~ COMPOSITAE &
* NEWSLETTER

Number 34 21 December 1999

Scientific Editor: BERT. NoRDENSTAM

Technical Editor: GUNNEL WirENIUS NOHLIN

Published -and distributed by The Swedish Museum of Natural History,
Department of Phanerogamic Botany,

P.O. Box 50007, SE-104 05 Stockholm, Sweden

(Director: Professor Berti. NoRDENSTAM) ISSN 0284-8422

CONTENTS

J. F. Pruski: Novelties in Angelphytum and Oyedaea (Compositae:
Heliantheae: Ecliptinae) from South America

M. S. AYopELE : Studies on the reproductive biology of Vernonia SCHREB.
(Asteraceae). V. Fruit production and fruit-set strategies among growth forms

M. S. AYopeLe: Studies on the reproductive biology of Vernonia ScuREB.
(Asteraceae). VI. Seed germination strategies among growth forms

B. Norpenstam: New combinations in Monticalia (Compositae-Senecioneae)
from Colombia

M. Rar & D. Acuarya: Screening of some Asteraceous plants for antimycotic
activity

M. Rar & D. Acuarya: Diversity of arbuscular mycorrhizae in naturally
growing plants of family Asteraceae in India

H. BELTRAN: New combinations in Dendrophorbium and Pentacalia
(Senecioneae - Asteraceae) from Peru

A. N. Sennikov & I. D. ILLarionova: Reclassification of Prenanthes pendula
(Asteraceae: Lactuceae)

J. Kayope: Phytosociological investigation of Compositae weeds in abandoned

farmlands in Ekiti State, Nigeria

New taxa and combinations published in this issue

62
69

-

Comp. Newsl. 34, 1999 1

Novelties in Angelphytum and Oyedaea
(Compositae: Heliantheae: Ecliptinae)
from South America

JOHN F. Prusk1
United States National Herbarium
Department of Botany, MRC-166
Smithsonian Institution
Washington, DC. 20560-0166, U.S.A.

Abstract

Oyedaea cuatrecasasii from Colombia and Oyedaea neei from Bolivia are described
as new and their relationships are discussed. The new combinations Angelphytum
apense and Angelphytum herzogii are made for species formerly placed in Zexmenia.

Introduction

Ecliptinae LEssinG, a subtribe of Compositae: Heliantheae has been delimited by Rosin-
SON (1981), although most of its genera were more recently treated under the name
Verbesininae by Karis & Rypinc (1994), who did not subtribally place Eclipta. Gen-
eric additions and adjustments of Ropinson (1984a, 1984b, 1992, and literature cited
therein), STROTHER (1991 and literature cited therein), PRUsKI (1996 and literature cited
therein), and most of the suggestions in PANgRO et al. (1999) are accepted as emendations
to the Ecliptinae as defined by RoBinson (1981). The subtribe Verbesininae treated as
asynonym of the Ecliptinae by Rosinson (1981), STROTHER (1991), and Prusk1 (1996),
was recently resurrected by PANgRO et al. (1999) and includes only Verbesina and
three small Mexican and Central American genera. The following novelties in genera
with winged fruits are the result of routine identifications of species mostly from Andean
South America.

Angelphytum

Angelphytum Barroso was originally described for a disciform species, but the generic
concept has been expanded by Rosinson (1984b) to include several species with winged
fruits and fertile ray florets from South America once referred to Zexmenia LA LLAvE.
Angelphytum differs from Zexmenia most notably (RoBINSON 1984b) by being largely

2 Comp. Newsl. 34, 1999

xylopodial herbs, less commonly shrubs, by abaxially glanduliferous, strongly recurved
(vs. non-glandular, gradually curving) style branches, and by non-constricted (vs.
apically constricted) cypselas. Dimerostemma Cass. in broad disk corollas, winged
fruits, and leafy involucres is similar to Angelphytum, but differs in sterile ray florets.
The two radiate species transferred here fall within Angelphytum as delimited by Rosin-
SON (1984b). The generic limits of Angelphytum are currently being studied by MARTA
D. Morass (UEC), a student of Dra. GRAZIELA Barroso. As presently circumscribed,
Zexmenia contains only two species, both Central American (STROTHER 1991). The
two new combinations proposed here raises to 16, the number of species in
Angelphytum.

Angelphytum apense (Cuopat) Pruski, comb. nov. Basionym: Aspilia apensis CHODAT,
Bull. Herb. Boiss., Ser. 2, 3: 721. 1903. Zexmenia apensis (CHODAT) HASSLER, Rep-
ert. Spec. Nov. Regni Veg. 14: 177. 1915. Syntypes: PARAGUAY: Prope Valenzuela
(protologue), in regione cursus superioris fluminus Y-acé (label), Feb 1900, HassLer
7096 (syntype: G n.v., photograph: US!); In regione cursus superioris fluminus
Apa, Nov 1901, HassLer 7731 (syntype: G n.v., photograph: US!).

This species is a shrubby herb with fertile ray florets and occurs in Paraguay and
adjacent Mato Grosso do Sul, Brazil. It and the one following are similar by opposite,
broad, long-petiolate leaves. Angelphytum apensis differs most noticeably from the
closely related A. herzogii by serrulate to weakly serrate (vs. serrate to dentate) leaves,
by cypselas with poorly (vs. well) developed pappus awns, and by smaller capitula
commonly with shorter leafy outer phyllaries. Lectotypification is differed until both
syntype collections are examined.

Angelphytum herzogii (HASSLER) PRusKI, comb. nov. Basionym: Zexmenia herzogii
HassLer, Repert. Spec. Nov. Regni Veg. 7: 357. 1909. Type: BOLIVIA. Santa
Cruz: Halbstrauch im Bergwald von Samaipata (Ostkordillere), ca. 1700 m, Dec
1907, Herzoc 704 (G n.v., HBG?).

This Bolivian shrub with fertile ray florets was known only from the type, until plants
matching the protologue were recently rediscovered by MICHAEL NEE (NY) in several
localities, these and the type all from Depto. Santa Cruz, Bolivia. Also other material
from Depto. Santa Cruz, Bolivia under the name Zexmenia apensis is determined here
as Angelphytum herzogii. The type of Zexmenia herzogii was not seen, but a photocopy
and a drawing of the flowers by GisELA SANCHO (LP) of the type number in G on loan
to ALcIDES SAENZ (LP) has been recently sent to me. Examination of this material and
other collections from Bolivia show this taxon to be a species of Angelphytum. The
specimen of Zexmenia herzogii in G is fragmentary and no specimens of this collec-
tion were found in Z. Thus, it seems that a full sheet (the holotype?) is elsewhere,
perhaps in HBG or as a second sheet in G.

Comp. Newsl. 34, 1999 3

Oyedaea

Oyedaea DC., revised by BLakE (1921) and characterized in part by winged fruits and
sterile ray florets, is largely Andean, although at present not known from Ecuador
(Pruski 1997, H. Rosinson, pers. comm.). One species (O. verbesinoides DC., the
type) extends from the Andes eastward into the Venezuelan coastal ranges and north
into Central America, and three others are endemic to the Guayana Highland (Prusk1
1996, 1997). The other Central American species of Oyedaea (O. steyermarkii S.F.
BLAKE, O. mexicana RzepowskI, O. ovalifolia A. Gray, and O. lundellii H. Ros.,) have
been transferred to Lundellianthus H. Ros. (RoBINSON 1979, STROTHER 1989), Oto-
pappus BENTH. (RoBINSON 1979), Perymeniopsis H. Ros. (ROBINSON 1978, STROTHER
1991), and Zyzyxia STROTHER (STROTHER 1991), respectively. The Brazilian species
once placed in Oyedaea have been removed to Dimerostemma Cass. by RoBINSON
(1984a), the latter differing by its leafy involucre and broader disk corollas. Oyedaea
was recently treated by Pruski (1996, 1997) as containing about 16 species (including
O. neei), this number plus O. cuatrecasasii now increases to 17, the number of species
in the genus.

Oyedaea cuatrecasasii Pruski, sp. nov. Type: COLOMBIA. Meta: La Serrania entre
los rfos Ariari y Meta, Los Churrubayes, 300 m, 24 Nov 1939, J. CuaTRECASAS
7853 (holotype: US-1774514!; isotypes: BC n.v., F n.v.).

Oyedaeae verbesinoideae affinis, sed ab ae lamina foliorum basi rotunda vel acuta,
capitulisque paucis diversa.

Subshrubs to shrubs 0.8—3 m tall; stems subterete, strigose, the internodes 2-11 cm
long, commonly slightly shorter than the subtending leaves. Leaves simple, opposite;
petioles 2-8 mm long, strigose; blades elliptic-lanceolate, (2.5) 3.5-9.5 x 14.5 cm,
chartaceous, eglandular, weakly to moderately triplinerved from well above base,
venation strongly reticulate, basally rounded to acute, apically acute to sometimes
narrowly so, the margins subentire to serrulate, adaxially green, scabrous, abaxially
paler green, strigose, especially on larger veins. Capitulescences corymbiform to rarely
monocephalous, somewhat compact and not loose and spreading, terminal or sometimes
also axillary from uppermost leafy node, with 1-5 capitula held above the subtending
leaves, less commonly overtopped by uppermost leaves; peduncles 0.54.5 (6) cm
long, densely hispid to strigose. Capitula radiate, 45—119-flowered; involucres
campanulate, 8-11 x 8-12 mm; phyllaries 16-19, in 2-3 series, subequal, 8-11 x 2—
3.5 mm, rigid, the outer series herbaceous, lanceolate to oblanceolate, strigose, the
apex acute to obtuse, the inner series lanceolate to elliptic-lanceolate, strigose or only
so apically, scarious at base, the apex acute, herbaceous; receptacles flat, 3-4 mm
diam., paleate, the pales conduplicate, lanceolate, to ca. 7.8 x 1 mm, firm, scarious

4 Comp. Newsl. 34, 1999

with a brownish-greenish strigose central nerve extending to near tip, the apex acute.
Ray florets 10-12, sterile, lacking styles; corollas yellow, ca. 10-19 x 3 mm, the tube
1.5-2 mm long, glabrous, the limb 8.5-17 x 3-5.6 mm, 8-13-veined with 2 veins
larger than the others, glabrous or weakly abaxially puberulent, shortly 2-lobed at
apex, apical lobes ca. 0.6 mm long. Disk florets 35-107, bisexual; corollas
actinomorphic, yellow, tubular-funnelform, 5.3—6.6 mm long, the tube 1.2—2 mm long,
glabrous, the throat 3.3-3.8 mm long, glabrous, the lobes ca. 0.8 mm long, triangular,
weakly puberulent; anthers included, 2.5—-3 mm long, the thecae black, the apical
appendages deltoid, becoming cream-colored, glanduliferous, the connectives black,
the filaments ca. 1.5 mm long; style branches ca. 1.5 mm long, papillose, slightly
spreading to recurved, with paired stigmatic lines, the apices sterile, lanceolate. Ray
ovaries sterile, 2—2.6 mm long, obconic, 1—3-awned, the awns (0.8) 2-3 mm long,
subequal to unequal. Disk cypselas (immature, thus wings not formed) compressed,
ca. 3 mm, the body brown to black, weakly puberulent, the margins ciliate; pappus
stoutly 2-awned, the awns scabridulous, subequal to unequal, 2.6—-4 mm long, these
about as tall as the involucre and pales, the squamellae between the awns toca.0.5 mm
long.

Paratypes: COLOMBIA. Meta: Mun. La Macarena, Reserva de La Macarena, 20 km
NO en la via a Conejos, 450 m, 2°15’N, 73°45’ W, 7 Aug 1988, R. CALLEIAs &
O. Marutanpa 6966 (US!); Mun. La Macarena, 5 km O de La Macarena, via a Cone-
jos, 410 m, 2°16’N, 73°11’ W, 7 Aug 1988, R. CALLEyAs & O. MARULANDA 7052 (US!);
San Juan de Arama, valle a la izquierda del rio Giiejar frente a la parte norte de la
Sierra de La Macarena, 500 m, 23 Aug 1950, J.M. Iproso 467 (US!); Reserva Biolégica
de la Macarena, rio Guayabaro 4 km al suroeste del Refugio, 350 m, 26-28 Oct 1976,
R. Starr 74 (US!); 35 km al E de San Martin, en ruta hacia, 16 Jun 1989, F.0. ZuLoAGA
4009 (US!).

Distribution and ecology: This species is known only from savannas and gallery
forests in the region of Sierra de la Macarena, Meta, Colombia. It occurs from 300-
500 m in elevation and has been collected in flower in June, August, October and
November.

This species is named for its discoverer, the late (RoBINSON et al. 1996) Dr. José
CUuATRECASAS. Don José was the foremost Colombian taxonomist of modern times,
and he specialized in Compositae. It is thus fitting and proper that I dedicate this new
species to him.

Oyedaea cuatrecasasii is very similar in leaf shape and size to extra~Guayana Venezu-
elan O. maculata S.F. BLake and O. obovata S.F. BLAKE, both of which differ from the
new species, however, by pinnately veined leaves. In its generally triplinerved leaves

Comp. Newsl. 34, 1999 5

the new species is similar to the northern Andean and Guayanan group including
O. huilensis Cuatr., O. reticulata S.F. BLAKE, O. tepuiana (V.M. BapILLo) PRusKI,
O. verbesinoides DC., and O. wurdackii Prusxt. The new species generally has longer
pedunculate capitula and less pubescent ray limbs than the other species. Additionally,
Oyedaea cuatrecasasii differs from O. verbesinoides by acute to rounded (vs. attenuate)
leaf bases and by few (vs. several) capitula per branch. By leaf shape Colombian
O. huilensis and O. reticulata are similar to O. verbesinoides but they differ from it
and the new species by leaves abaxially harshly pilose-hispid and by unequal phyllaries.
Additionally, O. huilensis and O. reticulata differ from the new species by larger leaves
and by a larger capitulescence with many capitula. Oyedaea cuatrecasasii differs from
O. wurdackii by yellow (vs. white) ray corollas and from O. tepuiana by larger and
longer pedunculate capitula. Oyedaea scaberrima (BENTH.) S. F. BLAKE from the Gu-
ayana Highland differs from the new species by unequal phyllaries and pinnately
veined leaves.

Oyedaea neei PRuskI, sp. nov. Type: BOLIVIA. Santa Cruz: Prov. Florida, 4 km N of
center of Samaipata, 18°08’S, 63°52’ W, 2000—2100 m, 31 Dec 1992, M. Nee & I.
Varaas C. 43442 (holotype: US-3362069!; isotypes: LPB n.v., NY!, USZ n.v.).
Fiz. 1.

Oyedaeae bolivianae affinis, sed ab ea foliis minoribus ellipticis acutis, capitulis paucis,
pedunculisque longioribus diversa.

Low shrubs; stems subterete or subhexagonal, strigose, the internodes 0.5—3.5 cm
long, usually shorter than the subtending leaves. Leaves simple, opposite; petioles 2—
3 mm long, strigose, stout and much broadened at base, those of the same node often
connected by a narrow rim; blades elliptic,1-2.5 x 0.4-1.3 cm, chartaceous,
eglanduliferous, triplinerved from well above base, basally cuneate to attenuate, apically
acute, the margins serrulate to subentire, adaxially green, scabrous, abaxially pale
green, weakly strigose especially on larger veins, nearly glabrescent with age.
Capitulescence monocephalous or cymose, open, terminal, of 1 or less commonly 2
capitula held above the subtending leaves; peduncles 1.5—3.5 cm long, strigose. Capitula
radiate, ca. 30-flowered; involucre campanulate to hemispherical, 6-7.5 x 9-13 mm;
phyllaries ca. 17, in 2-3 series, subequal or weakly graduate, lanceolate to oblanceolate,
6-7.5 x 1.5-3 mm, rigid, broad and scarious at base, the apex broadly acute to obtuse,
herbaceous, strigose; receptacles flat, ca. 5 mm diam., paleate, the pales conduplicate,
often keeled near the apex, ca. 8.5 mm long, firm, scarious throughout, attenuate. Ray
florets ca. 11-13, sterile, lacking styles; corollas yellow, the tube ca. 1.6 mm long,
glabrous, the limb ca. 9.5 x 34 mm, minutely 2- or 3-lobed at apices, ca. 17-veined
with ca. 4 veins larger than the others, smaller veins often anastomosing, abaxially
glabrous or weakly puberulent to weakly pilose near tube. Disk florets ca. 15-19,

6 Comp. Newsl. 34, 1999

bisexual; corollas actinomorphic, yellow, tubular-funnelform, 44.5 mm_ long, the
tube ca. 1 mm long, glabrous, the throat 2.5—3 mm long, glabrous, the lobes ca. 0.5
mm long, triangular, puberulent-papillose to densely so; anthers slightly to much-
exserted, ca. 2.5 mm long, the thecae black, the apical appendages ovate, tan, eglandular;
style branches ca. 1 mm long, densely papillose or weakly so, laxly recurved, with
paired stigmatic lines, the apices acuminate to attenuate. Ray ovaries sterile, ca. 2.5
mm long, obconic, 3-awned, the awns 1.5—2.5 mm long, subequal to unequal. Disk
cypselas compressed, slightly winged, ca.4 1.5 mm, the body black, puberulent, the
wings ciliate, cream colonred, brown-spotted; pappus stoutly 2-awned, the awns weakly
scabridulous, subequal to unequal, 2—3 mm long, these exserted from the involucre
and often reaching to the top of the pales, the squamellae between the awns 7-10, to
ca. 0.8 mm long.

