenever I found them. Oil analysis discloses a high epoxy acid content in the achene

oil of V, afromontana, which may or may not

be an indication of relationship. This complex

of species is generally separated from Stengelia

because the phyllary tip is not expanded, although it is herbaceous.

Until a major revision of all the African species of Vernonia is undertaken, section Stengelia will have to remain a heterogeneous mixture. My own experience suggests that the vast

number of species of Vernonia will have to be

reduced to biologically valid taxa that take

into account all of the more recent tests for

species validity. Morphological characteristics

alone will no longer provide the sole evidence

for their separation. However, the number of

described species is large, and the revision of

the sections of Vernonia cannot be accomplished without a prior consideration of the

species.

Chemical constituents of stengelioid Vernonia species

Interest in section Stengelia was originally

sparked by the discovery of epoxyoleic acid in

the oil from achenes of V. anthelmíntica. Therefore, so far as it was possible, I collected samples of achenes from native populations in eastern Africa. To the south, the season was late

and I was too early, so collections for oil analysis were made by D. B. Fanshawe in Zambia

(7 samples analyzed), Mrs. Mell in Transvaal,

and W. Duthie in Natal (1 sample each). All

analyses were made by the Northern Utilization Research and Development Division staff

at Peoria, 111. Unfortunately, the figures obtained are badly skewed by immature and

insect-damaged achenes. Until new supplies are

grown and analyzed, the oil analysis figures

cannot be published as they may be very misleading. However, some samples contained more

than 70-percent epoxyoleic acid.

Among samples of species of Vernonia submitted to Cancer Chemotherapy, National Service Center, National Cancer Institute, U.S.

Department of Health, Education, and Welfare,

by R. E. Perdue, botanist in ARS, USDA, V,

hymenolepis was found to have an antitumor

constituent. The compound, vernolepin, a sesquiterpene dilactone, was extracted and isolated

from leaves by S. M. Kupchan and his colleagues (Kupchan and others, 1968). When

tested against Walker intramuscular carcinosarcoma 256 in rats, 12 mg. of the vernolepin

per kg. of body weight produced significant inhibition of the cancer. Subsequent experimentation (Sequeira and others, 1968) has shown

that vernolepin acts as a plant growth inhibitor

against wheat coleoptiles. The growth-inhibiting effects can be reversed by treatment with

Índole acetic acid.

Although the initial interest in the achene

oil chemicals was centered in epoxyoleic acid,

two other constituents have been isolated from

achenes of F. a7ithelmintica. (Krewson and

others, 1962). Krewson and his colleagues isolated trivernolin and 1, 3 divernolin which,

along with epoxyoleic acid, it was suggested

might be of interest for experimental determination of their utility in plastic formulations

and in the preparation of chemical derivatives.

Ecology of Vernonia species in eastern Africa

The Ethiopian highlands are largely treeless

grasslands or brushlands except where they

are under cultivation. In Eritrea, V. abyssinica

is often a prominent component of the low,

weedy vegetation of the much mistreated land.

(See fig. 17, p. 30.) On the scarp rising from

the Red Sea, V. abyssinica is found near the

top in areas which have been denuded of forest
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA

and where the soil is disturbed. The plant forms

a scraggling open shrub with small, scattered

leaves and, unless it is in fruit, is difficult to

differentiate from other vegetation. The flowering heads are small and are not colorful enough

to be conspicuous. Wherever cultivation of the

soil is abandoned for several years, plants of

V. abyssinica are apt to be found, but they do

not appear to persist where natural vegetation,

including scattered trees, becomes well grown.

Farther south, and southwest of Addis

Ababa, in areas of open Acacia forest and

grassland, V. adoensis (fig. 5A) appears to

prefer disturbed soil. While it is perennial from

a large root crown, the thickest colonies occupy

areas along road grades. It may be growing

in almost open grassland or it may compete

with other perennials and shrubs of about the

same stature, but I did not find it under shade.

In east Africa, V. adoensis occupies similar

habitats, but to the southward in Zambia and

South Africa, I have seen it in situations where

soil moisture is somewhat greater. Here the

stems become much taller and the leaves are

usually somewhat larger. Even in these areas,

though, it grows in open sunlight, usually where

the soil has been disturbed (fig. 5, B). Unless

the white phyllary tips are well developed, the

plant is inconspicuous among other vegetation.

Vernonia hymenolepis is a plant of somewhat

higher rainfall areas than the species which

were discussed above (fig. 6). Around Jimma,

Ethiopia, in an area where much of the land

has been disturbed, this species grows in hedgerows and on ground which has been abandoned

for some time. The lanky shrub may become

more heavily branched because of constant

trimming by passers-by. In a more natural

habitat with other shrubs in lightly wooded

areas and often on slopes, the shrub stands

upright to equal or overtop other vegetation

(fig. 6,2?).

To the south in east Africa, V. hymenolepis

remains much the same in habitat preference

and in habit. When it is in flower, the white

phyllary tips are the most conspicuous part of

the inflorescence. I have not seen it growing

where it was overtopped by trees (fig. 6, A).

The several collections which I have made of

V. lasiopus were all growing in areas which

i / .-

^Ä-<

it^-*^.

PN-2025

Figure 5.—Vernonia adoensis growing in disturbed soil.

A, Among roadside vegetation near Adamitullo, Ethiopia, the plant is conspicuous even though most of its

leaves have dried and fallen (Smith, Desta, & Sesai,

4566). B, Near Duiwelskloof, Transvaal, South Africa,

V. adoensis is a component of the vegetation along a

well-watered roadside (Smith, 4675).

were less disturbed than those generally occupied by the species discussed so far. The plant

is persistent in pastures where the brush is not

cut, but colonies are often found in little dis-
AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

m

'5'*i JÍ.fe ¿à^ARI*'?'

PN-2026

Figure 6.—Vernonia hymenolepis growing in eastern Africa. A, Just off the road north of Hoey's Bridge, Kenya,

V. hymenolepis forms an open shrub in the disturbed soil (Smith & Njeroge, 4632). B, Near Sheshemene, Ethiopia,

where the forest has been high-graded for wood but the undergrowth is little disturbed, V. hymenolepis becomes

a tall shrub in competition with surrounding species (Smith, Desta, & Sesai, 4564).

turbed, open areas of native shrubs and grass

where there is adequate moisture. The plant is

somewhat spreading and forms a mass among

competing vegetation, but it apparently does

not tolerate shading (fig. 7, B). V. lasiopus

seems to require more available moisture than

the species previously discussed. I have found

it only in topographical situations where moisture accumulates from nearby slopes. The label

data on herbarium specimens bears this out.

Although V. afroniontana is not a member of

the section as currently constituted, I collected

it several times in Kenya. I have found it only

in upland areas of higher rainfall. It is abundant on the slopes of Mount Kenya and also

grows on the slopes of Mt. Elgon. Often, the

colonies are in thick brush where they are difficult to distinguish because the colonies do not

seem to overtop the surrounding plants. It is

shade tolerant, being found under the open

canopy of forested areas where individual

plants are directly exposed to sunlight for only

a short period each day. In such situations, the

blue flowers and sordid pappus of mature heads

are very difficult to see.

I found that the relatively open Br'achystegia woodlands of Zambia are rich in stengelioid Veynonia species early in the year (see

fig. 7, A). In flat areas in the region of Abercorn, V. sciaphila is showy and abundant. On
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA

better-drained slopes, V. guineensis is less common and scattered. V. subaphylla and V. filipéndula are infrequent, occurring generally as

individual plants. In the Abercorn area, V.

superba, which does not belong to section Stengelia but may be closely related to V. afromontana, grows with V. sciaphila and is fairly

common.

All of these species are partially shade tolerant, growing with associated grasses, herbs,

and shrubs among the scattered forest trees.

They grow only in little disturbed sites away

from roadside banks and cultivated patches.

While some sorting of the species by habitat

preference occurs, the factors responsible are

not immediately evident.

From Ethiopia to the Transvaal, all of the

species of section Stengelia which I observed

shared a common habitat characteristic, with

the exception of those growing on permanently

moist, montane slopes. During the dry season,

most of the habitats are subject to fires. At two

localities, achenes of V. adoensis were gathered

at the margin of a recent burn. V. lasiopus may

generally escape burning because it is in areas

of higher moisture. The others, particularly

the species growing annually from a perennial

rootcrown, are subject to periodic, low intensity fires. Burning obviously does not seriously

damage the underground portions of the plants,

but it could well damage achenes that are still

attached or that have dispersed and fallen

among the nearby grasses and herbs. At the

same time, achenes may well be caught in the

updraft from a fire and become widely distributed over many miles. Thus, the same agent

that destroys propagules can also be beneficial

in maintaining a favorable habitat for the species and may also serve to disseminate achenes.

Unfortunately, I was not in Africa at an appropriate time to observe and collect species of

the group with thin, long-lobed florets and,

usually, a basal rosette of leaves. From the label

data, I find that these species are restricted to

savannas and plains over most of the continent.

In some species, the flowering and fruiting

heads remain on ground level ; other species

have heads elevated on scapes and are no taller

PN-2027

Figure 7.—A, Vemonia guineensis growing in open Brachystegia woodlands in the Luano Forest Reserve near

Chingóla, Zambia. B, Individual plants of V. lasiopus near Mbarara, Uganda, are full and overtop surrounding

vegetation (Smith & Wood, 4627.)
10 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

than the grasses in which they grow. Thus,

the plant is difficult to find, which accounts for

the paucity of herbarium specimens. Also because these plants are so inconspicuous except

when in flower, several of the species have not

been collected with mature achenes. All of these

species grow in habitats strongly influenced by

fire, as is shown by the specimens themselves

and the label data for many of them. The wide

distribution of a number of these species may

well be the result of airborne achenes that were

lofted by the updraft from fires.

Biological units within section Stengelia

The arrangement of the species in this paper

does not imply any phyletic order. Naturally,

those species which are most similar morphologically are grouped. The association of species

is otherwise strictly fortuitous.

I have begun the presentation with F.

adoensis, the type species for section Stengelia,

primarily because it is the one species I am sure

belongs to the section. Actually, morphologically similar species which follow V, adoensis

probably are members in good standing of the

section. For many of the species, my only reason for listing them is that an earlier botanist

said that they belong to the section. I shall

leave the task of defining the section to someone with sufficient time to investigate in detail

the bulk of the African species of Vernonia.

Only in this way can the sectional lines be

drawn with assurance.

Until recently, no chromosome counts have

been reported for Vernonia species belonging

to section Stengelia. The most recent atlas

known to me (Bolkhovskikh, Z., et al., 1969)

lists the following counts :

V. anthelmintica 2n =: 20 Mehra, P. N., et al., 1965.^

2n = 54 Parameswar, 1960.^

V, guÍ7ieensis 2n —: 20 Mangenot, S. & G.

Mangenot. 1958 & 1962.i

V. kotschianai^ V. adoensis)

2n= 18 Miège. 1960a.i

V. prhnulinai— V, gei^heriformis)

2n = 20 Mangenot, S. & G.

Mangenot. 1958 & 1962.^

1 Complete citations may be found in the compendium

cited.

Three of the five reported counts show that

n=10. The count of 2n=:54 for a plant identified as V. anthelmintica may be due to misidentification of the plant counted. It is also

possible that it was a count on a plant from a

polyploid population, but this seems unlikely

in view of counts made recently. The count of

2n=18 for V, kotschiana (sic F. kotschyana)

may represent aneuploidy, but, again, it may

be based on incorrectly identified plant material.

In 1969, S. B. Jones (personal communication) counted several species of section Stengelia. All of these counts have proven to be

n=::10. One of these was a recount of V. anthelmintica, strengthening further my feeling that

the count of 2nr=54 is based on an incorrectly

identified plant. The remainder of Jones's count

are as follows :

V. adoensis n = W

Ethiopia. Perdue 6441. Smith, Desta & Sesai 4566.

South África. Duthie s. n.

y. abyssinica n =r 10

Ethiopia. Smith & Andemeskal 4531.

V. anthelTnintiea n = 10

Pakistan. P. I. 283729.

V. lasiopus n := 10

Uganda. Smith, Wood & Perdue 4612.

V. filipéndula n r= 10

Zambia. Fanshawe 10,018.

Enough counts have now been made over a

wide enough representation of section Stengelia

to assure me that the basic chromosome number

for the species of this section must be n—10.

However, I suspect that polyploidy may account

for some observations which I have made. For

instance, F. p7^ocera, which others have submerged in V, guineensis, is a much larger and

more robust plant with the same basic morphology whose size may be attributable to a

higher ploidy level. Both the size of the plant

parts and the pollen characteristics indicate

the possibility that the entities formerly called

F. calvoana and F. insignis may be polyploid

members of the group in which they have been

placed.

Section Stengelia appears to be composed of
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 11

currently evolving complexes mixed with a few

relatively stable taxa of limited distribution.

The latter are easily defined geographically

and morphologically. It is the complexes in a

state of ñux which have bothered earlier botanists and which bother me today. It is the usual

story of variability almost without limit. Variation in Vernonia section Stengelia is amply

illustrated in figure 8. The tendency among

some earlier workers, with far fewer specimens

to indicate the overlapping of morphological

characteristics, has been to name relatively

small variants. I tend to another extreme of

placing all morphologically similar specimens

in the same group.

The latter course has its pitfalls as surely

as does the naming of all variants. However,

I feel that it more nearly represents the actual

PN-2028

Figure 8.—Variation in a single collection of V. filipéndula Hiern. Habit from E. A. Robinson J^OSG in the Federal

Herbarium of Rhodesia and Nyassaland. Details are from the same collection from specimens deposited as follows :

A, SRGH; B,K\ C, EA; and D, PRE. Details of pubescence (left), leaf size and marginal variation and achene

variation (right).
12 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

biological situation. My impression of Africa

today, on the eastern side of the continent, is

that the whole vast area from the Red Sea to

Capetown is ecologically disturbed. In Ethiopia,

apparently the disturbance dates far in the

past over much of the plateau area. Disturbance

of the natural vegetation in the area south of

Ethiopia appears to be more recent, but sufficiently long ago to have opened up new habitats

into which the vegetation has differentiated.

Species of section Stengelia frequently grow in

obviously disturbed areas or in areas of second

growth vegetation.

A prime example is the type species, V.

adoensis, which I have seen growing only where

the native vegetation has obviously been disturbed. Colonies of this species exhibit a notable

amount of variation which is perhaps more

striking toward the southern end of its geographical range than at its northern end in

Ethiopia. Once it has been seen in the field,

there is no doubt that the southern and northern variants are of the same species, in spite of

the rather wide differences in the plants at the

margins of the range. This led to their recognition as different species at an early date.

Another factor lending variability to populations of stengelioid Vemonia species may be

nearly complete interfertility. While I have no

experimental evidence to support my speculations, I have seen miany areas of Brachystegia

woodland in Zambia in which the populations

of Vernonia species are unusually variable.

Among these plants, it was easy to find characteristics (alien to the species as a whole) which

suggest a species growing sympatrically. This

appears to be particularly true of V, guineensis

and related species, many of which appear to

have interbred with non-stengelioid species of

Vernonia such as F. swperha and V, glabra.

When I visited central and northern Zambia,

I saw many of the herbaceous perennial species

blooming at nearly the same time so that crossing would have been readily possible if no barrier to the reception of foreign pollen were

present.

Even among the shrubby species, variation

by any other mechanism in populations of V,

hymenolepis and V. lasiopus is difficult to explain. Generally, the variation is completely

intergrading within a species population so it

would not seem to be a question of misinterpretation of species lines. Rather, the intergradation of V, hymenolepis and V. lasiopus is probably due to hybridization whenever plants of

the two species grow in close proximity. This is

readily seen in western Uganda, where I found

variable populations of V, lasiopus in which

some plants are difficult to distinguish from

V. hymenolepis. Of course, the ecological disturbance in eastern Africa has provided ample

habitats for the development of intergrades.

Preparation of the report

This series of observations on the species

of section Stengelia is based partly on field observations and partly on a study of herbarium

specimens and pertinent literature. Field work

was carried out in eastern Africa from Eritrea

in Ethiopia in the north to Durban, Natal,

South Africa, in the south. Vernonia populations were observed in the field, herbarium

specimens were prepared and achene samples

were collected for oil analysis whenever mature

achenes were available. During most of the

year following my visit to Zambia, Dr. Dennis

B. Fanshawe, Principal Scientific Officer, Forestry Department, Ministry of Natural Resources and Tourism, continued to collect herbarium specimens and achene samples for this

study, I am most grateful for his wholehearted

cooperation.

Herbarium specimens were borrowed from

several European collections and from many

African herbaria. In addition, when it became

apparent that I must deal with many more

names than I had originally thought necessary,

many institutions were very cooperative in

furnishing photographs of type collections in

their care. All of the type collections were verified insofar as authors cited specimens. The

specimens were studied in detail and finally

sorted into my interpretation of the biological

units.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 13

In preparing the following descriptions, herbarium specimens, chosen for their relative

completeness, were used. Characteristics and

measurements were tabulated and the descriptions were written from the tables. This is not

a comprehensive monograph on the species of

section Stengelia, and the descriptions are intended only to differentiate one species from

another.

Certain characteristics are common to all or

most of the species accepted as belonging to the

section. In all of the species, the phyllaries are

differentiated into a basal, usually chartaceous,

portion and an herbaceous tip. The flowers of

the cauline species are generally clearly differentiated into an elongate tube and an enlarged

limb with triangular lobes. Most of the species

of the rosette-leaved group differ in that the

flowers enlarge gradually from the base and

the lobes are lanceolate and thin. Achenes of

all of the species are costate, tapered from the

base and usually are pubescent or pilose. Pappus ranges from short on the outside to long

on the inside; the setae are numerous, usually

cream color and variously pubescent or ciliate

for the most part. Setae are generally more or

less deciduous although the shortest outer setae

are sometimes slightly broader and tend to persist after the longer setae have been lost. These

characteristics will not be noted in the descriptions unless some feature differs sufficiently

to be notable.

Measurements reported here were all made

on dried herbarium specimens which had not

been restored by soaking or boiling. The measurements for fresh material may be somewhat

greater than I have reported. Leaf length includes the apex and petiole; leaf width was

measured at the widest part of the blade, regardless of shape. Head height was measured

from the top of the peduncle to the top of the

pappus and did not include phyllary tips or

flowers which often extended beyond. Head

diameter was less accurate because I attempted

to adjust it for the lateral expansion caused

by pressing. Flower length is the length of the

corolla exclusive of the anther tube or style

which is often exserted; amount of exsertion

of reproductive structures is largely dependent

upon the age of the flower and is very unreliable

for comparison purposes. Achene length is from

the point of basal attachment to the point of

attachment of the setae ; pappus length is that

of the longest setae on the achene measured.

I have reported maximum lengths primarily.

Many species of this group appear to respond

markedly to environmental conditions and all

plant parts will be much smaller when conditions are poor.

Selection of lectotypes

This study is not intended as a definitive

monograph ; it cannot be definitive until related

sections have been thoroughly studied. Yet,

some of the early authors of species cited several collections without designating a type.

From among these I have selected the specimen

most representative of the description from

those cited specimens which I have seen. However, in many instances, I have not seen all of

the collections cited with the original description, and specimens may exist among which

one may be a better lectotype.

Where I have been reasonably certain that I

have seen the material studied by the author

of a new species, I have designated the holotype

by placing an exclamation mark (!) after the

initial designating the herbarium. The type collections of species described by Spencer Moore

provided a problem in that Moore worked in

London and the same collection is often in the

herbarium of the British Museum as well as in

the herbarium at Kew Garden. Moore probably

had both specimens in hand and did not designate the holotype (the concept is relatively

young). I have probably done an injustice to

the British Museum, but I have designated the

Kew specimen as holotype when it was available to me and the specimen in the British Museum was not. In other instances, the only

specimen is in the British Museum and the

museum has kindly furnished photographs for

my use. These specimens, of course, are the

holotypes.
14 AGRICULTURE HANDBOOK NO. 896, U.S. DEPT. OF AGRICULTURE

The destruction of many collections of the photographs disclosed only a few specimens in

Berlin (Dahlem) herbarium has posed another type collections, but I am sure that others rest,

problem. Type collections deposited there may unrecognized, in general herbaria where the

, ,, . , . .r 1 1 . staff has been unable to spare the time to look

be represented by isotypes m other herbaria, ^^^ ^^^^^ Unfortunately, I was unable to underbut this will require an on-the-spot search of ^^j^^ the search myself. I would recommend

thousands of specimens in all of Europe's major that future students of African Vernonia plan

herbaria. My request to many herbaria for type to spend considerable time in such a search.

Systematic treatment

Vernonia Schreb., Section Stengelia (Sch.-Bip. ex Walp.) Bentham & Hooker. Gen. PI. 2:227

1873.

Subsection Stengelia Sch-Bip. ex Walp. Repert. Bot. Sy^t. 2 : Suppl. 1, 946.1843.

Inner involucral bracts with membranous, colored appendages; heads medium large; outer

pappus bristles or scales short, inner sometimes scattered.

Lectotype species : Vernonia (Stengelia) aífoeriszs Sch.-Bip. ex Walp.

As currently constituted, the section includes annual herbs, perennial herbs, shrubs, and trees to

10 m. tall. Leaves whorled (species with a basal rosette) opposite, or generally, alternate. Heads

borne singly, terminal, to many, terminal or lateral, aggregated in paniculate to corymbose groups ;

phyllaries various, but the inner always composed of a coriaceous basal portion and an herbaceous

tip which may be green, white, or variously colored ; flowers inseparable from those of species of

Vernonia in other sections; achenes generally with basal callus light in color, the fruit increasing

in size upward, often costate, sometimes variously pubescent and/or glandular, with pappus in

several series, the outermost of which is sometimes very short, scalelike and persistent, inner

generally long, flattened or fililform, usually variously pubescent, sometimes early deciduous.

Geographical distribution : Continental Africa south of the Sahara Desert to Natal, South Africa,

one species also in Yemen, one species restricted to the Indian subcontinent.

The following key to species described in this paper is completely artificial. In spite of the

similarities between some of the species, they frequently appear widely separated in the key

because of contrast in some characteristic which may otherwise be unimportant.

1. Plants with erect stems.

2. Trees or shrubs with woody bases and branches.

3. Trees to 10 m. tall with single trunk, sometimes shrubby.

4. Heads to 3.0 cm. hig-h; phyllaries to 3.5 cm. long.

4'. Heads to 1.5 cm. high; phyllaries to 1.3 cm. long.

3'. Shrubs. 5.

5. Upright, moderately branched shrubs to as much as 5 m. tall; heads to about 3,0 cm. in diameter;

achenes not noticeably glandular, 6.

6. Leaves predominantly lanceolate; achenes to 0.45 cm. long, generally black, glabrous; pappus

markedly early deciduous. 4. V. hymenolepis

6'. Leaves predominantly ovate; achenes to 0.35 cm, long, medium brown, moderately pubescent;

pappus persistent. ' 2. V. hracteosa

5'. Shrubs to 2.5 m. tall or less; heads to 2.0 cm. in diameter; achenes glandular or so densely pilose

that the surface is not visible. 7.

7. Shrub not more than 1.0 m. tall, densely branched; leaves crowded; heads strict, taller than

broad; achenes densely white or golden pilose. 14. V. vallicola

7'. Shrubs 1.5 to 2.4 m. tall, moderately to openly branched; leaves moderately to openly spaced;

heads as broad or broader than tall; achenes glabrous to pilose, glandular surface visible

through hairs. 8.

2.

3.

4.

3. V. adolfi-frederici

11. y. oxyura
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 15

8. Openly branched; leaves to 5.0 cm. long, scattered on branches; tube and limb of flower about

equal. 6. V. abyssinica

8'. More compact shrubs; leaves more than 5.0 cm. long, stems and branches leafy; tube up to

twice as long as limb. 9.

9. Stems to 1.5 m. tall or less, perhaps more herbaceous than woody; phyllary tips often

attenuate and conspicuously spreading. (Known only from Ethiopia.)

9. V. petitiana

9'. Stems to 2.4 m. tall, sometimes more, woody; phyllary tips small to petaloid, lanceolate,

sometimes reflexed or spreading. 10. V. lasiopus

2\ Annual herbs or plants with herbaceous shoots produced annually from a perennial root crown. 10.

