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“COMPOSITAE &
* NEWSLETTER

July 1998

Number 32

Scientific Editor: Berta. NORDENSTAM
Technical Editor: GUNNEL WirENtUS NOHLIN

Published and distributed by The Swedish Museum of Natural History,
Department of Phanerogamic Botany, LIBR A RAR Y

P.O. Box 50007
S-104 05 Stockholm, Sweden
(Director: Professor Berta. NoRDENSTAM)
ISSN 0284-8422 SEP if 8 1998
NEW YORK
BOTANICAL GARDEN

CONTENTS

H. Tunon: Alkaloids in Achillea millefolium L. - confusion in the literature

P. Suort: Notice of type speciemens of Dahlia Cav. in the National Herbarium

of Victoria (MEL) 8
M. Denarteis: Chromosome studies on Vernonia flexuosa and V. lithospermifolia 10 ~
A. Sotpano: New names and combinations and other nomenclatural notes for
Compositae of various countries 19 Meet
M. Ipu & L.S. Git: Nature of ergastic substances in some Asteraceae seeds - VII 23
M. Ibu & C. A. Omonuinmw: Allelopathic effect of Calotropis oe on the
germination of Helianthus annuus seeds 31
M. Ipu & C. A. OMonHINMIN: Germination control of Helianthus annuus L.

37

using two growth regulators - thiourea and coumarin

NEW NAMES AND COMBINATIONS PUBLISHED IN THIS ISSUE

Centaurea aksamica SOLDANO, nom. nov.: p. 17

Inula stenocalathia (REcu. F.) SOLDANO, comb. nov.: p. 19

Senecio linaresensis SOLDANO, nom. noy.: p. 18

Senecio neoviscidulus SOLDANO, nom. nov.: p. 18

Senecio nublensis SOLDANO, nom. nov.: p. 19

Senecio pemehuensis SOLDANO, nom. nov.: p. 19

Seriphidium densiflorum (Viv.) SOLDANO, comb. nov.: p. 19
Stemmacantha caulescens (Cosson et BALANSA) SOLDANO, comb. nov.: p. 19
Stemmacantha imatongensis (PHILIPSON) SOLDANO, comb. nov.: p. 19
Tanacetum musili (VELEN.) SOLDANO, comb. nov.: p. 20

Wedelia ciliata (ScHuM.) SOLDANO, comb. nov.: p. 20

Wedelia kotschyi (Scu. Bir. ex Hocust.) SOLDANO, comb. nov.: p. 20

Comp. Newsl. 32, 1998 1

Alkaloids in Achillea millefolium L.
— confusion in the literature

HAKAN TUNON
Division of Pharmacognosy
Department of Pharmacy
Biomedical Centre
Uppsala University
P.O. Box 579, S-751 23 Uppsala, Sweden

Abstract

There is some confusion in the literature about alkaloids in Achillea millefolium L.
(Asteraceae) and the purpose of this article is to clarify and correct this situation. In
summary there are so far only four alkaloids that have been isolated from A. millefo-
lium; betaine, betonicine (= achilleine), choline (or choline chloride) and stachydrine
(= leonucardine). A fifth alkaloid, trigonelline, has been detected by paper chroma-
tography but not isolated. A sixth, homostachydrine (moschatine or N-methy] piperi-
din-2-carboxylic acid methylbetaine), has not been isolated from A. millefolium, but
from A. moschata WuLF.

Keywords: Achillea millefolium, alkaloids, achiceine, achilleine, achilletine, betaine,
betonicine, choline, homostachydrine, moschatine, stachydrine, trigonelline.

Introduction

Yarrow, Achillea millefolium, is a common medicinal plant that grows all over the
northern hemisphere, primarily throughout the temperate and boreal zones. It has
tradtionally been used for a variety of medical purposes, such as treating wounds and
curing different kinds of stomach ailments’. Extracts from the plant have shown
different types of pharmacological activites, including anti-inflammatory and
antispasmodic effect. However, in the literature that deals with the alkaloids of A.
millefolium, there is some contradictory information. One review article lists some 8
alkaloids’ but a more careful approach to the literature would indicate that this is an
overestimation. The purpose of this brief survey is to correct the literature.

There are only four alkaloids that have been isolated from Achillea millefolium

"References, se page 4-5

2 Comp. Newsl. 32, 1998

according to the literature: betaine, betonicine, choline and stachydrine. Trigonelline
has been detected by paper chromatography in a crude mixture. The structures of
these compounds are shown in Fig. 1.

Survey of literature

The different studies dealing with alkaloids from Achillea millefolium are summarized
in Table 1 which is arranged in chronological order.

One paper by Zirvi & Ikram’ claims that A. millefolium contains the following
alkaloids: achiceine (C,,H;;NO,), achilletine (no formula presented), achilleine
(C,4H,3NO;), choline chloride (C;H,,NOCI), homostachydrine (C,gsH,;NO>, this must
be a misprint and should be CsH,;NO,) and stachydrine (C;H;;NO,). The same paper
also reports that moscatine (C,,;H27NO;) and trigonelline (C;H;NO,) are constituents
of Achillea species. According to a more recent review’ A. millefolium contains:
achiceine (C,,H,,;NO,), betaine (C;H,,NO,), betonicine (C;H,;NO;), choline
(C;H,,NO), homostachydrine (CsH,;NO;), moschatine (or moscatine) (C,;H27NO;),
stachydrine (C;H;;NO;) and trigonelline (C;H;NO>)

Many similarities are found in the reference lists of these authors. Zirvi & Ikram’
have mainly used different kinds of handbooks and reviews*’ but also some original
papers*’. The handbooks and reviews are often referring to each other and have
failed to consult the primary literature. CHANDLER et al.' on the other hand are using
original papers*” with two exceptions *’. However, much of the material mentioned
in Zirvi & Ikram?’ is neglected by CHANDLER et al.’

The first paper that describes isolation of an alkaloid from A. millefolium is ZANON”,
i.e. the isolation” of achilleine with a yield of 6.5 %. vON PLANTA-REICHENAU™ was
the first to question whether this was a pure substance or not. That report also describes
the isolation of two alkaloids from A. moschata and these were named achilleine (in
German achillein) (Cy>H3gN.O;;) and moschatine (in German moschatin) (C,,Hy;NO,).
When achilleine was hydrolyzed, it formed the alkaloid achilletine (C;,Hi7NOs,),
ammonia and a reducing sugar. This reaction is described in several alkaloid hand-
books (i.e. 6, 14, 15, 16). In 1928 H. SCHALLER in Ziirich explained that he could not
show the presence of achilleine in A. millefolium or A. moschata in spite of repeated
experiments’. According to the review article by Zirvi & Ikram’ both achilleine and
achilletine are constituents of A. millefolium and moschatine is considered to occur
only in A. moschata, but this is only based on literature data from different hand-
books.