Distribution and ecology: This species is known only from the type collection, from
grazed ridge tops in the area near Samaipata in Depto. Santa Cruz, Bolivia. It occurs
from 2000-2100 m and was collected in flower in December.

I am happy to name this new Oyedaea for the collector of the type material, my friend
Dr. MicHAgL Nee of the New York Botanical Garden. Micuaev’s fine collections from
Depto. Santa Cruz, Bolivia also resulted in the rediscovery of Zexmenia herzogii,
transferred here to Angelphytum.

By its triplinerved leaves Bolivian O. neei is most closely related to Oyedaea boliviana
Brirron. However, the new species clearly differs from shrubby arborescent O. boliviana
by low stature, by very small elliptic (vs. lanceolate) acute (vs. acuminate) leaves, by
fewer or solitary capitula with shorter ray limbs, and by longer peduncles. Bolivian
O. bullata Koster is another species with triplinerved leaves, but differs from the two
former species by its involucre. The other Andean species of Oyedaea from Peru and
Bolivia have pinnately veined leaves and are less closely related.

Acknowledgements

I thank PEprRo ACEVEDO, RUPERT BARNEBY, MICHAEL NEE, DAN NICOLSON, and HAROLD
Rosinson for reviewing this paper; Rupert BARNEBY for correcting the Latin diagno-
ses; GISELA SANCHO for her drawing of the type of Zexmenia herzogii; GIsELA SANCHO
and AtcibEs SAENz for sending me a xerox of the type collection of Zexmenia herzogii;
and Don Hur.Bert, VicTOR KRANTZ, and JOHN STEINER for the photograph of the type
of Oyedaea neei.

Comp. Newsl. 34, 1999

ae
y, q

a A %

nA

xv

———

“ly )
z=
otanica
f BOLIVIA
l
; He lot ype
per Osedaca ncei Pruski
det. Pprken Prussk:, 06 -IE ~/t9-
mia Cruz, Prov lorida, 4 km of
os ita. Grazed ridgetop with
UNITED: STATES et of forest in ravines. Aine:
Vata, nstroemia gsymmetrica, Eacaticonis
culnta. i8°O8'S, G3I°S2"W. alt, 2000-2100 m
acone
3362069 ery low "shrublets". Dise and rays bright
Fetiow, +
uh: M. Bee &°0 Vargas C. $3442

© a *
: 31 Bec. 1992

NATIONAL HERBARIUM

Fig. 1. Photograph of the holotype of Oyedaea neei (NEE & VarGAS 43442, US).

8 Comp. Newsl. 34, 1999

References

BLAKE, S. F. 1921. Revision of the genus Oyedaea. Contr. U.S. Natl. Herb. 20: 411-
422.

Karis, P. O. & O. Rypinc 1994. Chapter 22, Tribe Heliantheae. Jn: K. BREMER
Asteraceae: Cladistics & Classification, pp. 559-624. Timber Press, Portland,
Oregon.

PaneERO, J. L., JANSEN, R. K. & J. A. CLEVINGER 1999. Phylogenetic relationships of
subtribe Ecliptinae (Asteraceae: Heliantheae) based on chloroplast DNA restriction
site data. Amer. J. Bot. 86: 413-427.

PruskI, J. F. 1996. Compositae of the Guayana Highland-XI. Tuberculocarpus gen.
nov. and some other Ecliptinae (Heliantheae). Novon 6: 404-418.

PruskI, J. F. 1997. Asteraceae. In: J. A. STEYERMARK et al. (eds.), Flora of the Venezu-
elan Guayana, Volume 3. Araliaceae—Cactaceae, pp. 177—393. Missouri Botanical
Garden, St. Louis.

Rosinson, H. 1978. Studies in the Heliantheae (Asteraceae). XIII. A new genus,
Perymeniopsis. Phytologia 40: 495-496.

Rosinson, H. 1979. Notes on Oyedaea in Central America (Heliantheae: Asteraceae).
Wrightia 6: 43-45, pl. 82.

Rosinson, H. 1981. A revision of the tribal and subtribal limits of the Heliantheae
(Asteraceae). Smithsonian Contr. Bot. 51: i-iv, 1-102.

Rosinson, H. 1984a. Studies in the Heliantheae (Asteraceae). XX XI. Additions to the
genus Dimerostemma. Proc. Biol. Soc. Wash. 97: 618-626.

Rosinson, H. 1984b. Studies in the Heliantheae (Asteraceae). XXXIV. Redelimita-
tion of the genus Angelphytum. Proc. Biol. Soc. Wash. 97: 961-969.

Rosinson, H. 1992. New combinations in Elaphandra StrotuHer (Ecliptinae-
Heliantheae-Asteraceae). Phytologia 72: 144-151.

Rosinson, H., Funk, V. A., Pruski, J. F. & R. M. Kine 1996. José CuATRECASAS
Arumi (1903-1996). Comp. Newsl. 29: 1-30.

STROTHER, J. L. 1989. Expansion of Lundellianthus (Compositae: Heliantheae). Syst.
Bot. 14: 544-548.

STROTHER, J. L. 1991. Taxonomy of Complaya, Elaphandra, logeton, Jefea,
Wamalchitamia, Wedelia, Zexmenia, and Zyzyxia (Compositae-Heliantheae-
Ecliptinae). Syst. Bot. Monogr. 33: 1-111.

Comp. Newsl. 34, 1999 9

Studies on the reproductive biology of
Vernonia SCHREB. (Asteraceae)

V. Fruit production and fruit-set strategies
among growth forms

M.S. AYODELE
Biological Sciences Department
University of Agriculture
P.M.B. 2240
Abeokuta, Nigeria
e-mail: segmuy @ unaab.edu.ng

Abstract

Fruit production performance was found associated with a number of vegetative and
reproductive characteristics of Vernonia plants. A prolific production of small size
achenes was recorded on individual plant basis. This cuts across different growth forms,
but the values were higher in shrubs than in herbaceous species.

The incidence of good quality fruits (high fruit-set % per capitulum) was recorded
consistently among the herbs. Habitats with moisture stress encouraged the production
of small size fruits which were characteristic of the herbs. These traits were noted as
those of weedy plant colonizers.

Introduction

Fruit/Seed production is important, because according to JANZEN (1969), it provides
the plant with the prominent functions of escape, genetic recombination and dispersal
for colonization. Each seed is an independent colonizer and seeds are adapted in many
ways for dispersal (INGROUILLE 1992).

The condition and exposure of a plant during the flowering and fruiting phase determines
much the quality of fruits produced and the seeds set. In fact, Burtt (1975) emphasized
that the demands of the flowering and fruiting phase of a plant interact. AYODELE
(1994b) highlighted the protectoral role of some floral parts in the flowering and fruiting
phases of the plant life.

10 Comp. Newsl. 34, 1999

The call for more studies, earlier solicited for by Burtt (1975) on the co-evolution of
these two phases, has remained very relevant even now for a deeper understanding of
the reproductive strategies in the array of plants dotting our terrain.

The genus Vernonia is widely reported for its versatility with respect to its morphological
characteristics, ecological distribution and extensive speciation (Faust 1972, JoNEs
1979, and AYopELe 1995). This recommends the genus for a comparative observation
of the trends in fruit production and fruit-set quality among its species and within the
different growth forms.

AYODELE (1997) identified five of such growth forms in Nigeria, among the 15 species
collected from different ecological zones. These were: - Erect tall woody tree, Erect
woody shrub, Straggling woody shrub, Erect annual herb and Erect perennial herb.

The objective of this paper is to put on record the fruit production potentials of some
investigated species of Vernonia in Nigeria. The quality of fruits produced by the
different species is also being elucidated with a view to identifying any plant survival
strategies associated with fruit production and quality of fruits in Vernonia.

Materials and Methods

Estimation of Fruit Production Performance in Vernonia Plants

An overall estimate of fruit production per plant for each of the species investigated,
was made by collecting data on:

a. the number of branches per plant

b. the number of terminal capitulum-clusters per branch (ex canopy view) or the
number of inflorescences per branch, for some species

c. the number of capitula per terminal cluster [or the number of ‘spikes’ (AYODELE
1994a) per inflorescence branch for some species]

d. the number of set (filled) fruits per capitulum.

The data were collected from plants with luxurious growth and deliberate pruning or

accidental breakage of any of the branches had not tampered with. Calculated values

for fruit production indices in species of Vernonia were obtained from the multiplication
of the values for a-d above 1.e. (a)x(b)x(c)x(d).

Comp. Newsl. 34, 1999 11

Assessment of Fruit Quality in Species of Vernonia

‘Seed-lots’ in batches or in single heads (depending on the mode of harvest) were
inspected for mature fruits filled with embryo. Vernonia fruits (achenes) were classified
into two groups for the purpose of this study namely: -the large (2.5—5.5 mm length)
and small (0.5—2.5 mm) fruit types (AYODELE 1987).

The large fruit type was assessed by visual/physical observation of fruit plumpness.
Each fruit was held between the thumb and the first finger, or by means of a pair of
forceps and pressed to fell whether fruits were filled or empty. The smaller fruit type
was observed in batches under a dissecting microscope by means of a pair of forceps.

Filled and empty/shriveled fruits were separated and counted by means of a tally
counter.

Analysis of Data

The mean of the Fruit Production Potential Index obtained for each plant among the
species was estimated. Any significant differences among the species in fruit production
were analyzed by Duncan Multiple Range Test. The fruit set values for each of the
species were obtained as the percentage of all the fruits examined which were filled
with embryos. Fruits observed were obtained from at least 25 randomly selected ripe
capitula per species.

Observations and Results

Achenes Production Potentials in Some Species of Vernonia

The calculated values of achenes produced on individual plant basis for each of the
species investigated, were derived from a combination of floral attributes and vegetative
traits (Table 2) associated with flower production in the plants. The values can be
separated into four distinct but overlapping groups (Table 1) on the basis of the
significant differences obtained (Table 3).

While group A was clearly distinct from the other three groups, the difference between
group B and C, group B and D, and group C and D, were not statistically significant
(P=0.0% level). The shrubby and arborescent species were clearly the most prolific
fruit (achenes) producers on individual plant basis (Table 1). The herbaceous species
were relatively poor in achene production. They were mainly the constituents of group
D (i.e. the fourth group) with the lower range of achenes production index values.

Comp. Newsl. 34, 1999

Quality Evaluation of Achenes Produced by Some Species of Vernonia

Achenes that contained embryos were considered “‘filled” or “set” (i.e. of good quality).
These were hard to touch and plump. The bad achenes were shriveled and chaffy and
contained no embryos when dissected. Many of the herbaceous species were observed
to be among the species producing good quality (i.e. filled) achenes. The special ob-
servation recorded on the herb V. galamensis was that the incidence of well-set achenes
was higher in plants growing under water stress condition.

The general trend on achene quality was that species producing the small size fruits
recorded a higher incidence of set fruits. This observation however, cuts across the
different growth forms especial as in V. biafrae, a shrub, and V. migeodi, an herb
(Table1) .

The values for V, colorata, V. nestor, V. purpurea and V. glaberrima were discountenanced
because their achene production rate could not be monitored continuously. These spe-
cies were difficult to raise in the garden and were mostly restricted to their habitats.
The results obtained from the field and the garden for the other species were however,
observed to be compatible.

Comp. Newsl. 34, 1999 13

Table 1. Mean Fruit Production Potential Index & % Fruit-Set in Vernonia

_ % Fruit-Set
() No. Growth Form
| Examined | 7 Fruit-Set

100 Straggling shrub

79.6 Erect Shrub

: am V. conferta

Erect Shrub
/Arborescent+

Arborescent

42.0 Arborescent

oo V. perrottetii 1,400 97.1 Erect Herb
oS Le V, tenoreana — 4,050| 92.9 |Erect Shrub
- ae V. migeodi Erect Herb

: J vcinerea | : - : : . eae
: (D) |v gclamensis | 3601) 2870 67.1 | Erect Herb
ee +1 fo on : =| Herb
= =a =
57 Rares Erect Shrub

* Duncan Multiple Range Test
+ Depending on age of plant & level of growth disturbance

Comp. Newsl. 34, 1999

Table 2. Mean Values of Vegetative/Reproductive Attributes Associated with Fruit
Production in Vernonia

Species

V. biafrae

V. stenostegia

Mean # Stem
branches/plant
—-

Mean # Terminal|Mean # Capitula/|Mean # Fruit-set
capitula clusters | terminal cluster | per capitulum

4.2800
9.4000,

V. colorata

V. amygdalina

ale

| V. conferta

| V. migeodi

V. cinerea
EEE

| V. galamensis

aa |

L—
V. ambigua

=!

5.1500

| V. perrottetii
| V. tenoreana 4.4000 6.0000 1.360} 128.0 ona

4.5600

4.4400
5.0000
5.5200

468.760 34.0 (34.9)*

5.200

14.1200 1.520 35.0 (38.0)

4.4000 7.240 29.0 (29.4)

5.5200 7.400 20.0 (21.8)

6.4000) 1.880 55.0 (81.0)
4.5600 2.240 38.0 (39.2)

2.0800 1.7200 2.720 42 (146.2)

-

V. kotschyana
Seite

7.3200} 5.520} - (17.9)
3.3600; 1.800 - (98.2)

V. nestor 1.0000
V. purpurea 1.0000
8.0400

V. glaberrima
—

6.4000 3.920 - (4.1)

*Numbers in parenthesis are values for # Florets/Capitula

Comp. Newsl. 34, 1999 15

Table 3. Analysis of Variance (Anova) in Fruit Production/Species of Vernonia

1925441323703.59

Estimated | Model
Fruit

Production/ | Error 781071404367.60
Species oeniatan

Corrected

Total te 2706512728071.19

—

* Significant at 0.05 % level.
Discussion

Type of Fruits and Factors Associated with Fruit Production Performance
in Vernonia

The type of fruits produced in Vernonia is achenes. Fruit production performance in
the species investigated was observed to be associated with a number of vegetative
and reproductive characteristics of the plants. These include the type of inflorescence;
the incidence of stem branching and number of such branches in a plant; the number
of terminal capitulum clusters on a plant or the number of inflorescence branches and
the number of capitula per such clusters or inflorescence branches.

AYODELE (1994a) reported eight distinct types of capitula arrangement found among
plants of 15 species of Vernonia. The various arrangements of capitula on Vernonia
plants were reported to enhance terminally clustered small-size capitula as found in
the arborescent species, while clustering of capitula was not too conspicuous in the
big-size capitula-producing herbs or shrubs. The variation in the pattern of inflorescence
was also reported to be associated with the type of habitat.

The herbaceous species of Vernonia growing in humid habitats were observed to have
more stem branching than similar species growing in drier habitats. Flowering and
fruit development were slower during the rainy season. On the other hand, capitula
produced during the dry season or under moisture stress condition were maturing
faster. These had a greater value of good quality fruit production index in the herbs.
The faster maturing capitula of shrubs and arborescent species usually contained fewer
well-filled fruits. This was particularly true of those shrubs which have the big size
fruits e.g. V. kotschyana and to some extent V. stenostegia.

16 Comp. Newsl. 34, 1999

Hastened fruit development and ripening due to the onset of the dry season usually
resulted in the production of many malformed fruits in the shrubby species with the
big-size fruits. The fruits of V. galamensis, an herb, ranged in size among those of the
shrubs with big fruits. However, fruits produced by plants of this species when under
moisture stress condition were usually smaller but with a higher incidence of well-set
fruits.

In fact, the quality of fruits especially in V. galamensis, was observed to be associated
with the stage of growth of the plant. The early-developed capitula had few and well
formed fruits. The plants within a community had more set fruits in the capitula than
those in the periphery. The issue about the centrifugal development of heads (i.e.
capitula) on a plant and the centripetal development within a capitulum earlier reported
in Vernonia by Jones (1977) becomes very helpful in explaining some of these
observations. The capitula more centrally positioned on a plant in a community had a
better fruit-set quality. These observations also suggest outcrossing characteristics in
most Vernonia species (AYODELE 1977).