10. Annual herbs. 11.

11. Outer phyllaries not conspicuously long. Plant of India, Ceylon, and Pakistan. 8. V- anthelmintica

IV, Outer phyllaries often conspicuously long, spatulate. Widely distributed in southern and eastern

Africa. 7. V. stenolepis

10'. Perennial root crowns from which herbaceous tops are produced annually. 12.

12, Heads borne on scapes separate from and before leafy stems. 28. V. pumila

12'. Heads borne on leafy stems. 13.

13. Leaves densely lanate beneath. 14.

14. Heads single or few, to 2.5 cm. high, 4.0 cm. in diameter. 13. V. procera

14'. Heads usually numerous, to 1.5 cm. high, 2.0 cm. in diameter. 12. V. guineeiisis

13'. Leaves pilose or pubescent, but not lanate beneath. 15.

15. Outer phyllaries to 4.0 cm. long, lanceolate, yellow-green without, dark red within.

18. y. nigritana

15'. Phyllaries less than 2.5 cm. long, not bicolored as above. 16.

16. Basal callus confluent with costae on achenes. 19. V, sciaphila

16'. Basal callus discreet. 17.

17. Phyllary tips conspicuously veiny. 18.

18. Stem foliage closely investing the several heads. 17. V. campicola

18'. Stem foliage separated from the single or few heads by a short peduncle.

5. V. buchingeri

17'. Phyllary tips various, but not conspicuously veiny. 19.

19. Phyllaries with marked midribs, 20. V. sclerophylla

19'. Phyllaries various, but lacking marked midrib. 20.

20. Branches originating at leaf axils shortly below heads so that the older heads

are partially hidden, heads terminal on main stem and branches.

1, V. adoensis

20'. Branching shorter and the upper smaller so that the several heads are crowded,

but none are hidden or the heads long pedunculate. 21.

21. Leaves lanceolate, glandular above and below; heads to 1.5 cm. high, 1.5 cm.

in diameter. 15. V. benguelensis

21'. Leaves oblanceolate, glandular beneath only; heads to 2.5 cm. high, 3.5 cm.

in diameter. 16. V. filipéndula

1'. Rosette plants from perennial root crowns (except 21. V. gerberiformis sometimes caulescent). 22.

22. Flowers produced before leaves. 23.

23. Single scape produced from each root crown. 27. V, praecox

23'. Several scapes produced from each root crown. 24.

24. Only one head borne on each scape. 29. V. chthonocephala

24'. Several heads in an open panicle borne on each scape. 26. V. denudata

22'. Flowers borne above a rosette of foliage (except 21. V. gerberiformis sometimes caulescent). 25.

25. Several heads borne on each scape. 26.

26. Heads borne in an open panicle; phyllary tips scarious. 25. V. subaphylla

26'. Heads closely crowded at summit of scape; phyllary tips herbaceous. 24. V. praemorsa

25'. A single head borne on each scape. 27.

27. Leaves largely immature at time of anthesis; phyllaries usually nearly 1.0 cm. wide.

23. V. acrocephala

27'. Leaves well developed at time of anthesis; phyllaries lanceolate, narrow. 28.

28. Leaves 23.0 cm. long by 6.0 cm. wide; scapes to 0.4 m. tall; heads to 3.0 cm. in diameter;

plants sometimes caulescent. 21. V. gerberiformis

28'. Leaves to 9.0 cm. long by 1.5 cm. wide; scapes to 0.15 m. tall; heads to 2.0 cm. in diameter;

always acaulescent. 22. V. nyassae
16 AGKICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

1. Vernonia adoensis Sch.-Bip. ex Walp. Rep. Bot Syst. 2:946. 1843. (Figs. 9, 10, and 11, See also

figs. 1, 2A, and 5.)

Type Coll : Schimper 318, In inferiore et media regione septentrionali montis Scholoda

[Seelloda Mt.] 1 Dec, 1837. (BM, CAL, FI, G, K, US).

Stengelia adoensis Seh.-Bip. ex Höchst. Flora 24: Intelligenzbl. 1841. I (2) : 26. 1841.

Nomen nudum,

Vernonia kotschyana Sch.-Bip. ex Walp. Op. cit. 947.

Type Coll: Kotschy 290, In valle prope pagam Cordofanum Millees. 4 Dec, 1839.

(BM, BR, CAL, G, GH, K, NY).

V. macrocephala A. Rich. Tent. Fl. Abyss. 377. 1847. (non Less.) Nom. illeg, (later

homonym).

Type not seen. Synonymy fide Oliver & Hiern, Fl. Trop. Africa 3:291. 1877.

Ascaricida adoensis (Sch.-Bip. ex Walp.) Steetz in Peters. Reise nach Mossambique, etc.

358.1862^64.

A. richardi Steetz in Peters. Loc cit.

Vernonia grantii Oliv. Trans. Linn. Soc London 29:92.1873.

Type Coll : Grant s. n. In Unyoro forests. Nov., 1862. (K !).

Candidea grantii (Oliv.) Stapf, Bot. Mag. 149 : t. 5689, 7255.1923.

F. tenoreana Oliv. Loc cit.

Type not seen.

F. polymorpha var. accedens Vatke. Linnaea 39:477. 1875.

Type Coll: Schimper 817. Auf Bergen, Amba Sea 6,500^ über Meer. 22 Nov. 1862.

(BM,E).i

F. polymorpha var. adoensis (Sch.-Bip. ex Walp.) Vatke. Op. cit. 476.

F. polymorpha var. ambigua Vatke. Op. cit. 477.

Type Coll: Schimper 817? Ebene Hamedo dicht am Mörebb Thel 4,200' über Meer.

(BM).

F. shirensis Oliv. & Hiern in Oliver. Fl. Trop. Africa 3:291. 1877.

Type Coll : Meiler s. n. Lov^er valley of river Shire. Zambesi. May, 1861. (K !).

F. tigrensis Oliv. & Hiern in Oliv. Op. cit. 290.

Type Coll : see F. polymorpha var. accedens Vatke.

F. woodii 0. Hoffm. Bot. Jahrb. Syst. 38:198. 1906.

Lectotype Coll : Wood 8155, Near Pinetown, Natal. Apr., 1901. (K, NH).

F. integra S. Moore. J. Bot. 46:39. 1908.

Type Coll: Eyles ^77 Rhodesia. Mazoe. Bernheim Hill 4,300-4,800 ft. Mar. 1906.

(SRGH).

Vernonia latisquama Mattf. Bot. Jahrb. Syst. 59 : Beibl. 133 :5.1924.

Type Coll : Milbraed 836-Í. Kamerun Übergangsund Kampfgebeit die Savanne an der

Nordgrenze der Hylaea südlich des Sanaga zv^ischen Jaunde und Dengdeng unweit der Vereinung von Lom (Sanaga) und Djerem. Etwa 155 km. N. 0. Jaunde.

Feb., 1914. (K).

F. fulviseta S. Moore, J. Linn, Soc. Bot. 47:266. 1925-27.

Type Coll : Buchanan i^P7. Nyassaland. 1891. (Photo — BM !).

Candidea stenostegia Stapf. Bot Mag. 149 : t. 8981. 1923.

Type Coll: Bobbins s. n, Narawassa Prov., Nigeria (Grown from seed at Edinburgh

Botanic Garden). (K !).

Vernonia stenostegia (Stapf) Hutch. & Dalz. Fl. W. Trop. Africa 2:164, in clavi, 166.

1931.

1 Also the type of Vernonia tigrensis Oliv. & Hiern.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 17

/'.N

^ 7

'7 ^

l.im|ii-n ilrr Al.y%.iiilfuin.

31*«. MionrrlU jM|»4>wiU

< . it. .«'ftHn* n,r.,( („„f.,,

SI

Í > ^

Figure 9.-

PN-2029

-Type collection of V. adoensis Sch.-Bip. ex Walp. A, Habit (US) ; B, detail of head (G) ; C, achene (US) ;

D, flower, dissected and E, unopened (expanded in water) (G). Scales in millimeters.
18 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

Herbaceous perennial from a large root crown; stems growing to 2.5 m. tall during the wet

season. Stems rarely glabrate to, usually, thinly to densely pubescent, branching at the top as heads

are produced. Leaves to 26.5 cm. long, to 8 cm. wide, usually lanceolate to ovate-lanceolate, rarely

ovate ; blade scattered pilose to moderately pilose, sometimes shortly so, above, moderately to densely

pubescent below ; margin irregularly dentate and often undulate ; apex obtuse to, generally, acute ;

base acute, rarely abruptly acute, usually cunéate, the blade occasionally decurrent along the sides

of the petiole to the stem; petiole to 2.5 cm. long, frequently 1 cm. long or less. Heads terminal on

the main stem and on branches produced immediately below the first head so that the heads are

ii*i«

'S^-è. tU^À

PN-2030

Figure 10.—A, Plants of V. adoensis along the roadside at the edge of a plain near Kasese, Uganda (Smith, Wood &

Perdue 4626). B, Ato Desta and Ato Sesai collecting achenes from a roadside colony near Adamitullo, Ethiopia

(Smith, Desta & Sesai 4566). C, Flowering heads of a plant near Abercorn, Zambia (Smith & Richards 4673).

Note the similarity to (D) flowering heads of plants in the colony shown in B.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 19

• 1. V. adoensis

200 400 600 800 MILES

_J I I I I I I

~~1 r400

~1 1

800 1200 KILOMETERS

Figure 11.—Distribution of V. adoensis.

PN-2031

often cymosely arranged, few to many, to 2.5 cm. high, to 3.5 cm. in diameter; longest phyllaries to

2.4 cm. long, lanceolate, outer usually green, uniformly herbaceous, pubescent, inner chartaceous,

glabrous below, herbaceous, rarely puberulent above and the latter portion frequently broadly

ovate, white, rarely pink or green ; flowers to 2.3 cm. long, tube long and generally glandular, rarely

scattered pilose, where the corolla abruptly expands, lobes triangular-lanceolate to lanceolate;

achenes to 1.7 cm. long, usually ca. 1.4 cm. long, light to dark brown, scattered to thickly pubescent,

usually 20 costate, sometimes with alternate 10 costae prominent, sometimes light in color ; callous

tip discreet ; pappus to 1.4 cm. long, flat, obtuse to usually acute, often ciliate at tip, sometimes finely

pilose overall ; phyllaries extending laterally in drying.
20 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

In the northern part of its range, F. adoensis tends to be somewhat shorter and more pubescent

with the outer phyllaries more evidently green. Southward, a greater proportion of the phyllaries

are broad or bear colored tips. The most noticeable feature in the field is the large, light-colored

phyllaries around the heads.

For those species growing in Zambia, D. B. Fanshawe has provided notes which are too valuable

to be edited. Thus, I shall quote these verbatim. His note for V. adoensis is the following :

**A stout, woody-based, perennial herb or shrub, usually multistemmed with erect spreading

branches. Bark grey-brown, smooth, lenticillate, slash green, thin, soft, sappy. Flowering and

fruiting virtually all the year around. Distribution territory wide. A secondary species on the

fringes of riparian and swamp forest, miombo, chipya, munga and Kalahari woodland as well as

roadsides, waste places, etc. In Malawi, a decoction of roots and pounded leaves are used to treat

indigestion, a root infusion with roots of Cissampelos mucronata to treat VD and leaves with those

of the same Cissampelos rubbed into the skin to reduce swellings (authority Williamson). The

Zulus use an infusion to treat influenza and stomachache, a leaf infusion for chronic cough and

a root decoction for pains in the chest. The Zulus inject the infusion as an enema in feverish conditions and use the plant as an antiparasiticide. The Jindwe rub a powder made from the burnt wood

into incisions to treat rheumatism (authorities Watt and Brandwijk. ) ''

Several native uses for F. adoensis have been recorded. Grover reports (herbarium label) that

the natives use the hollow stems for drinking pombe (native beer) near Port Victoria, Kenya. In

Tanzania, it has been reported several times that the roots are used as treatment for abdominal

pains and tuberculosis (R. E. S. Tanner). The leaves (Nioka, Congo) and the roots (Masai, East

Africa) are used as an emetic. The Tanzanians also use the roots to treat gonorrhea; and they cut

the roots into small pieces, soak them in water, and use the water to wash children with skin disease

(Tanner). White reports that the Chewa (Zambia) make a salt called '*chikungu" from the ash

of the plant. The species is reported to be cultivated by the natives as a medicinal plant in Natal,

South Africa (Wood).

F. adoensis is generally distributed along the eastern side of the African continent from Eritrea

in Ethiopia to southern Natal, South Africa. Through the equatorial region, it is usually found at

elevations of 1,000 m. or higher. It has been collected infrequently westward through the Republic

of the Congo, Nigeria, and as far west as Gambia (see fig. 11, p. 19).

Selected Collections: ETHIOFIA: Eritrea; Hamasen, Abrascido. A, FiorL 1729, Feb. 16, 1909. (FI).

Da Halibanet ad Amba Derho. A, TellinilOlO. Dec, 1902. (FI). Scioa; Ambo. 2,000 m. G. Giordano

2126 (FI). Addis Ababa-Shashemene road near Adamitullo. Ca. 5,400 feet. C. Earle Smithy Jr.,

Desta Hundesa & Ato Sesai. 1^566. Dec. 28, 1966. (BR, EA, FI, K, MO, US). Amhara; Dembia.

Boscaglia rocciose presso Asosa. G. Chiovenda 2726. Oct. 27, 1909. (FI). Gojam; Boscaglia ad

ouest del villagio di Selcién. R. Pichi-Sermolli 962, Feb. 8, 1937. (FI). SUDAN: South of El Obid,

Nuba Mts. C. E. Ltigard U. Aug. 11, 1921. (BM, MO). Darfur Prov, ; Kebkabiya-Zalingei Road.

900-1,000 m. /. E. Dandy 29. Jan. 1, 1934. (BM, MO). Equatoria; near Upper Nile. Along the

Bahr elJebel between Nimule and Gondokoro. E. A. Mearns 3051. Feb. 4-25, 1910. (GH, K, NY,

US). KENYA: Eldoret District, Elgeyo Escarpment. 8,000 feet. R. L. Harger s. n. 1926. (BM).

Central Kavirondo District, Escarpment, Kisumu-Mutet Road about 4 mi. N. of Kisumu. R. B.

Drummond & J. H. Hemsley U82. Sept. 27, 1953. (K). Western Region, Port Victoria. G. P,

Glasgow Jf5/S5. Sept. 28, 1945. (EA). UGANDA: West Nile Distr. Maracha County, in valley % mi.

S.E. of Maracha rest camp. 4,600 feet. R. J. Chancellor 126. Aug. 7, 1953. (K). Mbale Distr.,

Eastern Prov., 1 mi. S. of Nsola River in W. Bugwe L. Forest Reserve. 3,850 feet. G. H. S. Wood

234. May 5, 1951. (BR, K). Western Prov., south of Mubuku on the Kasese road. 3,500 feet.

C. E. Smith, Jr., D. Wood & R. E. Perdue 4626. Jan. 19, 1967. (BM, EA, FI, K, PRE, US). Livengo,

Buddu. A. P. G. Michelmore 1301. Mar. 12, 1936. (BR, EA). RUANDA: Biumba. Region du Mutara,

environ de Nyakagenge, colline Nyakagenge. 1,420 m. G. Troupin 7124. Apr. 17, 1958. (MO). Pare

National de la Kagera. /. Lebrun 9813.1938. (BR). BURUNDI : Rive orientale du Lác Kivu, a Mushao.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 21

1,460-1,500 m. H. Humbert 8U8. May, 1929. (BR). TANZANIA: Bugene, Bukoba Distr., 5,000 feet,

A. E, Haarer 2335. Oct., 1931. (EA, K, NY). Mara Region, Musoma Distr. Hambe ya Mwakenylui,

S. boundary. 4,800 feet. P. J, Greemvay & M. Turner 10370, June 7, 1961. (EA, PRE). Tabora

Region, Mwanza Distr., Usagara, Bukumbi. 3,700 feet. R, E. S. Tanner 1217, Feb. 19, 1953.

(BR, K). Ufipa, L. Kwela. 5,100 feet. A. A. Bullock 2968, July 5, 1950. (K). Iringa Prov., Mporoto

Mts., 7,000 feet. C, G. McGregor 23. Aug. 1, 1936. (EA). Rungwe Distr. Near Materna village and

Lake Nyassa. 900 m. H. M. Richards 98U. May 21, 1957. (K). ZAMBIA: Northern Prov. In canon

near Abercorn. C. E. Smith, Jr. & H. M. Richards Í673. Feb. 22, 1967. (EA, K, SRGH, US). Kitwe

Distr., Kitwe. D. B. Fanshawe 10141. July 16, 1967. (NDO, US). Eastern Prov., Fort Jameson,

Ndolo Forest Reserve, 12 mi. S. of Katete. F. White 2U6. Aug. 22, 1952. (BR, K). Broken Hill

Distr., Broken Hill. 4,000 feet. F. A. Rogers 8124. May, 1909. (SRGH). Mt. Makulu, Makulu Res.

Sta., 10 mi. S. of Lusaka. M. Lister 92. Dec. 14, 1957. (SRGH). Southern Prov. Ca. 12 mi. W. of

Livingstone. C. E. Smith, Jr. 6k7U. Mar. 1, 1967. (K, NDO, SRGH, US). RHODESIA: Vicinity of

Umvukwe Mts., 5 mi. N. of Banket. R. J. Rodin 4389. Apr. 23, 1948. (K, PRE, SRGH). Sebungwe

Distr., near Gunyankas Pan. 0. West. 2352. May 28, 1947. (SRGH). Inyanga Distr. Pungwe River

near P. E. A. border. 3,300^ N. C. Chase 6451. Apr. 19, 1957. (K, SRGH). Inyanga Distr. Honde

Valley. /. B. Phipps 1125. Apr. 18, 1958. (BR, SRGH). Sabi-Lundi Junction near Lundi River.

800^ H. Wild 3444^ June 7,1960. (BR, PRE, SRGH). MALAWI : Kota Kota Distr. Nchisi Mt. 1,600 m.

L. /. Brass 17059. July 31, 1946. (BR, K, SRGH). Central Prov. Namitete River above bridge on

Lilongwe-Ft. Jameson Road. 1,150 m. A^. K. B. Robson 1476. Feb. 5, 1959. (BM, K, SRGH). ZombaBlantyre Road. P. 0. Wiehe N/446. Mar, 17, 1950. (SRGH). MOZAMBIQUE: Niassa-Massangulo.

Pedro i Pedrogâo 3555. June 15, 1948. (EA), Delgoa Bay, Lourenço Marques. H. Junod 12. 1890.

(BR, G). Umbeluzi flood plain. A. J. W. Hornbij 3822. (PRE). SOUTH AFRICA: Transvaal; Between

Duiwelskloof and Tzaneen. C. E. Smith, Jr. 4676. Mar. 5, 1967. (K, PRE, SRGH, US). Transvaal;

Letaba. 3,250 feet. /. C. Scheepers 129. Feb. 22, 1958. (EA, K, SRGH). Transvaal; Barberton Distr.

Barberton. 1,300 feet. /. Thorncroft 94. Apr. 20, 1956. (EA, K, PRE, SRGH). Swaziland. Mbabane.

Black Mbuluzi Valley, 4,000 feet. R. H. Compton 25707. Mar. 3, 1956. (MBA, K, PRE). Natal;

Durban. Spring Grange. J. M. Wood 441. Apr. 21, 1887. (CAL). CHAD: Between Lake Chad &

Bornu. P. A. Talbot 1015. Feb., 1911. (BM). REPUBLIC OF CONGO: Mt. Adjigu (Terr. Mahagi).

1,800 m. Taton 447. Mar. 18, 1947. (BR). Entre Faradje et Aba. /. Lebrun 3414. July, 1931,

(BR, K). Kibali-Ituri Prov. 10 km. S. de Mahagi, tete de source de TOri. R. Germain 4223. Aug. 10,

1945. (BM, BR, K). Kivu-Central. Rulenga. L. Lily 810. Oct. 22, 1953. (BR). Plaine de la Ruzizi,

Luvungi. R. Germain 6190. Oct. 20, 1950. (BR). NIGERIA: Bauchi Prov. Sabon Gari, Tama, Lame

Distr. 2,800 feet. G. V. Summerhayes 34. Aug. 24, 1954. (K). Bauchi. H. V. Lely P160. Feb., 1929.

(K). Oyo Prov. Along the Ibadan-Ijebu Ode motor road. S. Tamajong FHI 19512. Aug. 5, 1946.

(K). GUINEA: Timbo. H. Jacques Felix 1911. Oct., 1937. (K, P). PORTUGUESE GUINEA: Bambadinca.

Bafata. Espirito Santo 3833. Dec. 20, 1945. (K). Falacunda. Sao Joâo. Espirito Sarito 2225. Oct. 24,

1945. (BR, K). Buba, Lagoa de Cufada. /. d^Orey 214. Jan. 24, 1954. (K). GAMBIA: Abeokuta.

Rowland s. n. Jan., 1893. (K).

V. adoensis is the taxon on v^hich the section Stengelia is based. The name, Stengelia adoensis,

was published as a nomen nudum in a list advertising Schimper's collections in 1841. It v^as validated as Vernonia adoensis by Walpers in 1843 who defined the section (as a subsection) at the

same time. Steetz transferred the name to the segregate Ascaricida in 1862-64, but this placement

was not accepted by other workers. The type of A. Richard's V. macrocephala (illegitimate because

it is a later homonym) was examined by Oliver and Hiern and placed in synonymy of V. adoensis

in 1877, This automatically placed Steetz' Ascaricida richardi (a nomen novum for V. macrocephala) in synonymy. Vatke's conglomerate V. polymorpha of 1875 consists largerly of V. adoensis.

The remaining synonyms were erected on the basis of smah morphological differences and

geographical isolation which was valid at the time they were proposed. All of them are now known

to be a part of a large complex of intergrading plants which cannot be satisfactorily kept apart.
22 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

This species is not liable to be confused with related species, although snippets from the branch

ends of V. bracteosa and the tops of V. adoensis from the southern end of the range of the latter

may resemble one another. V. bracteosa is a shrub and V. adoensis grows annually from a perennial

rootcrown. Furthermore, the lower portion of the outer phyllaries of V. adoensis are usually green

and strongly curved giving the heads an appearance which is duplicated by no other species known

to me.

2. Vernonia bracteosa. 0. Hoffm. Bot. Jahrb. Syst. 30:421.1901. (Figs. 12 and 13.)

Type Coll: Goetze 793. Uhehe. (Photo —BM).

Shrub to 4.5 m. tall. Stems generally scattered pilose, rarely glabrate, often more densely

pubescent on specimens from the southern end of the range ; leaves to 17 cm. long, 5.5 cm. wide,

ovate to long ovate ; margin irregularly dentate to, rarely, few dentate ; blade scattered pilose above,

generally moderately pilose, rarely scattered pilose and glandular below ; apex acute, base frequently

abruptly acute, rarely nearly cunéate, blade decurrent along the petiole as a very narrow flange

of tissue, thus the free petiole seldom more than 1 mm. long. Heads terminal, cymosely arranged

for the most part, smaller, more closely crowded and irregularly arranged in plants to the southward, generally ca. 2 cm. high, 2.5 cm. in diameter, rarely to 3 cm. by 3.5 cm. ; phyllaries to 2.1 cm.

(exceptionally, 2.8 cm.) white to pink, mauve or purple tipped; flowers to 1.4 cm. long, white,

mauve or purple ; achenes to 0.35 cm. long, medium brown, moderately pubescent for the most part,

only rarely with alternate costae larger and lighter in color, callus discreet; pappus to 1.3 cm. long,

flattened, long acute, ciliate, usually without single very short outer row.

"'3 4'. 5'

.11! Illl ¡11! 11' ii.iiul

Figure 12.—Type collection of V. bracteosa O. Hoffm. A, Habit; and B, detail of heads. (Photos, BM.)
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 23

• 2. V. bracteosa

* 3. V. adolfi-frederic

0 200 400 600 800 MILES

I \ \ I I I I L_J

0 400 800 1200 KILOMETERS

.^ ^''^ :• .TrElisabethvi

PN-2033

Figure 13.—Distribution of V. bracteosa and V. adolfi-frederici.