In the 1950s Mier & Cuow’* isolated on alkaloid from A. millefolium and named it

Comp. Newsl. 32, 1998 3

achilleine, having the formula C,,H2sN2O,. The formula was later revised when PAILER
& Kume"' isolated an alkaloid with the same composition judged from the elemental
analysis buth with only half the molecular weight. In that paper the alkaloid was
studied both chemically and by infrared-spectroscopy and it was judged that achilleine
was identical to betonicine (C;H;;NQO;).

Pater & Kump’ isolated further alkaloids from both A. millefolium (betaine
[C5H,,NO,], choline [C;H,,NO] and stachydrine [Cj;H,;NO,]) and A. moschata
(homostachydrine [C,H,;NO,; N-methyl] piperidin-2-carboxylic acid methylbetaine]).
Homostachydrine is later called moschatine by HEGNAUER®. Homostachydrine is
also mentioned in KaRRER": ”L(-)-Homostachydrin ... aus Kraut von Achillea millefo-
lium L. und A. moschata WuLF. isoliert” [L(-)-Homostachydrine ... isolated from the
herbs of Achillea millefolium L. and A. moschata WutF.] with PALER & Kump’ as
reference. Merck Index" from 1989 does not have any entries on achiceine, achilleine,
achilletine, homostachydrine or moschatine; only betonicine is mentioned in connec-
tion with A. millefolium.

A subsequent study on the alkaloid content in A. millefolium was made by Ivanov &
YANKov™ who used an ion exchanger for a crude separation and paper chromatography
with reference compounds for the analysis. They managed to detect the already
reported alkaloids: betaine, betonicine, choline, stachydrine and another alkaloid
with an Rf value similar to trigonelline.

Both CHANDLER et al.' and Zirvi & Ikram’ refer to an alkaloid named achiceine.
CHANDLER et al. have used Zirvi & Ikram’ and Soxo.ov’ as sources while Zirvi &
IkraM only used SokoLov. Soxo.ov has on the other hand used two different sources’”””
written in Russian and these two references have not been available to me. However,
the most recent source” from 1939 is described in Chemical Abstracts as a screening
for alkaloid content in 259 different plants and no isolation work is mentioned.
SoKko ov’ writes that A. millefolium”’ contains the glucoalkaloid achilleine Cy9H3sN20}5
and the amorphous basic achiceine C,,H;;NO,”. The alkaloids isolated by ZaNoN””
and von PLANTA-REICHENAU” achiceine and achilletine are described as amorphous
bases which have the same molecular formula as the compounds described by
SoxoLov. It might be possible that SoxoLov’s two sources, LAZUREvski & SADYKOV™
and SHACK” are quoting ZANON and Von PLANTA-REICHENAU, but this, at present is
uncertain.

A more recent study demonstrated, by TLC-comparison with reference compounds
and the use of Dragendorff spray reagent, the presence of betaine, betonicine, sholine,
stachydrine and trigonelline in A. millefolium, while homostachydrine could not be
detected”’. A. millefolium is now considered to be a group of several closely related

4 Comp. Newsl. 32, 1998

species and a very recent article described the pattern of different betaines in species
of the aggregate. They investigated the presence of betaine, betonicine, choline, and
stachydrine in 46 individuals from 11 species of the A. millefolium group”.

However, some of the earlier papers are still quoted in scientific studies. Recently
errors concerning the alkaloids from A. millefolium appeared in different papers”.
Eran et al.” have studied A. santolina and have performed a microanalysis on an
isolated alkaloid. They determined the formula C,,H2.N2O, and reported that it was
”an expected formula similar to that of the alkaloid isolated from Achillea mille-
folium, named Achilleine (MILLER & CHow 1954). Therefore, it could be suggested
that the separated alkaloid in this work from Achillea santolina may be achilleine’”™.

PAILER & Kump" showed that achilleine was identical with betonicine, C;H,;NO3.

Conclusion

In conclusion the alkaloids that have been found in A. millefolium are until now:
betaine, betonicine, choline, stachydrine and probably trigonelline. All the other
alkaloids mentioned in the literature have either been isolated from other species
(e.g. A. moschata) or are synonyms for some of the above mentioned alkaloids.
Some of the earliest reported alkaloids might have been crude fractions and not pure
compounds.

References

CHANDLER, R. F., Hooper, S. N., Harvey, M. J. Econ. Bot. 36: 203—223 (1982).

2. NEWALL, C.A., ANDERSON, L. A., PaiLuirson, J. D. Herbal medicines - A guide
for health-care professionals. The Pharmaceutical Press, London (1996).

Zirvi, K. A., Ikram, M. Pakistan J. Sci. Ind. Res. 18: 93-115 (1975).

Bort, H. G. Ergebnisse der Alkaloid-Chemie bis 1960. Akademie-Verlag, Ber-
lin (1961).

5. H&GENAUER, R. Chemotaxonomie der Pflanzen. Birkhauser-Verlag, Basel and
Stuttgart (1964).

Henry, T. A. The Plant Alkaloids. 4" ed. J. & A. Churchill Ltd. London (1949).

Soxo ov, V. S. Alkaloid Plants of the U.S.S.R. Akad. Nauk., Moscow and Lenin-
grad (1952) [in Russian].

8. Miter, F. M. , Cuow, L. M. J. Am. Chem. Soc. 76, 1353-1354 (1954).

Comp. Newsl. 32, 1998 5

23.

24.

Pater, M., Kump, W. G. Arch. Pharm. 293, 646-654 (1960).

. Ivanov, C., YANKov, L. God. Vissh. Khimikotechnol. Inst., Sofia 1967. 14, 195-—

222 (1971). (Through CA 77:111471p.).

. Pamer, M., Kump, W. G. Monatschr. Chem. 90, 396-401 (1959).

. ZANON, B. Annal. Chem. Pharm. 21, 21-36 (1846).

. PLANTA-REICHENAU, V. A. Chem. Pharm. 155, 145-160 (1870).

. Grassy, J. S. Encyclopedia of the Alkaloids. Vol. 1. Plenum Press, New York

and London (1975).

. Trizr, G. Die Alkaloide - Eine Monographie der natiirlichen Basen. 2™ ed.

Verlag von Gebriider Borntraeger, Berlin (1931).

. WOLFFENSTEIN, R. Die Pflanzenalkaloide. 3" ed. Verlag von Julius Springer,

Berlin (1922).

. Karrer, W., CHERBULIEZ, E. EUGsTer, C. H. Konstitution und Vorkommen der

organischen Pflanzenstoffe. Vol. 1. Birkhauser Verlag, Basel und Stuttgart
(1977).

. BupavarI, S. (ED.). The Merck Index. 11th ed. Merck & Co., Inc. Rahway, N. J.,

USA (1989).

. Lazurevskul, G. V., SavyKov, A. S. Trudy Uzbekskogo Gosudarst, univ. Sbornik

Trudov Khim. 15, 182-196 (1939). (Through CA 35:4154[7]).

. SHacku, E. A. Studies on plant alkaloids, glucosides and ptomaines. Uch. zab.

Kazansk. veterin. inst. T. VI. Kazan (1889). (Not seen and studied).