The Relationship Between Fruit Size and Quantity/Quality of Fruits Produced

The small fruit-size achenes are produced in large quantities on individual plant basis.
This is evident in the performance of V. biafrae, a straggling shrub which ranked
topmost in both fruit production capacity and the quality of fruits produced (Table 1).
A higher incidence of well-set fruits was also noted in species producing small fruits
especially among the shrubby and arborescent species like V. amygdalina, V. conferta
and V. stenostegia. The quantity of such fruits produced on individual plant basis by
the herbaceous species was lower, as evident in V. perrottetii, the best performer among
the herbs (Table 1).

It is however pertinent to mention that the attribute of small fruit-size coupled with a
high fruit-set % cuts across different growth forms (Table 1). Obviously small fruits
(or seeds) require less metabolic energy for their production and hence less food is
stored in them (SALISBURY 1942). Generally, a high fruit-set % seems advantageous for
plants to survive and produce in moisture-distress situation. This is a property of weedy
colonizers (BAKER 1965). The observations on V. stenostegia and V. galamensis are
two interesting cases in this regard.

The former, a shrub, is domicile in the dry savanna zone while the latter can thrive in
both dry and humid vegetations. However, V. stenostegia in its natural environment
produces smaller achenes and a higher fruit-set % than when raised in the humid zone.
Vernonia galamensis displays a more luxuriant vegetative growth in humid habitats
but a higher fruit-set % is only recorded among its plants in the dry savanna.

Comp. Newsl. 34, 1999 17

Clearly the shrubby and arborescent species are the most prolific fruit producers on
individual plant basis. This was observed as enhanced by the branching attributes of
the plant stem and the inflorescence upon which numerous capitula are borne ((Table
2). It is usually presumptuous of reproduction biologists to associate small seed size
with a higher number of seed productions in herbaceous species (SALISBURY 1942,
JANZEN 1969 and Hix 1977). The observation in this study cannot ascribe such an
attribute to the herbs at least on individual plant basis. The usually dense populations
of the herbs resulting from the highly efficient though relatively few seeds, seem to
overshadow the fruit production potentials of the other plant growth forms.

References

AYODELE, M. S. 1987. Cytological and Morphological Studies on some species of
Vernonia Scures. in Nigeria. M. Sc. Thesis Obafemi Awolowo University, Ie-Ife,
Nigeria. 98 pp.

AYoDELE, M. S. 1994a. Studies on the reproductive biology of Vernonia ScCHREB.
(Asteraceae). I. Types of inflorescence among different habits. Comp. Newsl.
25: 15-23.

AYODELE, M. S. 1994b. Studies on the reproductive biology of Vernonia ScCHREB.
(Asteraceae). II. Flowering and post-pollination developments in the capitulum.
Comp. Newsl. 25: 24-30.

AYODELE, M. S. 1995. Studies on the reproductive biology of Vernonia ScureB.
(Asteraceae). III. Floret shapes/number, pappus types and surface structure among
different growth habits. Comp. Newsl. 26: 26-39.

AYopELE, M. S. 1997. Studies on the reproductive biology of Vernonia ScuREB.
(Asteraceae). IV. Seasonal flowering sequence among plant forms of Vernonia in
Nigeria. Comp. Newsl. 30: 5-14.

Baker, H. G. 1965. Characteristics and modes of origin of weeds. Jn: Baker, H. G. &
G. L. StepBins (eds.), Genetics of colonising species, pp. 147-168. Academic Press,
New York.

Burtt, B. L. 1975. Patterns of structural change in the flowering plants. Trans. Bot.
Soc. Edinb. 42: 133-142.

Faust, W. Z. 1972. A biosystematic study of the Interiores species group of the genus
Vernonia (Compositae). Brittonia 24: 363-378.

INGROUILLE, M. 1992. The study of diversity and evolution of land plants 1: 3-10.
Chapman & Hall, London, UK.

18 Comp. Newsl. 34, 1999

JANZEN, D. H. 1969. Seed-eaters versus seed size, number, toxicity and dispersal.
Evolution 23: 1-27.

Jones, S. B. Jnr. 1977. Vernonieae - Systematic review. Jn: HEywoop, V. H., HARBORNE,
J.B. & B.L. Turner (eds.), The Biology and Chemistry of the Compositae, pp.
503-521. Academic Press, London & New York. Jones,

Jones, S. B. Jnr. 1979. Synopsis and pollen morphology of Vernonia (Compositae:
Vernonieae) in the New World. Rhodora 81: 425-447.

Saissury, E. J. 1942. The reproductive capacity of plants. Studies in quantitative
Biology. G. Bell & Sons Ltd. 244 pp.

Comp. Newsl. 34, 1999 19

Studies on the reproductive biology of
Vernonia SCuHREB. (Asteraceae)
VI. Seed germination strategies among
growth forms

M.S. AYODELE
Biological Sciences Department
University of Agriculture
P.M.B. 2240
Abeokuta, Nigeria
e-mail: segmuy @ unaab.edu.ng

Abstract

Four germination-related components of fitness and three general trends in the
relationship between fruit-set attribute and seed germination performance were recorded
among the species of Vernonia investigated.

Adaptations for safety and variable dormancy of propagules were identified as important
germination strategies in Vernonia. The discrepancy between the observed high seed
production potentials and scanty populations of some species in the wild is accounted
for largely by seed wastage arising from the non-availability of suitable germination
sites.

Introduction

The ecological versatility of Vernonia has been widely reported and the genus
conclusively described as capable of occupying varying vegetation zones, by many
workers (Faust 1972, Jones 1979 and AYopDELE 1995, to mention a few). AYODELE
(1997) reported that some of the species of Vernonia in Nigeria also display some
considerable degree of ecological preferences to which populations of such species
are commonly restricted. These observations seem to confirm the tremendous diversity
that has occurred during the evolution of the genus (Jones 1979).

The pollen efficiency and the high rate of seed (achenes) production among some
species of the genus have also been reported (AYODELE 1999). These various attributes
of Vernonia presuppose high populations, especially of the prolific seed-producing

20 Comp. Newsl. 34, 1999

taxa in their various domains compared with other plants in the community. The absence
of such high populations calls for an investigation on the efficiency or otherwise, of
the seed germination performance among the species of the genus.

There are reports on plant seed-production capabilities in relation to seed longevity
attribute as well as the plant strategy towards seed wastage due to predators on one
hand, and the immediate lack of appropriate environment/substrate for germination,
on the other hand (SALIsBuryY 1942, JaNzEN 1969). Another possible motive for high
seed production in plants has been expressed as a major component attribute of an
aggressive colonizing species (e.g., JANZEN 1969). None of the plants on which these
conclusions were drawn is in the genus Vernonia.

A couple of reports noted the prevalence of initial seed germination and seedling
growth rate problems in some species of Vernonia (BERRY & LESSMAN 1969 and PERDUE
1986 & 1987). Saispury (1961) had earlier identified the variability in the mode of
seed germination even if conditions are favourable. Generally, the recorded observations
on seed germination behaviour from the point of view of the plant strategy for survival
are rather meagre except for some economic plants. Even for such, they are liable to
be based upon common strains in the particular areas, although strains with other
physiological features may occur elsewhere. More emphasis is therefore required on
seed germination studies to reflect some continuity in biological research in this area.

The objective of this paper is to document the seed germination performance studies
on some species of Vernonia in Nigeria. This is with a view to providing a clearer
understanding of the various strategies involved in the observed high seed production
and fewer plant populations among different growth forms of Vernonia species.

Materials and Methods

Field Collection and Processing of Fruit (Achenes) Specimens

Ripe capitula were harvested from field plant populations and bagged in sachets labelled
according to their accession numbers. The fruits (achenes) were separated from each
capitulum by manual threshing. This also detached the pappi from the achenes. The
threshed material was then winnowed to separate the seeds from the chaffs of involucre
bracts and detached pappi.

Germination Trials on Vernonia Achenes

Three disposable plastic Petri dishes were cleaned and the inside lined with moist
filter paper. Twenty-five to fifty seeds (the higher number of seeds being for the small-

Comp. Newsl. 34, 1999 Ai

sized seeds) were put on the moist filter paper in each Petri dish. This was done for
each of the species investigated. The dishes were kept on laboratory benches at room
temperature for germination. The filter paper was kept moist (but not wet) regularly
by adding more distilled water using a wash bottle.

Germination observations were recorded daily for up to 30 days. The experiment was
repeated three times at different intervals of the study. The cumulative percentage
germination values for each species were estimated at 7, 14, 21 and 28 days after
sowing respectively. A pre-trial sowing was conducted to identify any species that
may require any pre-treatment aid for germination to occur. Such pre-treatment included
soaking achenes in water; soaking in 10 ml concentrated sulphuric aid in a staining
block for 3-10 min., depending on seed size and seed coat toughness. Seeds soaked in
acid were thereafter rinsed in three changes of tap water contained in a staining block.

Seeds were also pre-treated for germination by first soaking in water for 1-3 days,
followed by embryo excision through manual removal of the outer coat by means of a
mounted needle. Twelve of the 16 species collected were used for the germination
trials. Those species, whose seeds were not sufficiently available or could not be raised
in the experimental garden, were not included in the trials.

Results

Features of the Ripe Vernonia Capitulum

A ripe capitulum is usually dry and crowned with the characteristic pappus of the
species. This gives the capitulum the appearance of a full bloom flower. The colour of
the pappus varied among the species from cream to black, with shades of the two
extremes.

Germination Performance at 7 Days after Sowing

Vernonia cinerea seeds were quick in germination (2-4 days after sowing) and had the
highest germination performance (69.9 %). This was followed by V. amygdalina (45.6
%) and V. perrottetii (42.3 %) respectively (Fig. 1). The germination of the seeds of V.
ambigua and V. migeodi did not commence during this period. Seed germination in V.
stenostegia and V. conferta was considerably poor at 6.0 % and 2.7 %, respectively.

Germination Performance at 14 Days after Sowing

Vernonia cinerea and V. perrottetii maintained an increase in the cumulative number
of seeds which germinated. However, while the increase in V. perrottetii was steady,

22 Comp. Newsl. 34, 1999

there was a slight fall in the rate of increase of germination performance in V. cinerea
(Fig.1.). Vernonia tenoreana showed a conspicuously sharp increase in the number of
seeds which germinated between 7 and 14 days; rising from 10.8 % at 7 days after
sowing to 63.0 % at 14 days after sowing (Fig. 1.).

There was a steep fall in the rate of increase in germination observed in V. amygdalina.
The germination performance in V. perrottetii (76.0 %) was now considerably higher
than that in V amygdalina (51.3 %), a significant departure from the occurrence between
the two species during the first seven days after sowing. The performance in V.
stenostegia and V. conferta remained poor, even though there was a steady rise in the
number of seeds that germinated (Fig. 1). The germination of the seeds of V. ambigua
and V, migeodi did not commence until after 7 days with an initial poor value of 3.3 %
for V. ambigua, despite the relative lateness in germination commencement. This
occurrence was repeated in all the trials conducted.

Germination Performance at 21 Days after Sowing

Vernonia cinerea remained at its top-ranking germination performance during the
period of the trial. There was a continuous and steady increase in the number of seeds
which germinated between 14 and 21 days and later. In some trials 100 % germination
was attained just before or by 21 days. Even in those trials with less than 100 % at 21
days, there was no further germination of such seeds thereafter.

For V. perrottetii, V. biafrae, V. migeodi and V. conferta, seeds that were viable did
germinate by or just after 14 days. There was a sudden upsurge in the number of V.
amygdalina seeds germinating between 14 and 21 days after sowing (Fig. 1). This
case of asudden upsurge also occurred and in a greater dimension, in the germination
of V. kotschyana seeds. In fact, the sudden increase was so much as to give this species
the second best performance (89 %) next to V. cinerea (95 %) in the overall germination
performance. The sudden increase in the number of V. tenoreana seeds germinating
between 7 and 14 days slowed down in tempo after 14 days (Fig. 1). The poor
germination rate of the seeds of V. migeodi, V. stenostegia and V. conferta remained so
throughout the experiment.

Discussion

A summary of Observed Seed Germination Trend in Vernonia

Generally, varying modes of seed germination were encountered, cutting across growth
forms and associated with seed-size type in species of Vernonia studied. There were

Comp. Newsl. 34, 1999 23

those seeds that commenced germination almost as soon as a favourable environment
for germination prevailed, e.g. V. cinerea and V. perrottetit. For such species, germination
progressed steadily until all the viable seeds had germinated.

For some species, germination was characterized by an initial flush within the first
seven days or thereabout of exposure to favourable conditions. This was soon followed
by adrop, after which another flush of germinating seeds occurred, e.g. V amygdalina.
This multi-peaked germination performance continued until all the viable seeds in the
seedlot had germinated. The next mode of germination was that characterized by an
initial slow rate lasting up to 7 days after sowing. A flush of germination then followed
and was steadily maintained until all the viable seeds had germinated, e.g. V. tenore-
ana and V. kotschyana. Other species manifested poor germination performances in
form of late commencement of germination and subsequent relatively poorer rate and
number of seed germinated. Vernonia ambigua, V. migeodi, V. conferta and V. stenostegia
belong to this group (Fig. 1).

This explains the poor colonization ability of these species, especially V. ambigua and
V. stenostegia. They were also the species found restricted only to their natural habitats
— V. conferta in the rain forest; V ambigua and V. stenostegia in the savanna and
usually in sparse populations.

Fruit-set Attribute and Germination Performance

There were three general trends in the relationship between fruit-set attributes and the
overall seed germination performance observed in the species investigated. Some spe-
cies noted for the production of well-set fruits, had poor germination performance
recorded for the fruits e.g. V. stenostegia. A seedlot of this species with fruit-set value
of 76.9 % had only 17.6 % seed germination performance. Other notable species in
this regard were V. migeodi (100 % & 14 %), V. galamensis var. ethiopica (85.3 % &
49.2 %); the values in parenthesis being for fruit-set and seed germination respectively.
When some non-germinating, but plump seeds of V. stenostegia were dissected to
excise the embryos, a corky or woody substance was found instead of a fleshy em-
bryo. The cause and the degree of the incidence of corky tissue in place of an embryo
in some seeds of V. stenostegia deserve some investigation.

The second trend was that some species had poor fruit-set attributes, but efficient
germination of the few quality seeds produced e.g. V. kotschyana (30.9 % & 89.0 %).
The third trend was that in some species fruit-set attribute and germination performance
were both good, e.g. V. cinerea (95.2 & 95.0 %). Vernonia cinerea is a fast colonizer
of open fields and lawns.

24 Comp. Newsl. 34, 1999

Adaptive Significance of Reproductive Strategies Related to Germination

The species of Vernonia which produce small-sized fruits were observed to have a
high incidence of well-set fruits. This cuts across growth forms e.g. V. cinerea, V.
migeodi, V. biafrae and V. amygdalina. V. galamensis from habitats with different
moisture status demonstrated in their fruit-set and fruit-size attributes relationship, the
emphasis on the balance between the advantage of more (but smaller) seeds in open
habitats and less (but larger) seeds with greater competitive capacity in the closer
communities of the rain forest. The difference in germination performance of the two
V. galamensis materials (Fig. 1) corroborates the report that the Nigerian accession
was a better performer in seed germination and seedling development than the mate-
rial from Kenya (PERDUE, pers. comm.).

The four germination-related components of fitness observed among the species of
Vernonia are adaptations for ensuring the safety, longevity and effective utilization of
propagules. These are all relevant to the individual species seed-production capabilities
and the efficiency of germination of its seeds. The immediate high peak germination
in V. cinerea is good for the species that produce small fruits. Such fruits have highly
limited food stored in them (SALISBURY 1942). This stored food must be utilized as
soon as the seeds find a favourable environment. Finding a favourable environment is
an important determinant of the germination performance of the small-seed-size spe-
cies of Vernonia. This will explain the lack of relatively large populations of the prolific
small-seed-species like V. biafrae and V. conferta in the natural habitats, whereas, V.
biafrae seedlings were observed to be common pollutants (’ weeds’) in other pots in
the screen house. This conclusively indicates that the small-seed-size species do not
readily find suitable environment for germination in the wild and hence loose their
viability.

The intermittent burst or spurts in the germination of V. amygdalina seeds are favourable
adaptation for such a widely dispersed species in an environmental mosaic. Vernonia
amygdalina is a cultivated species that is effectively raised from both seeds and stem
cuttings. The deductions from these observed strategies related to germination show
that the herbaceous weedy colonizers among the species of Vernonia, combine the
good qualities of a high fruit-set value with an effective germination performance as in
V. cinerea, V. perrottetii and to some extent V. galamensis.