V. bracteosa an eastern African species which, to judge from label data, grows only where water

is generally available. Toward the southern end of its range in Rhodesia and Mozambique, specimens become quite variable with smaller heads irregularly arranged and the bottoms of the leaves

more densely tomentose. Without experimental data, this is impossible to prove, but it looks as

though V. lasiopus may be strongly influencing the phenotype.

The roots of V. bracteosa are reported to be used as medicine for elephantiasis in Tanzania

(Carmichael).

V. bracteosa is restricted to highland areas south of the equator in eastern Africa with its greatest

concentration in Rhodesia. (See fig. 13.) I may have misplaced the Congo collection because of

scant label information. It apparently is not abundant anywhere.

Selected Collections: UGANDA: Muhavura-Mgahinga Saddle. G. Schaller W. Nov. 28, 1959. (EA).

TANZANIA: Arusha Distr. Mt. Meru, south slope. 5,800 feet. P. J. Greenway 1039Í. July 2, 1961.

(EA). Lushoto Distr. Shagai Forest near Sunga, West Usambaras. 1,950 m. R. E. Drummond &

J. H. Drummond 2578. May 17, 1953. (BR, EA, SRGH). Shume Forest Reserve, Usambara Mts.

5,000-6,000 feet. E. D. Mabes H9. July, 1924. (K). Southern Highlands Prov. Iringa Distr. Isata.

6,000 feet. W. Carmichael 393. Mar., 1954. (EA). Kyimbila Distr. N. of Lake Nyassa. A. Stoltz

2^9. (GH, K, NY, SRGH). Songea Distr. Matengo Hills. About 1.5 km. N. of Miyau on River

Utili. E. Milne-Redhead & P. Taijlor 9003. Mar. 2, 1956. (EA, K). REPUBLIC OF CONGO: Kapanda.
24 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

(Katanga). W, Robyns 1805.1926. (K). RHODESIA: Umtali. Stapleford Forest Reserve. F. L. Orpes

11/56, Aug. 18,1956. (SRGH). Chirinda outskirts. 3,700-4,000 feet. C. F. M, Swynnerton273. 1906.

(K, SRGH). Melsetter. Mt. Belinda. Queen Victoria Memorial Herbarium 6778 Dec. 23, 1931.

(SRGH), Melsetter Distr. 2 mi. N.W. of Melsetter. 5,000 feet. L. Methuen 13i a&b. Aug. 24, 1961.

(K). Chimanimani Mts. R, C. Munch 126. June 2, 1948. (SRGH). MOZAMBIQUE : Vumba Mt, region

of Manica. 900 m. A. Gomez e Sousa Í783. Aug. 24, 1962. (PRE).

3. Vernonia adolfi-frederici Muschler. Bot. Jahrb. Syst. 46:68. 1911. (See fig. 13.)

Type Coll : Not selected. Specimens not seen.

Trees to 10 m. tall, or shrubs. Stems glabrate to usually velvety, sometimes with trichomes intermixed; leaves to 31 cm. long, 6.5 cm. wide, ovate-lanceolate, rarely ovate; margin apiculate-dentate;

blade nearly glabrous to generally moderately pilose to rarely densely pilose above, scattered

glandular, moderately pilose, particularly along the veins and venules to densely floccose or

pubescent below; blade sometimes decurrent along the petiole and forming auricles at the stem;

otherwise, petioles to 4.0 cm. long, pubescent to densely velvety. Heads borne terminally, rarely few,

frequently closely crowded in a cymose arrangement, to 3.0 cm. high, to 3.5 cm. in diameter,

phyllaries to 3.5 cm. long, rarely longer, often lanceolate acute and externally pilose or puberulent

or ciliate, the shorter sometimes more abruptly acute and proportionally broader, white to pink ;

flowers to 1.7 cm. long, glandular where the tube expands near the top, petals lanceolate, often

with glandular tips without, white to mauve, rarely purple; achenes to 0.45 cm, long, brown or

dark brown, scantily pubescent to densely so, often lightly glandular, costae about 15, rarely lighter

in color, frequently topped with a flaring margin of light colored tissue, callus discreet; pappus to

1.4 cm. long, flat, acute, ciliate, without short single outer ring.

V. adolfi-frederici is accepted on the basis of 2 specimens from Kew Herbarium which were

annotated as having been compared with the type which I have not seen. Habitat data indicate that

this is a montane species which seems to have been collected primarily at Ngorongoro Crater, the

Rumenzori Mountains and Mount Bambutu in the Cameroons at elevations from 6,500 to 9,800 feet

above sea level (see ñg. 13). It is separable from F. bracteosa by its large shrub to tree habit and

the apiculate dentations of the leaf blades ; from F. hymenolepis by its pubescent achenes,

Dunlap has reported that the leaves of 7. adolfi-frederici nre eaten by the natives in Cameroons.

Selected Collections: CAMEROON: Mt. Bambutu. 2,450 m. M. R. Letouzey 87, Nov. 3, 1946. (P).

Cameroon Mts,, above Buea. 1,800 m. /. Milbraed 10817. Dec. 21, 1928. (AAH). UGANDA: Western

Prov. Toro Distr. Ruwenzori, Nyamleju. 3,200 m. H. A. Osmaston 8681. Jan., 1951. (K). Western

Prov. Kigezi Distr. Virunga Mts. Mghinga. 10,300 feet. G. Taylor 1988. Nov. 22, 1934. (BM). Kigezi,

Bufumbira. 9,800 feet. Morrison 86. Sept. 9, 1952. (EA). Ngorongoro Crater, west and south

summits. 7,500-8,000 feet. B. D. Burtt Í2H. Sept. 23, 1932. (EA, K). S. of Meru. 6,500 feet.

G, W.Irens 761^. (EA).

4. Vernonia hymenolepis A. Rich. Tent. Fl. Abyss. 1:378. 1847. (Figs. 14, 15, and 16. See also figs.

land 7.)

Type Coll : Petit s. n. Choa (Shoa, Ethiopia). (K).

Stengelia calvoana Hook. f. Proc. Linn. Soc. Bot. 7:199.1864.

Type Coll : Mann 1288. Camer. Mount. Elevât. 2,500-7,000 ft. Dec, 1861. (K !).

Stengelia insignis Hook, f. Loc. cit.

Type Coll : Mann 1925. Camer. Mount. 3,000-7,000 feet. Dec, 1862. (GH, K !).
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 25

&.* 'V

«^

^

•!"-3B^

PN-2034

Figure 14.—Type collection, V. hymenolepis A. Rich. A, Habit; B, detail of heads; C, broken flower unopened and

dissected (expanded in water) ; and D, achene (immature, but dehiscence of pappus is typical). (K). Scales in

millimeters.
26 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

r^ *.^ "••. _ . >'. -^í ,

'»:-'-t

í.«í.

T¿?f*î{ y/fvf'f^^^^^i

PN-2035

Figure 15.—A, A well-developed shrub of V. hymenolepis in little-disturbed forest on the slope of Mt. Elgon, Kenya

(Smith & Njeroge 4638). B, Plant heads in bloom near Shashemene, Ethiopia (Smith, Desta, & Sesai 4565).

C, Plant heads in bloom near Borie, Ethiopia (Smith, Desta, & Sesai 4559). The phyllaries in this specimen are

reflexed; those in B are upright. D, Colony of upright, open plants (as in C) in a disturbed roadside habitat near

Borie; and E, mature heads on a plant in the same colony.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 27

Vernonia calvoana (Hook, f.) Hook. f. Bot. Mag. 94 :t. 5698. 1868.

V. insignis (Hook, f.) Oliv. & Hiern in Oliv. Fl. Trop. Africa 3:292.1877.

V. rothii Oliv. & Hiern in Oliver, Fl. Trop. Africa 3:293. 1877.

Type Coll: Roth 3^6. Ankober. Nov., 1841. (K!).

V. leucocalyx O. Hoffm. Bot. Jahrb. Syst. 30:422. 1901.

Type Coll: Goetze 928. Deutsch-Ostafrika: Ukinga-Berge. 1898-1901. (K).

V. homilocephala S. Moore. J. Linn. Soc. Bot. 35:322. 1902.

Type Coll : Elliot 7058. Mau to Nologoseri. 7,000 feet. Dec. (K !).

V. calvoana var. microcephala Adams. J. W. African Sei. Assoc. 3:118. 1957.

Type Coll: Lightbody FHI26259. Cameroon: mile 110, Nkambe-Misage road. Alt.

6,000-7,000 feet. Sept. 1950. (K!).

V. leucocalyx var. acuta Adams. J. W. African Sei. Assoc. 3 :120. 1957.

Type Coll : Keay & Lightbody FHI28366. Cameroon : Bamenda Prov. Bafut-Ngemba

F. R. near Lake Bambulue. Alt. c. 6,200 feet. Jan., 1951. (P).

• 4. V. hymenolepis

• 6. V. abyssinica

A 7. V. stenolepis

0 200 400 600 800 MILES

_L_J \ I I I I I

1 T -

PN-2036

Figure 16.—Distribution of V. hymenolepis, V. abyssinica, and V. stenolepis.
28 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

Shrub to about 5 m. tall, usually less. Stems densely short pubescent to glabrate, rarely with

intermixed triehomes ; leaves to 20 cm. long, often less, to 7.0 cm. wide, lanceolate to ovate-lanceolate,

infrequently ovate; margin apiculate dentate, blade glabrous to scattered puberulent or pilose

(rarely glandular) above; often densely pubescent, sometimes scattered pilose or pilose only on

veins, then sometimes glandular beneath ; apex generally acute, sometimes apiculate, base cunéate

acute or acute, usually decurrent on petiole, sometimes strongly so and forming auricles at stem.

Heads terminal in more or less crowded cymose or paniculate arrangement; to 2.5 cm. high, to

3.0 cm. in diameter (most frequently about 1.5 by 1.5 cm.), phyllaries upright or the herbaceous

tip reflexed, to ca. 2.3 cm. long, usually 1.2 to 1,5 cm. long, white, lanceolate to (less frequently),

broadly lanceolate, acute, sometimes somewhat pubescent or lightly glandular without; flowers

to 1.5 cm. long, tube only about % of total length, glandular and/or slightly pilose at top of tube,

white to mauve ; achenes turbinate, to 0.45 cm. long, glabrous, rarely with a few hairs, generally

nearly black, less often medium brown, occasionally with lighter costae, sometimes glandular, callus

discreet, usually small; pappus to 1.4 cm. long, usually 0.7 to 0.9 cm. long, flat, ciliate, strongly

deciduous.

F. hymenolepis is most nearly uniform in Ethiopia and Kenya; and it becomes more widely

variable southward and westward. Not only are the individual heads less showy than those of the

species previously discussed, but often the coriaceous basal portion of the phyllaries is exposed.

In the larger-headed species, the basal portion of the phyllaries is generally well concealed by the

herbaceous portion of the bracts.

Perhaps the most general characteristic that separates V. hymenolepis from other nearby species

is the ease with which the pappus detaches from the turbinate achene. The achene is often nearly

black, glabrous, small in diameter at the small callus and increasing in diameter gradually upward

until the diameter suddenly decreases just before the attachment of the pappus.

Johnson reports that the wood and leaves of this species are burned, water is strained through

them and evaporated, and the residue is mixed with tobacco to make snuff in central Africa.

F, hymenolepis is frequent in the uplands of central Ethiopia and is distributed southward

through Tanzania. Some collections have been made in the uplands of Ruanda and eastern Congo.

While the collections from Cameroon are larger overall, they are not morphologically distinct from

the eastern collections (see ñg. 16). The high incidence of aborted pollen and the large pollen

diameters of about half of the grains indicate that the Cameroon populations may well be polyploids.

Selected Collections: ETHIOPIA: Uallaga. Ghimbi. 1,820 m. ñ. Milchersich 112, Dec. 12, 1908. (FI).

Anñlo near Dembidollo. 6,300 feet. H ,F. Mooney 6881. Mar. 5,1957. (FI). Keffa Prov. Near Agaro.

C. E. Smith, Jr, Desta Hundesa &Ato Sesai\í560. Dec. 25, 1966. (BM, EA, FI, K, US). Near Borie

close to Beleta Forest on Bonga road. C, E. Smith, Jr., Desta Hundesa & Ato Sesai Í559. Dec. 25,

1966. (BM, EA, FI, K, US). Arussi. About 10 km. from Shashemene. C. E. Smith, Jr., Desta

Hundesa & Ato Sesai Í56J,, Dec. 27, 1966. (BM, EA, FI, K, US). Irgallem. Sidamo, 5,770 feet.

H. F. Mooney 5321. Dec. 29, 1953. (K). Tra Dorzé en il Lago Regina Margherita. A, Vatova 1773,

Feb. 7, 1938. (FI). Agheremariam. 1,750-2,100 m. /. B. Gillett 1U82. Nov. 30, 1952. (BR, EA, FI,

K, PRE). KENYA: Marakwet Hills. 8,500-9,500 feet. /. R. Dale 3U3. June, 1935. (BR, EA, K).

Trans Nzoia Distr. Mt. Elgon. 7,800 feet. C, E. Smith, Jr. & D. Njeroge Í638. Jan. 27, 1967, (BM,

EA, FI, K, US). 5 mi. N. of Hoey's Bridge. C. E. Smith, Jr, & Z). Njeroge 4632. Jan. 25, 1967. (BM^

EA, FI, K, US). North Kavirondo Distr. Kakamega Forest. R. B. Drummond & J. H. Hemsley 4,71,8.

Oct. 15, 1953. (BR, EA, FI, K). Kîsumu. Victoria Nyanza. Tindoret. P. Davoli 16. 29 Nov. 1936.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 29

(FI). Mt. Nyiri. 8,000 feet. 0. Kerfoot 201,1. July 27, 1960. (EA). UGANDA: Northern Prov.

Karamoja Distr. Moroto Mt. M. S, Phillip 762. Dec. 1954. (K). Mt. Debasien. 8,200 feet. W, J.

Eggling 2807. Jan., 1936. (BR, K). Near Bulucheke, S. W. Elgon. 7,000 feet. /. R. Dale U87. Dec,

1939. (BR, EA). Rubaya S. Kigezi 7,000 feet. /. W. Purseglove 2571. Dec, 1947. (K). TANZANIA:

Morogoro, Uluguru Hills. 1,500 m. H. C. Schlieben 4226. Aug. 11, 1933. (BM, BR, G). Northern

Prov. Ngorongoro. 6,500 feet. R. E. S. Tanner 3258. Nov. 17, 1956. (K). Iringa Prov. Idodi. E. G.

Ward 18. Oct. 1936. (EA). Western Prov. Kigoma Distr. Lumbye River, Mahali Mts. 2,800 feet.

/. Newbould & T. G. Jefford 1179. July 30, 1938. (K). Tanga Prov. Lushoto Distr. Shume-Magamba

Forest. 6,000 feet. /. Benedicto 68. Sept. 29, 1955. (EA). Southern Highlands Prov. Rungwe Distr.

Maroka Crater. Geilinger 2617. Sept. 25, 1932. (K). Mbozi. 5,000 feet. H. Horsbrugh-^Porter s. n.

June, 1935. (BM). MALAWI: North Nyassa Distr. Nyika Plateau. 2,300 m. L. /. Brass 17174. Aug.

11, 1946. (K). RUANDA: Shangugu Terr. Astrida-Bukavu road near Uwinka. 2,400 m. G. Troupin

11126. Sept 23, 1959. (EA). REPUBLIC OF CONGO: Kivu. Rumangabo. R. Germain 3018. Dec. 15,

1924. (BR, K). Mts. W. of Lake Kivu, Tshibinda. 2,000 m. H. Humbert 7362. March, 1929. (K).

Mt. Kahuzi, Bukavu-Walikah highway. G. Schaller 129. June 9, 1959. (EA). Van Geysel to Pepa,

Section de Mulongoshi. Thienpont 23. Aug., 1946. (BR). NIGERIA: Ogoja Prov. Obundu Div.

Sonkwala area north of Koloishe. 5,000 feet. H. Savory & R. Keay FHI25069. Dec. 20, 1948. (K).

Oban. P. A. Talbot 396. 1911. (BM). CAMEROON: Adamwa Div. Mambila Distr. Mambila Plateau.

1500 m. F. N. Hepper 1671. 1958. (K). Bamenda Distr. Lakora. 6,000 feet. H. S. Maitland H85.

Apr., 1931, (K). Victoria Distr. Cameroon Mts. 5,700-6,000 feet. R. W. J. Keay FHI286J,7. Jan. 15,

1951. (K). Musake above Buea. 6,000 feet. F. W. H. Migeod i9. Nov. 4, 1927. (BM). FERNANDO

Poo: Near Musola. E. Guinea 1638. Jan. 20, 1938. (K).

5. Vernonia buchingeri (Sch.-Bip. ex Steetz in Peters) Oliv. & Hiern in Oliver. Fl. Trop. Africa

3:289.1877. (Figs. 17 and 18.)

Ascaricida buchingeri Sch.-Bip. ex Steetz in Peters. Reise nach Mossambique, etc. 359.

1862-64.

Type Coll : Schimper 386. Abyssinia. (BM, G, K, P !).

Vernonia nimbaensis C. D. Adams. J. W. African Sei. Assoc. 8 :139. 1964.

Type Coll: Chevalier 21023. Guinée française: Pays de Guerzes. Montagne de N'Zo,

entre 1,200 m. et 1,400 m. d'alt. Mar., 1909. (K, P !).

Herb to 1 m. tall from thickened perennial rootstock. Stems scattered pilose, glandular. Leaves

sessile, to 8.5 cm. long, 2.5 cm. wide, ovate or ovate-lanceolate; margin blunt or apiculate dentate;

blade scattered pilose, glandular above (one collection strongly raised reticulate), veins lightly

pilose, surface glandular below (one collection strongly raised reticulate) ; apex subacute, base

obtuse. Heads terminal, single or several, to 2.5 cm. high, to 3.0 cm. in diameter, phyllaries to 2.3

cm. long, herbaceous tip broad and strongly veined in both type collections ; flowers to 2.2 cm. long,

tube about 2.3 times longer than urceolate top; achenes to 0.6 cm. long, brown, densely pubescent,

pappus broadly flat, stiffly outcurved, to 1.5 cm. long.

I have no doubt about the conspecificity of V. nimbaensis and V. buchingeri in spite of the disjunction in distribution. The former has strongly raised reticulate venation in the foliage, but this

is hardly sufficient to distinguish it. The paucity of collections of this species may be an indication

that it is only an atypical variant of a more common species. Until more specimens are available

and more is known of its biology, I prefer to recognize it as a separate species.
30 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

PN-2037

Figure 17.—Type collection of V. buchingeri (Sch.-Bip. ex Steetz in Peters) Oliv. & Hiern. A, Habit; B and C, flower,

unopened and dissected (expanded in water) ; and D, detail of heads. (K). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 31

V ^ñ/ ^

PN-2038

Fi^re 18.—Holotype of V. huchingeri in the Museum d'Histoire Naturelle, Laboratoire de Phanerogamie, Paris.

A, Detail of heads; and B, habit.

6. Yernonia abyssinica Sch.-Bip. ex Walp. Rep. Bot. Syst. 2:947. 1843. (Figs. 19 and 20. See also

figs. 2, 3A, and 16.)

Type Coll: Schimper 389. In vallibus prope Adoam. Sept., 1837. (BM, BR, CAL, FI,

G, GH, K, NY).

Teichostemma fruticosum R. Br. ex Salt. Voy. to Abyss., etc. Append. IV:lxv. 1814.

Nomen nudum.

Type Coll: Salt s. n. Abyssinia. (BM!).

Vernonia polymorpha var. microcephala Vatke. Linnaea 39 :477. 1875.

Type Coll: Schimper 581. An sterilen Erden auch auf Brat Akern von 4,000-7,500'

... Höhe. Anadehr 7,300' über Meer. 20 Oct. 1862. (BM, NH).

V. abyssinica var. vestita Cuf.

Lectotype Coll: Corradil966. Neghelli, Ethiopia. 26-29 Sept., 1939. (FI!).
32 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

^

|p3

PN-2039

Figure 19.—Type collection of V. abyssinica Sch.-Bip. ex Walp. A, Detail of heads (NY) ; B, flower, entire and

dissected (expanded in water) (NY) ; C, achene (GH) ; and D, habit. (BM). Scales in millimeters.

Upright, branched shrub to 2 or more m. tall. Stems striate, pubescent or puberulent, older stems

with tan bark longitudinally split. Leaves gray-green, sessile, to 5.0 cm. long, to 1.4 cm. wide,

lanceolate; margin dentate; blade puberulent, scattered glandular above, pubescent to floccose,

glandular beneath ; apex and base acute. Heads borne terminally, paniculately to cymosely arranged ;

to 1.4 cm. high, to 2.0 cm. in diameter; phyllaries to 1.1 cm. long, herbaceous tip small, lanceolate,
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 33

often strongly reduced, gray-green to yellowish green, herbaceous tip often strong glandular

without, coriaceous basal portion prominent in more mature heads; flowers to 1.2 cm. long, tube

and limb about equal, rose-violet to lilac to white, often glandular at the top of the tube and on

the tips of the petals without; achenes to 0.5 cm. long, brown, alternate costae sometimes lighter

in color, frequently glandular, moderately to densely softly pilose ; pappus to 0.9 cm. long, flat.

V. abyssinica is an open shrub that is relatively common in the weedy flora of field margins and

abandoned land of the drier areas in eastern Ethiopia and northern Kenya on the slopes and

f

r'

t'^'

%

PN-2040

Figure 20.—A, A straggling shrub of Y. abyssinica in the much-disturbed land at Scitchity, Eritrea, Ethiopia.

The heads (B), when flowering are inconspicuous; but the plants (C) are frequently recognizable when they are

in fruit. (All Smith & Andemeskal 4531.)
34 AGRICULTURE HANDBOOK NO. 396, U.S. DEFT. OF AGRICULTURE

plateaus (see fig. 16, p. 27). Except for the conspicuous balls of pappus on mature heads, it is

relatively inconspicuous. This is the only species of Vernonia in this section that is strikingly

distinct, although some specimens from the southern portion of its range closely related to

V. glabra have been mistakenly identified as F. abyssinica.

Selected Collections: SOMALI REPUBLIC: Ourano. E, Chedville 831, 561, June 6, 1953. (FI). Allurioni

del Baidoa. G, Paoli 1257. Oct. 29,1913. (FI). ETHIOPIA : Eritrea. Asmara. Mt. Lesa. A. Pappi J,905,

Apr. 25, 1902. (AAH, BM, FI, G). Ansaba valley, 15 mi. N. of Keren. 4,000 feet. P. R. 0. Bally

86640, Mar. 21, 1949. (EA). Sarae. Gaza Gebo. 1,800-2,000 m. A. Pappi 209, Oct. 7-10, 1902. (FI).

Mensa. Aba Maitan, Dada. A. Terracciano & A, Pappi 1962, Jan, 8, 1893. (FI). Along Massaua road

to Nefasit. 6,000 feet. C, E, Smith, Jr. & Ato Andemeskal 4536, Dec. 18,1966. (BM, EA, FI, K, US).

About 1/2 way between Asmara and Adi Ugri at Scitchity. 7,700 feet. C, E, Smith, Jr. & Ato

Andemeskal 4531, Dec. 17, 1966. (BM, EA, FI, K, US). Sidamo. Neghelli. H. F. Mooney 7352, July

11,1958. (EA). KENYA: Marsabit, road to Lake Paradite. P. R, O. Bally 35477, (EA). TANZANIA:

Near Manyoni N. of Kilimatinde. T. B, Kiwnaga 336. Feb., 1929. (BM, GH).

7. Vernonia stenolepis Oliv. Trans. Linn. Soc. London Bot. 2:337. 1887. (Fig. 21. See also fig. 16.)

Type Coll : Johnston s. n, Kihmanjaro. 5,000 feet. (K !).