. Tunon, H., THoRSELL, W., BOHLIN, L. Econ. Bot. 48, 111-120 (1994).

MEHLFUHRER, M., TROLL, K., JURENITSCH, J., AUER, H., KuBELKA, W.
Phytochem. 44, 1067-1069 (1997).

Erman, M. A., Hassan, A. B., Atra, A. H., Fanmy, G. E., Yousser, A. K.,
Nani, A. E. B. Egypt. J. Vet. Sci., 24, 181-189 (1987).

Miticu, L. W. Weed Technol. 4, 451-453 (1990).

6 Comp. Newsl. 32, 1998

Table 1. A summary of the literature dealing with the betaine-type alkaloids
from Achillea millefolium.

Achilleine isolated from A. millefolium (ZANon 1846)”.

Claimed that ZANon’s achilleine was not a pure substance and isolated two alkaloids
from A. moschata, named achilleine (C..H3,;N,O,;) and moschatine
(C,,Hx,;NO;). By hydrolysis of achilleine an alkaloid named achilletine
(C,,H,;7NO,) is formed (von PLANTA-REICHENAU 1870)”.

Failed to show the presence of achilleine in A. millefolium or A. moschata (SCHALLER
1928)”.

Literature-review. A. millefolium contains achilleine (C,)H;3N,O,;) and achiceine
(C1,;Hi;NO,) (SoKoLov 1952)’.

Isolated achilleine (C,,H2,N,O,) from A. millefolium (MILLER & Cow 1954)’.

Isolated achilleine and find that the formula is (C;H;;NO;). Spectroscopical studies
show that it is identical with betonicine (PAmER & Kump 1959)".

Betaine (C,H, ,NO,), choline (C,H, ,NO) and stachydrine (C,H,,NO,) from A. mille-
folium and homostachydrine (C,H, .NO,) from A. moschata (PAILER & Kump

1960)’.

Homostachydrine is named moschatine but still isolated from A. moschata (PAILER
& Kump 1960)’ according to HEGNAUER (1964)?.

Detected betaine, betonicine, choline, stachydrine and a new alkaloid trigonelline
(C,H;NO,) (Ivanov & YANKov 1971)".

Literature review. Claims that A. millefolium contains achiceine (SoKoLov 1952)’,
achilleine (MILLER & Cuow 1954)*, achilletine, betonicine, choline chloride,
stachydrine, (Handbooks), moscatine and trigonelline in Achillea sp. (Hand-
books) according to CHANDLER et al. (1982)'.

15

Comp. Newsl. 32, 1998 7

Fig. 1. The structures of the alkaloids definitively isolated from or identified in
Achillea millefolium and homostachydrine isolated from A. moschata.

a
-

Betonicine Stachydrine

“on

SA+

N

Trigonelline Homostachydrine

8 Comp. Newsl. 32, 1998

Notice of type specimens of Dahlia Cav. in the
National Herbarium of Victoria (MEL)

Pair SHORT
Northern Territory Herbarium
P.O.Box 496, Palmerston, Northern Territory 0831
Australia

Prior to moving from Melbourne to northern Australia I had reason to examine MEL’s
collection of Dahlia. As I recall there are only 20-30, mostly unmounted, specimens,
none of which was examined by SorENSEN (1969) during his revision of the genus.
All collections are from last century or earlier this century and some are undoubtedly
type specimens or of historical interest. My notes made at the time were not elaborate
and the status of each specimen was not thoroughly checked but for anyone interested
in the taxonomy and nomenclature of the genus Dahlia I draw attention to the follow-
ing specimens.

Dahlia barkerae KNowLes & Westc. — Type not seen by Sorensen. I recorded,
surprising as it may seem, that MEL has a ’likely type’ specimen.

Dahlia brevis SORENSEN — isotype at MEL.
Dahlia dissecta S. Wats. — isotype at MEL.

Dahlia merckii Leno. Specimen received from Hamburg Botanical Garden, the label
bearing the date 1844. The type specimen was originally grown in the aforementioned
garden, the species being described by LEHMANN in 1839. MEL’s specimen is perhaps
a type, depending on whether the date 1844 is the date of collection or the date of
receipt of the specimen.

Georgina coccinea (Cav.) WiLD. var. flava WILLD. — old specimen, perhaps type.
SORENSEN referred to WILLD., Hort. Berol. 2: pl. 96 (1809) as the lecotype specimen.
This may be an extant type specimen.

Georgina variabilis WuLp. — there are several old specimens, one ex ’herbario Dr H.
van Hemck’, with this name. One specimen is labelled as var. purpurea WILLD. So-
RENSEN made reference to published plates as being the lectotype specimens of formally
named varieties of G. variabilis. There is a possibility that there are extant types at
MEL.

Comp. Newsl. 32, 1998 9

The presence of the Dahlia specimens, at least those obtained last century from
Germany, is undoubtedly a result of the connections and efforts of the renowned
botanist, FERDINAND MUELLER. During his term as Government Botanist of Victoria, a
position he held from 1853 to 1896, MuELLER was responsible for the acquisition of
a number of private herbaria, including those amassed by Orro WILHELM SONDER and
JOACHIM STEETZ (SHortT 1990), and as such MEL often houses important collections
which may be overlooked by taxonomists.

References

Suort, P. S. 1990. Politics and the purchase of private herbaria by the National
Herbarium of Victoria. Jn: P. S. SHort (ed.), History of systematic botany in
Australasia (Australian Systematic Botany Society Inc.) pp. 5-12.

SORENSEN, P. D. 1969. Revision of the genus Dahlia (Compositae, Heliantheae —
Coreopsidinae). Rhodora 71: 309-416.

10 Comp. Newsl. 32, 1998

Chromosome studies on Vernonia flexuosa
and V. lithospermifolia

MASSIMILIANO DEMATTEIS
Instituto de Botanica del Nordeste
Casilla de Correo 209
3400 Corrientes, Argentina

Abstract

Meiotic and mitotic chromosomes of Vernonia flexuosa Sms and V. lithospermifolia
Hieron. were analyzed in detail. V. flexuosa presented 2n=4x=40 (28m + 12sm) and
V. lithospermifolia showed 2n=4x=20 (10m + 2m-sm + 8sm). Meiotic behaviour
was regular in both species and supports an allotetraploid origin of V. flexuosa. The
results obtained in this work could contribute to a better taxonomic identification of
the two species.

Introduction

Vernonia flexuosa and V. lithospermifolia are two closely related species distributed
in Brazil, Paraguay, Argentina and Uruguay. The two species are morphologically
similar, differing mainly in the head size and the leaf disposition and shape (Ca-
BRERA & KEIN 1980). They are easily distinguished from other species of the genus
by the combination of seriate-cymose inflorescence, sessile heads, yellowish-
brownish pubescence on leaves and stems, and anther appendages with glands.

The two species belong to CABRERA’S (1944) subsect. Flexuosae, or maybe better to
genus Chrysolaena of Rosinson (1988), which includes seven other taxa from South
America, centralized geographically in southern Brazil and northern Argentina.