A combination of the strategy of seed dispersal (AvopELE 1995) and germination in
Vernonia serve more of a colonization function than escape from pests. There were no
significant or noticeable insect pest or any plant diseases of the seeds (fruits) observed
during this study. The bitter chemicals in Vernonia plant parts (including the fruits)
that Burtt (1977) reported, would not make the fruits vulnerable to predators.

Comp. Newsl. 34, 1999 25

The larger seeds are dispersed close to the parent plants and could remain in the soil
pending an opening in the community before germinating as in V. tenoreana. The
small size seeds are dispersed far from the parent plants to suitable sites for colonization.
Such favourable sites may, however, not be readily available for the longevity of the
seeds. Even where the seeds germinate, not many of the seedlings reach adult stage
because of the low rate of seed germination or growth of seedlings earlier mentioned.
Other plants in the community soon overtake them and they get smothered.

The survival strategy against this problem in V. migeodi is a pertinent example. This
species usually takes off either from rootstocks or through seed germination as soon
as the rains commence after the dry season. The other plants in the community, especially
the tall grasses, are usually in an active vegetative growth stage when V. migeodi would
have started flowering and fruiting. By so doing the species is able to overcome the
competition from other plants in its community and so ensure the continuity of existence
and spread of the species to other opened-up areas. This might explain the readily
observed preponderance of the species on the roadsides of newly constructed highways,
particularly those highways adjourning previously identified habitat locations.

26 Comp. Newsl. 34, 1999

References

Ayope.e, M. S. 1995. Studies on the reproductive biology of Vernonia SCHREB.
(Asteraceae). III. Floret shapes/number, pappus types and surface structure among
different growth habits. Comp. Newsl. 26 -39.

Ayope.e, M. S. 1997. Studies on the reproductive biology of Vernonia SCHREB.
(Asteraceae). IV. Seasonal flowering sequence among plant forms of Vernonia in
Nigeria. Comp. Newsl. 30: 5-14.

AyopELE, M. S. 1999. Studies on the reproductive biology of Vernonia ScHREB.
(Asteraceae). V. Fruit production and fruit-set strategies among growth forms. Comp.
Newsl. 34: 9-18.

Berry, C. D. & K. J. LessMAN 1969. Dehulling and seed germination in Vernonia
anthelminitica (L.) WILLD. Crop Science 9: 247-248.

Burtt, B. L. 1977. Aspects of diversification in the capitulum. Jn: HEywoop, V. H.,
HARBORNE, J. B. & B. L. Turner (eds.), The Biology and Chemistry of the
Compositae, pp. 41-59. Academic Press, London & New York.

Faust, W. Z. 1972. A biosystematic study of the Interiores species group of genus
Vernonia (Compositae). Brittonia 24: 363-378.

JANZEN, D. H. 1969. Seed-eaters versus seed size, number, toxicity and dispersal.
Evolution: 23: 1-27.

Jones, S. B. Jr. 1979. Synopsis and pollen morphology of Vernonia (Compositae:
Vernonieae) in the New World. Rhodora 81: 425-447.

PERDUE, R. E. Jr. 1986. Vernonia galamensis Foreign Travel Report: Ethiopia,
Zimbabwe and Kenya. June 18 - July 28, 1986. USDA Agricultural Research
Services. Plant Genetics and Germplasm Institute, Maryland, USA.

PERDUE, R. E. Jr. 1987. Response of Vernonia galamensis to poorly drained soils or
excessive precipitation. Update Report Ref. 081. USDA Germplasm Introduction
and Evaluation Laboratory, Beltsville, Maryland, USA. 2 pp.

Sa.isBury, E. 1942. The reproductive capacity of plants. Studies in quantitative Biology.
G. Bell & Sons Ltd. London. 244 pp.

Sa ispury, E. 1961. Weeds and aliens. St. James Place, London. 384 pp.

Comp. Newsl. 34, 1999

Table 1. Key to nomenclatural abbreviations in Fig. 1.

*

*x

*** Collected in Kenya (PERDUE’s accession)

: V. amygdalina
teno V. tenoreana
ia ba
4. steno V. stenostegia Shrub
; V Shrub

Collected in Nigeria

Vernonia conferta

/ kotschyana

Growth
form

Tree/Shrub*

V, biafrae

V. migeodi

V. cinerea

V. galamensis**

ambi V. ambigua Herb
i

Determined by level of plant growth disturbance through topping

27

28

Comp. Newsl. 34, 1999

Germination percentage(%)

N w p= a a pp a
Oo Oo oO Oo Oo Oo Oo

o
o

OOL

Buimos saye skeqg

Iyja —y— |
ued
Ique—o—|
eje6 ——
a6iu —+— |
Je1q—e—
$}0} —K—
oQua}s —<—
oua} —y—

BAwe —g—

ju0o—@—

euo———

Fig. 1. Cumulative seed germination trend in some species of Vernonia

Comp. Newsl. 34, 1999 29

New combinations in Monticalia
(Compositae-Senecioneae)
from Colombia

Bertit NORDENSTAM
Department of Phanerogamic Botany
Swedish Museum of Natural History
P. O. Box 50007, SE-104 05 Stockholm, Sweden
e-mail: bertil.nordenstam @nrm.se

Abstract

In a recent treatment of a part of the Senecioneae of Colombia, covering the genera
Dendrophorbium and Pentacalia (Diaz-PiEDRAHITA & CUATRECASAS 1999), the genus
Moniticalia was included in Pentacalia as subgenus Microchaete. However, Monticalia
is better treated as a distinct genus, and hence a number of new combinations are
required for the Colombian taxa of this group. In this paper 24 such new combinations
are published.

Introduction

The genus Monticalia was established by Jerrrey (1992) for a widespread group of
Central and South American senecioid species. The genus comprises ca. 70 species of
erect shrubs or shrublets with usually small leaves which are closely set to imbricated,
and white to yellow capitula which may be radiate or discoid. The genus has been
widely accepted as separate from Senecio and Pentacalia (e.g., BREMER 1994, JORGENSEN
& Utioa ULtoa 1994, Norpenstam 1996, 1999), although some authors prefer to
have it as a subgenus under Pentacalia, viz. subgenus Microchaete.

In a recent excellent treatment of Colombian Senecioneae (Part I), Diaz-PIEDRAHITA &
Cuatrecasas (1999) have adopted the wider generic concept of Pentacalia, but accepted
Dendrophorbium as a distinct genus. They account for 44 species of Pentacalia sub-
genus Microchaete and some infraspecific taxa. Many of these have valid names un-
der Monticalia, but several new combinations are required when the genus is kept
separate.

30 Comp. Newsl. 34, 1999

New Combinations
(The sequence of taxa follows that of Diaz-Prepranita & Cuarrecasas 1999)

1. Monticalia romeroana (Diaz & BuENO) B. Norp., comb. nov.

Basionym: Pentacalia romeroana Diaz & Busno, Rev. Acad. Colomb. Cienc. 21(80):
202-204 (1997). - Orig. coll.: Colombia, Magdalena, municipality of Ciénaga, Santa
Marta region, W side of Sierra Nevada, 16.1.1954, R. Romero 4551 (COL holotype).

Ref.: Diaz-PrEDRAHITA & CUATRECASAS 1999: 188 ff.

2. Monticalia subarachnoidea (Wevv.) C. JEFFREY var. pauciflora (Diaz &
Bueno) B. Norp., comb. nov.

Basionym: Pentacalia subarachnoidea (Scu. Br.) Cuatr. var. pauciflora Diaz & BUENO,
Rev. Acad. Colomb. Cienc. 21(80): 202 (1997). - Orig. coll.: Colombia, Magdalena,
Sierra Nevada de Santa Marta, village of San Sebastian, 2400 m, 5.XII.1978, O. RANGEL,
H. Sturm & E. WEDLER 1868 (COL holotype).

Ref.: Diaz-PIEDRAHITA & CUATRECASAS 1999: 198.

3. Monticalia juajibioyi (Diaz & Cuatr.) B. Norp., comb. nov.

Basionym: Pentacalia juajibioyi (’juajibioy”) Diaz & Cuatr., Rev. Acad. Colomb.
Cienc. 17(67): 688 (1990). - Orig. coll.: Colombia, Magdalena, Sierra Nevada de
Santa Marta, surroundings of rio Sevilla township, 3360-3410 m, 26.1.1959, H. BARCLAY
& P. Juampioy 6709 (COL holotype, US isotype).

Ref.: Diaz-PreEDRAHITA & CUATRECASAS 1999: 198 ff.

Note: The original epithet, formed from the name of a person, is here changed to the
genitive case.

4. Monticalia schultzei (Cuatr.) B. Norp., comb. nov.

Basionym: Pentacalia schultzei Cuatr., Phytologia 57: 169 (1981). - Orig. coll.:
Colombia, Magdalena, Sierra Nevada de Santa Marta, Siminchucué, 3400 m, VI.1928,
A. SCHULTZE 1285 (B holotype, B isotype).

Ref.: Diaz-PrmpRAHITA & CUATRECASAS 1999: 201 ff.

Comp. Newsl. 34, 1999 31

5. Monticalia schultzei (Cuatr.) B. Norp. subsp. sevillana (Cuatr.) B. Norp.,
comb. nov.

Basionym: Pentacalia schultzei Cuarr. subsp. sevillana Cuatr., Phytologia 57: 171
(1985). - Orig. coll.: Colombia, Magdalena, Sierra Nevada de Santa Marta, W peaks,
rio Sevilla townships, 3495 m, 20-30.1.1959, H. G. BARcLay & P. Juampioy 6594 (US
holotype, COL isotype).

Ref.: Diaz-PIEDRAHITA & CUATRECASAS 1999: 203 f.

6. Monticalia harrietae (Cuatr.) B. Norp., comb. nov.

Basionym: Senecio harrietae Cuatr., Proceed. Biol. Soc. Washington 74: 19 (1961).
- Syn.: Pentacalia harrietae (Cuatr.) Cuatr., Phytologia 49: 255 (1981). - Ong. coll.:
Colombia, Magdalena, Sierra Nevada de Santa Marta, rio Sevilla townships, W peaks,
3400 m, 1.1959, H. BARcLay & P. Juampioy 6690 (US holotype, COL isotype).

Ref.: Dfiaz-PIEDRAHITA & CUATRECASAS 1999: 205 ff.

7. Monticalia abietina (WiLLD. Ex WEDD.) C. JEFFREY var. aciculata (Cuatr.)
B. Norp., comb. nov.

Basionym: Senecio aciculatus Cuatr., Notas Fl. Colombia 6: 24, Trabajos Comis.
Bot. Secret. Agric. Valle, Cali (1944). - Syn.: Pentacalia abietina (WILLD. EX WEDD.)
Cuatr. var. aciculata (Cuatr.) Cuatr., Phytologia 49: 252 (1981). - Orig. coll.:
Colombia, Norte de Santander, Cordillera Oriental, paramo de Tama, above La Cueva,
3100-3200 m, 27.X.1941, J. CuarrecasAs, R. E. Scuures & E. SmirH 12612 (COL
holotype, COL, US isotypes).

Ref.: Diaz-PIEDRAHITA & CUATRECASAS 1999: 213 ff.

8. Monticalia elatoides (WeEvp.) B. Norp., comb. nov.

Basionym: Senecio elatoides Wevv., Chloris Andina 1: 101 (1856). - Syn.: Pentacalia
elatoides (WEDD.) Cuatr., Phytologia 49: 254 (1981). - Orig. coll.: Colombia,
Santander, Prov. Pamplona, paramo near La Baja, 3900-4000 m, J. Scuim 2 (P?).

Ref.: Diaz-PIEDRAHITA & CUATRECASAS 1999: 215 ff.

32 Comp. Newsl. 34, 1999

9. Monticalia carupana (Diaz) B. Norp., comb. nov.

Basionym: Pentacalia carupana Diaz, Rev. Acad. Colomb. Cienc. 22(83): 187-192
(1998). - Orig. coll.: Colombia, Cundinamarca, municipality of Carupa, paramo de
Guargua, W part, 3720 m, 5.11.1987, Roberto SANCHEZ 071 (COL holotype).

Ref.: Diaz-PrEDRAHITA & CuaTRECASAS 1999: 223 f.

10. Monticalia carrikeri (Cuatr.) C. JEFFREY var. macrophylla (Diaz)
B. Norp., comb. nov.

Basionym: Pentacalia carrikeri (Cuatr.) Cuatr. var. macrophylla Diaz, Rev. Acad.
Colomb. Cienc. 22(83): 192 (1998). - Orig. coll.: Colombia, Magdalena, Sierra Ne-
vada de Santa Marta, Buritaca transect”, edge of La Cumbre, 3700 m, 15.VIII.1977,
O. RancEL, A. M. CueeF et al. 985 (COL holotype, COL isotype).

Ref.: Diaz-PrepRAHITA & CuaTrecasas 1999: 240 f.

11. Monticalia pulchella (H.B.K.) C. JEFFREY subsp. guantivana (CuatTR.)
B. Norpb., comb. nov.

Basionym: Senecio guantivanus Cuatr., Notas Fl. Colombia VI: 23, Trab. Comis. Bot.
Secret. Agric. Valle, Cali (1944). - Syn.: Pentacalia pulchella (H.B.K.) Cuatr. subsp.
guantivana (Cuatr.) Cuatr., Phytologia 49: 258 (1981). - Orig. coll.: Colombia, Boyaca,
paramo de Guantiva, Alto de Canutos, 3200-3400 m, 3.VIII.1940, J. CuATRECASAS
10344 (COL holotype, NY, US isotypes).

Ref.: Diaz-PrepRAHITa & CUATRECASAS 1999: 255 ff.

12. Monticalia vaccinioides (H.B.K.) C. JEFFREY var. microdentata (CvuatR.)
B. Norp., comb. nov.

Basionym: Senecio vaccinioides H.B.K. var. microdentatus Cuarr., Notas a la Flora
de Colombia VI: 23 (1944). - Syn.: Pentacalia vaccinioides (H.B.K.) Cuarr. var.
microdentata (CuatrR.) Cuatr. & Diaz in Diaz & Cuatr., Asteraceas de la Flora
Colombia, Senecioneae-I: 266 f. (1999). - Orig. coll.: Colombia, Santander, cordillera
Oriental, paramo de Almorzadero, NE slope, 3800-3900 m, 20. VII. 1940, J. CUATRECASAS
& H. Garcia 10027 (COL holotype, US isotype).

Ref.: Diaz-PEmRAHITA & CuaTRECASAS 1999: 266 f.

Comp. Newsl. 34, 1999 33

13. Monticalia iguaquensis (Diaz & Cuatr.) B. Norp., comb. nov.

Basionym: Pentacalia iguaquensis Diaz & Cuatr., Rev. Acad. Colomb. Cienc. 19(73):
254-256 (1994). - Orig. coll.: Colombia, Boyaca, municipality of Villa de Leyva,
Reserva Natural de Iguaque, near la Laguna, 3400-3600 m, 27.III.1993, O. RANGEL et
al. 9804 (COL holotype).

Ref.: Diaz-PIEDRAHITA & CUATRECASAS 1999: 267 ff.

14. Monticalia summapacis (Cuatr.) B. NorbD., comb. nov.

Basionym: Senecio summapacis Cuatr., Feddes Repert. 55(2/3): 150 (1953). - Syn.:
Pentacalia summapacis (Cuatr.) Cuatr., Phytologia 49: 256 (1981). - Ong. coll.:
Colombia, Cundinamarca, summit of San Juan highlands, 18 km E Cabrera, paramo
de Sumapaz, 4200 m, 1943, F. R. Fosserc 20791 (F holotype).

Ref.: Diaz-PIEDRAHITA & CUATRECASAS 1999: 284 ff.

15. Monticalia almorzana (Diaz & Cuatr.) B. Norb., comb. nov.

Basionym: Pentacalia almorzana Diaz & Cuatr., Rev. Acad. Colomb. Cienc. 19(72):
22 (1994). - Orig. coll.: Colombia, Santander, paramo de Almorzadero, moist meadows,
3700-3750 m, km 50-51, 20.1X.1969, J. Cuarrecasas & L. RopRiGUEZ 27888 (COL
holotype, US isotype).

Ref.: Dfiaz-PIEDRAHITA & CUATRECASAS 1999: 298 ff.

16. Monticalia ledifolia (H.B.K.) C. JEFFREY var. lanata (H.B.K.)
B. Norp., comb. nov.

Basionym: Cacalia lanata H.B.K., Nov. Gen. Sp. Pl. 4: 127, ed. folio: 162 (1820). -
Syn.: Senecio lanatus (H.B.K.) DC., Prodr. 6: 422 (1837); Pentacalia ledifolia (H. B.
K.) Cuatr. var. lanata (H. B. K.) Diaz & Cuatr., Asteraceas de la Flora de Colombia,
Senecioneae-I: 311 (1999). - Orig. coll.: Colombia, Cundinamarca, Monserrate ’Crescit
locis frigidis in declivitate montis Chingasa juxta sanctuarium Montserratense, alt.
1500 hex” (2920 m), A. HumsBotpt & A. BonpLanp s.n. (P holotype).