Annual herb to 2 m. tall, stems single or openly branched. Stems with scattered hairs or, rarely,

pubescent; leaves ovate to ovate-lanceolate, thin, to 18 cm. long, 5.0 cm. v^ide; margin scarcely

apiculate to sharply serrate; blade few pilose, sometimes glandular above, few pilose, particularly

along the veins, glandular below; apex acute, base cunéate; petiole generally none, sometimes to

0.3 cm. long. Heads few to several in an open panicle, to 1.7 cm. high, 2.0 cm. in diameter ; phyllaries

to 1.6 cm. long, outer often long, spatulate, sometimes scattered pilose or pubescent, green to pink

or reddish; flowers to 1.2 cm. long, tube often 3 times longer than limb, lobes long, lanceolate, thin,

white to purple; achenes to 0.6 cm. long, dark brown to black, surface often pappilose, few to

moderately pilose, glandular, callus small, discreet, yellow to brown or black; pappus to 1.0 cm.

long, filiform or narrowly flattened, acute.

V, stenolepis is apparently found in damp, shady habitats in local colonies. Under poor conditions,

the plants reach maturity when only a few centimeters tall. The long spatulate outer phyllaries

clearly distinguish this species from other African species in the section. They are very similar

to the phyllaries on some of the collections of Vernonia anthelmintica from Asia. This species cannot

be confused with the strikingly difi'erent V, nigritana which has deeply colored phyllaries= '

"Occurs in Baikiaea dry deciduous forest in large colonies to 8' high and %/' diameter. Flowering

February, March, April. Fruiting March, April, May. Also on fringes of evergreen thicket as you

saw at Chisamba. Apparently always associated with forest or thicket." D. B. Fanshawe Zambian

notes.

F. stenolepis is confined to eastern Africa in the equatorial zone but it grows westward in

Rhodesia and the westernmost collection is from South West Africa (see fig. 16, p. 27). Colonies

of this species probably only appear in years in which the weather conditions are optimal for

germination of the achenes.

Selected Collections: KENYA: Isiolo to Mathews Range and Mt. Nyiru. 01 doinyo Lengiyo. 5,000

feet. J, G. Neivbould 3515, Dec. 20, 1958. (K). Ukamba-Garissa road at the turning to Sosoma.

2,000 feet. P. R. 0. Bally 9480. Jan. 24, 1954. (EA). Macjakos Distr. Nairobi-Mombasa road.

/. Ossent 536. Dec. 26, 1960. (EA, K). Yatta Plateau, Ndalani Ranch. Hale s, n. Apr. 11, 1954.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 35

|niiiiiii|iiiiiiiimni

3 4 5

PN-2041

Figure 21.-—Type collection of V. stenolepis Oliv. A, Flower, dissected and unopened (expanded in water)

B, Achene (immature) ; C, habit; and D, detail of heads. (K). Scales in millimeters.
36 AGRICULTURE HANDBOOK NO. 396, U.S. DEFT. OF AGRICULTURE

(EA). UGANDA: Lodo Keteminit, Moroto, Karamoja. 4,200 feet. 0. Kerfoot H3. Apr 26 1959

(EA). Loyoru, Karamoja. 4,000 feet. A. S. Thomas 3771. June 9, 1940. (EA, K) TANZANIAKilimanjaro, Pangani. G. Volkens m. July, 1893. (BM, G). Kondoa Distr. Sambala Hills. 5,000

feet. B. D. Burtt 2061. May 21, 1929. (EA, K). Iringa Prov. Iringa. H. E. Emson 564. (EA)

Kongwa. 4,500 feet. B. Anderson U27. Apr. 3, 1949. (EA). ZAMBIA : Abercorn. H. M. Richards 1611.

Apr. 9, 1952. (SRGH). Másese. D. B. Fanshawe 5460. Mar. 11, 1960. (SRGH). Eastern Prov River

Nyamadzi. A. W. Exell, F. A. Mendonça & H. Wild 1172. Mar. 25, 1955. (BM, SRGH). Barotseland

Distr. Shangombo, near Mashi River. 3,400 feet. L. E. Codd 7425. Aug. 7, 1952. (BM K PRE

SRGH). RHODESIA: Hartley. Poole. 4,000 feet. H. Wild 1004. Apr. 3, 1946. (K, SRGH) Urungwe'

Zambesi Valley, Rifa River. H. Wild 4059. Feb. 25, 1953. (K, MO, SRGH). Victoria Falls C E F

Allen 131. Nov., 1906. (SRGH). REPUBLIC OF CONGO: Luapula. 970 m. W. Robyns 1934. Apr 9

1926v (BR, GH, K). SOUTH AFRICA: Southwest Africa. 6 mi. W. of Ohopoho! B. de Winter &

Leistner 5215. Mar. 27, 1957. (K).

8. Vernonia anthelmintica (L.) Willd. Sp. PI. 3:1634. 1803.

Conyza anthelmintica L. Sp. PI. (ed. 2) 1207. 1763.

Branched herbaceous annuals. Stems glabrate to moderately pubescent; leaves sessile, usually

ovate, to 13.0 cm. long. 5.0 cm. wide; margin dentate ; blade few pilose above, glandular, veins pilose

below; apex acute, base cunéate, acute. Heads borne in a loose paniculate arrangement on the top of

the plants closely subtended by leaves, heads to 1.5 cm. high, 1.5 cm. in diameter; phyllaries to 1 2

cm. long, sometimes scarcely pubescent, green, inner sometimes with reddish margin lanceolateflowers to 0.9 cm. long, tube 2 or more times longer than the limb, mauve or lavender ; ächenes to

0.5 cm. long, dark brown or black, moderately pubescent, callus small, yellow, discreet, pappus to

0.9 cm. long, filiform.

V. anthelmintica is obviously very similar, morphologically to V. stenolepis. Inasmuch as all other

species of the section are African, I suspect that V. anthelmintica originated in Africa and was

transferred to Asia as a semicultivated plant for medicinal use. It may well have been derived from

V. stenolepis.

I have checked other Indian Vernonia species and find that V. gardneri, V. ornata, V. pulneyensis

V. setigera and V. thwaitesii resemble V. anthelmintica only superficially. They cannot be considered

synonymous. This strengthens the supposition that V. anthelmintica is, indeed, a foreigner in India.

I attempted to check the early synonymy of this species. Ascaricida indica Cass. Diet Sei Nat 3

(suppl.) : 38 apparently is well placed here. The specific name was apparently taken from the

Vaillant reference to Conyza indica. Linnaeus cites "Vaill. act. 358. Conyza indica." This must be

an erroneous page reference. A search at Houghton Library, Harvard University, disclosed a series

of papers by Vaillant, Sebestien. Établissement de noveaux caractères de trois familles ou classes

de plantes, etc., torn from Mem. Acad. Roy. Sei. Paris, consisting of the pages - 143-191 1718

277-318. 1719 and 277-339, 1720. I found the only reference to Comjza indica on page 300 of the

section published in 1719. To correct the record,the citation would appear to be Mem. Acad. Roy.

Sci. 1719 :300. This, of course, is largely academic with our Linnean starting date for nomenclature,

but it may help someone who attempts to follow the same course.

V. anthelmintica is widespread in India from the lower slopes of the Himalayas to the southern

end and Ceylon and from Pakistan in the west to Burma in the east.

Selected Collections: PAKISTAN: Azad Kashmir. Nawal Nadi. 3,500 feet. A. Rashid E Nasir &

R. R. Stewart 1064. Sept. 11, 1953. (RAW). Punjab. Rawalpindi Distr. Saidpur 1900 feet

R. R. Stewart 13670. Sept. 1951. (RAW). NEPAL: lia, Bheri River. 6,000 feet. O. Pohmin W R
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 37

Sykes & L. H. J. Williams 3250. Sept. 7, 1952. (BM). INDIA: Punjab Biari, Kulu, Kangra. 3,500

feet. W. Koelz 7A69. Nov. 15, 1933. (NY). Azad Kashmir. Poonch, Kotli. A. Rashid 26987. Sept.

18, 1954. (BM). Punjab. Pampur, Kulu. 4,500 feet. W. Koelz 3150. Nov. 10, 1931. (NY). Dehra

Dun. 2,200 feet. U. Singh U32. Sept. 21, 1928. (NY). Bihar. Chota Nagpur, Sirguja. 1,500-2,000

feet. /. /. Wood 277. Dec. 2, 1890. (NH). Karwar, Lejvi^ad. L. /. Sedgtvick & T. R. D. Bell 672i.

Oct., 1919. (K). Rajauri Prov. Puch via Kotli to Islamabad. 2,000-4,000 feet. Schlagintweit 12557.

Nov. 10-15, 1856. (BM). Chittagong Hill Tracts. Fenoa Hill, 29 mi. from Chittagong. B. Khan i5.

Oct., 1886. (CAL). Ganpur. Orissa, Sundargarh. 800 feet. H. F. Mooney 1593. Oct. 29, 1940. (K).

Bombay Presid. Near Padkhumb. A. P. Young s. n. Jan., 1879. (BM). Kangra, Dharmsala. 5,000

feet. G. A. Gammie 18760. Sept. 27, 1896. (K). Madras. Marmigudi. 3,000 feet /. S. Gamble 18^70.

Nov., 1886. (BM). Madras. Coimbatore. Anonymous. (NY). Madura Distr. Periyakolam riverside.

A. G. Bourne s. n. Jan. 2, 1899. (K). CEYLON: Kandy. A. Moon 102. Feb. 4, 1819. (BM). BURMA:

S. S. S. Thamakhan. A. Khalil s. n. 1896. (BR).

9. Vernonia petitiana A. Rich. Tent. Fl. Abyss. 1:373. 1847. (figs. 22 and 23.)

Type Coll: Petit s. n. Yedjou. (Apparently region around Amba Samber—see Salt map

of 1814.) (P!).

V. filigera Oliv. & Hiern in Oliver. Fl. Trop. Africa 3:288. 1877.

Type Coll: Schimper 1530 (?) (K). The Kew specimen marked as type is obviously

authentic material of this species (see notes on sheet), but it does not bear a

label listing either of the localities cited.

Figure 22.—Type collection of V. petitiana A. Rich. A, Habit; and B, detail of head (P).
38 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

• 9. V. petitiana

• 10. V. lasiopus

All. V. oxyura

0 200 400 600 800

J \ \ \ I I L

PN-2043 PNFigure 23.—Distribution of V. petitana, V. lasiopus, and V. oxyura.

Shrub (perhaps more nearly herbaceous than woody?) to 1.5 m. tall. Leaves to 19.5 cm. long,

7.0 cm. wide, ovate to ovate-lanceolate ; margin irregularly apiculate dentate ; blade glabrous,

scattered glandular or scattered pilose above, generally floccose or arachnose, densely glandular

beneath; apex acute, base usually decurrent-cuneate, rarely acute; petiole 0.4 to 2.5 cm. long,

puberulent, pubescent or floccose. Heads terminal, usually crowded, corymbosely (cymosely?)

arranged, to 1.5 cm. high, 2.0 cm. in diameter; phyllaries to 1.5 cm. long, herbaceous tips often

attenuate, spreading or reñexed, sometimes merely lanceolate, outer phyllaries pubescent to often

arachnosely pubescent; flowers to 1.2 cm. long, generally tube twice as long as urceolate top, often

glandular at top of tube ; achenes to 0.4 cm. long, usually medium brown, glabrous, highly glandular,

rarely dark brown, moderately pilose and densely glandular ; pappus to 0.8 cm. long, flattened.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 39

y. petitiana seems to be restricted to Ethopia (see fig. 23). Richards' description and the

type specimen (Mus. Paris) belong with several collections with less crowded heads and somewhat

pilose achenes which look suspiciously like introgression from 7. ahyssinica, I have no proof for

this. The material formerly identified as 7. filigera forms the bulk of this species. While the more

extreme of these might be worthy of distinction in some way, the presently known range of variation and the lack of geographical separation argues against this at the present time.

Selected Collections: ETHIOPIA: Eritrea. Asmara. Filfil. 700 m. A. Fiori 1735, Mar. 31, 1909. (FI).

Carnescin, Valle Gerbabit. 1,000-1,300 m. A. Pappi 137, Apr. 6, 1924. (FI). Amhara. Dembia,

Gondar. G. Chiovenda 2169, Oct. 1, 1909. (FI). Lago Tana, presso il villaggio de Bachianà (Tueur

Dinghia-Gondar). R. Pichi-Sermolli 2U7. Jan. 24, 1937. (FI). Amhara. Dembia, valle Sciuta sopra

Asoso. G, Chiovenda 2563, Oct. 17, 1909. (FI). Scioa. Addis Ababa. H, F, Mooney 6337. Nov. 16,

1954. (FI, K). Neghelli. P, G, Piovano 131, Oct. 27, 1937. (FI).

10. Vernonia lasiopns 0. Hoffm. in Engl. Planzenw. Ost-Afrika. 403. 1895. (Figs. 24 and 25. See

also figs. 7B, and 23.)

Lectotype Coll : Volkens UUh^ Marangu. 1,550 m. June, 1893. (BR, K).

F. iodocalyx 0, Hoffm. Loc. cit.

Lectotype Coll: Hoist 1^327, Usambara. (K).

V, massaiensis S. Moore. J. Linn. Soc. Bot. 35:320. 1902.

Type Coll : Elliot 6780, Masai. 7,000 feet. (Photo—BM !).

V. kaessneri S. Moore. J. Bot. 40:340. 1902.

Type Coll : Kaessner 635, Brit E. Africa, Simba River. April, 1902. (K!).

y. ruwenzoriensis S. Moore. J. Linn. Soc. Bot. 35:321.1905.

Type Coll : Elliot 7673, Ruwenzori, Kivate. 6-8,000 feet. May. (BM, K !).

V, braunii Mnschler, Bot. Jahrb. Syst. 46:73. 1911.

Type Coll : Braun 1979, Tanganyika Terr., Amani. Aug., 1908. (EA, K).

V, albo-violacea De Wild. Feddes Report. 13:205. 1911. (non Muschler). Norn, illeg,

(later homonym).

Lectotype Coll : Bequaert Í92, Shinsenda. June, 1912. (BR !).

V, brownii S. Moore. J. Bot. 54:252. 1916.

Type Coll : Brown 2656, Uganda. 100 miles N. W. of Kampala on the Umbendi Road.

June, 1915. (K!).

V, dumicola S. Moore. J. Bot. 54:252. 1916.

Type Coll: Dümmer 8i, Uganda: Karirema. 4,000 feet. Aug., 1913. (Photo BM!).

y. ringoetti De Wild. Contrib. FI. Katanga. 228. 1921. (Nom. nov, for V, albo-violacea

De Wild.).

V, mokaensis Milbr. Wiss. Ergeb. Deutsch Zentral. Afrika-Exped. 1910-11. 2:195. 1922.

Lectotype CoW: Milbraed 7077. Fernando Poo. 1,200-1,800 m. (Photo HBG!).

V, saltiiarii S. Moore. J. Linn. Soc. Bot. 47:264. 1925-27.

Type Coll: Sivijnnerton 827, Tanganyika Terr. Madolema. June, 1921. (BM!).

V, tuberculata Hutch. & Burtt. Rev. Zool. Bot. Africaines. 23:39. 1932.

Type Coll: de Witte i63, (Not seen) ; 500, Congo. Kasiki. June, 1931. (K) (Syntype).

Shrubs to 2.4 m. tall, rarely to 4.0 m. tall. Stems seldom glabrate to generally densely puberulent

or pubescent; leaves to 25 cm. long, 9.5 cm. wide, usually about 10 cm. by 2.5 cm., ovate to ovatelanceolate to lanceolate; margin usually apiculate dentate; blade pilose along the veins to generally

moderately pilose, rarely with scattered glands above, pilose along the veins to scattered pilose to

often densely lanate, glandular beneath; apex acute, base sometimes decurrent and sessile, sometimes cunéate, often acute; petiole, when present, to 3.0 cm. long, usually 0.5 to 1.5 cm. long,
40 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

PN-2044

Figure 24.—Lectotype collection, V. iodocalyx 0. Hoffm. = V. lasipous 0. Hoffm. A, Habit; B, flowers, unopened and

dissected (expanded in water) ; C, detail of heads; and D, achene (immature). (K). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 41

f A^-

-".^'^ :

^^¿^p

PN-2045

Figure 25.—A, Plants of V. lasiopus, scattered through the grass and shrubs in open pasture near Fort Portal,

Uganda; B, flowering heads of a plant at Fort Fortal (Smith, Wood & Perdue 4612); C, mature heads, and

D, flowering heads of plants near the Tombs of Karambi, south of Fort Portal (Smith 4624).

densely pubescent. Heads terminal, generally crowded, corymbosely or paniculately arranged, sometimes so crowded as to form a ball; heads to 1.5 cm. high, to 1.5 cm. in diameter; phyllaries to

1.4 cm. long, coriaceous base usually exposed, herbaceous tip frequently small, ovate to lanceolate,

occasionally larger and conspicuous, sometimes moderately puberulent without, white to rose, rose

color sometimes persistent in dried specimens; tips sometimes reflexed; flowers to 1.4 cm. long,

tube from V2 to % the length of the flower, white to mauve or purple; achenes to 0.4 cm. long

(one collection to 0.7 cm. long), often plump, medium to dark brown, (costae occasionally lighter),

rarely glabrous to usually scattered to moderately pilose, often glandular, callus discreet ; pappus

to 0.9 cm. long, generally only slightly variable in length, narrowly flattened.

V. lasiopus appears to be generally found in wet or well-watered habitats across the equatorial

portion of Africa and into Zambia and Mozambique on the east. The foliage is widely variable and

often undulate on the plants which I have seen in the field.

The specimens which have well-developed phyllary tips are often found at higher elevations.

These can be readily confused with specimens of V. bracteosa with small heads, but the latter species
42 AGRICULTURE HANDBOOK NO. 396, U.S. DEFT. OF AGRICULTURE

is consistently larger in head size, flower length, achene length and pappus length. The specimens

of V, lasiopus are usually readily recognized whether the arrangement of the heads be more open

or densely crowded into a ball

Confusion between F. lasiopus and V. hymenolepis is more difficult to define as the heads of the

species may be the same size and have similar large, white phyllary tips. Generally, though, the

coriaceous phyllary bases are exposed, achenes are sparsely to moderately pubescent and glandular

and the pappus is more persistent in F. lasiopus, I have found F. hymenolepis more frequently in

disturbed habitats, but they both are found in similar undisturbed habitats. Because they also

occupy similar ranges and may be found in close proximity, I suspect that there may be some gene

exchange between them which would account for the many intermediate specimens which are

difficult to assign to a species.

In Zambia, D. B. Fanshawe notes, "Shrub with erect-spreading branches and loose corymbose

crown, to 2'' diameter. Bark grey, smooth with coarse brown lenticels, slash pale green, thick, soft,

sappy. Flowering recorded for September, October and fruiting for October and November, but

could be much more extensive (I have only seen the plant twice). Locally frequent on dambo

margins and stream sides, occasional in montane forest. Apparently confined to the eastern district on both sides of the Luangwa Valley from Abercorn southwards."

F. lasiopus is reported to be used as a remedy for a number of ills. In Kenya, the leaves are used

in making soap, the branches are used to strengthen hut walls, the juice of the leaves is used as

a medicine for goats, and the ashes of the plant are used as medicine (Napier). In Tanzania, a tonic

made from the plant is reported to be used in pregnancies (Tanner) ; the leaves and roots are used

to treat stomach pains (Braun and Carmichael) or as a purgative (Koritschorner). A decoction

of the whole plant is given for epilepsy, the root is given to aid birth (Bally), and the plant is even

used as medicine for cows (de Wulf). In both the Congo and Ruanda-Urundi, extract from the

leaves is used as an anthelmintic remedy (Becquet and Gille), and in Ruanda-Urundi the stem bark

is used to make sores suppurate (Elkens).

F. lasiopus has been generally collected in the uplands of eastern Africa from Ethiopia to

Rhodesia and Mozambique and westward in the Republic of the Congo and Angola (see fig. 23, p. 38).

Selected Collections: ETHIOPIA: Mt. Tulu Wallel. 8,000 feet. F. Piffard 57, Mar. 26, 1958. (K).

SUDAN: Lado, Yei River. F. Sillitoe 196. Nov. 10, 1919. (K). KENYA: Endebess. 6,400 feet. M. 7.

Webster 8899, (K), Cherangani. 6,400 feet. M. V, Webster 8898. (EA). Rift Valley near Eldama

Ravine. 700 feet. P. H. Irwin 288. June 30, 1956. (EA). Nieri. R. E. & C. E. Fries 150, Dec. 19,1921.

(BR, MO). Mau Forest. 8,000 feet. Q. C. Ivens 1003. July, 1957. (EA). Nairobi. Thika Road House.

6,500 feet. B. Verdcourt Ui, Jan. 14, 1951. (EA, MO, PRE). Nairobi. 5,450 feet. R, A. Dümmer

2030, Feb. 1915. (BM). Chyulu Hills center. 5,500 feet. P. R. 0. Bally 1030. June 11, 1938. (EA).

TANZANIA: West Usambaras. Lushoto to Megamba. 2,000-2,300 m. June 3, 1926. A. Peter V251.

(EA, K). Arusha Distr. Embagai. G. W. St. Clair-Thompson 1235. Feb. 5, 1932. (EA, K). West

Lake Prov. Muganza/Bushubi, Ngara. 4,500 feet. R. E. S. Tanner 4555. Dec. 1, 1959. (K). Kihmanjaro Distr. Lyamungo. P. /, Greenway 3019. Aug. 13, 1932. (EA). Between Bahati and Galappo in

Endanok Ravine. 5,000 feet. J. R. Welch 109. Aug. 26, 1951. (EA). Eastern Usambaras. Mt.

Bambole, Amani. 3,200 feet. B. Verdcourt 297. July 28, 1950. (BR, EA, MO, PRE). Western Prov.

Kigoma Distr. Kasangazi. Mahali Mts. C. D. Mgaza 15Í. July 16,1958. (K). Mahenge Distr. Issongo.

900 m. H. J. Schlieben2181, May 5, 1932. (K, PRE). Songea Distr. Matengo Hills, Miyau. 1,620 m.

E. Milne-Redhead & P. Taijlor 10252. May 19, 1956. (EA, K, SRGH). UGANDA: Mubende Distr.

Road from Bukumi to Madudu. 4,200 feet. Lind 211,2. (EA). Mt. Elgon, Butandiga. 7,000 feet.

R. A. Dümmer 3677. (K). Entebbe region. 3,842 feet. T. D, Maitland 1,56. (K), Western Prov.

Mubuku valley ridges. 4,500-5,000 feet. C. E. Smith, Jr. 1,608. Jan. 16, 1967. (BM, EA, FI, K, US).

Along Kasese-Mbarara road about 40 mi. from Mbarara. C. E. Smith, Jr. & D. Wood 4627. Jan. 20,*
OBSERVATIONS ON gTENGELIOID SPECIES OF VERNONIA 43

1967. (BM, EA, FI, K, US). Kachwekano Farm, Kigezi. 6,700 feet /. W, Purseglove 2776, Apr.,

1949. (EA, K). ZAMBIA : Abercorn Distr. Kwimbe. H. M. Richards 9619. May 8,1957. (K). Serenje.

D. ß. Fanshawe 6732. Sept. 27, 1961. (K). Malawi. Mt. Mlanje, Malosa Valley. E. /. Newman &

T. C. Whitmore 565. Aug. 24, 1956. (SRGH). RHODESIA: Inyanga. Trias Hill. F. Eyles 5199. Dec,

1919. (SRGH). MOZAMBIQUE: Niassa Prov. Maniamba Div. Malulo. Tessi Mt. Pedro e Pedrogao

Jp056. May 29, 1948. (EA). RUANDA: Ngoma. Lake Kivu, 20 km. S. of Kibuye. Ritland 316. (EA).

Biumba. Mutara, environs de Mimuli. 1,400 m. G. Troupin 5199. Dec. 14,1957. (K). Kigali. 1,500 m.

A. Becquet 379. Feb., 1933. (BR). Terr. Astrida. Entre Astrida et la Riv. Kamahembe, environ

Gisagara. 1,750 m. T. Bazarusanga ^0. Jan. 16, 1957. (K). BURUNDI: Kisozi. 1,800 m. J. B. H.