The cytology of this natural group is relatively well known. All the species have the
basic chromosome number x=10, contrasting with most of related New World taxa,
which commonly present x=17 or x=16 (DEmarTTEIs 1997). Previous studies have re-
ported the chromosome number of V. lithospermifolia (DEMATTEIS 1998) and described
the karyotype of V. flexuosa (Ruas et al. 1991). However, to date no comparative
study of the somatic and meiotic chromosomes of both entities has been made.

In this paper V. flexuosa and V. lithospermifolia are analyzed cytologically in detail
with the purpose to provide data for an accurate identification of the two species.

Comp. Newsl. 32, 1998 11

Materials and methods

Voucher specimens were deposited in the herbarium of the Instituto de Botanica del
Nordeste (CTES). The material examined is as follows.

V. flexuosa Sms: Uruguay. Dept. Artigas. Tomas Gomensoro. DEMATTEIS et al. 491
(CTES).

V. lithospermifolia Hizron.: Argentina. Corrientes. Dept. Saladas. Rio San Lorenzo.
DEMATTEIS & SOLis NeFFA 503 (CTES, LP).

Chromosome studies were made from root tips of germinating seeds. After a
pretreatment of about 4 hours in 8-hydroxyquinoline 0,002 M, the roots were fixed
in ethanol-acetic acid (3:1) and stained according to the Feulgen technique.
Nomenclature used for karyotype description is that proposed by Levan et al. (1964).
Chromosome morphology was characterized using the centromeric index (ci= short
arm x 100/total chromosome length). The idiograms and measures were obtained
from the mean of ten metaphase plates for each species.

The fertility and diameter of the pollen grains were estimated from herbarium speci-
mens by stain with carmin-glicerina (1:1).

Results

Chromosome number, total chromosome length, mean chromosome length,
centromeric index and pollen diameter of V. flexuosa and V. lithospermifolia are
summarized in Table 1.

The somatic chromosome number of V. flexuosa was found to be 2n=40 (Fig. 1).
Karyotype was composed of 28m + 12sm (Fig. 3). Chromosomes ranged in length
from 1,50 to 3,20 um. The pair 2m showed a macrosatellite in the short arm, while
the pair 12m presented a microsatellite in the short arm.

In V. lithospermifolia 2n=20 chromosomes were observed (Fig. 2). The karyotype
formula was composed of 10m + 2m-sm + 8sm (Fig. 4). The pair 3m showed
macrosatellite in the long arm and the pair 6m presented microsatellite in the short
arm.

Meiotic behaviour was regular in the two species, showing always bivalents, 20II in
V. flexuosa and 1011 in V. lithospermifolia. Fertility of pollen grains was 95,30 % in
V, flexuosa and 96,36 % in V, lithospermifolia. Results indicate that V. lithospermifolia
is diploid on base x=10, while V. flexuosa in view of its meiotic behaviour would be
considered allotetraploid with the same basic chromosome number.

12 Comp. Newsl. 32, 1998

Discussion

Most taxonomic treatments on the Flexuosae group of Vernonia distinguish V. flexuosa
and V, lithospermifolia according to the head size, number of flowers, and leaf shape
(CABRERA & KLEIN 1980, Jones 1981, RoBinson 1988). However, in several cases it is
difficult to make a clear distinction between the species, due to the wide variation
and apparent overlapping in these characters.

V, flexuosa shows a great variation in head size, ranging from (6) 7 to 12 (14) mm in
length. This was first noted by Hieronymus (1897), who established four varieties
according to this feature. V. lithospermifolia is particularly similar to V. flexuosa var.
microcephala Hizron., with which it may be confused due to the reduced heads of
the latter taxon.

Results obtained here indicate that it is possible to perform a further distinction of
these species considering the cytological information. From this viewpoint, V. flexuosa
and V, lithospermifolia are certainly very distinct. Besides the evident difference in
ploidy level, the karyotypes differ in number of metacentrics and submetacentrics,
symmetry level and total length per haploid genome.

The difference in ploidy level between the species is also evident in the diameter of
the pollen grains, which is relatively constant within each species. The pollen size
might be effectively used to separate herbarium material that cannot be distinguish-
ed otherwise.

Despite the differences noted above, the two species have the same number and
morphology of satellites. Most members of subsect. Flexuosae have constantly one
or two chromosome pairs with a microsatellite (Dematteis 1997). V. flexuosa and V.
lithospermifolia are the only two taxa of this group with both macro- and micro-
satellites, which supports the close relationship between these species.

Acknowledgements
This work was supported by grants of the SGCyT-UNNE.

Comp. Newsl. 32, 1998 13

References

Caprera, A. L. 1944. Vernonieas Argentinas (Compositae). Darwiniana 6: 19-379.

Casrera, A. L. & R. M. Kien 1980. Compostas. 3. Tribo Vernonieae. In: R. Remrz
(ed.), Flora Illustrada Catarinense 1: 225-408.

Dematteis, M. 1997. Cromosomas en Vernonia platensis y especies afines
(Asteraceae). Bonplandia 9 (3-4): 259-264.

Dematteis, M. 1998. Chromosome studies of some Vernonia species (Asteraceae).
Brazil. J. Gen. (in press).

Hizronymus, G. 1897. Erster Beitrag zur Kenntnis der Siphonogamenflora der Ar-
gentina und Angrenzenden Lander, besonders von Uruguai, Paraguai, Brasi-
lien und Bolivien. Bot. Jahrb. Syst. 22: 672-704.

Jones, S. B. 1981. Revision of Vernonia series Flexuosae (Compositae: Vernonieae).
Brittonia 33: 214-224.

Levan, A., FrepGa, K. & A. A. SANDBERG 1964. Nomenclature for centromeric
position on chromosomes. Hereditas 52: 201-220.

Rosinson, H. 1988. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). V.
The new genus Chrysolaena. Proc. Biol. Soc. Wash. 101 (4): 952-958.

Ruas, P. M., Ruas, C. F., Viera, A. O. S., MATZENBACHER, N. I. & N. S. Martins
1991. Cytogenetics of the genus Vernonia SCHREBER (Compositae). Cytologia
56: 239-247.

14 Comp. Newsl. 32, 1998

Table 1. Somatic chromosome number (2n), total chromosome length (TCL),
mean chromosome length (ML), centromeric index (CI), and pollen
diameter in pm (9) of V. flexuosa and V. lithospermifolia.

40 | 94.60 + 3.70 | 2.36] 41.10+0.50 (39) 41-42 (46)

V. lithospermifolia | 20 | 40.54 + 2.30 | 2.02 | 38.88 + 0.23 (31) 32-33 (36)

Comp. Newsl. 32, 1998 15

Figs. 1-2. Somatic chromosomes of V. flexuosa, 2n=40 (1), and V. lithospermifolia,
2n=20 (2). Scale = 5 pm.

Figs. 3-4.

Comp. Newsl. 32, 1998

Idiograms of V. flexuosa, 28m + 12sm (3) and V. lithospermifolia,
10m +2m-sm + 8sm (4). Scale = 2um.