Ref.: Dfaz-PreDRAHITA & CUATRECASAS 1999: 311 ff.

34 Comp. Newsl. 34, 1999

17. Monticalia ledifolia (H.B.K.) C. JerFrey var. glabrata (Diaz & MENDEZ)
B. Norp., comb. nov.

Basionym: Pentacalia ledifolia (H.B.K.) var. glabrata Diaz & MENDEz in Diaz & Cuatr.,
Asteraceas de la Flora de Colombia, Senecioneae-I: 313 (1999). - Orig. coll.: Colombia,
Cundinamarca, Neusa dam, 3000 m, 4.]1.1966, L. Ure U. 5527 (COL holotype, US

isotype).
Ref.: Diaz-PrEDRAHITA & CUATRECASAS 1999: 313 f.

18. Monticalia albi-panquei (Diaz & Cuatr.) B. Norb., comb. nov.

Basionym: Pentacalia albi-panquei Diaz & Cuatr., Rev. Acad., Colomb. Cienc. 17(67):
689 (1990). - Orig. coll.: Colombia, Boyaca, road Soata-Onzaga, San Antonio ravine,
3140 m, 3. VIIT.1958, R. JARAMILLO et al. 826 (COL holotype).

Ref.: Dfiaz-PIEDRAHITA & CUATRECASAS 1999: 316 ff.

19. Monticalia cacaosensis (CuatTR.) B. Norp., comb. nov.

Basionym: Senecio cacaosensis Cuatr., Phytologia 31: 324 (1975). - Syn.: Pentacalia
cacaosensis (CuaTR.) Cuatr., Phytologia 49: 253 (1981). - Orig. coll.: Colombia,
Boyaca, paramo de Pisba, road Socha-La Punta, Cacaos, km 77, subparamo, 3430 m,
20.V1.1972, A. M. CLEEF 4776 (US holotype, COL, U isotypes).

Ref.: Diaz-PreDRAHITA & CUATRECASAS 1999: 318 ff.

20. Monticalia hammenii (Diaz & Cuatr.) B. Norp., comb. nov.

Basionym: Pentacalia hammenii Diaz & Cuatr., Rev. Acad. Colomb. Cienc. 19(73):
256 (1994). - Orig. coll.: Colombia, Cesar, Sierra Nevada de Santa Marta, valley of rio
Donachuy, road Corisa-Lago Naboba glacial valley, 4200-4300 m, 17.X.1958, Tu.
VAN DER HAMMEN 1197 (COL holotype, US isotype).

Ref.: Diaz-PrepDRAHITA & CUATRECASAS 1999: 322 ff.

21. Monticalia taironae (Diaz & Cuatr.) B. Norp., comb. nov.

Basionym: Pentacalia taironae Diaz & Cuatr., Rev. Acad. Colomb. Cienc. 17(67):
687 (1990). - Orig. coll.: Colombia, Magdalena, Sierra Nevada de Santa Marta,

Comp. Newsl. 34, 1999 oS

surroundings of rio Sevilla townships, 3520-3770 m, 28.1.1959, H. G. BaRcLay & P.
Juampioy 6766 (COL holotype, US isotype).

Ref.: Diaz-PrEDRAHITA & CUATRECASAS 1999: 322 ff.

22. Monticalia guadalupe (Cuatr.) C. JEFFREY subsp. caucana (Cuatr.)
B. Norp., comb. nov.

Basionym: Pentacalia guadalupe (Cuatr.) Cuatr. subsp. caucana Cuatr., Phytologia
52: 163 (1982). - Orig. coll.: Colombia, Cauca, Central Colombian massif, paramo de
las Papas, hills SE of Magdalena lagune, slopes at Magdalena river, 3350-3450 m,
12.IX.1958, J. M. Ipriwo, P. Pinto & H. BiscHLer 3250 (COL holotype, COL isotype).

Ref.: Diaz-PrEDRAHITA & CUATRECASAS 1999: 334 ff.

23. Monticalia fimbriifera (Diaz & MENDEz) B. Norp., comb. nov.

Basionym: Pentacalia fimbriifera Diaz & MENpEz, Rev. Acad. Colomb. Cienc. 21(81):
404 (1997). - Orig. coll.: Colombia, Narifio, El Tablén municipality, Doha Juana
volcano, 3200-3500 m, 3.IV.1997, G. NaRVAEz & F. Rosero 06 (COL holotype).

Ref.: Diaz-PIEDRAHITA & CUATRECASAS 1999: 336 ff.

24. Monticalia tolimensis (Cuatr.) C. JEFFREY var. colombiana (Cuatr.)
B. Norp., comb. nov.

Basionym: Senecio colombianus Cuatr., Caldasia 1(1): 9 (1940). - Syn.: Pentacalia
colombiana (Cuatr.) Cuatr., Phytologia 49: 253 (1981); Dendrophorbium
colombianum (Cuatr.) C. JEFFREY, Kew Bulletin 47: 66 (1992); Pentacalia tolimensis
(Scu. Br. ex Webb.) Cuatr. var. colombiana (Cuatr.) Diaz in Diaz & Cuatr., Asteraceas
de la Flora de Colombia, Senecioneae - I: 343 (1999). - Orig. coll.: Colombia,
Cundinamarca, pdéramo de Cruz Verde, Chipaque gap, 3150 m, 7.X.1938, J.
Cuarrecasas 422 (COL holotype, US isotype).

Further syn.: Senecio pavonicus BADILLO, Bol. Soc. Venez. Cienc. Nat. 10: 318 (1946).
- Orig. coll.: Venezuela, Tachira, J. STEYERMARK 57379 (VEN holotype).

Ref.: Diaz-PrepRAHITA & CuaTRECASAS 1999: 343 ff.

36 Comp. Newsl. 34, 1999

References

Bremer, K. 1994. Asteraceae. Cladistics and Classification. Timber Press, Oregon.

Diaz-Prepranit, S. & J. Cuarrecasas 1999. Asteraceas de la Flora de Colombia,
Senecioneae-I. Géneros Dendrophorbium y Pentacalia. Santafe de Bogota, D. C.

Jerrrey, C. 1992. Notes on Compositae, VI. The tribe Senecioneae (Compositae) in
the Mascarene Islands with an annotated world check-list of the genera of the tribe.
Kew Bulletin 47: 49-109.

JORGENSEN, P. M. & C. Uttoa ULLoa 1994. Seed Plants of the High Andes of
Ecuador - a checklist. AAU Reports 34, Aarhus.

NorbenstaM, B. 1996. New combinations in Ecuadorean Senecioneae. Comp. Newsl.
29: 47-50.

NorbENSTAM, B. 1999. Senecioneae (except Gynoxys). In: JORGENSEN, P. M. & LEON-
YAneEz: Catalogue of the Vascular Plants of Ecuador. Missouri Bot. Garden,
St. Louis.

Comp. Newsl. 34, 1999 37

Screening of some Asteraceous plants for
antimycotic activity

MaAHEnpRA Rar & DEEPAK ACHARYA
Department of Botany
Danielson College
Chhindwara-480 001, M. P., India

Abstract

In vitro acitivy of extract of 31 plant species of family Asteraceae was evaluated against
Fusarium oxysporum and Trichophyton mentagrophytes. The sensitivity of both fungi
was tested by dry weight method. Both fungi were sensitive to extracts of all the
plants, except Launaea acaulis and Elephantopus scaber, extracts of which accelerated
the rate of growth. The maximum antimycotic activity against F oxysporum and T.
mentagrophytes was exhibited by flower extract of Tagetes erecta followed by whole
plant of T. patula and leaf extract of T: erecta. The minimum inhibition of F’ oxysporum
was shown by extracts of Spilanthes acmella followed by Emilia sonchifolia and
Vernonia divergens, whereas extracts of Emilia sonchifolia, Tridax procumbens and
Cichorium intybus exhibited the minimum inhibitory effect on T. mentagrophytes.

Introduction

There are many antimycotics available in the market including some plant-based
antifungal agents. Increasing interest in plant-derived antimycotics all over the world
has attracted botanists/pharmacologists to discover newer herbal antimycotics. One
clear reason behind it is that most of synthetic drugs are only useful for local treatment
because of their toxic nature. They are broad spectrum antibiotics which are effective
in the treatment of the most important groups of fungi that infect human beings. In the
treatment of mycosis, this kind of active agent is preferred because here a broad spectrum
therapy is required. Increasingly, allergic reactions of the skin are observed today. The
reason is high rate of sensitization power of these antifungal agents. Moreover, synthetic
antifungals are costly and exhibit side-effects. Therefore, the awared people are turning
towards herbal antimycotics.

The literature survey on the antifungal therapy reveals that a lot of contribution has
been made in India and abroad (BHakunret al. 1971, DHAR et al. 1974, Ray & MaAsuMDAR
1975, Jain & AGRAWAL 1976, DHAWAN et al. 1977, Misra & Doar 1979, Wanas et al.

38 Comp. Newsl. 34, 1999

1981, SincH & DESHMUKH 1984, Mares 1989, Mares & FasuLo 1990, RAHALISON et
al. 1994, ViLLaRREAL 1994, Rat 1995, Apams et al. 1996, NARAYANARAO et al. 1996,
GOPALAKRISHNAN et al. 1997, Rar et al. 1999).

Asteraceae constitutes one of the largest vascular plant families with about 30,000
species and over 1100 genera. Usually, the members of the family exhibit antimicrobial
activity. Phytochemical research in the recent past revealed that sesquiterpene lactones
are the principal secondary metabolites responsible for antimicrobial activity (GOREN
et al. 1996).

The present investigation was aimed to screen the plant/plant-parts of family Asteraceae
against an opportunistic human pathogen Fusarium oxysporum and a potential hu-
man fungal pathogen Trichophyton mentagrophytes.

Materials and Methods

The Test Fungi

The test fungi were isolated from local patients suffering from skin infections. These
fungi include F- oxysporum and T. mentagrophytes. The former is basically a plant
pathogen but now it has become human pathogen while the latter is a potential human
pathogen.

The Test Plants

The Asteraceaous plants of Chhindwara and suburbs were selected for antimycotic
activity. These include: Acanthospermum hispidum, Ageratum conyzoides, Blainvillea
acmella, Blumea balsamifera, B. mollis, Caesulia axillaris, Carthamus tinctorius,
Cichorium intybus, Cyathocline purpurea, Dahlia pinnata, Echinops echinatus, Eclipta
alba, E. prostrata, Elephantopus scaber, Emilia sonchifolia, Erigeron asteroides,
Eupatorium triplinerve, Galinsoga parviflora, Gnaphalium luteo-album, G. purpu-
reum, Launaea acaulis, Parthenium hysterophorus, Spilanthes acmella, Synedrella
nodiflora, Tagetes erecta, T. patula, Tithonia diversifolia, Tridax procumbens, Vernonia
cinerea, V. divergens and Xanthium strumarium.

Plant Extracts

Ten g sample (leaves, flowers or whole plant) of each test plant was collected and
washed first in tap water and thereafter in distilled water. The test samples were cut
into small pieces and boiled in 100 ml of distilled water for 5 minutes in an autoclave
at 15 lb. pressure and 121° C. The extracts were filtered and centrifuged at 5,000 rpm

Comp. Newsl. 34, 1999 39

for 10 minutes. The supernatant was taken as extract sample and used for test.
Determination of CFU/ml

Sabouraud dextrose agar medium was prepared and F. oxysporum and T. menta-
grophytes were inoculated on it. After one week, spores were washed in sterilized
normal saline solution by glass-wool filter apparatus. The suspension thus obtained
was treated as parent suspension. The latter was utilized for tenfold serial dilution.
Nine ml of normal saline solution was taken in 5 test tubes. In first test tube, 1 ml of
spore suspension (PS) was poured by pipette under aseptic conditions. The solution of
first test tube was homogenized with the help of pipette and 1 ml of this solution was
transferred to second test tube containing 9 ml of normal saline solution. This process
was repeated up to Sth test tube. In each case, sterilized pipette was used. From every
test tube (for each dilution) 0.1 ml suspension was transferred to sterilized SDA petri-
plates. Triplicates of each dilution were maintained. The SDA seeded petri-plates were
incubated at 28 + 2°C for 24 hours. The colonies formed in each petri-plate were
counted in colony counter. The average of 3 petri-plates was taken in each case. The
test inocula was adjusted between 1.5 x 10° spores/ml.

Inoculation of Test Fungus

The test fungus was grown on Sabouraud broth (40 g glucose, 10 g peptone, 0.5 mg/
ml chloramphenicol, pH 6.5). Each 250 ml flask was filled with 50 ml of broth and 5
ml of extract. Triplicate flasks were run for each test plant. After the sterilization of
flasks at 15 Ib pressure for 10 minutes, all flasks were inoculated by test inocula (1 ml)
of definite CFU/ml. The inoculated flasks were kept for a week at 28 + 2° C. Mycelia
from each flask was harvested on a preweighed Whatman filter paper No. 42. The
actual mycelial growth of each test fungus was calculated and is given in Table 1.

Results and Discussion

It is evident from Table 1 that both species of Tagetes, viz., T. erecta and T. patula were
found to be the most effective inhibitors of mycelial growth of Fusarium oxysporum
and Trichophyton mentagrophytes. This suggests a strong antimycotic potential in
especially T. erecta, as also stated by KHAN & Evans (1996). The plant has been in use
as a traditional medicine in India since time immemorial. Among the two species of
Tagetes, T. erecta was found to be more effective mycelial inhibitor as compared to T.
patula. In the former, extract of the flowers showed the maximum inhibition, whereas
the leaves exhibited the least growth. It is probably due to presence of more bio-active
principles in flowers as compared to other parts of the plant.

40 Comp. Newsl. 34, 1999

The other Asteraceous species after T. erecta and T: patula which were found to be
effective inhibitors of mycelial growth of F oxysporum include: Cyathocline purpurea,
followed by Parthenium hysterophorus, Blumea balsamifera, B. mollis, Eupatorium
triplinerve, Tithonia diversifolia, Eclipta prostrata, Ageratum conyzoides, and Galinsoga
parviflora, whereas the minimum inhibition was exhibited by extracts of Spilanthes
acmella followed by Emilia sonchifolia and Vernonia divergens. In case of T.
mentagrophytes, other plant species containing potential for mycelial inhibition were
E. triplinerve, followed by P. hysterophorus, G. luteo-album, B. balsamifera, B. mollis,
C. purpurea and E. alba, while the minimum inhibition was represented by extracts of
E. sonchifolia, T. procumbens, and C. intybus. However, it was interesting to note that
extracts of Elephantopus scaber and L. acaulis enhanced the growth of T.
mentagrophytes.

Table 1. In vitro evaluation of different extracts of plants of Asteraceae against
F. oxysporum and T. mentagrophytes

Dry weight in mg

part/used

T™M
Acanthospermum hispidum 358.6 (14.65 %) | 266.4 (23.83 %)
Ageratum conyzoides 255.2 (27.89, %)
Blainvillea acmella 290.5 (16.9 %)
Blumea balsamifera WP 236 (32.52 %)

B. mollis WP 233.5 (44.42 %) | 239.3 31.57 %)
Caesulia axillaris WP 332.4 (20.88 %) | 252.2 (27.89 %)

Carthamus tinctorius WP 271.5 (22.37 %)
Cichorium intybus 295.3 (15.58 %)
Cyathocline purpurea WP 245 (29.95 %)
Dahlia pinnata WP 267 (23.65 %)
Echinops echinatus WP 285 (18.51 %)
Eclipta alba WP 248.4 (28.97 %)
E. prostrata WP 280 (19.94 %)
Elephantopus scaber WP 398.5 (-13.93 %)
Emilia sonchifolia WP 323.5 (7.5 %)
Erigeron asteroides WP 247 (29.37 %)

Plant

Name of plant

Comp. Newsl. 34, 1999

Eupatorium triplinerve WP 231.4 (44.92 %) 225.5 (35.52 %)
Galinsoga parviflora WP 265.2 (36.88 %) | 285.5 (18.37 %)
Gnaphalium luteo-album WP 315.2 (24.98 %) | 235 (32.8 %)

=

G. purpureum WP 310.5 (26.09 %) —_| 255.5 (26.94 %)

Launaea acaulis WP 332.5 (20.86 %) — | 432.5 (-23.65 %)

Parthenium hysterophorus WP 220.8 (47.44 %) | 230.7 (34.03 %)
WP

eee 266.4 (23.8 %)
348 (17.17 %) | 252 (27.94 %)
Reee ein 280.2 (19.88 %)
Tagetes erecta 232.2 (44.73%) | 220 (37.09 %)
239 (31.66 %)
201 (42.6 %)
211 (37.67 %)
237.8 (32%)

aa

3%

05
285.4 (18.4 %)
315.3 (9.84 %)
WP 285 (18.5 %)

V divergens WP 387.2 (7.84 %) 270 (22.8 %)
Xanthivon strumarium 311.5 (25:86 %) | 239.5 (31.52%)
420.16 349.75

Z

Tithonia diversifolia
Tridax procumbens

Vernonia cinerea

dul
ua

Note: WP - Whole Plant L - Leaf F - Flower
Percentage inhibition is given in parentheses

42 Comp. Newsl. 34, 1999

Acknowledgements

We are grateful to Professor Berti. NorDENSTAM, Director, the Swedish Museum of
Natural History, Stockholm, Sweden for encouragement and to Professor S. A. Brown,
Principal, Danielson College, Chhindwara, M.P. State, India for providing laboratory
facilities. The financial assistance provided by University Grants Commission, New
Delhi is thankfully acknowledged.