Lejeune 17i. 1934. (BR, MO). Malagarasi, Keifo Nello. Michel & Reed 95Jf. Dec. 12, 1951. (BR).

REPUBLIC OF CONGO: Bogoro. Bequaert 4952. July 9, 1914. (BR). Mayólo. Béni. Bequaert 3493.

Apr. 9, 1914. (BR). Lake Kivu. Entre Kiotshe et Bobandana. D. Van der Ben 201. Mar. 10, 1953.

(BR). Remera Mission, 30 km. from Kigali road to Usumbura. R. Germain 1195. Jan. 11, 1942.

(BR). Lualu. P. Quarre 2762. Nov., 1931. (BR, K). Lukonzolwa-Moero. P. Quarre 3309. July, 1933.

(BR). Kando. G. F. de Witte 112. Apr. 4, 1931. (BR, K). Kanda-Kanda. Luxen 336. 1934. (BR, K).

Ivumu, Mulungu. F. L. Hendrickx 3423. Oct., 1945. (BR, EA). Mutombo-Mukulu. P. Quarre 2520.

Nov., 1931. (BR, K, MO). ANGOLA: Malange region, Kela, River Lui. Herb. Serv. Agri. Ang. 9577.

Dec. 27, 1930. (K).

11. Vernonia oxyura 0. Hoffm. in. Engl. Pflanzenw. Ost-Afrika. C: 403. 1895. (Fig. 26. See also

fig. 23.)

Lectotype Coll : Buchanan 41- Nyassaland. 1891. (BM !).

F. polyura 0. Hoffm. Bot. Jahrb. Syst. 30:422.1901.

Type Coll : Goetze 866. Langenburg. Elev. 500 m. (BR).

7. cirrifera S. Moore. J. Linn. Soc. Bot. 35:320. 1902.

Type Coll : Buchanan 370. Nyassaland. (Photo—BM !).

V. longipetiolata Muschler. Bot. Jahrb. Syst. 46:74. 1911.

Type Coll : Kassner 2746. Congo, Kundelungu. Apr., 1908. (K).

V. seyrigii Humbert. Notul. Syst. (Paris). 13:324. 1949.

Lectotype Coll : Decary 8577. Madagascar : Ambovombe. Mar., 1931. (K).

Tree to 10 m. tall. Branchlets moderately to densely puberulent, rarely lenticellate ; leaves to

20 cm. long, 8.5 cm. v^ide, ovate; margin often undulate, rarely apiculate; blade scattered to

moderately pilose above, scattered to moderately pilose and glandular belov^ ; apex obtuse to acute,

base acute; petioles to 4.0 cm., generally less, densely puberulent. Heads in terminal, open to

generally crowded, corymbose aggregations; heads to 1.5 cm. high, 1.5 cm. in diameter, phyllaries

to 1.3 cm. long, lanceolate acute to abruptly attenuate, outer generally pilose without and frequently

ciliate; ñowers to 1.2 cm. long, tube about equal to the limb or slightly longer, white, pink, mauve

or purple ; achenes to 0.45 cm. long, medium brown or, rarely, yellow, glabrous, glandular ; callus

distinct ; pappus to 0.9 cm. long, narrowly flattened or nearly filiform, generally rufous-tawny.

V. oxyura is readily distinguished from most species of this group by its arboreal form. It is also

distinguished by the glabrous achenes which are rare in the section Stengelia.

Fanshawe notes that, in Zambia, this species is **Semi-deciduous shrub or small tree to T'

diameter. Bark pale brown, reticulately cracked, flaky, flakes 1.5 cm. diameter. Slash green, thin,

soft, sappy. Flowering April, May, June. Fruiting May, June. Rare on granite kopjes.''

In Tanzania, the root is used for influenza (Koritschoner) and a root decoction is given for

stomach ache and gonorrhea (Tanner). Malawi natives are also reported to use the leaves as

medicine and Robyns reports that in the Congo the species is used for wounds.

V. oxyura is largely confined to eastern Africa below the equator and it is rarely found westward
44 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

^

A

Figure 26.—Lectotype collection of V. oxyura O. Hoflfm. A, Dissected flower and B, unopened flower

(expanded in water) ; C, habit; and D, detail of heads. (BM). Scales in millimeters.

PN-2046

in the Congo (see fig. 23, p. 38). I did not borrow specimens of Vernonia from Madagascar, except

types whose descriptions seemed to fit Vernonia section Stengelia, and I cannot tell how common the

species is on the island. It would not appear to be abundant anywhere in its range.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 45

Selected Collections: KENYA: Lumi River Forest. Taveta. H, M. Gardner 2965, (K). TANZANIA:

Moshi Distr. Mashame. 1,500 m. M. Mücke 221. Aug. 28, 1909. (K). Usambara. Maschana. C. Hoist

3556. July, 1893. (BM, K). Tanga Distr. Pangani, Madanga, Pombwe. R, E. S, Tanner 2952, July 4,

1956. (BR). Eastern Prov. Morogoro Distr, Mtibwa Forest Reserve. S. R, Semsei 907. Aug., 1952.

(EA, K), Kyimbila Distr. N. of Lake Nyassa. A. Stolz K7. (BM, BR, K, MO, PRE). Tunduru Distr.

11 km. E. of Songea Distr. boundary. 840 m. E. Milne-Redhead & P. Taylor 10710, June 7, 1956.

(BR, EA, SRGH). Songea Distr. Kwamponjore Valley about 9.5 km. S.W. of Songea. 1,000 m.

E, Milne-Redhead & P. Taylor 9829, Apr. 26, 1956. (BR, EA, K, SRGH). ZAMBIA: Eastern Prov.

Fort Jameson Distr., Fort Jameson to Lundazi, mi. 30. F. White 2h76, Apr. 26, 1952. (BR, K, MO).

MALAWI: Langenburg. W, Goetze 86i. (K). North of Bua River, Nyansi Hills. /. B. P. Evans &

/. Erens 616, June 2, 1938. (PRE). Kasungu Hill. 1,100 m. L. /. Brass 17Í52, Aug. 28, 1946. (BR,

MO, NY, SRGH, US). Zomba Distr. 23 mi. N. of Zomba. L. C. Leach & Brinton 9888. May 2, 1960.

(SRGH). Mlanje Distr. Mlanje Mt. /. D, Chapman 355. Apr. 15, 1957. (K, PRE). Likubula Gorge.

840 m. L. /. Brass 1637Jf. June 20, 1946. (BM, BR, K, MO, NY, PRE, SRGH, US). MOZAMBIQUE:

Nyassa. Massaneulo. 1,100 m. A, Gomes e Sonsa 1397. Apr., 1955. (K). REPUBLIC OP CONGO: Kivu

Prov. Walikale Terr. Near Mutongo, 52 km. N.W. of Masisi, 1,100-1,200 m. G. Troiipin 3006. Jan.

22, 1957. (K). Between Shiebeie and Kitabi, road from Tweto to Baudouinville. 1,100 m. W. Robyns

2063, Apr. 21, 1926. (BR, G).

12. Vernonia guineensis Benth. in Hook. f. & Benth. Niger PL 427. 1849. (Figs. 27, 28, and 29.

See also fig. 31, ß.)

Type Coll: Don 70. Sierra Leone. (Photo—BM!).

F. firma Oliv. & Hiern in Oliv. Fl. Trop. Africa 3 :290. 1877.

Type Coll : Schweinfurth 3153. Linduku. May, 1870. (BM !, K).

y. ulophylla 0. Hoffm. Bol. Soc. Brot. ser. 1. 13 :13. 1896.

Lectotype Coll: Welwitsch 3279. Angola. (K).

F. cardiolepis O. Hoffm. Op. cit. 12.

Type Coll: Welwitsch 3280 (cited 2380). Angola. (K).

V. temnolepis O. Hoffm. Op. cit. 11.

Lectotype Coll: Welwitsch 3273. Angola: Huila. Inter Nene et Humpata. Apr., 1860.

(Photo—BM).

V, prolixa S. Moore. J. Linn. Soc. Bot. 35:322. 1905.

Type Coll: Elliot 8383, Urundi hills. Sept. (K!).

V, rotundisqnama S. Moore. J. Linn. Soc. Bot. 37:312. 1906.

Type Coll: Gossweiler 1228. Angola. Malange. (K!).

V, chevalieri 0. Hoffm. Bull. Soc. Bot. France 55 :Mem. 8b :39. 1908.

Type Coll: Chevalier 5505. Haut Oubangui. Entre lo Tomi et la Kémo. Sept., 1908.

(BR).

7. hierniana S. Moore. J. Bot. 52:335. 1914.

Type Coll : Welwitsch 3278. Angola : Pungo Andongo. (K !).

y. incompta S. Moore. J. Bot. 56:210. 1918.

Type Coll : Kassner 2261. Rhodesia. Katuba Stream. Dec, 1907. (K !).

V. gnineensis var. earneroonica Adams. J. W. African Sei. Assoc. 6 :154.1960.

Type Coll : Latilo & Daramola FHI3U67. Cameroons. Yashsaka Distr. Apr., 1955.

(BR).

Perennial herb from a woody root crown; stems to 1.5 m. tall, usually about 1 m. Leaves sessile,

to 12 cm. long, 4.5 cm. wide, lanceolate, rarely ovate-lanceolate; margin blunt apiculate-dentate;

blade generally glabrous, scattered glandular, rarely scurfy, above, raised reticulate, densely lanate
46 AGRICULTURE HANDBOOK NO. 396, U.S. DEFT. OF AGRICULTURE

\

^

. ^ ^ "^ »(» Â j

,>,s fliÉi

If ^y;:fy

mUPlE

ß

PN~2047

Figure 27.—Type collection of y. guineensis Benth. in Hook f. & Benth. A, Habit, and B, detail of heads.

(Photos, BM.)

and glandular beneath. Heads terminal, borne in an open paniculate arrangement ; heads to 1.5 cm.

tall, 2.0 cm. in diameter, phyllaries to 1.5 cm. long, tips white to rose; flowers to 1.5 cm. long, tube

twice as long as the limb or more than twice as long, white to mauve or lilac, rarely purple ; achenes

to 0.4 cm. long, usually less, medium brown to, rarely, dark brown, glandular, pubescent; pappus

to 1.0 cm. long, usually less, flattened, apex obtuse.

V. guineensis is a variable species which has been given numerous names on the basis of minor

morphological differences. When sorted by leaf shape, the narrow-leaved forms have been collected

primarily along the eastern side of the range of the species in eastern Congo, Mozambique, Malawi,

Zambia and Tanzania. Broad-leaved plants have been collected primarily in the Congo, but a few

come from Sierra Leone and Togo. The intermediate leaf forms are widely distributed throughout

the range of the species and overlap both extremes.

Fanshawe, in Zambia, furnishes the following note : "Subshrub, unbranched or with a few erectspreading branches. Bark grey-black, smooth, slash green, thin, soft, sappy. Flowering JanuaryApril. Fruiting February-May. Locally frequent in miombo woodland, occassional on granite kopjes.

Distributed throughout the high rainfall belt."
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 47

Although V. guineensis had once been reported as poisonous (Homblé), Congo natives used

a decoction of the boiled roots for hernia (Lecompte) and the leaves as a purgative (Devs^ulf).

In Sierra Leone, the plant is used as a wash for cravi^craw or paralysis (Elliot). In this area, the

raw roots are also eaten as an aphrodisiac (Deighton). An infusion of the plant is used to treat

"garli," a cattle disease, in Ghana (Williams).

V. guineensis ranges from western to eastern Africa in mid-continent from about 15° N. to 15° S.

at medium elevations. (See fig. 29, p. 48). It is a species of open deciduous woods and savannas.

Selected Collections: GUINEA: Diaguma. Maclaud 169. Jan. 12, 1898. (P). Conakry, Boffa.

G. Roberty 17506. Sept. 17, 1955. (G). Near Beyla. H. J. Félix 15Í6. Feb., 1937. (K, P). SIERRA

LEONE: Near Mussaia. F. C. Deighton 5388. Apr. 2, 1951. (K). Northern Prov. Mambolo. F. C.

Deighton 979. Jan. 5, 1928. (BM). Near Magbile (Maforki). ÍT. D. Gordon 893. May 26, 1953. (K).

Near Buyabuya, Scarcies River. G. F. S. Elliot U8k2. Feb. 13. (BM). Binkolo. 570 feet. N. W.

Thomas 1884. Aug. 25, 1914. (K). IVORY COAST: Baoulé. H. Pobequin 169. Oct., 1895. (P). 30 km.

N. of Bouaké. 300 m. A. J. M. Leeuwenberg 2073. Nov. 20, 1958. (K, PRE). Kokoumbo. G. Roberty

13996. Feb. 24,1951. (G). GHANA : Tamale. /. K. Morton 8778. Mar. 27, 1953. (K). Between Ejura

^ >-^-

PN-2048

Figure 28.—A, Plants of V. guineensis, nearly hidden in the grasses and herbs. B, Flowering heads, conspicuous

because of their white phyllary tips. (Smith & Fanshawe 4662.) (See also fig. 7, A, p. 9.)
48 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

• 12. V. guineensis

* 13. V. procera

A14. V. vallicola

0 200 400 600 800 MILES

I I I I I I I I I

~' I r400 800 1200 KILOMETERS

Figure 29.—Distribution of V. guineensis, V. procera, and V. vallicola.

and Kintambo. 500 feet. P. R. 0. Bally 175. Apr. 27, 1947. (EA). Afram Plains. W. Johnson 715.

Mar. 15, 1900. (K). TOGO: Kpando. /. M. Robertson 117. Apr. 24. (BM). Agome. R. Schlechter

1296A. Mar., 1900. (BM, BR, G). NIGERIA: Zaria Prov. Near Jengre, 37 mi. from Los. 3,000 feet.

D. E. Coombe 83. Dec. 28, 1954. (K). Naraguta. 4,000 feet. H. V. Lely 2U2. May 30, 1921. (K). Oyo

Prov. Oyo Distr. 9 mi. E. of Igboho. R. W. J. Keay FHI22508. Feb. 29, 1948. (K). CAMEROON:

Kongola, Mbussa. 800-900 m. J. Milbraed 9063. 1914. (BM, K). Bamenda. 5,000 feet. F. W. H.

Migeod 295. Jan. 14, 1928. (BM). Bamenda Div. Near Bafut-Ngemba Forest Reserve above Lake

Bambuluwe. 2,110 m. F. N. Hepper 2152. Feb. 23, 1958. (K). Sansane, Kongola. 750-800 m.

/. Milbraed 8973. Apr., 1914. (BM, K). GABON: Mouila. A. Walker s. n. Mar., 1938. (P). ANGOLA:

Cuanza Norte. Malange, Cainbaxe. 1,000 feet. /. Gossiveiler 8875. (K). Benguella. Forte P. Amelia,

Cubango. /. Gossweiler 19^0. (K). Kirima, near River Jombo-Luando. /. Gossweiler 9503. Feb. 22,

1931. (K). Alto Catumbela. Ganda. 3,500-4,500 feet. H. G. Faulkner 1^8. Feb., 1942. (K). Near

Humpata. 6,000 feet. H. H. W. Pearson 2096. May 6, 1909. (K). REPUBLIC OF CONGO: Tukpwo.

P. Gerard 1273. Apr. 17, 1954. (BR). Basape, Rivier Nanzaie, tributaire de Duma. 700 m.

R. Germain 70h- Jan., 1946. (BR, K). Km. 32, Yakoma-Banzyville road. R. Germain 8550. Apr. 3,
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 49

1955. (BR). Near Bambesa, Uele. Pittery 252. 1936. (BR). Region Api, near Likwangili. W. Robyns

1275. Dec. 30, 1925. (BR). Between Isoro and Mt. Nambunga. Uele. J. Louis 1726. Apr. 20, 1936.

(BM, BR, K). Yalinga, haut Oubangui. G. Le Testu 2U95. (P). Kivu Prov. Kalehe Terr. BukavuWalikae road. 2,320 m. J. Petit 70. Mar. 18, 1960. (EA). Baise Ruzizi. Paysannat Bulamata.

R. Germain 6565. Mar., 1950. (BR). Wombali. H. Vanderyst 2195. Sept., 1913. (BR). Vakpudu

(Doruma). P. A. M. de Graer 225. Mar., 1934. (BR). Pare Heenen, Lubumbashi. P. Quarre 79U.

Apr., 1946. (BR). Mayaba, entre Tshobo et Kai Baku. 350 m. V. Goossens U05. Aug., 1919 (BR)

Boitsfort, Elisabethville, 1,250 m. W. Robyns 1637. Mar. 11, 1926. (BR, K, MO). Kiayaka, Kwango.

R. Devred2860. Mar. 10, 1956. (BR, K). RUANDA: Mosso, Nyabitare. Michel & Reed 692. Nov. 21,

1951. (BR). TANZANIA: West Lake Prov. Keza, Bushubi, Ngara. 5,000 feet. R. E. S. Tanner 5069.

July 20, 1960. (K). Buha Distr. 86 mi. from Kibondo on road to Kasulu. 4,500 feet. B. Verdeourt

2850. July 15, 1960. (EA). Kigoma Distr. 36 mi. S. of Uvinsa. 5,700 feet. A. A. Bullock 8286. Aug.

31, 1956. (K). Ufipa Distr. Kito Mts. 1,800 m. H. M. Richards 11867. Dec. 3, 1959 (K) Mbeya

6,500 feet. R. M. Davies UU. Mar. 3, 1932. (EA, K, SRGH). ZAMBIA: Abercorn Distr. Nkali Dambo

C.E. Smith, Jr. & H. M. Richards i672. Feh. 21, 1967. (BM, EA, K, SRGH, US). Kasama Distr.

Ml. 46 on Kasama-Abercorn road. P. J. Greenway 8359. Aug. 3, 1949. (EA, K, SRGH) Ndola

D. B. Fanshawe 683. Jan. 18, 1954. (K). Western Prov. Luano Forest Reserve near Chingóla.

C. E. Smith, Jr. & D. B. Fanshawe 1,662. Feb. 12, 1967. (BM, EA, K, ND, SRGH, US). MALAWIMisuku Hills. 4,000 feet. E. A. Robinson 31U. Jan. 10,1959. (K, SRGH).

13. Vernonia procera 0. Hoífm. Bull. Soc. Bot. France 55: Mem. 8b:39. 1908. (Fig. 30 See also

fig. 29)

Lectotype Coll : Chevalier 7899. Chari. Ndelli. Apr., 1903. (K).

V. guineensis var. procera (O. Hoffm.) Adams. J. W. African Sei. Assoc. 6:154. 1960.

Perennial from root crown. Stems to 1.75 m. tall, glabrate to pubescent or lanate. Leaves sessile

to 14.5 cm. long, 5.0 cm. wide, lanceolate to ovate; margin apiculate dentate; blade glabrous to

pustulate (1 specimen) above, densely lanate, glandular below; apex obtuse to sub-acute, base acute.

Heads terminal, single to few on long peduncles; heads to 2.5 cm. high, 4.0 (?) cm. in diameter;

phyllaries to 2.2 cm. long; flowers to 2.2 cm. long, tube 2 to 3 times longer than the limb, white to

mauve; achenes to 0.5 cm. long, brown, short pubescent, sides nearly parallel, glandular; pappus

to 1.5 cm. long.

^ Although this species was recently reduced to a variety of V. guineensis, it is as morphologically

distinct as many of the other species of section Stengelia. Furthermore, it exhibits a coherent

geographical distribution within a small area rather than appearing sporadically throughout the

range of V. guineensis (see fig. 29, p. ). For these reasons, I prefer to maintain it. It is distinguished from V. guineensis by the consistently larger heads with flowers 2 or more cm. long, achenes

with nearly parallel sides and relatively few heads borne on long peduncles.

Collections Seen: NIGERIA: Bauchi. H. V. Lely 179. Mar., 1929. (K, MO). Bonn Kurmi, 18 mi. N.W.

of Abuja. E. W. Jones 157. June 10, 1958. (K). Oloke Meji. E. W. Foster 280. Aug. 7, 190?! (K).'

CAMEROON: Koutschya. H. J. Félix 3398. May, 1939. (P). Buar. 1,000 m. /. Milbraed 9376 Mav"

1914. (K).

^ A. Chevalier 7899 was selected as the lectotype because it is the only specimen of those originally

cited which was available to me. It is a complete specimen whose identity is undoubted.

14. Vernonia vallicola S. Moore. J. Bot. 52:95. 1914. (Fig. 31. See also ñg. 29).

Type Coll: Gossweiler 31,81. Angola: Benguela. Valley of Tiengo, Cuito. (K!).
50 AGRICULTURE HANDBOOK NO. 396, U.S. DEFT. OF AGRICULTURE

PN-20B0

Figure 30.—Lectotype collection of V. procera 0. Hoffm. A, Habit; B, dissected flower; C, achene; and

D, unopened flower (expanded in water). (K). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 51

i>»

. >

w-= *

>

%:'%é'

PN-2051

Figure 31.—Type collection of K vallicola S. Moore. A, Detail of heads; B, habit; and C, achene (K).

Scales in millimeters.
52 AGRICULTURE HANDBOOK NO, 396, U.S. DEPT. OF AGRICULTURE

Shrub to 1 m., densely branched. Stems light to gray lanate. Leaves sessile, closely set on stems,

to 11.0 cm. long, 4.0 cm. wide, ovoid; margin entire to apiculate dentate; blade scattered pilose,

slightly reticulate above, deeply reticulate, pilose, glandular beneath; apex obtuse, base acute.

Heads borne moderately densely in terminal corymbose arrangement ; heads much taller than wide,

strict, to 2.5 cm. tall, to 1.5 cm. in diameter; phyllaries to 2.0 cm. long, bract tips petaloid, white;

flowers to 2.1 cm. long, tube about twice as long as limb, light purple or mauve; achenes to 0.45 cm.

long, densely white or golden pilose ; pappus to 1.4 cm. long, flattened, obtuse.

V. valUcola is a distinctive species related to F. guineensis. It appears to be uncommon as I have

seen only the type collection and 2 others. Like V, guineensis, the roots of this species are fleshly

and spindle-shaped. At least one of the collections was made on deep sand. Restricted to Southwest

Africa and adjacent Angola (see fig. 29, p. 48).

Collections Seen: SOUTH APRICIA : South West Africa. Okavango Native Terr. 24.5 mi. N, of Tamso

on road to Kapupahedi Camp. S. de Winter & W, Marais ins, Feb. 17, 1956. (K, PRE, SRGH).

30 mi. N. of Gautcha Pan. K Story 6Jf31. Feb. 5,1958. (EA, K, SRGH).

15. Vernonia benguelensis Hiern. Cat. Welw. African PI. 1:536. 1898. (Figs. 32 and 35.)

Lectotype Coll : Welwitsch 3276h, Angola. Ad lacum de Ivantola. Feb., 1860. (Photo—

BM!).

V, limosa O. Hofïm. Warb. Kunene-Sambesi Exped. 400. 1903.

Lectotype CoU: Baum 624. Südwest Afrika. Am Longa unterh, Chijija. Jan., 1900.

(Photo—BM),

Perennial from a root crown. Stems to 0.75 m. tall, scattered pilose or puberulent, glandular,

densely leafy; leaves sessile, to 7.0 cm. long, 2.0 cm. wide, lanceolate; margin apiculate dentate;

blade few pilose and glandular above and below, secondary veins variably wide (appearing to

spread at intervals) ; apex and base acute. Heads single to several, moderately crowded, cymosely

arranged; heads to 1.5 cm. high, 1,5 cm. in diameter; phyllaries to 1.2 cm. long, tips red-purple

(fide Faulkner) ; flowers to 1.7 cm. long, tube twice as long as the limb, purple; achenes immature

in all specimens seen, light brown, moderately pubescent, glandular; pappus to 1,0 cm. long, flat,

obtuse.

Although I have only photographs of the types, V. bengtielensis and V. limosa certainly are the

same taxon. I am not so sure that one of the cited specimens belongs here, but it differ,s only in

bearing a single head which is insufficient grounds on which to separate a species in the group.