Comp. Newsl. 32, 1998 17

New names and combinations and other
nomenclatural notes for Compositae of
various countries

ADRIANO SOLDANO
Largo Brigata Cagliari, 6
13100 Vercelli, Italy

Abstract

The author proposes new names, or the proper synonym, for taxa of different countries
that hold the same names, new combinations in proper genera, and emphasizes older
priorities on names currently used.

Introduction

The recent publication of floras of different countries has shown that there are some
Compositae species holding the same name. Many of these are taxa with a narrow
distribution, and the lack of available synonyms necessitates some new names to be
established. Concerning other species, mainly of the Middle East or Mediterranean
Region, the appropriate combinations, following the latest taxonomic treatments (e.g.
BREMER 1994), have been traced. Furthermore, some combinations are reassigned to
authors earlier than those currently cited.

Abbreviations: FC = Flora Vascular de Chile (MARTICORENA & QUEZADA 1985), FE =
Flora Europaea (Tutw et al. 1976), FF = Index Synonimique de la Flore de France
(KERGUELEN 1993), FI= Flora d’Italia (Picnarti 1982), FO = Conspectus Florae
Orientalis (HELLER & HEyN 1993), FR = Vascular Plants of Russia (CZEREPANOV 1995),
FSA = Plants of Southern Africa (ARNOLD & DE Wet 1993).

New or correct names

Centaurea aksamica SoLDANO, nom. nov. = C. veneris B. L. Burtt et P. H. Davis,
Kew Bull. 4: 104-105 (1949), non (Sommigr) BEcuinor in Bécuinor et LANpI, Arch.
Bot. (Forli) 7: 90 (1931).

Ref.: FO, 8:120.

)

18 Comp. Newsl. 32, 1998

The same name of this Cyprus endemite was earlier employed for another restricted
taxon of NW Italy (Liguria) listed in FI.

Senecio incertus DC., Prodr. 6: 433 (1838) = S. tuberosus (DC.) Harv. in Harv. et
Sonb., Fl. Cap. 3: 375 (1865), non Scu. Bir. ex A. Ricu., Tent. Fl. Abyss. 1: 434
(1848).

Ref.: FSA, 768.

The synonym of Harvey’s illegitimate combination that replaces it, is listed in the
indicated reference. SCHULTZ BIPoNTINUS’ name validated by A. RICHARD concerns a
species recently included in Solanecio as S. tuberosus (Scu. Br. ex A. Ricu.) C.
JEFFREY (1986).

— Senecio linaresensis SOLDANO, nom. nov. = S. subdentatus PuL., Linnaea 28: 748

(1856), non Lepgs., Fl. Alt. 4: 110 (1837).
Ref.: FC, 39.

Puurrr’s plant is an endemite of Cordillera de Linares” in Chile and the LEDEBOUR
one, currently listed (FR, 98), grows from Russian Caucasus to Eastern Siberia.

Senecio neoviscidulus SOLDANO, nom. nov. = S. viscidulus Compton, Journ. S. Afr.
Bot. 33: 303 (1967), non SCHEELE, Linnaea 18: 480 (1844).

Ref.: FSA, 768.

CompTon’s species, an endemite of Natal and Swaziland, has the same name as the
hybrid between S. sylvaticus L. and S. viscosus L., a taxon of Central-North Western
Europe (Benorr et al. 1975) listed, for example, in the latest British Flora (STACE
1995).

— Senecio nublensis SoLDANO, nom. nov. = S. carnosus Puw., Anal. Univ. Chile 21:

382 (1862), non THunB., Prodr. Pl. Cap.: 158 (1800).
Ref.: FC, 38.

This endemite of Chillan Mountains in Chile (CABRERA 1949: 190-191) had the same
binomial as the earlier THUNBERG taxon, a species restricted to South Africa, listed in
FSA. Hovewer, it has also to be considered that THUNBERG’s name is a later homo-
nym of S. carnosus LAMARCK (1779) and it is therefore illegitimate (art. 53.1 of the
Code of Nomenclature; GrEuTER et al. 1994); but art. 56 of the same Code may be
used (a proposal is in preparation) for the rejection of LAMARCk’s name, considering
the long permanence of THUNBERG’s taxon in the literature and the illegitimacy of
LaMARCK’s name, a superfluous renaming of Senecio doria L.

Comp. Newsl. 32, 1998 19

’ Senecio pemehuensis SoLDANO, nom. nov. = S. scoparius Pu., Anal. Univ. Chile
88: 282 (1894), non Harv. in Harv. et Sonn. Fl. Cap. 3: 389 (1865).

Ref BC, 39:

Harvey’s earlier name concerns a South African endemite currently listed (FSA,
767). PHwipPr’s taxon is known only from a collection near Pemehue in south Chile
(CABRERA 1949).

Senecio variifolius DC., Prodr. 6: 393 (1838) = S. lyratus L. fil., Suppl. 369 (’1781”,
publ. 1782), non Forssk., Fl. Aeg. Arab. 148 (1775).

Ref.: FSA, 765.

ForssKAL’s earlier name refers to a currently listed (FO, 67) Eritreo-Arabian/East
African species.

New combinations

Inula stenocalathia (RECH F.) SOLDANO, comb. nov. = Codonocephalum steno-
calathium REcH. F., Fl. Iranica 145: 74 (1980).

Ref.: FO, 8: 30.

The inclusion of Codonocephalum FeEnzz in Inula L. is supported by ANDERBERG’S
(1991; see also ANDERBERG 1994) accurate study on Inuleae, where other
”*Codonocephalum’ of the same country (inuloides and peacockianum) are listed
under Inula. I. stenocalathia is an Iranian endemite.

Seriphidium densiflorum (Vv.) SoLDANO, comb. nov. = Artemisia densiflora Vw..,
Fl. Cors. App. Alt.2, 4 t. 2 (1830).

Ref.: FI, 3: 108.

This species is endemic to Northern Sardinia/extreme South Corsica. The specific
rank adopted in FI, by Corrias (1986) - with accurate taxonomy and typification -
and by Geuu et al. (1989) is lowered to the subspecific one in FF.

Stemmacantha caulescens (Cosson et BALANSA) SOLDANO, comb. nov. = Rhaponticum
caulescens Cosson et BALANSA, Bull. Soc. Bot. France 20: 251 (1873). = Leuzea
caulescens (Cosson et BALANSA) Hotus, Folia Geobot. Phytotax. 8(4): 391 (1973).

Ref.: JAHANDIEZ & Mare 1934: 820.

Stemmacantha imatongensis (PHILIPSON) SOLDANO, comb. nov. = Centaurea imaton-
gensis Puirson, Jour. Bot. (London) 77: 232 (1939). = Rhaponticum imatongense

20 Comp. Newsl. 32, 1998

(Pumirson) SosAK, Novit. Hort. Bot. Univ. Carol. Prag. 1962: 48 (1962). = Leuzea
imatongensis (PHILIPSON) Hous, Folia Geobot. Phytotax. 8(4): 391 (1973).

Ref.: Lisowski 1991: 594.