References

Apams, S., Kunz, B. & M. WEIDENBORNER 1996. Mycelial deformations of
Cladosporium herbarum due to the application of Eugenol or Carvacrol. J. Essent.
Oil Res. 8: 535-540.

Buakuni D. S., Duar, M. L., Doar, M. M., Doawan, B. N., Gupta, B. & R. C.
Srimav 1971. Screening of Indian plants for biological activity, part III. Indian J.
Experimental Biol. 9: 91-102.

Duar, M. L., Doawan, B. N., Prasap, C. R., Rastoat, R. P., Sincu, K. K. & J. S.
Tanpon 1974. Screening of Indian plants for biological activity, part IV. Indian J.
Experimental Biol. 12: 512-523.

Duawan, B. N., Patnaik, G. K., Rastoct, R. P., Sincu, K. K. & J. S. TANDON 1977.
Screening of Indian plants for biological activity, part V & VI. Indian J. Experi-
mental Biol. 15: 208-219.

GoOPALAKRISHNAN, G., BANUMATHI, B. & G. SurREsH 1997. Evaluation of the antifungal
activity of natural xanthones from Garcinia mangostana and their synthetic
derivatives. Journal of Natural Products 60 (5): 519-524.

Goren, NEZHUN, WOERDENBAG, H. J. & CANDAN Bozox-JOHANSSON 1996. Cytotoxic
and antibacterial activities of sesquiterpene lactones isolated from Tanacetum
praeteritum subsp. praeteritum. Planta Medica 62: 419-422.

JAIN, P. C. & S. C. AGRawa 1976. Activity of plant extracts against some keratinophilic
species of Nannizzia. Indian Drugs 13 (12): 25-26.

Kuan, M. T. & F. J. Evans 1996. Clinical evaluation of Tagetes erecta in the treatment
of parakeratosis. Phytotherapy Research 10: 186-188.

Mares, D. 1989. Electron microscopy of Microsporum cookei in vitro treatment with
protoanemonin: a combined SEM and TEM study. Mycopathologia 108: 37-46.

Comp. Newsl. 34, 1999 43

Mares D. & Maria P. Fasuto 1990. Ultrastructural alterations in Epidermophyton
floccosum and Trichophyton mentagrophytes exposed in vitro to protpanemonin.
Cytobios 61: 89-95.

Misra, S. S. & S. N. Doar 1979. Antifungal activity of leaf-extracts of some higher
plants. Acta Bot. Indica 7: 147.

NARAYANARAO, A., PApMasRi, K. & R. C. ANANADKUMAR 1996. Antifungal activity of
extracts from Azadirachta indica Juss. against Drechslera oryzae BREDA DE HANN.
Geo-bios new Reports 15: 126-128.

RAHALISON, L., HAMBURGER, M., Monon, M., FRENK, M. & K. HoSTETTMANN 1994.
Antifungal tests in phytochemical investigations: Comparison of bioautotrophic
methods using phytopathogenic and human pathogenic fungi. Planta Medica 69
(1): 41-44.

Ral, M. K. 1995. Comparative antimycotic activity of different parts of Parthenium
hysterophorus. World Weeds 2 (1): 45-52.

Rat, M. K., Quresui, S. & A. K. Panpey 1999. Jn vitro susceptibility of opportunistic
Fusarium spp. to essential oils. Mycoses 42 (in press).

Ray, P. G. & S. K. Mayumpar 1975. New antifungal substance from Alpinia officina-
rum. Indian J. Exp. Biol. 13 (4): 489.

Ray, P. G. & S. K. Mayumpar 1976. Antifungal flavonoid from Alpinia officinarum.
Indian J. Exp. Biol. 14 (6): 712-714.

Sincu, K. V. & S. K. DesHMuku 1984. Volatile constituents from members of Liliaceae
and spore germination of Microsporum gypseum complexes. Fitoterapia 55 (5):
297-299.

VILLARREAL, M. L., ALVAREZ, L., ALonso, D., Navarro, V., GarciA, P. & G. DEL-
capo 1994. Cytotoxic and antimicrobial screening of selected terpenoids from
Asteraceae species. J. Ethnopharmacology 42: 25-29.

Waunag, S., TANDON, R. N., JACOB, Z., SAGAR, P. & O. P. Srivastava 1981. In vitro
actitivity of a phytochemical alantolactone from /nula racemosa Hook. f. against
some pathogenic and opportunistic fungi. J. Indian Bot. Soc. 60: 278-281.

44 Comp. Newsl. 34, 1999

Diversity of arbuscular mycorrhizae in naturally
growing plants of family Asteraceae in India

MAHENDRA Rat & DEEPAK ACHARYA
Microbiology Research Lab.
Danielson College
Chhindwara - 480 001, M. P., India

Abstract

Twenty-six plants of family Asteraceae were screened for occurrence of AM fungi.
The percentage of mycorrhizal colonization was determined. All plants screened in
the present investigation were found to be associated with Glomus pallidium, G.
macrocarpum, G. mosseae, G. pulvinatum, Acaulospora laevis and Sclerocystis. The
maximum colonization (90 %) was recorded in the roots of Spilanthes calva; mode-
rate (60-70 %) in Aster amellus, Echinops echinatus, Emilia sonchifolia, Guizotia
abyssinica and Tagetes patula, whereas the minimum percentage of colonization was
observed in Gnaphalium luteo-album and Zinnia elegans (30 %).

Introduction

Most plants on the earth including Angiosperms, Conifers (except Pinaceae),
Pteridophytes, and Bryophytes are associated with arbuscular mycorrhizal fungi. AM
fungi infect fine feeder roots and colonize the cortical region from where they extend
their mycelia in rhizosphere soil. Increasing interest in AM fungi has focused on the
effectiveness of various species to promote plant health. Many researchers have pro-
ved that plants harbouring AM fungi are more efficient in acquiring phosphorus than
non-mycorrhizal plants (Mosse 1973, O’ Keere & Sytvia 1991, Durca & Gupta 1995,
VARMA & JAMALUDDIN 1995).

A survey of literature vouches that there are few reports on occurrence of AM fungi on
medicinal plants (Govinp Rao et al. 1989, SULocHANA et al. 1991, SAMBANDAN 1995) in
general and family Asteraceae in particular (WaRcup & McGee 1983, Warcup 1990).
In the present investigation, plants of family Asteraceae have been screened to assess
the colonization of various arbuscular mycorrhizal fungi in roots.

Comp. Newsl. 34, 1999 45

Materials and Methods

The root samples of 26 Asteraceous plants were collected from various parts of
Chhindwara district. The plants were recovered from different types of soil, such as,
cultivated fields, wastelands, riverbanks, ponds, sandy areas, garden soil, etc. The
samples of fine feeder roots were thoroughly washed to remove soil, organic matter,
and debris. Roots were cut into small pieces of 1 cm length and stored in individual
vials containing Formalin, Acetic acid and Alcohol (1:1:18, V/V/V). In some plants,
quick processing was done and hence samples were not preserved in FAA. The root
samples were autoclaved at 120°C after addition of 10 % KOH. Thereafter, the roots
were covered with 1 N HCI which is poured off after 3 minutes. Washing of roots was
avoided as an acidic medium is required by the stain. Later, roots samples were stai-
ned with 0.05 % trypan blue in lactophenol (PHituirs & HayMaAN 1970).

The percentage of the root colonization was evaluated as per method suggested by
READ et al. (1976).

a Serre No. of AMF infected roots x 100
Total No. of root-bits examined

The percentage frequency of AMF was calculated as per method suggested by RAUNKIER
(1934). AMF were classified into 5 classes on the basis of percentage frequency, viz.,
class A = 10-20 %; B= 21-40 %; C = 41-60 %; D = 61-80 %; E= 81-100 %.

46

Table 1.

Ageratum conyzoides L.
Aster amellus L.

Caesulia axillaris Roxs.
Echinops echinatus Roxs.
Eclipta prostrata L.
Elephantopus scaber L.
Emilia sonchifolia (L.) DC.
Eupatorium triplinerve V AHL
Galinsoga parviflora Cav.
Gnaphalium luteo-album L.
Guizotia abyssinica Cass.
Lagascea mollis Cav.
Launaea nudicaulis L.
Parthenium hysterophorus L.
Sonchus asper L.

Spilanthes acmella Hook.

S. calva DC.

Tagetes erecta L.

T. patula L.

Tridax procumbens L.
Vernonia cinerea L.

Vicoa indica L.

Xanthium strumarium L.
Zinnia elegans Jaca.

Note: FC = Frequency Class

Blumea lacera (Bur. f.) DC.

Garden soil
Sandy soil
Wasteland
Riverbank
Wasteland
Bank of pond
Wasteland
Garden soil
Garden soil
Wasteland
Sandy soil
Cultiv. field
Wasteland
Wasteland
Wasteland
Sandy soil
Near nullah
Cultiv. field
Garden soil
Garden soil
Wasteland
Garden soil
Cultiv. field
Wasteland
Garden soil

AMF association with roots of Asteraceous plants

ete [ome

Acanthospermum hispidum DC.) Sandy soil

Bharatadeo
College garden
Bharatadeo
Shukludhana
Bodri river
Atarwada
Chhota talaab
Bharatadeo
College garden
Bharatadeo
Chandangaon
Bharatadeo
Bhanadehi
Chandangaon
Chhindwara
Kundalikala
Gangiwada
Vivek. Colony
Choubitkar Farm
College garden
Geet colony
Chandangaon
College garden
Bhanadehi
Bhanadehi
Ekta park

Numbers in parenthesis are Standard Deviation

Comp. Newsl. 34, 1999

%
Colonization

42.7 (+5.353)
40.0 (+5.812)
71.5 (+7.140)
35.3 (43.365)
55.6 (+4.117)
69.3 (+5.108)
46.9 (+8.654)
46.8 (+2.498)
68.8 (+5.398)
54.8 (+2.412)
56.9 (46.776)
34.7 (+5.498)
67.3 (45.32)
39.9 (+6.462)
62.6 (43.543)
35.6 (+5.303)
54.2 (+2.229)
62.6 (+5.877)
90.0 (+8.5)
54.0 (44.837)
71.9 (45.134)
50.6 (+5.86)
67.8 (+2.4)
62.3 (+3.653)
51.5 5.792)
31.1 (44.311)

DO; SF OS Osis Ow So BW See OOO © Sua Bo

Comp. Newsl. 34, 1999 47

Results and Discussion

It is obvious from Table 1 that roots of all the Asteraceous plants screened in the
present study were found to harbour arbuscular mycorrhizal fungi. The maximum
colonization (90 %) was exhibited by Spilanthes calva followed by Aster amellus
(71.5 %), Echinops echinatus (69.3 %), and Emilia sonchifolia (68.8 %), while the
minimum by Zinnia elegans (31.1%) followed Gnaphalium luteo-album (34.7 %)
and Parthenium hysterophorus (35.6 %). KEHRI et al. (1987) reported that roots of
Ageratum conyzoides were colonized by 40 %. Similarly, in the present study, A.
conyzoides was found to be associated with 40 per cent AMF. Moreover, Tagetes
erecta was reported to have 61 per cent colonization while in the present survey it
showed 54 per cent colonization. KEur! et al. (1987) stated that soil and environmental
factors are likely to affect the plants as well as their association with symbiont. RAUNKIER
(1934) suggested a frequency class system for ecological distribution to evaluate
homogeneity and heterogeneity in an ecosystem. In the present studies, frequency
class E showed higher value which was represented by only one species. The present
findings are in agreement with earlier reports on occurrence of AMF in roots of
medicinal plants (STREZEMSKA 1975, GovinD Rao et al. 1989, SuLocuana et al. 1991).
On the other hand, Kumar, MOHAN & MAHADEVAN (1984) did not find AMF colonization
in roots of medicinal plants. They stated that absence of AMF in roots of medicinal
plants is due to the presence of secondary metabolites in them. It seems that the absence
of colonization is not due to the presence of secondary metabolites in roots, as it has
been found that plants containing AMF in their roots were also reported to exhibit
antimycotic activity (Rar 1993, Rar & Upapuyay 1988, Rat & VASANTH 1995).
SULOCHANA et al. (1991), however, remarked that the chemical substances present in
plants may not affect AMF colonization in roots. However, absence of AMF in roots
of medicinal plants may be due to non-availability of these fungi in particular soil
samples. Recent findings provide evidence that non-mycotrophic plant species lack
molecular signals necessary for stimulating AM fungi (SCHREINER & Kome 1993). It
seems that stimulatory compounds essential for AMF penetration of roots largely depend
upon the environmental factors, particularly pH, temperature, moisture, etc.

It can be concluded that the biotechnology of AMF may be applied not only for the
promotion of plant health, but also for the conservation of rare and endangered
Asteraceous plants by mycorrhization of tissue-culture-derived plants. This will help
to develop tolerance against transient transplant shock.

48 Comp. Newsl. 34, 1999

Acknowledgements

We thank Professor S. A. Brown, Principal, Danielson College, Chhindwara, for provi-
ding laboratory facilities, and SivaAtaH Karta for help in collection of plants.

References

Durca, V. V. K. & S. Gupta 1995. Effect of Vesicular arbuscular mycorrhiza on the
growth and mineral nutrition of Teak (Tectona grandis Linn. f.). Indian Forester
121 (1): 518-526.

Govinp Rao, Y. S., SurREsH, C. K., Suresu, N. S., MALLIKARJUNAIAH, R. R. & D. J.
BaGyaRAJ 1989. Vesicular arbuscular mycorrhizae in medicinal plants. Indian
Phytopath. 42: 476-478.

Keuri, H. K., CHANpRA, S. & S. MAHESHWARI 1987. Occurrence and intensity of
VAM in weeds, ornamentals and cultivated plants at Allahabad and areas adjoining
it. Mycorrhizae Round Table, Proceedings of the workshop held at Jawaharlal Nehru
University, New Delhi, 13-15 March, pp. 273-283.

Kumar Monan, V. & A. MAHADEVAN 1984. Do secondary substances inhibit
mycorrhizal association? Current Science 53: 377-378.

Mossg, B. 1973. Advances in the study of Vesicular Arbuscular Mycorrhiza. Annual
Review of Phytopathology 11: 171.

O’ Keer, D. M. & D. M. Sytvia 1991. Mechanisms of the vesicular arbuscular
mycorrhizal plant growth response. In: D. K. Arora et al. (eds.), Handbook of
Applied Microbiology, pp. 35 ff. Marcel Dekker, New York.

Pures J. M. & D. S. HayMan 1970. Improved procedure for cleaning roots and
staining parasitic and VAM fungi for rapid assessment of infection. Trans. British
Mycol. Soc. 55: 158-161.

Rat, M. K. 1993. Laboratory evaluation of fungitoxic activity of crude extract of
Parthenium hysterophorus. J. Environ. Biol. 14 (1): 41-44.

Rat, M. K. & Susum Urapuyay 1988. Screening of medicinal plants of Chhindwara
District against Trichophyton mentagrophytes, a causal organism of Tinea pedis.
Hindustan Antibiotics Bull. 30 (1-2): 33-36.

Rai, M. K. & S. Vasantu 1995. Laboratory evaluation of sensitivity of three
keratinophilic fungi to some vicolides. Hindustan Antibiotics Bull. 37 (1-4): 48-
50.

Comp. Newsl. 34, 1999 49

Raunkigr, C. 1934. The life form of plants and statistical plant geography: being the
collected papers of RAUNKIER, C. Clarendon Press, Oxford, England.

Reap, D. J., Koucnekt, H. K. & J. Hopcson 1976. Vesicular Arbuscular Mycorrhiza
in natural vegetation systems. New Phytologist 77: 641-653.