One of the specimens has roots with tuberous swellings: In general, the sum of the characteristics

indicate that the species is related to V. guineensis.

Collections Seen: ANGOLA: Alto Catumbela. Ganda, if. G. Faulkner A20. Dec, 1941. (K). H. G,

Faulkner Ai02, Nov., 1940. (K). Dondi. L. S. Tucker 49, Oct. 2,1924. (GH).

16. Vernonia filipéndula Hiern. Cat. Welw. African PI 1:536. 1898. (Figs. 33, 34, and 35. See also

fig. 8.)

Type Coll : Wehvitsch 3275, Angola. Catumba. Apr., 1860. (Photo—BM !).

V, lancibracteata S. Moore. J. Bot. 46:293. 1908.

Type Coll : Eyles 291: Rhodesia. Mazoe. Bernheim Hill and Iron Mask. Mar., 1906.

(SRGH).

F. longepedunculata De Wild. Feddes Repert. 13 :207. 1914.

Type Coll : Homble 881. Congo. Katanga. Plateau de Biano. Nov., 1912. (BR !),
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 53

PN-2052

Figure 32.—Lectotype collection of V. benguelensis Hiern. A, Habit; and B, detail of heads. (Photos, BM.)

V. descampsii De Wild. Bull Jard. Bot. État. 5:97. 1915.

(E. descr., type collection not seen).

V. manikensis De Wild. Op. cit. 99.

{E. descr., type collection not seen).

V. pleiotaxoides Hutch. & Burtt. Rev. Zool. Bot. Africaines. 23:38. 1952.

Type Coll : Quarre 265i. Congo. Katanga. Luniemu. Aug., 1931. (PRE).

V. wittei Hutch & Burtt. Rev. Zool. Bot. Africaines. 23 :39. 1932.

Type Coll: de Witte 5^3. Congo. Kangenge (sic. Kanzenze). Aug., 1931. (K).

Perennial from a root crown, roots tuberous. Stems to 1 m. tall, generally less, usually pubescent

to scarcely puberulent, leaves moderately crowded, generally sessile, rarely with petiole to 0.5 cm.

long, oblanceolate to lanceolate, to 15 cm. long and 9 cm. wide, but more often to 12 cm. long and

4.0 cm. wide or less ; margin apiculate dentate, sometimes irregularly so ; blade glabrous to pilose

to scabrous above, scattered pilose to densely pubescent, glandular below; apex obtuse or rounded

to subacute, base usually cunéate acute. Heads borne singly or at most 3 or 4 together, sometimes

long pedunculate, sometimes short pedunculate and somewhat crowded; heads to 2.5 cm. high,

3.5 cm. in diameter; phyllaries to 2.2 cm. long, herbaceous tip often broad, sometimes pubescent
¡M AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

„il 1;_->J_

):ii:;:^:V:Avï.:d:.^o.

/

';,'-..;;!y

Figure 33.—Type collection of V. filipéndula Hiern. A, Habit; and B, detail of heads. (Photos, BM.)

without, rose or green; flowers to 2.2 cm. long, tube sometimes three times longer than the limb,

white to lilac or purple (blue fide Bullock) ; achenes to 0.5 cm. long, stout, sides nearly parallel,

usually medium brown, short pubescent to pubescent, often glandular, callus discrete, very small;

pappus to 1.4 cm. long, flat, tips obtuse.

V. filipéndula has received several names on the basis of length of peduncles or the size and shape

of the leaves, none of which appear to be particularly reliable characters. It appears to be related

to V. guineensis but is shorter, lacks lanate pubescence on the leaves beneath, and has larger heads

and flowers.

V. filipéndula is restricted to medium and lower elevations across Africa below the equator and

above 22° S. latitude (see fig. 35, p. 56).

Selected Collections: REPUBLIC OF CONGO: Leopoldville Prov. Bayaka Sud Terr. Sanzi-Mawenga

Kwango. R. Devred 1670. Mar. 18, 1955. (K). Kipaila. T. Kassner 25Jf8. Mar. 2, 1908. (K) Katanga.

Kamina. J. Becquaert 85. Aug. 2, 1934. (GH, US). Kamina, Haut Lomani. W. Mullenders 1785.

Dec. 18, 1947. (BR). Katanga. Lunumba. R. Desenfans 2209. (BR). Kando. G. F. de Witte 89.

Apr. 4, 1931. (K). ANGOLA: Dist. Cuanza-Sul. Seles Cucul. 1,000 m. /. Gossweiler 9353. Oct., 1930.

(K). Libólo. M. T. Dawe 317. Jan., 1922. (K). 130 mi. E. of Ching, W. of Cuanza River. Chinguar.

A. G. Curtis 37U. Sept. 25, 1923. (GH). ZAMBIA: Abercorn Distr. Abercorn. A. A. Bullock 1018.

Sept. 20, 1949. (K). Abercorn Distr. Mpulungu, Abercorn road near Kasama turning, 5,000 feet.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 55

H. M. Richards 2011. Oct. 11, 1954. (SRGH). Kawambwa. D. B. Fanshawe 3519. Aug. 22, 1957.

(K). Mwinilungu Distr. 16 mi. W. of River Katompo. E. Milne-Redhead 1102. Sept. 11, 1930. (K).

Luano Forest Reserve near Chingóla. C. E. Smith, Jr. & D. B. Fanshawe Jt66i. Feb. 12, 1967.

(SRGH, US). Kitwe. D. B. Fanshawe 10017. Apr. 10, 1967. (ND, US, S). 6 mi. N.W. of Ndola.

C. /. Doloway 8W92. Mar. 27, 1958. (SRGH). RHODESIA: Urungwe Distr. Zuoipani. R. Goodier &

T. B. Phipps 260. Mar. 5, 1968. (SRGH). Marandellas Distr. Shortlands Farm. /. M. Rattray 603.

Mar. 1, 1933. (SRGH). Salisbury. H. G. Flanagan SOU. May 1, 1906. (PRE). Miami. H. Wild 182A.

Mar. 4, 1947. (SRGH). MALAWI: Dzalanyama Forest Reserve, near Chiungiza, 1,550 m. N. K. B.

PN-20B4

Figure 34.—A plant of V. filipéndula in Luano Forest Reserve near Chingóla, Zambia. (Smith & Fanshawe 4660.)
56 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

V ) 'v r^^^-^

Leopoldv¡lle-¡)

• 15. V. benguelensis

• 16. V. filipéndula

17. V. campicola

0 200 400 600 800 MILES

I I I I I I I I I

I ' 1 ' 1 r—I

0 400 800 1200 KILOMETERS

PN-2055

Figure 35.—Distribution of V. benguelensis, V. filipéndula, and V. campicola.

Robson 1519. (SRGH). Southern Prov. Ncheu Distr. Lower Kirk Range, Chipusiri. 1,460 m. A. W.

Exell, F. A. Mendonça & H. Wild 95i. Mar. 17, 1955. (SRGH). MOZAMBIQUE: Niassa. Vila Cabrai.

Pedro e Pedrogâo 3708. May 20, 1948. (EA).

17. Vemonia campicola S. Moore. J. Bot. 52 :97.1914. (Fig. 36). See also fig. 35.

Type Coll : Kassner 28U5a. Congo. Lufongo. May, 1908. (Photo—BM !).

V. retifolia S. Moore. J. Linn. Soc. Bot. 47:264. 1925-27.

Type Coll : Jelf U. Rhodesia. Luwingu. May, 1922. (Photo.—BM !).

V. venosa S. Moore. J. Linn. Soc. Bot. 47:265. 1925-27.

Type Coll : Kassner 25Í5. Congo. Kipaila. Mar., 1908. (Photos.—BM !, HBG).

Of this species, I have only the photographs of type collections as cited and a single immature

achene of V. venosa which was sent to me by the Hamburg Herbarium. I am, therefore, preparing

no description nor do I see any value in repeating previously published descriptions. I have no doubt

that all of the specimens belong to a single species which is distinguishable from V. sclerophylla

by the lanceolate foliage ; the ñower, which is clearly composed of a tube twice as long as the limb ;

the limb which has short, triangular lobes ; and by the foliage closely investing the fewer crowded

heads. The ñower color note ("yellow") on the label for Jelf No. 44 may be in error as I know of

no other species in this complex with yellow flowers. If the flowers are, indeed, yellow, V. campicola

would not appear to belong to section Stengelia, but this I cannot credit as it otherwise appears

to belong here.

18. Vemonia nigritana Oliv. & Hiern in Oliv. Fl. Trop. Africa. 3:288. 1877. (Fig. 38, See also fig.

37.)

Type Coll : Barter s. n. Nupe (K !).

Perennial from a root crown. Stems glabrate to sparingly pubescent; leaves ovate to ovatelanceolate, to 15.5 cm. long, 5.5 cm. wide; margin scarcely apiculate to apiculate dentate; blade

harshly scabrous above, rarely scattered pilose, sometimes glandular below ; apex acute to obtuse,

base acute, sessile or petiole to 0.7 cm. long. Heads few, in a close or crowded panicle, to 3.0 cm.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 57

high, 3.0 cm. in diameter; phyllaries to 4.0 cm. long, lanceolate, pubescent to scabrous, generally

dark red within, yellowish green without, apex rounded, sometimes radiating or reflexed ; flowers

to 1.4 cm. long, tube enlarging graduating into limb, lobes long, lanceolate, purple?; achenes to

=—'f

tiiiiini mil iMiliiiilimLJuiliiiihni'

Figure 36.—Type collection of V. campicola S. Moore. A, Habit; and B, detail of heads. (Photos, BM.)

PN-2066

• 18. V. nigritana

• 20. V. sclerophylla

0 200 400 600 800 MILES

_l I \ I I I I I

~^ 1 ' 1 "—I

0 400 800 1200 KILOMETERS > ,V^'l\f ^

S.i /OElisobethville,-,^

Figure 37.—Distribution of V. nigritana and V. sclerophylla.

PN-2057
58 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

PN-2058

Figure 38.—Type collection of V. nigritana Oliv. & Hiern in Oliv. A, Detail of heads; B, habit; and C, achene. (K).

Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 59

0.9 cm. long, sometimes curved near narrow base, dark brown, moderately pubescent to pilose,

callus small, discreet ; pappus to 1.4 cm. long, filiform.

F. nigritana is very distinct from other species of section Stengelia. Although the morphology

of the flowers and achenes clearly places this species with the rosette-leaved group, the heads are

borne in panicles on a leafy stem. Tubercles on the underside of the leaves are surrounded at the

base by a rosette of specialized cells which, from the top, look much like a rose window in a Gothic

church. I have not been able to investigate these tubercles in detail, but they appear to have large

lumens and thin walls and may be secretory in function. Mineral deposits are common on the

underside of leaves of the species.

In Sierra Leone, milk calabashes are washed with a decoction of the leaves or roots to curdle milk

(Deighton).

V. nigritana is restricted to the western portion of Africa below the Sahara west of 15° E.

longitude (see fig. 37, p. 57).

Collections Seen: GAMBIA: Abeokuta. Irving s. n. (K). PORTUGUESE GUINEA: Pitche, Pansor.

Espirito Santo SJfSO. Sept. 3, 1955. (K). GUINEA: Fouta-Djallon. Madaud 272. (K). Near Siguiri.

H, J. Felix 500, Nov., 1935. (P). Near Kissidougou. H. J. Félix 2083. July, 1937. (P). SIERRA

LEONE: Musaia. F. C. Deighton U70. Dec. 19, 1946. (K). IVORY COAST: Cercle de Baoulé Nord.

Entre Bouaké et Langouassou. A. Chevalier 22H7. July 21, 1909. (K). Baoulé, Toumodi. H.

Pohequin 179. Oct., 1896. (P). GHANA: Near Babili Mts. K. 0. Darko 1051. (K). 14.6 mi. N. of

Chiraa on Sunjani-Wenchi road. R. R. Innés 312. Sept. 27, 1956. (K). Bame Pass. Eastern Prov.

/. K, Morton 9316. Oct. 25,1953. (K). NIGERIA : Bauchi Prov. Lame Distr. Dababe. 3,000 feet. G. V.

Summerhayes 27. Aug. 19, 1954. (K). Jemaa Div. Dogon Kurmi near Jagindi. 520 m. F. N. Hepper

1033. Oct. 16, 1957. (K). Minna Div. Near Minna. R. D. Meikle 11,80. Dec. 3, 1949. (K). Ilorin Prov.

Ilorin Distr. Share Forest Reserve E. Ujor FHI31613, Jan. 16, 1952. (K). CAMEROON: Adamawa

Prov. Jamtari on Jamtari-Karamti road. M. G. Latilo & B. 0. Daramola FHI28930. (K).

19. Vernonia sciaphila S. Moore. J. Bot. 56:208.1918. (Figs. 39 and 40.)

Type CoU: Kassner 2106. Rhodesia. Sangolo stream. Dec, 1907. (Photos—BM!, HBG).

Perennial from a root crown. Stems to 1.5 m. tall, densely pubescent, moderately leafy. Leaves

sessile, to 12.0 cm. long, 2.5 cm. wide, lanceolate; margin nearly entire; blade moderately pilose

above, raised reticulate, densely pilose, glandular below. Heads not crowded, few, corymbosely

arranged ; heads to 2.0 cm. high, 2.0 cm. in diameter; phyllaries to 1.8 cm. long, generally pubescent

without, particularly toward the tip, green, apex sometimes apiculate ; flowers lilac, to 1.3 cm. long,

limb sometimes longer than the tube, texture thin, lobes of limb long, lanceolate; achenes to 0.8 cm.

long, thin, glabrous, yellowish, occasionally glandular, callus confluent with the costae; pappus

filiform, to 0.9 cm. long, sometimes with spatulate tips.

V. sciaphila is readily recognizable from the photographs of sheets of the type collection at British

Museum and Hamburg and from the achene kindly sent to me by Walther from Hamburg. This may

well be the taxon described by De Wildeman and Muschler as V. luembensis of which I have seen

no authentic specimens. However, their description says that V. luembensis is a shrub with strong

branches while all of the plants I have seen of V. sciaphila are typically annual shoots rising from

a perennial root crown which branch only near the summit to bear several heads.

In Zambia, Fanshawe notes, "A multistemmed subshrub with erect branches. Bark grey-black,

smooth, velvety, slash green thin, soft, sappy. Flowering, January, February. Fruiting, February,
60 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

PN-2059

Figure 39.—Type collection of V. sciaphila S. Moore.

A, Habit, and B, detail of heads. (Photos, BM.)

PN-2060

Figure 40.—A, V. sciaphila is abundant in relatively

undisturbed Brachystegia woodland near Abercorn,

Zambia. (Smith & Richards 4665). B, Note the longer

stems beneath the heads of these plants near Kitwe,

Zambia (Smith & Fanshawe 4660).
OBSERVATIONS ON SfENGELIOID SPECIES OF VERNONIA 61

March. Locally common in miombo woodland. Abercorn, Mporokoso, Kawamba, Luwingu, Fort

Roseberry, Samyfa, Copperbelt. Which means that it almost certainly occurs in the Congo pedicle."

The glabrous achenes of this species are distinctive, but the flowers are indistinguishable from

those of the rosette-leaved species.

Because I did not specify that I wished to see this species, I did not see sufficient specimens to

delimit the geographical range of the species. I know it only from Zambia as follows : Western Prov.

Between Kitwe and Ndola. C. E, Smith, Jr. & D. B. Fanshawe Í660. Feb. 11, 1967. (BM, EA, K,

SRGH, US). Northern Prov. Near Abercorn on road to Ndingi Pans. C. E, Smith, Ji\ & H, M,

Richards 1,665, Feb. 17, 1967. (BM, EA, K, SRGH, US).

20. V. sclerophylla O. Hoffm. Bol. Soc. Brot. 13:13.1896. (Fig. 41. See also fig. 57.)

Lectotype Coll: Welwitsch S277, Angola. Pungo Andongo. (K).

y. lafukensis S. Moore. J. Bot. 52:94. 1914.

Lectotype Coll : Kassner 2846a. Congo. Lufongo. May, 1908. (Photo—BM !).

Perennial from root crown. Stems to 1.2 m. tall, puberulent or glabrate; leaves crowded, to 6.5

cm. long, 3.5 cm. wide, sessile; margin sharply apiculate dentate; blade glabrate to scabrous above,

few to moderately pilose, glandular beneath ; apex sub-acute to obtuse, apiculate ; base subacute to

obtuse. Heads numerous, crowded, cymosely arranged; heads to 1.5 cm. high, 1.5 cm. in diameter;

phyllaries to 1.5 cm. long, herbaceous tip with prominent midrib which may extend into an

apiculum, lower often pilose; upper small leaves often subtending heads; flowers to 1.2 cm. long,

tube enlarging gradually into limb so that they are not clearly distinct, lobes of limb lanceolate,

thin, blue or light purple; achenes immature, to 0.3 cm. long, medium brown, densely pilose, callus

small ; pappus narrowly flattened, acute, nearly plumose.

V. sclerophylla, from the numerous crowded heads, is clearly distinct from the last species.

However, it remains to be proven that the two cited Tanzanian specimens are truly conspecific with

the type of 7. sclerophylla. The latter has broader herbaceous phyllary tips than the Tanzanian

specimens. All have the accentuated midrib on the phyllaries which is unusual in this group of

Vernonia species. For what it is worth, the foliage is essentially the same on all specimens. On the

other hand, if these specimens do not belong here, I have no other place to put them.

The flowers of this species clearly show its close relationship with the rosette-leaved species which

follow. In general aspect, it is also reminiscent of V. buchingeri, but it differs from the latter species

in many ways, not the least of which are the numbers and disposition of the heads.

I have seen so few collections of V. sclerophylla that I have no knowledge of the true geographic

range of the species. It apparently spans the African continent south of the equator (see fig. 37,

p. 57).

Collections Seen: TANZANIA: Without locality. H. G. Faulkner 1219. (K). Usambaras. Sangarawe.

R J. Greenway 829. Sept. 18, 1928. (K).

21. Vernonia gerberiformis Oliv. & Hiern in Oliv. Fl. Trop. Africa. 3 :285. 1877. (Figs. 42, 43, and

44.)

Type Coll : Schweinfurth 2688. Bei. Addai. Nov., 1869. (K!).

V. macrocyanus 0. Hoffm. Bol. Soc. Brot. 13:20.1896.

Type Coll : Welwitsch 3883. Angola. Huila. Prope Catumba et versus Ohay. Jan., 1860.

(Photo. BM).

V. macrocyanus var. ambacensis Hiern. Cat. Welw. PI. 1:532.1898.

Type Coll: Welwitsch 3882. (K!).

V. primulina 0. Hoffm. Kunene-Sambesi Expedit. 402. 1903.

Type Coll: Baum 13i. Am Chitanda unterh. Gondkapse. 1,100 m. Sept., 1892. (K,

(Photo. BM).
62 AGRICULTURE HANDBOOK NO. 396, U.S. DEFT. OF AGRICULTURE

*:'#:

1

/

r

B

Figure 41.—Type collection of V. sclerophylla 0. Hoffm.

A, Detail of heads; and B, habit. (K). Scales in millimeters.

.0-, ^ f.,/^./

i'tt Xu^^

PN-2062

Figure 42.—Type collection of V. gerheriformis Oliv. &

Hiern in Oliv. A, Habit (inset, label) ; and B, detail

of head. (K). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 63

V. nandensis S. Moore. J. Linn. Soc. Bot. 35:323. 1905.

Type Coll : Elliot 7032. Nandi. 7,000 feet. (Photo—BM !).

V. gossweileri S. Moore. J. Linn. Soc. Bot. 37:312. 1906.

Type Coll: Gossweiler 1167. Angola. Malange in Parinari and U'Gambo groves.

(Photo—P).

1r

N '

B

^

' ^

PN-2063

Figure 43.—Type collection of V. gerheriformis. A, Achene; B, pollen grain (average diameter 66.3/x) ;

C, unopened flower; and D, dissected flower (expanded in water). (K). Scales in millimeters.
64 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

• 2L V. gerberiformis

A 22. V. nyassae

* 23. V. acrocephala

0 200 400 600 800 MILES

I I I I I I l_l

"" 1 1 1 1 I

0 400 800 1200 KILOMETERS

PN-2064

Figure 44.—Distribution of V. acrocephala, V. gerberiformis, and V. nyassae.

Usually rosettes from a perennial root crown, becoming caulescent in the northwestern part of

its geographical range, flowers generally borne singly on an elongated scape. Stems and scapes to

0.6 m. tall, glabrous to usually densely pubescent or lanate, rarely with a dense lanate mass at the

apex of the root crown ; leaves sessile, spatulate, to 23 cm. long, 6.0 cm. wide, margin sometimes

undulate, rarely entire, usually apiculate, sometimes dentate; blade glabrous to rarely scabrid to

scattered pilose above, glabrous to moderately pilose, occasionally glandular below. Heads generally

borne singly, to 2.5 cm. high, 3.0 cm. in diameter; phyllaries to 2.3 cm. long, lanceolate, sometimes

pubescent without and ciliate, usually green, sometimes reddish ; flowers to 2.7 cm. long, with tube

equal to twice as long as limb, tube widening gradually into limb, limb thin, lobes long lanceolate,

purple or violet to frequently blue ; achenes to 0.6 cm. long, generally stramineus, usually densely

pubescent or pilose, rarely glandular, callus, discreet; pappus to 1.9 cm. long, narrowly flattened,

nearly plumose.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 65

V. gerberiformis is a very variable species. Much of this may be due to ecological conditions and

a certain amount can certainly be attributed to the specimens available which are almost all collected

before the plant is mature. I suspect that some of the variability may have a genetic base ; I know

of no chromosome counts for this species, but the wide variation is reminiscent of species of plants

in which the chromosome number varies.

The pollen grains of this species are distinctive. The surface reticulation is winglike giving the

grains a much diiïerent appearance from the pollen grains of other species in the complex. It is

this which emphasizes the conspecificity of V. gerberiformis and F. macrocyanus. The extremes in

size are completely bridged by intermediate specimens.

7. gerberiformis is widely distributed over much of Africa south of the Sahara with a single

collection known from Natal in South Africa (see fig. 44). Because this is a plant of the

savannas and plains where it is largerly hidden among the grass and other herbs, it is probably

more common than the number of available specimens indicates.

Selected Collections: GUINEA: Near Labe. F. /. Félix 658. Dec, 1935. (P). NIGERIA: Zaria Prov.

Anchau Distr. Anchau. S, C. Omjeagocha FHI76Í2, Mar. 6, 1944. (K). Bokkos Plateau. 4,000 feet.

D. £7. Coombe 73. Dec. 27, 1954. (K). CAMEROON : Adamawa Div. Belel Distr. Belel on Demasa road.

280 m. F. N. Hepper 1629. Jan. 1, 1958. (K). ANGOLA: E. of Cuanza River. A. G. Curtis SSI. Sept.

23, 1923. (GH). Sierra de Chella near Huila. 1,850 m. H. Humbert 16711. Aug., 1937. (P), Near the

Coculovu River between Huila and Chibia. H. H. W. Pearson 22S7. May 12, 1909. (K). Mossamedas.

Berthelot s. n. (P). Galangus Mission, Nova Lisboa. H. G. Faulkner A170. Aug., 1942. (K). KENYA :

Trans Nzoia Plain. 5,900 feet. D. R. Tweedie S56. Feb., 1937. (K). Kitale. A. Bogdan S721. Apr. 18,

1953. (K). UGANDA: Mulanda Hill, Bulemezi. T. D. Maitland 95. June 24, 1915. (K), Entebbe. 3,900

feet. P. Chandler 1159. June, 1934. (K). REPUBLIC OF CONGO: Ruzizi Plain, Luwingi. R. Germain

5905. Jan. 30, 1950. (K). TANZANIA: West Lake Prov. Mumwendo, Bugifi, Ngara. 4,500 feet.

R. E. S. Tanner 5235. Aug. 12, 1960. (K). Kakama Distr. Nyamgalika-Magende. A. A. Bullock

3185A. Aug. 21, 1950. (K). Mpanda Distr. Mugombasi, about 65 mi. S. of Kigoma. 3,000 feet.