Rhaponticum, as indicated in the references of these two species, is a controversial
abandoned name (cfr. Hous 1973, Dittrich 1984) and Stemmacantha Cass. is the
one currently listed (BREMER 1994; FO; FR etc.).

Tanacetum musili (VELEN.) SOLDANO, comb. nova = Pyrethrum musili VELEN., Sitz.-
Ber. Boehm. Ges. Wiss. 1911, 11:11 (1912).

Ref.: FO, 8: 60.

Pyrethrum ZINN is currently reduced to Tanacetum L. (cfr. BREMER & HUMPHRIES
1993). The species in argument is a Saharo-Arabian endemite.

Wedelia ciliata (SCHUM.) SOLDANO, comb. nov. = Verbesina ciliata ScHuM. in SCHUM.
& THONN., Beskr. Guin. Pl.: 391 (1827). = Aspilia ciliata (ScHUM.) WILD, Kirkia 6:
41 (1967).

Ref.: FO, 8: 39.

Wedelia kotschyi (Scu.Br. ex Hocust.) SOLDANO, comb. nov. = Dipterotheca kotschyi
Scu.Br. ex Hocust., Flora (Intell.) 25: 435 (1842). = Aspilia kotschyi (Scu.Br. ex
Hocust.) BentH. et Hook. ex Ouiv., Trans. Linn. Soc. London 29: 98 (1875).

Ref.: FO, 8:39.

The last two combinations for these species of Tropical Africa and Eritreo-Arabian
province is a consequence of the reduction of Aspilia to Wedelia.

Neglected priorities
Anthemis saxatilis Lam. et DC., Syn. Pl. Fl. Gall.: 291 (1806), ante DC. ex WiLLD.,
Enum. Pl. Horti Berol.: 910 (1809).
Ref.: FF, 16.

Crepis lyrata (Lepes.) Turcz., Bull. Soc. Nat. Moscou 11: 96 (1838), ante (LEDEB.)
FRoEL. in DC., Prodr. 7: 170 (1838).

Ref.: FR, 55.

TURCZANINOW’S paper listing this Siberian endemite antedates D—E CANDOLLE’s name
published in the same year (cf. STAFLEU & Cowan 1985).

Comp. Newsl. 32, 1998 21

Crepis suffreniana (DC.) SteupEL, Nomencl. Bot., ed. 1: 236 (1821), ante LLoyp,
Fl. Loire-Inf.: 155 (1844).

Ref. FE, 5:356; FI, 3: 280; FF, 53.
Leucanthemum halleri (Virman) Ducommun, Taschenb. Schw. Bot.: 383 (1869). =

Chrysanthemum halleri VirtmMan, Summa PI. 5: (1791), ante Surer, Fl. Helv. 2: 193
(1802).

Ref.: HEss et al. 1972: 574.

Virman’s description of this endemite of Switzerland and Austria Alps exactly
reproduces SuTEr’s one and both are based on HaLter (1768 n.97). Ducommun’s
combination based on Surer’s name is a bibliographic citation error that does not
invalidate it (art 33.3 of the Code).

References

ANDERBERG, A. A. 1991. Taxonomy and phylogeny of the tribe Inuleae (Asteraceae).
Plant Syst. Evol. 176: 75-123.

ANDERBERG, A. A. 1994. Tribe Inuleae. Jn: BREMER, K., Asteraceae: Cladistics and
Classification: 273-291. Portland Press, Oregon.

ARNOLD, T. H. & B. C. DE Wer (eds.) 1993. Plants of Southern Africa: names and
distribution. Mem. Bot. Surv. South Afr. No. 62. Pretoria.

Benorr, P. M., Crisp, P. C. & B. M. G. Jones 1975. Senecio L. In: Stace, C.A. (ed.),
Hybridization and the Flora of the British Isles. London and New York.

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

Bremer, K. & C. J. Humpurigs 1993. Generic monograph of the Asteraceae-
Anthemideae. Bull. Nat. Hist. Mus. Lond. (Bot.) 23(2): 71-177.

Casrera, A. L. 1949. El genero Senecio” en Chile. Lilloa 15: 27- 501.

Corrias, B. 1986. Le piante endemiche della Sardegna, 187: Artemisia densiflora
Vw. Bol. Soc. Sarda Sci. Nat. 25: 187-191.

CzEREPANOV, S. K. 1995. Vascular plants of Russia and adjacent states (the former
USSR). Cambridge.

Dittricu, M. 1984. Neukombinationen in der Gattung Stemmacantha Cass.
(Compositae) mit Bemerkungen zur Typisierung einiger ihrer Arten. Candollea
39: 45-49.

22 Comp. Newsl. 32, 1998

Geuu, J. M., GeHu-FRANCK, J. & E. Bronpi 1989. Synécologie d’ espéces littorales
cyrno-sardes rares ou endémiques: Evax rotundata Moris, Spergularia
macrorhiza (REQ. ex LoIsEL.)HEYNH. et Artemisia densiflora Viv. Bull. Soc.
Bot. Fr, Lettres Bot. 136: 129-135.

GreuTer, W., BARRIE, F. R., BurpET, H. M., CHALONER, W. G., DEMOULN, V.,
HawkswortH, D. L., JORGENSEN, P. M., Nicotson, D. H., Sitva, P. C., TRE-
HANE, P. & J. McNEwL 1994. International Code of Botanical Nomenclature
(Tokyo Code). Regn. Veg. 131.

Ha ter, A. 1768. Historia stirpium indigenarum Helvetiae. Bernae.
HELLER, D. & C. C. Heyn 1993. Conspectus Florae Orientalis 8. Jerusalem.
Hess, H. E., LANDOLT, E. & R. Hirzev 1972. Flora der Schweiz 3. Basel & Stuttgart.

Hovvs, J. 1973. Contribution to the Taxonomy and Nomenclature of Leuzea DC.
and Rhaponticum auct. Folia Geobot. Phytotax. 877-395.

JAHANDIEZ, E. & R. Marre 1934. Catalogue des plantes de Maroc (Spermatophytes
et Ptéridophytes) 3. Alger.

JeFFReEY, C. 1986. Notes on Compositae, IV: The Senecioneae in East Tropical Africa.
Kew Bull. 41: 873-943.

KERGUELEN, M. 1993. Index synonimique de la flore de France. Paris.
Lamarck, J. B. A. P. Monnet DE 1779. Flore francoise. Paris.
Lisowsk1, S. 1991. Les Asteraceae dans la flore d’Afrique Centrale. Krakow.

Marrticorena, C. & M. Quezapa 1985. Catalogo de la Flora Vascular de Chile.
Gayana 42.

Picnattl, S. 1982. Flora d’Italia 3. Bologna.
Stace, C. 1995. New flora of the British Isles. Cambridge.
STAFLEU, F. A. & R. S. Cowan 1985. Taxonomic literature 6. Utrecht & Antwerpen.