SAMBANDAN, K. 1995. Investigations on VAM association of Neem (Azadirachta indica).
Ph. D. Thesis, University of Madras, 140 pp.

ScHREINER, R. P. & R. T. Kome 1993. Antifungal compounds from the roots of
mycotrophic and non-mycotrophic plant species. New Phytologist 123: 99-105.

STREZEMSKA, J. 1975. Occurrence of mycorrhiza in cultivated plants. In:
Endomycorrhiza (Ed.: SANDERS et al.), pp. 537-543. Academic Press, New York.

SuLocuana, K. K., SHrIvapRASAD, P. & G. SHRIKANTAN Narr 1991. Occurrence of
Vesicular-Arbuscular Mycorrhiza in medicinal plants. Ancient Science of Life 1 (1
& 2): 38-39.

Varma, R. K. & JAMALUDDIN 1995. Associational activity of Arbuscular Mycorrhizae
of Teak (Tectona grandis) in Central India. Indian Forester 21 (6): 533-539.

Warcoup, J. H. 1990. The mycorrhizal association of some Australian Inuleae. Muelleria
7: 179-187.

Warcup, J. H. & P.A. McGEe 1983. The mycorrhizal association of some Australian
Asteraceae. New Phytologist 95: 667-672.

50 Comp. Newsl. 34, 1999

New combinations in Dendrophorbium and
Pentacalia (Senecioneae - Asteraceae) from Peru

HAMILTON BELTRAN
Museo de Historia Natural
Universidad Nacional Mayor de San Marcos
Av. Arenales 1256 Apartado 14-0434 Lima, Peru
e-mail: hamilton@musm.edu.pe

Abstract

Eight new combinations in Peruvian taxa of Dendrophorbium and Pentacalia are made.

Resumen

Se hacen ocho nuevas combinaciones para especies peruanas en los generos
Dendrophorbium y Pentacalia.

Introduction

Generic circumscription in Peruvian Senecioneae (Asteraceae) has been recently
modified (ROBINSON & CUATRECASAS 1992, 1993; BELTRAN & DE MERA 1997, 1998;
NorDENSTAM & Pruski 1995; Ditton & SacAstecut 1996, 1999). Due to the splitting
of the core genus Senecio into several segregates (JEFFREY 1992, a.0.), such as
Dendrophorbium, Aequatorium, and Pentacalia, a few Peruvian species in need of
transfer out of Senecio are here transferred to the correct segregate genera.

The Senecioneae tribe in Peru includes approximately 16 genera and 326 species,
with a high percentage of endemic species with a restricted distribution (VISION &
Ditton 1996). A tribal revision of Peruvian Senecioneae is currently being prepared
by Ditton & SAGASTEGUI (in prep.).

Study of the species of the Senecioneae from Peru was carried out in different Herbaria
such as US, F, USM, MICH, MO, and following to the classification of JEFFREY (1992)
it is necessary to transfer the following species of Senecio to Dendrophorbium and to
provide the correct name to Pentacalia balsasana according to ICBN.

Comp. Newsl. 34, 1999 51

New Combinations

Dendrophorbium argutidentatum (Cuatrec.) H. BELTRAN, comb. nov.
Basionym: Senecio argutidentatus Cuatrec., Collect. Bot. (Barcelona)
3: 281.1953. Typus: Peru; Cuzco, Urubamba, C. Varcas 2916A (F).

Dendrophorbium cosnipatense (CABRERA) H. BELTRAN, comb. nov.
Basionym: Senecio cosnipatense CABRERA, Notas Mus. La Plata, Bot. 18:
236. 1955. Typus: Peru; Cuzco, Paucartambo, O. VELARDE 1451 (LP).

Dendrophorbium goodspeedii (CuatrEc.) H. BELTRAN, comb. nov.
Basionym: Senecio goodspeedii Cuatrec., Fieldiana, Bot. 27(2): 51. 1951.
Typus: Peru; Huanuco, Pillao, A. Tuesta Diaz & F. WoyTKowski 34097 (F).

Dendrophorbium Ilewelynii (Cuatrec.) H. BELTRAN, comb. nov.
Basionym: Senecio llewelynii Cuarrec., Fieldiana, Bot. 27(2): 45. 1951.
Typus: Peru; Amazonas, Chachapoyas, L. WiLLiaMs 7589 (F).

Dendrophorbium trigynum (Cuatrec.) H. BELTRAN, comb. nov.
Basionym: Senecio trigynus CuaTREC., Brittonia 8: 189. 1956. Typus: Peru;
Cuzco, Cr. Gay 20351 CP):

Dendrophorbium vanillodorum (CABRERA) H. BELTRAN, comb. nov.
Basionym: Senecio vanillodorus CABRERA, Notas Mus. La Plata, Bot. 18: 238.
1955. Typus: Peru; Cajamarca, Celendin, C. A. Ripoutt 512 (LP).

Dendrophorbium vargasii (CABRERA) H. BELTRAN, comb. nov.
Basionym: Senecio vargasii CABRERA, Notas Mus. La Plata, Bot. 9: 225.
1944. Typus: Peru; Cuzco, La Convencion, C.Varcas 3481 (Herb. Cabrera).

Pentacalia petiolincrassata (CABRERA & ZARDINI) H. BELTRAN, comb. nov.
Basionym: Senecio petiolincrassatus CABRERA & ZARDINI, Bol. Soc. Argent.
Bot. 16: 383-385. Fig. 4. 1975. Typus: Peru; Cajamarca, Hualgayoc Tahona,
R. FERREYRA 8518 (LP). Pentacalia balsasana Cuatrec. & H. RoBINson,
Novon 3(3): 293. 1993. Typus: Peru; Cajamarca, Celendin, Canyon Balsas,
HuTcHINSON & Wricut 5212 (US, F, USM, MICH, UC).

Acknowledgements

I thank J. Pruski for the revision of this manuscript and H. Rosson for his valuable
comments on Pentacalia balsasana.

52 Comp. Newsl. 34, 1999

References

BELTRAN, H. & G. DE MERA 1997. Senecio icaensis sp. nov. (Asteraceae) un nuevo
endemismo de las lomas costaneras del Peru. Anales Jard. Bot. Madrid 55(1):
168-170.

BELTRAN, H. & G. DE Mera 1998. Senecio larahuinensis sp. nov. (Asteraceae) una
nueva especie de los andes peruanos. Anales Jard. Bot. Madrid 56(1): 168-169.

Ditton, M. & A. SacAstecur 1996. Revision of the dioecious genus Chersodoma
Put. (Senecioneae, Asteraceae), including a new species and status change. Brittonia
48(4) : 582-604.

Ditton, M. & A. SacAstecut 1999. Caxamarca, New Monotypic Genus of Senecioneae
(Asteraceae) from northern Peru. Novon 9: 156-161.

JEFFREY, C. 1992. The tribe Senecioneae (Compositae) in the Mascarene islands with
an annotated world check-list of the genera of the tribe. Notes on Compositae VI.
Kew Bull. 47(1) : 49-109.

NorpDENSTAM, B. & J. F. Pruski 1995. Additions to Dorobaea and Talamancalia
(Compositae-Senecioneae). Comp. Newsl. 27: 31-42.

Rosinson, H. & J. Cuarrecasas 1992. Additions to Aequatorium and Gynoxys
(Asteraceae: Senecioneae) in Bolivia, Ecuador and Peru. Novon 2: 411-416.

Rosinson, H. & J. Cuatrecasas 1993. New species of Pentacalia (Senecioneae:
Asteraceae) from Ecuador, Peru, and Bolivia. Novon 3: 284-301.

Vision, T. J. & M. Ditton 1996. Sinopsis de Senecio L. (Senecioneae, Asteraceae)
para el Peru. Arnaldoa 4(1): 23-46.

Comp. Newsl. 34, 1999 39

Reclassification of Prenanthes pendula
(Asteraceae: Lactuceae)

ALEXANDER N. SENNIKOV & IRENE D. ILLARIONOVA
Herbarium
Komarov Botanical Institute
Prof. Popov str. 2
Saint Petersburg, 197 376 Russia
e-mail: sennikov @herb.bin.ras.spb.ru

Abstract

The species with uncertain taxonomic position, Prenanthes pendula Scu. Bir., was
studied in respect of morphology, anatomy and surface ultrasculpture of achenes. The
structure of its pericarp is similar to that in species of Sonchus subgen. Dendrosonchus
to which this species is to be attributed.

Introduction

The taxonomic position of P. pendula, endemic to the Canary Islands, has always
been unclear. First described as a member of the genus Prenanthes (WEBB & BER
THELOT 1842-1850), this species remains static in the Lactuceae (Cichorieae), even
though the protologue of this name includes the remark by P. Wess on its possible
relationship to Sonchus: “Sonchus (Picrosonchus) pendulus Wess, in litt.” (1. c.: 421).

This characteristic species is the only member of Prenanthes from the Canary Islands,
inhabiting mountain cliffs on Gran Canaria (BRAMWELL & BRAMWELL 1974). This spe-
cies is characterized by a woody basal stem, pinnatisect leaves with large triangular
lobes, corymbose inflorescence with small calathidia of 5—6 florets (each), smooth, 4—
5-striate achenes and thin white subglabrous submonomorphic pappus setules. Con-
sidering its special habit, P pendula is rather similar to Canarian species of Sonchus
sect. Atalanthus (D. Don) DC. (syn. Taeckholmia Boutos) and S. tuberifer SVENT.
(sect. Tuberiferi BouLos).

In recent years doubt was thrown upon the former position of P. pendula. Some
morphological and biological data (PEREZ DE Paz 1976) facilitated the suggestion that
P. pendula is closely related to Sonchus and Sventenia (small monotypic segregate
close to Sonchus). A naturally occurring hybrid between Sventenia bupleuroides Font

54 Comp. Newsl. 34, 1999

Quer and P. pendula was reported by Svententus (1960). The relationships drawn
from the results of the study of sequences from the internal transcribed spacer region
of nuclear ribosomal DNA (Kim et al. 1996) connect P. pendula with Sventenia,
Babcockia platylepis (WEBB) BouLos (= Sonchus platylepis WEBB) as well as S. sect.
Tuberiferi, S. subgenus Dendrosonchus, and Taeckholmia. To further clarify the posi-
tion of P. pendula, we have undertaken studies of morphology and anatomy of its
fruits. Carpological method is well-established in systematics of Asteraceae (cf. Lavi-
ALLE 1912, and for example, Pak & Kawano 1990 a, b), and usually provides good
results in studies on Lactuceae.

Materials and Methods

The achenes of P. pendula were studied in respect of morphology, anatomy and sur-
face ultrasculpture. The achenes were taken from herbarium specimen, the syntype of
P. pendula: “In montibus Canariae, WEBB” (LE).

The achenes were treated in an alcohol-glycerine-water solution (1:1:1) during three
days and cut with freezing microtome through their middle (through cotyledons) and
basal parts. Sections were made at a thickness of 10-16 mu; prepared slices were
coloured with safranin. Drawings were made with drawing device RA-7 by the second
author. Surface ultrasculpture was studied with scanning electron microscope JSM-
35C.

Morphological and Anatomical Data
Fruit

The mature achenes of P. pendula are homomorphic, 44.5 mm long., 1.1—-1.2 mm
wide, stramineous, cylindrical, broadly conical at the base and rounded at the apex,
round in cross section, smooth, 4—5-striate with furrows (Fig. 1). Pappus 2.5-3 mm
long, uniseriate or partly biseriate, subhomomorphic. Pappus setules are 0.02—0.03
mm thick, most of them are thin with few slightly thickened ones, white, slightly
denticulate, rather fragile, slender.

Pericarp

Cross section of mature achenes shows more or less round outline (Fig. 2). The pericarp
is 80-100 mu in thickness. The layout of the achene is 4-5- merous (from achenes in
the same calathidium). Pericarp is clearly divided in section into 4—5 lobes with narrow
furrows. It is composed by exocarp and well differentiated mesocarp. Each mesocarp

Comp. Newsl. 34, 1999 55

lobe contains three strands of sclereids, which are separate, round and nearly equal in
the base of achene, and fused into trilobed band in the center of the achene. This band
is homogeneous, consisting of 10—12 layers of strands and of (1) 34 layers between
them, underlain by obliterated parenchyma cells (Fig. 3). One or two layers of the
elongated sclerified parenchyma with prominent pores in the walls overlay the
sclerenchyma bands and converge between them. Exocarp consists of a single layer of
epidermic tissue. The epidermic cells are rather narrow, 4—6 mu high, 8-12 mu wide,
cuticularized, with concave upper walls.

Seed Coat
There are some rows of the seed coat cells obliterated, being present as cell walls only.
Endosperm

The endosperm is two-layered, consisting of thin-walled cells with grained substance
within.

The study of ultrasculpture of achene surface shows the epidermic cells to be rather
narrow, 50-120 mu long, 8-12 mu wide, elongated in axial direction, without apical
outgrowths (Fig. 4). The cell surface is slightly rugose, without prominent cuticle
structures.

Discussion

Analysis of morphological and anatomical data provides more evidence in favour of
the inclusion of P. pendula into the subtribe Sonchinae K. BREMER (BREMER 1993,
1994), rather than keeping it within Prenanthes. All species of Sonchinae possess
achenes with pericarp divided in section into 4-5 lobes with rather prominent (at least
at the base of achenes) furrows, complemented with soft or slender, heteromorphic or
subhomomorphic pappus setules (SENNIKOv & ILLARIONOVA, in press) whereas species
of Prenanthes (SENNIKov 1997, in press) have achenes without furrows, with hard,
homomorphic pappus setules (SENNIKov & ILLARIONOVA, in press). Plant morphology
allows us to suggest P. pendula be compared with S. tuberifer by its pinnatisect leaves
with large lobes, as well as with S. sect. Atalanthus by small calathidia, inflorescence
shape and basally woody stem. Judging by the most distinguishing feature in Lactuceae,
S. platylepis (sect. Babcockia (BouLos) SENNIK.) is the species most closely similar to
P. pendula by the structure of its achenes. The achenes of S. platylepis have 4 pericarp
lobes without ribs (WEBB & BERTHELOt 1842-1850); every lobe possesses 5 subequal
sclerenchymatous strands (ALDRIDGE 1976). In our opinion, P. pendula is characterized

56 Comp. Newsl. 34, 1999

by arather primitive achene structure of Sonchus-type, because the pericarpium is not
differentiated into ribs though the number of sclerenchymatous strands is reduced
from 5 (primarily) to 3 (as in S. leptocephalus Cass. from sect. Atalanthus), and the
strands are partly fused. Thus, this species shares the most important features of some
species of subgen. Dendrosonchus and may belong there. We support the opinion to
unite some segregates close to Sonchus with Sonchus s. str. into a single genus according
to morphological and anatomical features, and to treat the subgenus Dendrosonchus
in broader sense (WEBB & BERTHELOT 1842-1850, ALDRIDGE 1976, REIFENBERGER &
REIFENBERGER 1996). To this treatment, P. pendula should be placed into subgen.
Dendrosonchus, forming a section of its own.

Nomenclatural Conclusions

Sonchus sect. Chrysoprenanthes (Scu. Br.) SENNIK., comb. nov.
Basionym: Prenanthes subgen. Chrysoprenanthes Scu. Br. in WEBB et BERTH.,
Hist. Nat. Iles Canar. 3, 2, 2: 420 (1849).
Type: Prenanthes pendula Scu. Br. (= Sonchus pendulus (Scu. Bip.) SENNIK.).

S. pendulus (Scu. Bir.) SENNIK., comb. nov.
Basionym: Prenanthes pendula Scu. Br. in Wes et BERTH., Hist. Nat. Iles
Canar. 3, 2, 2: 421 (1849). — Syntype: “In montibus Canariae, Wess” (LE!).

References

Avprince, A. E. 1976. Anatomy and evolution in the Macaronesian Sonchus sub-
genus Dendrosonchus (Compositae: Lactuceae). Bot. Journ. Linn. Soc. 76 (4):
349-285.

BRAMWELL, D. & Z. BRAMWELL 1974. Wild flowers of the Canary Islands. Stanley
Thornes (Publishers) Ltd., London.

Breme_r, K. 1993. New subtribes of the Lactuceae (Asteraceae). Novon 3: 328-330.

Bremer, K. 1994. Asteraceae: Cladistics and Classification. Timber Press, Portland,
Oregon.

Ku, S.-C., CRAWFORD, D. J. & R. K. JANSEN 1996. Phylogenetic relationships among
the genera of the subtribe Sonchinae (Asteraceae): Evidence from ITS sequences.
Syst. Bot. 21 (3): 417-432.

LAVvIALLE, P. 1912. Recherches sur le developpement de |’ovaire en fruit chez les
Composées. Ann. Sci. Nat. (Paris). Sér. 9, 15: 39-149.