ß. M. Harley 9A77. Aug. 31, 1959. (K). Ufipa Distr. Malonje. 8,000 feet. A. A. Bullock 1886.

Nov. 21, 1949. (K). Kyimbila Distr. N. of Lake Nyassa. A. Stolz 1001. (K). ZAMBIA: Abereorn.

Nkali Dambo. H. M. Richards 13181. Sept. 1, 1960. (SRGH). Muwosi Stream 42 mi. S. of

Mwinilunga on Kabompo road. /. P. Loveridge 706. May 31, 1963. (AAH). Lake Ishika. D. B.

Fanshawe 216. Aug. 9, 1953. (K). Solwezi Distr. Head of Mbulungo Dambo, W. of Mutanda Bridge.

E. Milne-Redhead 720. July 17, 1930. (K). Solwezi. E. Milne-Redhead 1195. Sept. 24, 1930. (K).

Ndola. D. B. Fanshawe 1019Í. Oct. 6, 1967. (ND, US). RHODESIA: Salisbury Distr. Hunyani Landing, F. C. Greatrex s. n. Sept. 10, 1946. (SRGH). 3 km. along Norton road from Salisbury-Beatrice

road. 1,800 m. R. Rutherford-Smith 20. Aug. 30, 1960. (K, SRGH). Marandellas Distr. 4,500 feet.

ß. M. Davies 2259. Nov., 1956. (SRGH). Hartley Distr. Lake Mcllwane. /. A. Whelan U27. Sept.

14, 1957. (SRGH). Melsetter Distr. N. W. of Musapa Mt. 6,700 feet. N. C. Chase 6702. Sept. 6,1957.

(K, SRGH). Mutsarara Farm. A. 0. Crook M190. Oct. 5, 1950. (SRGH). Chimanimani Mts. 5,000

feet. H. Wild 2964. July 9, 1949. (K, SRGH). SOUTH AFRICA: Natal. Zululand. J. M. Wood 7^63.

Dec. 20, 1898. (K).

22. Vernonia nyassae Oliv. Hooker's Icon. PI. 36, t. 1349. 1881. (Fig. 45. See also fig. 44.)

Type Coll : Thomson s. n. Higher plateau north of Lake Nyassa. (K !).

Small rosettes from a perennial root crov^n, sometimes with a lanate tuft at the top from which

the leaves arise. Leaves ovate-lanceolate or lanceolate to, usually, spatulate, to 9.0 cm. long, 1.5 cm.

wide, generally smaller; margin entire; blade glabrous to moderately pilose, glandular above,

glabrous to moderately pilose, glandular beneath; apex obtuse to acute, base cunéate. Scape to
66 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

O

^0 -^

It -4 W

B

PN-2065

Figure 45.—Type collection of V. nyassae Oliv. A, Habit; B, detail of head; C, achene; D, unopened flower, and

E, dissected flower (expanded in water). (K). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 67

0.15 m. tall, scattered pilose to pilose or pubescent, usually densely pilose just beneath the head,

sometimes with a bract midway on the scape; heads single, to 1.5 cm. high, 2.0 cm. in diameter;

phyllaries to 1.5 cm, long, lanceolate, often 3-nerved, glabrate to pilose without, generally reddish,

tip often attenuate acute; flowers to 1.4 cm. long, tube broadening gradually into the limb, lobes

thin, long, lanceolate, sometimes scattered pilose without, violet or purple; achenes (immature)

to 0.35 cm. long, light to dark brown, moderately pilose, callus discreet; pappus to 1.1 cm. long,

filiform, sometimes moderately plumose.

V, nyassae is easily separated from 7. gerberiformis by its small size, entire leaves and generally

by its overall pilosity. Contrary to the general pattern in this group, F. nyassae does not seem to

be highly variable although one of the specimens cited is remarkably glabrous throughout. I cannot

tell whether the foliage of many of the specimens has reached its full size. All of them were

collected before the heads were fully mature.

V. nyassae has been collected largely on the highlands immediately north of Lake Nyassa

although one collection is known from Zambia to the west and another collection, which is

apparently this species, comes from far south in Transvaal, South Africa (see fig. 44, p. 64).

Specimens Seen: TANZANIA: Sao Hill. 6,200 feet. A. M, Watermeyer 32. Feb., 1959. (K). Mbeya

Distr. Slopes of Mbeya Mt. 9,000 feet. B, D. Burtt 6S3Í. Sept. 25, 1936. (K). Southern Highlands

Prov. Njombe Distr. Elton Plateau. 8,500 feet. R, L. Willan 172. Oct. 6, 1954. (K). Near Njombe.

2,100 m. H. M. Richards 787Jf. Jan. 17, 1957. (K). ZAMBIA: Highlands, Ndola. C. F. E. Allen 868.

Oct., 1906. (SRGH). SOUTH AFRICA: Transvaal. 121/2 mi. E. of Lydenburg on the road to Sabie.

7,000 feet. L. E. Codd & B. de Winter 8291. Nov. 12,1947. (K).

23. Vernonia acrocephala Klatt. Ann. Naturhist. Hofmus. 7:100. 1892. (Fig. 46. See also figs. 4, A,

and 44.)

Type Coll : Mechow 278. Angola. Malange, Oct., 1879. (Drawing & fragments—GH).

V. extranea S. Moore. J. Linn. Soc. Bot. 47:263. 1902.

Type Coll : Kassner 2707. Congo. Kundelungu Mt. Apr. 1908. (K !).

Rosettes from perennial root crown with densely pilose apex from which the leaves and scapes

arise. Leaves (mostly immature) ovate to lanceolate or spatulate, to 14.5 cm. long, 9.0 cm. wide;

margin entire ; blade scattered pilose to pilose on veins only above, scattered pilose, often glandular

beneath ; apex rounded to acute, base generally cunéate. Scapes to 0.2 m. tall, moderately to densely

pubescent, more pubescent just beneath head, sometimes with small bracts midway; heads single,

to 2.0 cm. high, 2.0 cm. in diameter; phyllaries broad, generally acute, to 1.8 cm. long by 0.9 cm.

wide, often reddish, sometimes veined, often pubescent without; flowers to 1.1 cm. long, rose? or

mauve, tube widening gradually into limb, limb with long, thin, lanceolate lobes ; achenes to 0.4 cm.

long, glabrous to moderately pubescent, light brown, perhaps nearly square in cross section when

mature ; pappus to 0.9 cm. long, narrowly flattened, scarcely plumose, apex acute.

V. acrocephala is usually represented by immature specimens. Apparently the foliage is much

larger than that of the last species when it is mature. It is also clearly separated by the broad

phyllaries.

V. acrocephala has been collected on opposite sides of the African continent in Cameroon, where

it is apparently most abundant, and in Tanzania (see fig. 44, p. 64). Inasmuch as the type collection

was made in Angola, it is probably more widely distributed and abundant than the recorded collections indicate, but, due to its habit and habitat, it is apparently difficult to find.
68 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

ri

'^•l j&'-:-

/^

Dr. CM

.,/.,J...,t,U.)..,^ ../„ j<-.,

ty ,«.v. /T///yy, /f-.w /i?.«-.*. ^JU.

PN-2066

Figure 46.—Type collection of V. acrocephala Klatt. A, Achene; B, flower dissected; C, habit; and

D, flower unopened (expanded in water). (GH). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 69

Collections Seen: CAMEROON: Mt. Gotel. H. /. Félix 3388, Mar., 1939. (P). Bamenda Div. BafutNgemba Forest Reserve. 2,210 m. F. N, Hepper 2181, Feb. 25, 1958. (BR, K). Bamenda Sta. 5,000

feet. T. D. Maitland 15^9 bis, June, 1931. (K). Bamenda. 5,000 feet. T. D. Maitland 1U7. (K).

12 mi. E. of Bamenda. 5,200 feet. C, F, A. Onochie FHI3Í867, Mar. 20, 1955. (K). Njon, Tibali.

G. Tessman2726, Sept. 6, 1914. (K). TANZANIA: Unyika. W. Goetze H50, (BR, G). Ufipa. Sumbawanga. 7,100 feet. H. M. Richards 3U1. Nov. 27,1954. (K).

24. Vernonia praemorsa Muschler. Bot. Jahrb. Syst. 46:68. 1911. (Figs. 47 and 48. See also fig.

4,iS.)

Type Coll: Stolz lOJ^, Nyassa hochland, Station Kyimbila, Namulapi. Nov., 1907. (GH,

K).

V. castellana S. Moore. J. Bot. 52:95. 1914.

Type Coll: Gossweiler 2883, Angola: Benguela. Forte dom Affonso, R. Cului. (K!).

7. kuluina S. Moore. J. Bot. 56:209. 1918.

Type Coll : Gossweiler 290i, Angola : Benguela. (K !).

Rosettes from a perennial root crown, apex often v^ith a felty tomentum from v^hich the leaves

and scapes arise. Leaves ovate to lanceolate, to 24.0 cm. long, 10.0 cm, wide ; margin undulate to

apiculate dentate; blade moderately pilose to scattered scabrid above, sparsely to moderately pilose,

generally glandular beneath; apex obtuse, rounded or subacute; base cunéate, often extending

down petiole, petiole 1.5 to 5.0 cm. long. Scape sparsely pubescent to pubescent, often bracteolate

midway, bearing many closely crowded heads ; heads to 1.0 cm. high, 1.0 cm. in diameter ; phyllaries

to 0.9 cm. long, lower often pilose without, green or usually reddish; ñowers to 1.0 cm. long, tube

gradually broadening into limb, lobes thin, frequently pilose on the tip without, mauve or purple ;

only immature achenes seen, to 0.3 cm. long, medium brown, moderately pubescent, callus discreet?;

pappus to 0.7 cm. long, filiform, scarcely plumose.

V, praemorsa is readily distinguished by the numerous crowded heads on the long scapes. In

eastern Africa, it is apparently locally common, but, like many of the rosette species, is only found

when it is conspicuously in flower. None of the herbarium specimens bear mature achenes.

Fanshawe's note for Zambia is, '^Flowering December, January, February. Fruiting January,

February, March. Occasional in miombo woodland, dambo margins and plains. Distribution in

Zambia, Isoka, Abercorn, Copperbelt. This type of distribution usually means throughout the high

rainfall belt, but, owing to its rarity, it has not been collected in the intermediate districts.''

V, praemorsa is found in Angola in the west and in Tanzania. Zambia and Malawi in the east

(see fig. 48, p. 71). It will probably eventually be found in intervening stations when the midcontinent has been more thoroughly collected.

Collections Seen: TANZANIA: Iringa Prov. Msima Stock Farm. H, E, Emson 2Í7, 1932. (EA).

Sao Hill. 6,500 feet P. /. Greenway 6^09. Nov. 5, 1941. (EA). Ruhudje, Lupembe. 1,600 m. H, J,

Schlichen 1222, Sept. 25, 1931. (GH, K). ZAMBIA: Abercorn Distr. Nkali Dambo. H, M, Richards

3922. Jan. 5, 1955. (K). Ndingi Pans. 1,500 m. H. M, Richards 11993. Dec. 19, 1959. (K). MALAWI:

Rumpi Distr. Nyika Plateau. 2,400 m. H, M, Richards 101,86. Jan. 5,1959. (K).

25. Vernonia subaphylla Baker. Kew Bull. 1895:290. (Non Muschler). (Figs. 49 and 50. See also

fig. 48.)

Type Coll : Carson 10. Kalangwizi, Moero. 1894. (K !).

V. armerioides 0. Hoffm. Bot. Jahrb. Syst. 24:462.

Type Coll: Pogge 1315. (Not seen.)

V. agrícola S. Moore. J, Bot. 56:208. 1918.

Type Coll : Kassner 2136. Rhodesia. Lukanda River. Dec, 1907. (K !).
70 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

^

PN-2067

Figure 47.—Type collection of V. praemorsa Muschler. A, Habit; B, flower unopened and dissected

(expanded in water) ; C, detail of heads; and D, achene (immature). (K). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 71

• 24. V. praemorsa

A 25. V. subaphylla

• 26. V. denudata

0 200 400 600 800 MILES

_1_J \ I I I L

Figure 48.—Distribution of V. denudata, V. praemorsa, and V. subaphylla.

PN-2068

Rosette from a perennial root crown, apex frequently with dense tan tomentum from which the

leaves and scrapes emerge. Leaves lanceolate to, rarely, obovate, to 19.0 cm. long, 5.0 cm. wide;

margin entire to barely apiculate to apiculate dentate ; blade often nitid, glabrous to usually few

pilose, rarely moderately pilose above, scattered to densely pilose or hirsute, glandular beneath;

petiole to 2.2 cm. long. Scape to 0.8 m. tall, heads rarely single, usually several, in an open panicle ;

heads to 1.3 cm. high, 1.5 cm. in diameter; phyllaries to 1.2 cm. long, outer sometimes pubescent

or lepidote, tips rounded, scarious, usually purple or reddish; flowers to 1.1 cm. long, tube enlarging

gradually into limb, lobes long, lanceolate, thin, white to pink, mauve, or usually, purple ; achenes

to 0.45 cm. long, yellow to tan or medium brown, ridged-angled at maturity, glabrous to moderately

pilose, rarely glandular, callus light to medium yellow, discreet, small; pappus to 0.7 cm. long,

filiform.

V. subaphylla is largely distinguished by the scarious purple or reddish phyllary tips. It is similar

in habit and aspect to V. praecox, but the latter species has much more narrow phyllaries with

narrowed tips which can rarely be interpreted as scarious. No other species in the rosette-leaved

group is liable to be confused with V. subaphylla.

V. subaphylla is distributed across southern Africa from Angola to Tanzania with an extension

northward into Cameroon and Nigeria where it apparently is less common (see fig. 48).
72 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

rf ^

c>

■■^aÈ- 1

PN-2069

Figure 49.—Type collection of V. subaphylla Baker. A, Habit; B, flower (expanded in water) ;

C, detail of heads; and D, achene (immature). (K). Scales in millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 73

PN-2070

Figure 50.—A, Plants of V. subaphylla are barely seen in the surrounding grass and herbs at the base of a tree

in Ndola Forest Reserve near Ndola, Zambia. The shiny leaves belong to the Vernonia plant. B, Heads in flower

are very inconspicuous, but the pappus of mature achenes is easily seen.

Collections Seen: NIGERIA : Von. Bauchi Plateau. 3,000-4,500 feet. /. D. Young 139. Jan., 1922. (K).

CAMEROON: Nkambe Div. 23 mi. W. of Nkambe. 3,500 feet. F. N. Hepper 2827. Feb. 11, 1958. (K).

REPUBLIC OP CONGO: Elisabethville. F. A. Rogers 163. Oct. 22, 1911. (K). Tshinsenda. F. A. Rogers

20A. Oct. 18, 1911. (K). ANGOLA: Camaquenhe, Dundo. /. Gossweiler H021. Apr. 2, 1948. (K).

Benguella. /. Gossweiler 2921. (K). Longa near Minnesera. 1,200 m. H. Baum 659. Jan. 11, 1900.

(K). TANZANIA: Songea Distr. 1.5 km. S. of Gumbiro. 780 m. E. Milne-Redhead & P. Taylor 8^16.

Jan. 24, 1956. (EA, K, SRGH). ZAMBIA: Mporokoso Distr. 16 mi. from Mporokoso, KasamaMporokoso road. 1,200 m. H. M. Richards 1207A. Jan. 6, 1960. (K). Kasama Distr. Mungwi. E. A.

Robinson UA9. Dec. 4, 1960. (EA, K, SRGH). Western Prov. Between Ndola and Kitwe. C. E.

Smith, Jr. & D. B. Fanshawe Jt661. Feb. 11, 1967. (EA, SRGH, US). Chingóla. D. B. Fanshawe

2667. Dec. 19, 1955. (K). 15 mi. S. E. of Mufulira. 1,300 m. E. A. Robinson 3304. Jan. 26, 1960. (K).

Solwezi Distr. Chingólo to Solwezi. P. J. Greenway & /. P. M. Brenan 8127. Sept. 29, 1947. (EA,
74 AGBICULTURE HANDBOOK NO, 396, U.S. DEPT. OF AGRICULTURE

SRGH). Mwinilunga Distr. 10 mi. W. of Lunga, 40 mi. S. of Boma. E, Milne-Redhead 909, Aug. 15,

1930. (K). 25 mi. S. E. of Chôma. 4,200 feet.^. A. Robinson 1775. Dec. 17, 1956. (K, SRGH).

26. Vernonia denúdala B-utoh, & Burtt. Rev. Zool. Bot. Africaines 23:37. 1932. (Fig. 51. See also

fig. 48.)

Type Coll : de Witte 588, Congo. Kangenge. Aug., 1931. (K).

Rosette from a perennial root crown, apex densely tan, woolly. Leaves produced after inflorescence mature, broadly spatulate, to 32.0 cm. long, 13 cm. wide (only one herbarium specimen with

leaves seen); margin irregularly apiculate dentate; blade scattered pilose glandular above,

moderately pilose, glandular beneath; apex rounded, base cunéate acute. Inflorescences often

several from single root crown, open paniculate, heads on long peduncles. Scapes sparingly

pubescent, scantily bracteolate; heads to 1.5 cm. high, 1.5 cm. in diameter; phyllaries to 1.3 cm.

long, lanceolate, usually sparingly pubescent without, somewhat glandular, sometimes ciliate,

generally with reddish margin ; flowers to 1.0 cm. long, white to deep mauve or purple, tube broadening gradually from base into limb; achenes to 0.4 cm. long, yellow to, usually, medium brown, slim,

moderately to densely pilose, callus pappilose, discreet; pappus to 1.0 em. long, filiform, scarcely

plumose, clear white.

y. denúdala can be confused with no other species in section Stengelia, because of the distinctive

form of the inflorescences. In detail, the flowers are not unlike those of other rosette-leaved species,

many of which also have a dense tomentum at the apex of the root crown. Other species in this group

also produce the inflorescences before the leaves appear and, in all of these species, usually only

the flowering plants are collected.

Although the type collection of F. denúdala was made in the Congo, all subsequent collections

have come from Tanzania and Zambia in eastern Africa (see fig. 48, p. 71).

Collections Seen: TANZANIA: Kigoma Distr. Near Ulemba. 900 m. H, M, Richards 11713, Nov. 4,

1959. (K). Ufipa. Sumbawanga-Nkunde. 7,000 feet A, A. Bullock 1951. Nov. 28, 1949. (FI, K).

Utahya. H. S. Mahinde 215. Aug. 4, 1958. (EA). ZAMBIA: Abercorn Distr. Kalambo Falls road.

1,500 m. H.M.Richards 6502, 1U26. Sept. 15, 1956 & Sept. 4, 1959. (K, SRGH). Abercorn. 5,500

feet. B. Kafuli H. Sept. 28, 1955. (SRGH). Western Prov. Mwinilunga Distr. 8 mi. W. of Kakome.

A. Angus 567. Sept. 28, 1952. (K).

27. Vernonia praecox Welw. ex O. Hoffm. Bol. Soc. Brot. 13:16.1896. (Figs. 52 and 53.)

Type Coll : Welwitsch 3830. Angola. (K).

F. díwígafaS. Moore. J. Bot. 58:44. 1920.

Lectotype Coll : Fander7/si ^^5^5. Congo. Gamboni. (Photo—BM!)

Rosette from a perennial root crown with tan, densely lanate apex. Leaves produced after

inflorescence is mature, spatulate?, to 24.0 cm. long, 10.0 cm. wide, (only mature leaves seen

were ragged) ; margin apiculate or apiculate dentate; blade glabrous or with a few hairs along

the midrib, shiny above, scattered pubescent beneath; apex rounded; base cunéate acute; petiole

to 4.0 cm. long?. Inflorescence an open panicle on a tall single scape; heads short pedunculate,

to 1.0 cm. high, 1.2 cm. in diameter; phyllaries to 0.9 cm, long, lanceolate, occasionally scarious

tipped, sometimes reddish, barely pubescent without; flowers to 0.8 cm. long, mauve, glandular

without, thin, lobes moderately long; achenes (immature) to 0.3 cm. long, yellow to medium brown,

moderately pubescent to moderately pilose, callus distinct, small; pappus to 0.6 cm. long, filiform.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 75

PN-2071

Figure 51.—Type collection of V. denúdala Hutch. & Burtt. A, Dissected flower, and B, unopened flower

(expanded in water) ; C, achene (immature) ; D, detail of heads; and E, habit. (K). Scales in millimeters.
76 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

PN-2072

Figure 62.—Type collection of V. praecox Welw. ex 0. Hoffm. A, Flowers (expanded in water). Note that they are

about the same diameter from the base to the lobes. B, Habit; C, detail of heads; and D, achene. (K). Scales in

millimeters.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 77

• 27. V. praecox

A 28. V. pumila

• 29. V. chthonocephala

0 200 400 600 800 MILES

J_l I I I I l_l

-1 I I I r

0 400 800 1200 KILOMETERS

Figure 53.—Distribution of V. chthonocephala, V. praecox, and V. pumila.

V. praecox is readily distinguished from the last species by the smaller heads borne on much

shorter peduncles, giving the inflorescence a much different appearance. Foliage may be similar,

but perhaps is usually shiny and leathery in this species. As before, all of the collections are in flower

with no mature achenes and only one specimen has rather worn foliage.

V. praecox is known only from Angola, Republic of the Congo and Zambia (see fig. 53).

Collections Seen: ZAMBIA: Mwinilungu Distr. 10 mi. W. of River Lunga. E. Milne-Redhead 904-.

Aug. 13, 1930. (K). REPUBLIC OF CONGO: Lukeni River near Mutombo-Mukulu. P. Quarre 2533.

July, 1931. (K). Lualu. P. Quarre 2812. Nov., 1931. (K).

28. Vernonia pumila Kotschy & Peyr. PI. Tinn. 37. t. 17a. 1867. (See fig. 53.)

Type Coll : Heuglin s. n. Prés de Bongo. Dec. 1863. (Type not seen.)
78 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

Perennial from a root crown; flowers and stems with leaves appearing at different times. Foliagebearmg stems to 0.4 m. tall, glabrous to pubescent, generally moderately so; leaves lanceolate, to

14.5 cm. long, 5.0 cm. wide ; margin barely apiculate to apiculate dentate ; blade glabrous, sometimes

glandular, or veins scurfy or slightly pubescent above, densely lanate, glandular beneath; apex

obtuse to subacute, base acute, rarely slightly petiolate. Inflorescences to 11.5 cm. tall, heads single

or several in a close or crowded panicle which may have very short or longer peduncles; heads to

2.0 cm. high, 3.0 cm. in diameter; phyllaries to 1.4 cm. long, lanceolate, often pubescent on the tip

without, sometimes glandular without; flowers to 1.6 cm. long, reddish purple or purple, tube

usually 2 to 3 times longer than the limb, lobes sometimes with glandular tips without; achenes

(immature) to 0.3 cm. long, medium brown, moderately to densely pilose, glandular; pappus to

1.1 cm. long, narrowly flattened, apex obtuse.

In many of its characteristics, V.pumila is similar to V. guineensis. The separate appearance of

the flowering and foliar shoots clearly separate these species. Although V. pumila is obviously

related to the other species which bear flowers separately from the leaves, in this species, the leaves

are borne on an elongate shoot rather than in a basal rosette.

V. pumila is distributed from Guinea across Africa to Kenya generally north of the equator

(see fig. 53, p. 77). Sparsity of collections probably fails to provide a complete picture of the

geographic distribution.

Collections Seen: GUINEA : Boké, Guilde. J. Chülous.n. Jan. 9, 1924. (P). Cercle de Boké. J. Ckillou

s.n. June, 1924. (BR, K). NIGERIA : Kontagora. J. M. Dalziel 190. Jan. 20, 1906. (K). Niger Prov.

Minna Fuel Forest Reserve about 5 mi. from Minna on the Zungeru road. R. W. J Keaij FHI37325

Jan. 2, 1958. (K). KENYA : Mt. Elgon. 6,000 feet. D. R. Tweedie 12U. 1934. (K). Trans Nzoia Plains'

D. R. Tweedie 357. Feb., 1937. (K). Nandi. G. F. S. ElUot 7037. (BM, K). UGANDA: Mt. Debasien,

Karamoja. 5,000 feet. A. S. Thomas 2207. 3&n,^, 1937. (K). Senu Teso. 3,600 feet. P. Chandler 21.