Tutw, T. G., HEywoon, V. H., Burces, N. A., Moore, D. M., VALENTINE, D. H.,
Watters, S. M. & D. A. WesB 1976. Flora Europaea 4. Cambridge, Lon-
don, New York & Melbourne.

Comp. Newsl. 32, 1998 23

Nature of ergastic substances in some
Asteraceae seeds - VII

MacDonaLp Inu & L. S. Gti
Department of Botany
University of Benin
P. M. B. 1154, Benin City, Nigeria
macidu @ uniben.edu.ng

Abstract

Seeds of 136 species of Asteraceae distributed in 68 genera were examined for
alkaloids, fats and oils, inulin, protein, starch granules and tannin. All the examined
species are herbs. Fats and oils, inulin and protein were found to be present in all the
investigated taxa and alkaloids were absent in 48 species. 10 taxa indicated positive
for tannin, and starch grains were found to be absent in all the investigated species
except two (Cyathula prostrata and Vernonia macrocyanus).

Introduction

Seeds apart from being the chief source of propagation of plants are also principal
storage organ of ergastic substances. A study of the distribution of the stored food
products (often end products of metabolism) shows positive correlation with other
morphological characteristics and has proved to be of diagnostic taxonomic value.
According to GL & AYODELE (1986), the amount of cultivated crops is relatively in
sufficient to provide for the world food supply and hence the knowledge of stored
products in the seeds of wild plants cannot be over-emphasized and this can be done
with a view to harness the resources of the wild plants. Also the future energy needs
of man will rely heavily on renewable plant resources to replace the presently decreas-
ing fossil fuel reserve (ABELSON 1998).

CaLvIN (1983) observed that relatively few plants have been identified as potential
sources of fuel.

The importance of the nature of ergastic substances in plant systematics has been
stressed by various workers, e.g. TATTEOKA (1955, 1962) EarLe & Jones (1962),
MAHESHWARI & CHAKRABARTY (1967), SmirH (1976), Gu et al. (1980, 1984, 1991),
Omoicut & Gut (1988), and Git & Asi (1989).

24 Comp. Newsl. 32, 1998

The present paper reports on the investigations of ergastic substances in 136 taxa of
Asteraceae from 68 genera.

Materials and methods

Seeds of 136 taxa from 68 genera were obtained from Botanischer Garten und
Botanisches Museum, Berlin-Dahlem, Germany. Vouchers of the seeds examined
are kept in the Botany Department of the University of Benin, Benin City, Nigeria.
Chemical tests of various ergastic substances were carried out following the procedures
described by Gur et al. (1991).

Results

The results of taxa studied for their ergastic substances along with their habit are
summarized in Table 1. The taxonomic arrangement of the taxa under this family is
alphabetical.

Discussion

For more than three decades now, much attention has been focused on the comparative
studies of basic molecules in relation to taxonomic problems. BATe-SmiTH & LERNER
(1954) were probably the first to study the leuco-anthocyanins in flowering plants
and concluded that ligneous taxa showed positive tests for them whereas they are of
re-occurrence in herbaceous taca. De Wer & Scorr (1965) are of the opinion that
essential oil can be used as a taxonomic criterion and according to them, chemical
characters are often found to be more reliable that gross morphology in determining
taxonomic affinities. According to ERDTMAN (1956), these ergastic substances are
secondary products of plant metabolism which must have been formed in certain
metabolic processes and are retained when the taxon in question undergoes further
evolution. Knowledge of the principle and direction of chemical processes might
contribute to an understanding of the phylogenetic relations of present day plant
taxa.

All taxa studied indicated the presence of fats and oils, inulin and proteins. 48 taxa
were devoid of alkaloids and 10 taxa indicated positive for tannin, while 2 taxa, viz.
Cyathula prostrata and Vernonia macrocyanus indicated for starch. The presence of
starch in these two species shows their primitive character over those species that do
not have starch.

Comp. Newsl. 32, 1998 25

Eare & Jones (1962), Gt & AYODELE (1986) and Omoicur & Git (1988) earlier
reported the presence of various ergastic substances in 11 plant families. They did
not report any incidence of starch. However, forms of ergastic substance were the
same as in the present report.

Gm et al. (1980, 1984, 1991) have established a relationship between life forms and
the nature of ergastic substances. The observed that starch grains are often associated
with a herbaceous habit, however from the present study, it is obvious that no such
correlation exists in the family Asteraceae as only two investigated taxa showed the
presence of starch.

Taxa with the presence of fats and oils should be further investigated qualitatively to
determine the nature of fats and oils and suitability for commercial exploitation.

Acknowledgements

The authors are grateful to the Director, Botanischer Garten und Botanisches Mu-
seum, Berlin-Dahlem, Germany for providing the seeds of Asteraceae used.

References

ABELSON, P. H. 1978. Bioenergy. Science 204: 1161.

Bare-Smitu, E. C. & N. H. LERNER 1954. Leuco-anthocyanin. 2. Systematic distri-
bution of Leucoanthocyanin in leaves. Biochem. J. 48: 126-132.

Catvin, M. 1983. New sources for fuel and materials. Science 219: 24-26.

De Wet, J. M. J. & B. D. Scorr 1965. Essential oils as taxonomic criteria in
Bothriochloa. Bot. Gaz. 126: 209-214.

Ere, F. R. & Q. Jones 1962. Analysis of seed samples from 113 plant families.
Econ. Bot. 16: 221-260.

Erprman, H. 1956. Flavonoid heatwood constituent of conifers. Sci. Pro. Roy. Soc.
Dublin 24: 129-138.

Gut, L. S. & M.A. Asi 1989. Nature of ergastic substances in some angiospermic
seeds. V. Feddes Repert. 100: 71-79.

Gu, L. S. & J. RAYopELE 1986. On the nature of ergastic substances in the seeds
of some tropical and temperate angiosperms. III. J. Plant Anat. & Morphol.
3: 35-47.

26 Comp. Newsl. 32, 1998

Gu, L. S., OLUBANI, G. O. & S. W. H. Husaint 1980. On the nature of stored food
material in the seeds of some Nigerian legumes. Legume Res. 3: 67-70.

Gui, L. S., OLABANJI, G. O. & S. W. H. Husain 1984. On the nature of ergastic
substances in the seeds of some Nigerian Leguminosae 11. Feddes Repert.
102: 613-628.

Gui, L. S., NyawuAME, H. G. K., AIBANGBE, M. L & D. A. AcHo 1991. Nature of
ergastic substances in some Mediterranean angiospermous seeds. VI. Feddes
Repert. 102: 613-628.

ManesuwarI, J.K. & B. CHAKRABARTY 1967. Starch grains of leguminous seeds.
Phyton (Austria) 12: 191-199.

Omorcut, I. D. & L. S. Git 1988. Nature of ergastic substances in some West
African Compositae. Feddes Repert. 99: 143-145.

Smitu, P. H 1976. The Chemotaxonomy of Plants. London, 313 pp.

TarTEoKA, T. 1955. Further studies in starch grains of seeds in Poaceae from the
view point of systematics. Jap. J. Bot. 30: 109-208.