Comp. Newsl. 34, 1999 57

Pak, J.-H. & S. Kawano 1990 a. Biosystematic studies on the genus /xerits (Compositae
—Lactuceae), I. Fruit wall anatomy and its taxonomic implications. Acta Phytotax.
Geobot. 41 (1-3): 43-60.

Pak, J.-H. & S. Kawano 1990 b. Biosystematic studies on the genus Jxeris and its
allied genera (Compositae — Lactuceae), [I]. Fruit wall anatomy and karyology of
Crepidiastrum and Paraixeris, and their taxonomic implications. Acta Phytotax.
Geobot. 41 (3-6): 109-128.

PEREZ DE Paz, J. 1976. Observaciones sobre la biologia y relaciones de Sventenia
bupleuroides F. Q. Botanica Macaronesica 1: 51-65.

REIFENBERGER, U. & A. REIFENBERGER 1996. Vereinigung der Gattung Taeckholmia
BouLos mit der Gattung Sonchus L. (Compositae) und Emendation des Subgenus
Dendrosonchus Scu. Br. Feddes Repert. 107 (5-6): 311-319.

Sennikov, A. N. 1997. The species of the genera Prenanthes and Cicerbita group
(Asteraceae) in the Caucasus. Bot. J. (St. Petersburg) 82 (2): 106-114 (in Russian).

Sennikov, A. N. (in press). De generibus ex affinitate Prenanthes L. (Asteraceae).
Novit. Syst. Pl. Vasc. (St. Petersburg) (in Russian).

Sennikov, A. N. & I. D. ILLartonova (in press). Morphological and anatomical structure
of achenes in the genus Sonchus s. |. (Asteraceae). Bot. J. (St. Petersburg) (in Russ-
ian).

SvENTENIUS, E. R. S. 1960. Additamentum ad floram canariensem. Madrid, Spain.

Wess, P. & S. BERTHELOT 1842-1850. Histoire naturelle des Iles Canaries. 3 (2, sect.
2): 1-496, Pt. 37-136 b.

58

Comp. Newsl. 34, 1999

WLI KU X20

Fig. 1. An achene of P. pendula (SEM micrograph), x 20.

Comp. Newsl. 34, 1999 59

Fig. 2. Simplified cross section of achene of P. pendula. A: at the base part of the
achene. B: at the middle part of the achene. Sclerenchyma tissue is crossed.

60 Comp. Newsl. 34, 1999

C
EP
SP
SC ons
ADOBE
[ELIXSO
— 1eeSess
BNIB Osoee :
ese: COAL,
Sad NSCs es
SASSI
SPN SIO}SIG
meenagy SCORE
Veg. 23 FF ap
(EL. Yas es A
Vices Zz
PLETerX
i En (0}
fj (OXQ?
) eOGa
He To
oIO10)
Neos
Ae aoulo
VERS 0.025 mm
IES SoSoa

Fig. 3. Part of cross section of mature achene of P. pendula with anatomical details.
I— pericarp; II — seed coat obliterated; III - endosperm. Cuticle of epidermic cells (C);
epidermal cells of pericarp (EP); sclerified elongated parenchyma (SP);
sclerenchymatous cells of pericarp (SC).

Comp. Newsl. 34, 1999

Fig. 4. Structure of achene surface of P. pendula (SEM micrograph), x 200.

61

62 Comp. Newsl. 34, 1999

Phytosociological investigation of
Compositae weeds in abandoned farmlands
in Ekiti State, Nigeria

JosHuA KayopE
Department of Botany
Ondo State University

Ado Ekiti, Nigeria

Abstract

Weed species belonging to the family Compositae were studied among other plant
weeds present in three abandoned farmlands in Ekiti State, Nigeria. Tridax procumbens,
Chromolaena odorata and Ageratum conyzoides are the dominant Compositae weed
species in the study area.

Introduction

In southwestern Nigeria, the prevalent agricultural system has been, and will most
likely continue to be, shifting cultivation. After the fields have been cultivated for one
or two years, they are left to fallow, and the fallow areas are immediately invaded by
a dense mass of weeds, most of which are members of the family Compositae.

The various colonization strategies employed by weeds have been widely studied but
a limited number of quantitative studies on their phytosociology and ecology have
been published. In Nigeria some investigations so far reported are those of AFOLAYAN
(1988) and AFOLAYAN & AYODELE (1990).

The present study was undertaken to elucidate the phytosociology of Compositae
weeds in Ekiti State of southwestern Nigeria.

Materials and Methods

The study area is situated in Ekiti State, between latitude 7°25’ and 8°20’N and longitude
5°00’ and 6°00’E. This state has a contiguous land mass of about 7,000 sq. kilometers
and over 75 % of its 1.6 million inhabitants are farmers (EKSG 1997). There are two
climatic seasons, viz., a rainy season which lasts from March to October, and a dry

Comp. Newsl. 34, 1999 63

season which lasts from November to February, and the annual rainfall is about
1150 mm (KayopeE & Favuyi 1994).

Methods: Three study sites designated A, B and C were chosen within 20 km radius
from Ado Ekiti, the Ekiti State capital. Site A was an abandoned farm in Ado Ekiti,
Site B was an abandoned farm in Ikere-Ekiti about 20 km south of the state capital,
and Site C was an abandoned farm in Ifaki-Ekiti, about 20 km north of the state capi-
tal.

In each site, a 100 m by 100 m study plot was investigated in January, 1998. Prior to
the study, each of the sites had been cropped continuously for a period of two to three
years for cassava, yams, maize, rice or vegetables, alone or in mixtures.

Also in each site, a quadrat sized 50 cm by 50 cm was tossed fifty times, and Compositae
weeds in each quadrat were observed, identified and recorded on an abundance scale
defined by Boncers et al. (1988) as follows: Less than 5 individuals as rare (R); 5 to 10
as occasional (O), 11 to 30 as frequent (F), 31 to 100 individuals as abundant (A), and
over 100 individuals as very abundant (VA).

The importance value index (IVI), which is the sum total of the relative density, relative
frequency and relative basal area, was also determined for each Compositae weed
species.

The Index of Similarity (IS) between the sampling sites, i.e. Sites A and B, A and C,
and B and C, were determined as

IS = 2C x 100/A +B
where A is the number of species in one site; B is the number of species in the other
site; C is the number of species common to both sites.

Results and Discussion

The results obtained revealed that a total of 36 different weed species were sampled in
the three study sites, out of which 21 weedy species were members of Compositae.
Table 1 shows that Compositae made up of over 60 % of the total number of weed
species and the numbers of weed individuals in the study area.

The ten most dominant Compositae weed species in the study area are shown in Table
2. The species were ranked in order of their decreasing IVI value. Tridax procumbens,
Chromolaena odorata and Aspilia africana were the most dominant Compositae weeds
in the study area, with IVI values of 89.26, 46.62 and 44.19 respectively.

64 Comp. Newsl. 34, 1999

On the abundance scale, these dominant species were very abundant in all the three
sites sampled in this study (Table 2).

The indexes of similarity between sites A and B, A and C, and B and C were 82.05 %,
89.47 % and 92.68 % respectively (Table 3). These values revealed that the Compositae
weed species were common to all the serial communities sampled.

The predominance of Tridax procumbens in the study area could be a function of its
method of dispersal which occurs by wind. Previous investigation by AFOLAYAN (1988)
had also revealed that this weed easily adapts itself to the climatic and edaphic conditions
prevailing in the study area. Apart from this rain of seeds, it could also be a function of
Tridax seeds stored already in the soil during cultivation of the fields, and the dormancy
of which is easily broken by light.

Chromolaena odorata is an efficient colonizer of disturbed sites (Epp 1987). Its
predominance in the study area could be attributed to its rapid dispersal by wind which
usually occurs between January and March, and its ability to establish easily in the
study area (ETEJERE 1980). Also previous research efforts had established that a bank
of C. odorata seeds exists in the soil of early successional communities (RAMAKRISHNAN
& Misura 1982). Longevity of these seeds which is increased by enforced dormancy
after burial in the soil (Yapav & TripaTHr 1982) is broken in full sunlight (AULD &
Martins 1975).

Similarly, Ageratum conyzoides 1s widely known to appear in the month of January
(Gupta & Mukersi 1995), which is the month when this study was carried out. Thus its
appearance at this time of the year explains its predominant status.

The indexes of similarity between the study sites (Table 3) show that a lot of the
Compositae weeds were common to the three sites. Thus it could be inferred that areas
with the same climatic conditions will enhance the growth of related weed species. In
conclusion, phytosociological information on the Compositae weed species in a given
vegetation type will surely contribute to the comprehensive understanding of their
ecology.

Comp. News!l. 34, 1999 65

References

Aro.ayan,A. J. 1988. Phytosociological investigations of three abandoned farmlands
in Oyo, Ondo and Kwara States of Nigeria. Nig. J. Weed Science 1: 77-82.

Aro.ayaNn, A. J. & G. AYODELE 1990. Phytosociophysiological study of the weed
flora of cassava (Manihot esculenta CRANTZ) in Kwara State, Nigeria. Bioscience
Research Communications.

AuLp, B. A. & P. M. Martins 1975. The autecology of Eupatorium adenophorum
SprENG. in Australia. Weed Research 15: 27-31.

BoncERs, F., Popma, J., MEAVE DEL CasTILLo, J. & J. CARABIAS 1988. Structure and
floristic composition of the lowland rain forest of Los Tuxtlas, Mexico. Vegetatio
74: 55-80.

EKSG 1997. First Anniversary Celebrations of Ekiti State Government. 22 pp.

Epp, G. A. 1987. The seed bank of Eupatorium odoratum along a successional gradi-
ent in a tropical rainforest in Ghana. Journal of Tropical Ecology 3: 139-149.

ETEJERE, E. O. 1980. Viability of herbicide-treated seeds of Eupatorium odoratum L.
Weed Research 20: 361-368.

Gupta, R. & K. G. Mukeryi 1995. Studies on weeds of Delhi III - Compositae.
Compositae Newsletter 27: 1-6.

Kayope, J. & M.A. Favuyr 1994. Studies on self and cross compatibility in soyabeans
(Glycine Max) in a tropical environment. Nigerian Journal of Botany 7: 55-61.

RAMAKRISHNAN, P. S. & B. K. Misura 1982. Population dynamics of Eupatorium
adenophorum Sprena. during secondary succession after slash and burn agriculture
in Northeast India. Weed Research 22: 77-84.

Yapav, A. S. & R. S. Tripatut 1982. A study on seed population dynamics of three
weedy species of Eupatorium. Weed Research 22: 69-76.

66 Comp. Newsl. 34, 1999

Table 1. Demographic features of weeds in abandoned farmlands in

Ekiti State, Nigeria.
Total number weed species/site 29 33

% Composition of Compositae weed species

to the total number of weed species/site 62 % 64 %

a
Number of Compositae weed species /site 18 21

Total number of weed individuals/site 8387 8349 9200 | 26936

CO ge IS 12 5638 | 5677 | 5520 | 16565
individuals/site

Percentage of Compositae weeds of total 64% | 68% | 60% | 64%
number of weeds

67

Comp. Newsl. 34, 1999

‘A[UO spoom ae}IsOduso_ ay} UO peseq paye[No[ed sonjeA “xepuy anjeA sourylodwy =J[AI = +x*
‘o1ey = Y pue [euoised009 = O ‘juonbelJ = J Wuepunqy = Vy -juRpuNnqe AIQA =VA ae
quiys=S ‘19H = H **

2: NUgIH 3 AIO MYyjanfuiamyos snyouog
NO ‘5D (WIS) 0219909 D1IjIWy
aNVIg CAVY) VIV1]19 DsosuljVH
“TAC DUIDpsAwDd DIUOULIA
‘Od CT) Diofyouos Dip
"7 vsojid suapig

“BLIOSIN (2381S HPI Ul spuepuLiey pauopuege ut satvads paam aezIsoduio. yUBUTWIOp Ua} JO SaINqIIWAY §°7 PIQEL

68 Comp. Newsl. 34, 1999

Table 3. Index of similarity (IS) between the three sample sites.

Comp. Newsl. 34, 1999 69

New taxa and combinations
published in this issue

Angelphytum apense (Cuopat) PRuski, comb. nov.: p. 2.
Angelphytum herzogii (HASSLER) PRUsKI, comb. nov.: p. 2.

Dendrophorbium argutidentatum (Cuatrec.) H. BELTRAN, comb. nov.: p. 51.
Dendrophorbium cosnipatense (CABRERA) H. BELTRAN, comb. nov.: p. 51.
Dendrophorbium goodspeedii (CuatTrEc.) H. BELTRAN, comb. nov.: p. 51.
Dendrophorbium Ilewelynii (Cuatrec.) H. BELTRAN, comb. nov.: p. 51.
Dendrophorbium trigynum (Cuatrec.) H. BELTRAN, comb. nov.: p. 51.
Dendrophorbium vanillodorum (CABRERA) H. BELTRAN, comb. nov.: p. 51.

Dendrophorbium vargasii (CABRERA) H. BELTRAN, comb. nov.: p. 51.

Monticalia abietina (WiLLD. ex WeEpp.) C. JEFFREY var. aciculata (Cuatr.) B. Norb.,
comb. nov.: p. 31.

Monticalia albi-panquei (Diaz & Cuatr.) B. Norp., comb. nov.: p. 34.
Monticalia almorzana (Diaz & Cuatr.) B. Norp., comb. nov.: p. 33.
Monticalia cacaosensis (Cuatr.) B. NorD., comb. nov.: p. 34.

Monticalia carrikeri (Cuatr.) C. JEFFREY var. macrophylla (Diaz) B. Norp., comb.
nov.: p. 32.

Monticalia carupana (Diaz) B. Norpb., comb. nov.: p. 32.
Monticalia elatoides (WEvp.) B. Norp., comb. nov.: p. 31.
Monticalia fimbriifera (Diaz & MENDEZ) B. Norb., comb. nov.: p. 35.

Moniticalia guadalupe (Cuatr.) C. JEFFREY subsp. caucana (Cuatr.) B. Norp., comb.
nov.: p. 35.

Monticalia hammenii (Diaz & Cuatr.) B. Norp., comb. nov.: p. 34.

Monticalia harrietae (Cuatr.) B. Norb., comb. nov.: p. 31.
Monticalia iguaquensis (Diaz & Cuatr.) B. Norp., comb. nov.: p. 33.

70 Comp. Newsl. 34, 1999

Monticalia juajibioyi (Diaz & Cuarr.) B. Norp., comb. nov.: p. 30.

Moniticalia ledifolia (H. B. K.) C. JEFFREY var. glabrata (Diaz & MENDEZ) B. Norp.,
comb. nov.: p. 34.

Monticalia ledifolia (H. B. K.) C. JerFrey var. lanata (H. B. K.) B. Norp., comb.
nov.: p. 33.

Monticalia pulchella (H. B. K.) C. JEFFREY subsp. guantivana (Cuatr.) B. Norp.,
comb. nov.: p. 32.

Monticalia romeroana (Diaz & BUENO) B. Norp., comb. nov.: p. 30.
Monticalia schultzei (CuaTr.) B. Norp., comb. nov.: p. 30.

Monticalia schultzei (Cuatr.) B. Norp. subsp. sevillana (Cuatr.) B. Norp.,
comb. nov.: p. 31.

Monticalia subarachnoidea (WeEpp.) C. JEFFREY var. pauciflora (Diaz & BuENO)
B. Norp., comb. nov.: p. 30.

Monticalia summapacis (Cuatr. ) B. Norp., comb. nov.: p. 33.
Monticalia taironae (Diaz & Cuatr.) B. Norb., comb. nov.: p. 34.

Moniticalia tolimensis (Cuatr.) C. JEFFREY var. colombiana (Cuatr.) B. Norp., comb.
nov.: p. 35.

Monticalia vaccinioides (H. B. K.) C. JEFFREY var. microdentata (Cuatr.) B. Norp.,
comb. nov.: p. 32.

Oyedaea cuatrecasasii PRUSKI, sp. nov.: p. 3.

Oyedaea neei PRUSKI, sp. nov.: p. 5.

Pentacalia petiolincrassata (CABRERA & ZARDINI) H. BELTRAN, comb. nov.: p. 51.

Sonchus sect. Chrysoprenanthes (Scu. Brr.) SENNIK., comb. nov.: p. 56.

Sonchus pendulus (Scu. Bre.) SENNIK., comb. nov.: p. 56.

im: ; bovit head} au Gare i \ te H hiv at

| ’
ete, 6 i aor i (wi) Lae, Ve

Sony Ta aby nn Aj wees ve Spe oll Pelli edi Ey RAT TA

Oe ee

ae ew " | | -

LIBRARY

JAN 3 2000

NEW YORK
BOTANICAL GARDEN