Nov., 1931. (K). Hoima, Unyote. ?>,Omie&t. A.G.Bagshawe 9^7. Mar. 3, 1906. (BM) TANZANIAUgalla River, 120 mi. S.W. of Tabora. 3,800 feet. P. R. 0. Bally 71,96. Sept. 28, 1949. (K).

29. Vernonm chthonocephala O. Hoffm. Bol. Soc. Brot. 13 -17. 1896. (Fig. 54. See also ñg. 53.)

liectotyipe Coll: Welwitsch 3886. Angola. (K).

y. 2?eri3aTOa S. Moore. J. Linn. SOG. Bot. 35:324. 1905.

Type Coll : Elliot 8123. Karagwe. 4,000-5,000 feet. Sept. (K !).

Rosette from blanched perennial root crown, apex densely tan, woolly, flowers and leaves

produced at different times. Leaves to 50.0 cm. long, 8.0 cm. wide, lanceolate to spatulate; margins

undulate; blade scattered scabrous to scattered pilose above, pilose on the veins or moderately pilose

or hirsute, sometimes glandular beneath; apex subacute to usually obtuse, base cunéate; petiole

probably seldom longer than about 3.0 cm., pubescent. Scapes produced one to several from each

root crown, each bearing a single head, to 4.0 cm. tall; heads to 1.5 cm. high, 2.0 cm in diameterphyllaries to 1.5 cm. long, lanceolate, glabrous to usually pubescent or pilose sometimes ciliate'

without, usually reddish, sometimes glandular; flowers to 1.4 cm. long, tube expanding gradually

into limb, lobes long, thin, pale purple to purple; achenes to 0.4 cm. long, yellow to light brown

few pilose to usually densely pilose, glandular; pappus to 0.9 cm. long, filiform.

V. chthonocephala is unlike any other species in this group. The very short, single-flowered scapes

are unusual. Because the plant is usually collected only in flower, very few collections of leaves are

available.

The few collections available of 7. chthonocephala indicate that it is widely distributed from
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 79

Cameroon to Sudan in the north to Angola and Malawi in the south (see fig. 53, p. 77). As with

most species of this group of the plains and savannas, the collector must almost step on the plant

before he sees it. Thus, the species is probably more uniformly distributed and more common than

the following specimens indicate.

A

\- -

^ tut J^ n^ ^ //

PN-2074

Figure 54.—Type collection of V. chthonocephala O. Hoffm. A, Detail of head; B, flower (expanded in water) ;

C, habit, and D, achene (immature). (K). Scales in millimeters.
80 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

Collections Seen: CAMEROON: Adamawa Div. Mambila Distr. Karara. 1,500 m. F, N. Hepper 1775.

Jan. 25, 1958. (K). REPUBLIC OP CONGO: Kangenge. G, F, de Witte 556. Aug. 12-15, 1931. (K).

ANGOLA. Malange Distr. /. Gossweiler 1152. (K). Capacca, Membassoco Cubae. 3,500 feet. H. G.

Faulkner 109, July-Aug., 1941, (K). SUDAN: Equatoria Prov. Kagelu Sta. /. G. Myers 8321.

Jan. 26, 1938. (K). KENYA: Trans Nzoia. 6,200 feet. P. H. Kioni 2U6. Mar. 27, 1956. (EA). Hoey's

Bridge. 6,000 feet. Mainwaring K25. 1936. (K). ZAMBIA: Abercorn Distr. TOD of Kambole escarpment. 1,500 m. H. M. Richards 13222. Sept. 11,1960. (K). MALAWI : Nzama road. P. O. Wiehe N26Jf.

Oct. 7, 1949. (K).

The following species probably are not truly members of section Stengelia. However, the phyllary

tips are differentiated (the character on which most species are included in the section) and the

flowers are morphologically similar to the rosette-leaved species which have long been included in

the section. Furthermore, the epoxy acid constituent of the achene oil of V. afromontana is as high

as it is in undoubted members of section Stengelia. On the other hand, many of the species in this

group have lanceolate phyllaries with acute green tips totally unlike the enlarged petaloid phyllaries

of many species of the section. Often, the pappus is sordid, whereas stengelioid species always have

cream, or rarely, pure white pappus. Many of these species have startlingly bright, clear blue

flowers in contrast to the purple, mauve, or white flowers of species of section Stengelia. It is

entirely possible that the blue color is only a function of cell sap pH, because these species

photograph mauve on color film, an indication that much more red is present than the human

eye sees.

For the most part, I have insufficient collections of these species to treat them fully : those which

I have were only casually included in loans, except for my own collections. Where I am convinced

of the synonymy, I present it, usually with a note about the impression I have of the relationship

of morphologically similar taxa. They are not keyed out, because I am not at all sure that I have

recognized biologically valid units and I do not wish to give the impression that I have studied them

as fully as I have the preceding species.

Vernonia bainesii Oliv. & Hiern in Oliv. Fl. Trop. Africa 3:272. 1877.

Type Coll : Baines s. n. South African gold fields. (Type not seen.)

V. wildiiMerxm. Proc. Rhodesia Sei. Assoc. 43:72. 1951,

Type Coll : Dehn 248. Rhodesia. Marandellas. (M !).

F. wildii is distinguished from F. bainesii only by the width of the foliage, a distinction which

I cannot accept as significant. The flowers of this species are morphologically indistinguishable

from those of the rosette-leaved members of section Stengelia. Otherwise, this species is not a good

morphological fit with stengelioid species.

Collections Seen: ZAMBIA: Northern Prov. Kwimbe School. C. E. Smith, Jr. & H. M. Richards 4671.

Feb. 21, 1967. (EA, K, SRGH, US). RHODESIA: Inyanga Distr. Mare Dam. O. West 7266. Apr 25

1966. (SRGH, US).

Vernonia africana (Sond.) Druce. Bot. Soc. Exch. Club Brit. Isles. 1916: 651,1917.

Type Coll : Gueinzius 339. Port Natal. (Type not seen.)

Vernonella africana Sond. Linnea 23:62. 1850.

Vernonia vernonella Harvey. Fl. Cap. 3:53.1894. Norn, illeg. (Superfluous).

F. africana is probably allied to F. bainesii. All of the collections which I have seen are old.

I searched the type locality for living plants, but the area is now devoted to sugarcane fields and

the species may have been completely eliminated. It apparently was never very widely distributed.
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 81

Collections Seen: SOUTH AFRICIA: Natal. Grassy flat, Victoria County, 400 feet. /. M, Wood 753.

May, 1889. (K, US). Natal K. Saunders s. n. May, 1881. (K).

Vernonia glabra (Steetz in Peters) Vatke. Oesterr. Bot. Z. 27 :194. 1877.

Linzia glabra Steetz in Peters. Reise nach Mossambique, etc. 353. 1862-64.

Described with two varieties :

var. a confertissima Steetz in Peters. Op. cit. 354.

Type Coll : Peters. In Boror und Rios de Sene in Mossambique. (Type not seen.)

var. ß laxa Steetz in Peters. Loc. cit.

Type Coll : Peters. Bei Sena, und andere Exampiare am Ufer des Flusses Zambeze in Mossambique. (Specimens not seen and lectotype not selected).

Vatke did not specify which element he intended as the type of the new combination so we must

conclude that he embraced the entire concept described by Steetz including all of the variants

illustrated by the cited specimens. Unfortunately, I did not find a duplicate of any of the Peters

collections, but a more intensive search may reveal some in the European herbaria.

I have seen a number of collections from eastern Africa, from Ethiopia to Rhodesia, which

I would interpret as V. glabra. Included are specimens identified as V. melleri. However, I do not

know the type collections and I cannot be sure of the synonymy. Many of the specimens had blue

flowers, others mauve or purple. I do not think that the flower color is a valid distinction,

A number of varieties have been described for or transferred into V. glabra. My understanding

of this complex is so poor that I make no comment on varieties of V. glabra per se. However, it has

been my experience in taxonomy of tropical or primarily tropical groups that seldom is a clear

understanding achieved of the overall broad species. Recognition of infraspecific taxa is, then,

hardly profitable. I doubt that all of the divisions of V. glabra will stand under close scrutiny.

Vernonia pauciflora (Willd.) Less. Linnaea 4:292. 1829. (Fig. 55.)

Conyza pauciflora Willd. Sp. PI. (ed. 4) 3 :1634. 1803.

Type Coll: Richard s. n. Senegal. (Photo—B!)

V. senegalensis Desf. Cat. Hort. Paris ed. 3. 400.1829.

Type Coll : Jardin des Plantes Juillet 1828. (P !).

I have seen the type of Desfontaine's species in Paris and I have no doubt that it is synonymous

with V. pauciflora. However, my choice of correct name is arbitrary, because I do not know whether

the Lessing article was published prior to the Catalogue or not.

The eastern African specimens which I have seen are blue-flowered, herbaceous plants which

are similar to, but apparently distinguished from V. glabra by their large thin folia^ge.

A further complication is introduced by Lessing's V. senegalensis (Linnaea 4:265. 1829) which

appears to be a transfer of Baccharis senegalensis Pers. (Syn. 2:424. 1806). As the species name

is priorable from 1806, V. senegalensis Desf. becomes an illegitimate name (later homonym).

However, Lessing cites other names in synonymy, including Eupatorium coloratum Willd. (Sp. PL

3:1768. 1803). Whether we agree with Lessing's decision or not, under the Code, the new combination should have been made with the earliest species name coloratum, and V. senegalensis (Pers.)

Less, is illegitimate. If this is the case, V. senegalensis Desf. is now legitimate. Whenever this taxon

is critically evaluated, the investigator should attempt to locate the Persoon specimen and any

corroborative evidence as the chances are rather good that Persoon's specimen and the plants in

the Paris Garden came from the same source.
82 AGRICULTURE HANDBOOK NO. 396, U.S. DEFT. OF AGRICULTURE

PN-2075

Figure 55.—Type collection of V. pauciflora (Willd.) Less. (Conyza pauciflora Willd.).

A, Habit; and B, detail of heads. (Photos, B.)

Vernonia superba. 0. Hoflfm. in Engl. Pflanzenwelt Ost Afrika. 406. 1895. (Fig. 56.)

Type Coll : Stuhlman 3i29. Bukowa. (Type not seen.)

V. paludigena S. Moore. J. Bot. 52:91. 1914.

Type Coll : Kassner 2832. Kabinda, Congo. (Type not seen.)

Although I have seen neither type, from the descriptions and from specimens which I have seen

which were identified in Europe, I have little hesitation in putting these species together. In the

field, V. superba is a very handsome plant in flower. Although it is similar to the two preceeding

species, the heads of this species are usually single and much larger.

Vernonia afromontana R. E. Fries. Acta Horti Berg. 9:116.1928. (Fig. 57.)

Type Coll :R.E.& I. C. E. Fries 78h. Mt. Kenia. (Type not seen.)

In view of the abundance of this species in Kenya, Uganda, and Tanzania, I can hardly imagine

that it went undetected until 1928. However, I have not found an earlier name for it.

Y. afromontana achenes have a high epoxy acid content in the achene oil. In this regard, it is
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 83

very like species of section Stengelia. Morphologically, it is rather closely related to V. glabra,

although it is generally larger over all than that species.

Mature heads of V. afromontana plants are inconspicuous because the pappus is sordid and

blends v^fell vi^ith the surrounding shadows. Open-grown plants are shorter than those that grow

in competition with other vegetation. Flowers of V. afromontana are often clear, bright or light

blue. The plants are called machagene by the people near Meru, Kenya.

As with any study of this nature, this report ends up with several riddles. The most vexing of

these is the type specimen of V. swynnertonii S. Moore (J. Linn. Soc. Bot. 40:107. 1911) from Kew

which consists of a broken scape with a single head and several scraps in the packet. Moore must

have had a better specimen to write his rather full description. I have found no other collection to

match nor do I feel confident enough to accept Moore's species on the basis of the available specimen.

It may belong near V. subaphylla.

The following specimens cannot be comfortably placed with any of the species which I am

recognizing. H. Lynes D-16-Í belongs to the group with a rosette of leaves from a perennial root

crown. It is also near V. subaphylla, but I lack specimens with intermediate morphological

characteristics which would allow me to place the collection here. P. R. O. Bally 9203 and /. B.

Gillett mis from the same region of Ethiopia are the same species and certainly belong with the

upright-stemmed species of section Stengelia. The Bally collection has been identified as V. abyssinica which it isn't. They may represent a new species, but I haven't sufficient collections to

warrant a description in a group in which too many species have already been described.

Two collections from Somaliland, C. N. Collenette 330 and /. G. B. Newbould 1007, are certainly

a new species, if they are placed in section Stengelia. However, I feel that my knowledge of

neighboring sections is insufficient and they may well belong with species in another section with

which I am unfamiliar.

PN-207G

Figure 56.^A, Plants of V. superba are locally abundant near Abercorn, Zambia. B, The clear, bright blue of heads

in flower is very noticeable, but the flower color photographs pale mauve on color film. (Smith & Richards 4666).
84 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

PN-2077

Figure 57.—A, Mature heads of V. afromontana. (Smith & Njeroge 4640.) B, Open-grown plants in Kericho, Kenya.

(Smith & Mygogo 4570). C, Flowering plants of V. afromontana, well hidden among surrounding vegetation, near

Meru, Kenya. (Smith & Mygogo 4585.) D, Flowers of V. afromontana. (Smith and Mygogo 4570.)

In addition, I have many names for species described as belonging to section Stengelia for which

I have seen no authentic specimens. The descriptions, in spite of the thoroughness of some of those

by German authors, are inadequate to place the names in synonymy or in the ranks of recognized
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 85

species. The following species were placed in section Stengelia by the authors or the descriptions

or discussions lead me to believe that they may belong to the section.

F. anandrioides S. Moore. I have a specimen of Gossweiler No. 2132, the type collection, but it is

a depauperate plant, and I cannot confidently place it with any of the species previously discussed.

y. alho-violacea

Muschler

y. argutidens Chiov.

V. bequaertii De Wild.

V, bojeri Less.

F. brideliifolia O. Hoffm.

V. britteniana Hiern

V. bukamensis De Wild.

y. calvoana var

mesocephala Adams

V, descampsii De Wild.

y. drymaria Klatt

y. goetzei Muschler

y. graciliflora De Wild.

y. hiemiana S. Moore

y. hockii De Wild. &

Muschler

y. kapirensis De Wild.

y. kirungae R. E. Fries

y. lasiolepis 0. Hoffm.

*See note under y. sciaphilaf p. 59.

y. ledermannii Mattf.

*y. luembensis De Wild,

& Muschler

y. manikensis De Wild,

y. napus O. Hoffm.

y. neumanniana

O. Hoffm.

y. oehleri Muschler

y. otophora Mattf.

y. paludigena S. Moore

y. printzioides Muschler

y. punctulata De Wild.

y. pygmaea 0. Hoffm.

y. quangensis O. Hoffm.

y. retusa R. E. Fries

y. roseo-violacea De Wild.

y. rufuensis Muschler

y. ruwenzoriensis var.

glabra S. Moore

y. tolypophora Mattf.

y. towaensis De Wild,

y. ulugurensis 0. Hoffm.

y. verrucata Klatt

y. yatesii S. Moore

Literature cited

Bentham, G., and Hooker, J. D.

1873. GENERA PLANTARUM 2:227-231. London.

Bolkhovskikh, Z., Grif. V., et al.

1969. CHROMOSOME NUMBERS OF FLOWERING

PLANTS. Leningrad.

De Candolle, A. P.

1836. PRODROMUS SYSTEMATIS NATURALIS, ETC.

5:60. Paris.

Engler, A., and Prantl, K.

1889. DIE NATURLICHEN PFLANZENFAMILIEN.

4(5) 39 Lief.: 124-126. Leipzig.

Hochstetter, C. F.

1841. ERSTE LIEFERUNG DER VOM REISE VEREIN

AUSGEGEBENEN DURCH WILHELM SCHIMPER

GESAMMELTEN ABYSSINISCHEN PFLANZEN.

Flora 24: Intell. 1841. I No. 2:17.

Krewson, C. F., Ard, J. S., and RiemenSchneider, R. W.

1962. VERNONIA ANTHELMINTICA (1.) WILLD.

TRIVERNOLIN, 1, 3-DIVERNOLIN AND VERNOLIC (EPOXYOLEIC) ACID FROM THE SEED OIL.

J. Amer. Oil. Chem. Soc. 39:334-340.

Kupchan, M. S., Hemmingway, R. J., Werner, D., and

others.

1968. TUMOR INHIBITORS. XXXI. VERNOLEPIN, A

NOVEL ELEMANOLIDE DILACTONE TUMOR INHIBITOR FROM VERNONIA HYMENOLEPIS.

J. Amer. Chem. Soc. 90:3596-3597.

Lanjouw, J. Mamay, S. H., McVaugh, R, and others.

1966. INTERNATIONAL CODE OP BOTANICAL NOMENCLATURE. Utrecht.

and Stafleu, F. A.

1964. INDEX HERBARIORUM. REXÎ. VEG. 31,

Lessing, C. F.

1831. DE SYNTHEREIS DISSBRTATIO QUARTA.

neaea 4:624. et seq.

LinOliver, D.

1877. FLORA OF TROPICAL AFRICA 3:266-297.

London.

Peters, W. C. H.

1862-64. NATURWISSENSCHAFTLICHE REISE NACH

MOSSAMBIQUE AUF BEFEHL SEINER MAJESTÄT DES KÖNIGS FRIEDRICH WILHELM

IV. PART 6. BOTANIK. Berlin.

Salt, H.

1814. A VOYAGE TO ABYSSINIA ETC. London.

Sequeira, L. Hemingway, R. J., and Kupchan, S. M.

1968. VERNOLEPIN: A NEW, REVERSIBLE PLANT

GROWTH INHIBITOR. Science 161:789-790.

Smith, C. E., Jr.

1969. POLLEN CHARACTERISTICS OF AFRICAN SPECIES OF VERNONIA. J. Arn. Arb. 50:469-

477.

Stafleu, F. A.

1967. TAxoNOMrc LITERATURE. Utrecht.

Steudel, E. T.

1841. NOMENCLÁTOR BOTANICUS. (Ed. 2), p. 638,

752-755. Stuttgart & Tubingen.

Walpers, W. G.

1843. REPERTORIUM BOTANICES SYSTEMATICAE.

2:538-541, 945-948. Leipzig.

Wildeman, E. De

1915. DECADES NOVARUM SPECIERUM FLORAE CONGOLENSis III. Bull. Jard. Bot. Etat. 596.
86 AGRICULTURE HANDBOOK NO. 396, U.S. DEPT. OF AGRICULTURE

Appendix

List of vernacular names for stengelioid Vernonia species

Vernacular Tribe or Country and

Name languxige locality Reported by

Vernonia adoensis

Akavumavuma Kinyaruanda Ruanda-Urundi G. Troupin 4915

Akco Hlur Congo, Nioka

Dju-dju Bilendu Congo, Nioka

Ilumba lya shimba Kisukuma Tanzania R. Tanner 1217

Liuluya Mwera Tanzania Semsei 2148

Msohoso Bende Tanzania Semsei 2570

Tsoyo Chewa Zambia F. White 2446

Ukafuma Kirundi Congo Michel & Reed 315

Umuvumo Kinyaruanda Ruanda-Urundi G. Troupin 4915

V. afromontana

Machegene Kikuyu Kenya, Meru C. E. Smith 4585

Morerangamba Kikuyu Kenya, Mt. Kenya CE. Smith4644

V. bracteosa

Mtokutu Kihehe Tanzania, Iringa Carmichael 393

V. filipéndula

Kasingele Angola, Libólo M. Dawe 317

F. guineensis

Akaniamymya Kirundi Congo, Ruyigi

Akanianina Kirundi Ruanda-Urundi Evrard 1769

Funyembe Ma. (?) Sierra Leone

Kanyaminya Kirundi Ruanda-Urundi Michel & Reed 692

Koberafaira Sierra Leone G. F. S. Elliot 4842

Kumbiarona T. (?) Sierra Leone N. W. Thomas 760

Lwishote Mashi Congo, Kivu J. Petit 70

Mombakalu Azomde Congo, Bas Ebele A. Dewulf 184

Nambakalu Zande Congo, Tukpwo Lecompte 26

Nwagoe Lo. (?) Sierra Leone

Teti Congo, Haut Uele Nannan 521

F. hymenolepis

Fulucha Central Africa Johnson s. n.

Ivumo Kinyaruanda Ruanda-Urundi G. Troupin II126

Uvumovumo Kinyaruanda Ruanda-Urundi G. Troupin II126

F. lasiopus

Gihihahika Kihunde Congo, Rutshuru Spitaels 129

Imbassa Kichagga Tanzania, Moshi G. B. Wallace 948

Ivumavuma Congo, Tshumba Tramasure 620

Ivumera Kinyaruanda Congo, Rutshuru Spitaels 129

Ivumo Congo, Kitana de Wulf 82

Ivumu Tshiragaga Congo ?, Mulungu Hendryx 33

Kilugutu Tanzania, Morogoro E.R.Bruce 43

Mbaaza Kichag. ? Tanzania, Kilimanjaro P. J. Greenway 3019
OBSERVATIONS ON STENGELIOID SPECIES OF VERNONIA 87

Vernacular Tribe or

Name language

V, lasiopus—Continued

Mbasa

Mbasa Kimeru

Mbasa Chaga

Mnyatira Sukuna

Mshasha Shambea

Mucatha Kikuyu

Muchatha Kikuyu

Muhasha Kish

Mululusa Kihehe

Musambanyumu Kinande

Muvatha Kamba

Muvuma

Mwitanyonyi Haya

Npalagasha Kisambaa

Omujuma

Pasapasa Kisafwa

Rinkalikali Arusha

Sondja sondja

Umukurasi Kitundi

Umundu Limatinzo ?

Umuturumbuka Kishubi

Umuvumavuma

F. nigritana

Kundidi tuna Yalunka

Nonomorandakofin Mandingo

V. oxyura

Fusa Chinyanja

Futsa Nyanja

Mdulua

Msangusangu Kigao

Msangusangu Ayaua

Mteli Nguru

Mtugutu Nguru

Mululusia Kibembo

Ugua Msugumbili

V, pumita

Muhindabij umiro Lunyoro

Country and

locality

Tanzania, Tengeru

Tanzania, Arusha

Tanzania, Kilimanjaro

Tanzania, Bomboju

Tanzania, Bomboju

Kenya, Nairobi

Kenya, Nairobi

Tanzania, Usambaras

Tanzania, Mufindi

Congo, Beni

Kenya, Katumani

Ruanda-Urundi

Tanzania, Arusha

Tanzania, Lushoto

Uganda, Kashwekana

Tanzania, Mbeya

Tanzania, Arusha

Mozambique, Malulo

Ruanda-Urundi

Tanzania, Arusha

Tanzania, Nyakisasa

Ruanda-Urundi

Sierra Leone

Sierra Leone

Malawi, Likubula Gorge

Mozambique, Massaneulo

Tanzania, Makuyuni

Tanzania, Tunduru

Mozambique, Massaneulo

Tanzania, Mandeni

Tanzania, Mtibwa

Congo, Shiebale

Tanzania, Pangani

Uganda, Bunyoro

Reported by

Carmichael 511

R. L. Williams 53

P. J. Greenway 3019

Bally 200

Bally 200

E. R. Napier 295

C. W. Leakey 470

Koritschoner 743

C. G. McGregor 62

Gille 110

Thomas 723

0. Elskens 92

Bintabara 3

Mahwilo 9

A. S. Thomas 4204

G. Myembe 39

L. Matalu 3139

P. Pedrogat 4056

Carmichael 607

R. Tanner 5749

A. Becquet 379

F, C. Deighton 4470

F, C. Deighton 4470

Brass

Sousa 1397

Koritschoner 1286

Barter H131/55

Sousa 1397

S. R. Semsei 570

Semsei 5907

W. Robyns 2063

R. Tanner 3190

it U. S. GOVERNMENT PRINTING OFFICE-. 1971 O 4Ï3-934