TaTtTEOKA, T. 1962. Starch grains of endosperms in grass systematics. Bot Mag.
(Tokyo) 75: 377-383.

Comp. Newsl. 32, 1998 27

Table 1. Nature of ergastic substances in Asteraeae taxa studied.

Taxon Life Fats Pro- Tannin
form* at i tein
ie as a a a Sa a ra S| etree ieee

Achillea abrotanoides (Vis.) Vis.
Achillea ageratum L.
Achillea coarctata Por.
Achillea crithmifolia W Avpst. &
Achillea grandifolia FRIvALD
‘Achillea millefolium L.
Achillea ptarmica L.
Achillea teretifolia WiLD.
Ageratum conyoides L.
Ageratum houstonianum Mi.
Anaphalis triplinervis (Sis)
C. B. CLARKE
Antennaria alpina (L.) GAERIN.
Anthemis macedonica
Borss. & OrpH.
Anthemis tinctoria L.
Anthemis tinctoria L. subsp.
australis R. R. FERNANDES
Anthemis tinctoria var. pallida DC.
Anthemis triumfettii (L.) ALL.
‘Arctium lappa L.
?Arctium minus BERNH.
Arctium nemorosum LE}.
Arctotis venusta NORLINDH
Artemisia pedemontana BALBIS
ex Loss.

Aster alpinus L.

Aster amellus Bess. subsp. ibericus
(STEv.) AVETISYAN

Aster cordifolius L.

Aster foliaceus LINDL. var. cusickit
(A. Gray) CRONQ.

Aster pyrenaeus DC.

Aster tripolium L.

Asteriscus sericeus DC.

Bidens cernua L.

Bidens leucantha (L.) WILLD.

Bidens pilosa L.

Carduus crispus L.

Carduus nutans L.

w

(ttt te tet
++etetet+ert
tee tetet¢est
t++eeteetet+et

+ '
++ ++
++ ++
++ ++
'
' '

t++et¢t¢4++
t++teettet
++et¢¢4
1
'

+ +
++ ++
++ ++
++ ++
;
I

aopenpacgecgangenganga>genmemechoomeme piel mmmme Benge ppenkenken her memesteommmeogscmemcogeopengespcopanpenpaspacgen

++eeteerttet
t++eettetest
t++eteteret
t++eteetest

28 Comp. Newsl. 32, 1998

Me ne ae eee
Carduus tmoleus Boiss. subsp.
armatus (Boiss. & HELDR.)

FRANCO
Carlina vulgaris L.

Catananche caerulea L.
Centurea jacea L.

Centaurea ornata WILLD.
Centaurea salonitana Vis.
Chamomilla suaveolens (PuRSH.)

Rybs.

Chaptalia arechavaletai ARECHAV.
Chondrilla juncea L.
Chromolaena odorata (L.)

RM. Kinc & Rosinson
Chrysanthemum coronarium L.
Chrysanthemum segetum L.
Chrysopsis villosa (PursH.) Nutt.
‘Cirsium arvense (L.) Scop.
Cirsium candelabrum GRIsEB.
Cirsium flavispina DC.

Cirsium monspessulanum (L.) HILL
Cirsium palustre (L.) Scop.

Cirsium trachylepis Botss.

‘Cirsium vulgare (SAv1) TEN.
Conyza bonariensis (L.) CRONQ.
Conyza canadensis (L.) CRONQ.
Coreopsis lanceolata L.
Crassocephalum crepidioides

(Bentu.) S. Moore
Crepis commutata (SPRENG.) BABC.
Crepis foetida L. subsp. foetida
Crepis multiflora SM.

Crepis pyrenaica (L.) GREUTER
Crepis rhoeadifolia (M. Bis.)
CELAK.

1
t+++et+¢+
t++e+ttet
teettet

1

1

+++

+++

+++

+++
;

eee ee

tteteettetet+e+ett
+ttettetet+r+ett
+etetetetettetetet
i]
'

+ +

++teet
tee tet
++4+4++4
+++++

+

Cyathula prostrata (L.) BLUME
Dendroseris micrantha Hook.

& ARN.
Dimorphotheca pluvialis

(L.) MoENCH
Dittrichia graveolens (L.) GREUTER
Doronicum austriacum Jacq.
Dyssodia setifolia (LAGASCA)

ROBINSON var. radiata

(Gray) STROTHER H + |+ ] + = - -
Echinops albidus Boss. & SpruN. | H - + | + + - -

Tom hm OO GO Lee ee oe

Comp. Newsl. 32, 1998 29

Emilia sonchifolia L.
Erigeron speciosus (LinDL.) DC.
'Eupatorium cannabinum L.
Eupatorium purpureum L.
Gaillardia aristata Pursu.
Galinsoga ciliata (RAF.)
S. FE BLAKE
Galinsoga parviflora Cav.
Gerbera anandria (L.) Scu. Br.
Gnaphalium sylvatium L.
Helianthus annuus L.
Helichrysum rupestre (RaF.) DC.
Hieracium amplexicaule L.
Hieracium bornmuelleri FREYN.
Hieracium sabaudum L.
Hyochoeris maculata L.
Hyochoeris oligocephala
(SvenT. & BraMw.) LACK
Hypochoeris radicata L.
Hypochoeris unifora Viv.
Inula orientalis Lam.
Inula salicina L.
Inula thapsoides (Wi1D.) Spr.
Inula verbascifolia (WiLLD.)
HAUSSKN. subsp. aschersoniana
(JANKA) TUTIN
Jurinea alata (DEsF.) Cass.
‘Lactuca serriola L.
‘Lapsana communis L.
Leontodon autumnalis L.
Leontodon glabratus (W. Kocn)
BISCH.
Leontodon hispidus L.
Leuzea centauroides (L.) Hous
Matricaria maritima L.
Onopordum acanthium L.
Onopordum bracteatum
Boiss. & Hepr. subsp. ilex
(JANKA) FRANCO
Onopordum illyricum L.
Picris evae LACK
Porophyllum ruderale L.
Pulicaria dysenterica (L.) BERNH.
Pulicaria odora (L.) REICHENB.
Santolina chamaecyparissus L.
Santolina rosmarinifolia L.

(+++
t+++4+4
++++¢+
t++etet

!
++tetteteet
t++eteteteetet
t++eteteeteest

!

'

t++et¢4e4¢+
t++++¢+4++
t++ttett
1!
'

'
++4+4+4
++ +44
t++tet

1

!

Gocco OOO CO Pee Oe

1‘
+++4+4
t+++4++
+++4++4+

'

1

jaogangangacgangangengen)
+++4+5

t++etteeeest
t++etetetet
t+etetettet

30 Comp. Newsl. 32, 1998

Santolina squarrosa (DC.) NYMAN

Sanvitalia procumbens Lam.

Scorzonera cretica WILLD.

Senecio adonidifolius Loss.

Senecio appendiculatus (L.F.)
Scu. Br.

Senecio aquaticus Hh.

Senecio chrysanthemoides DC.

Senecio doria L.

Senecio thapsoides DC.
subsp. thapsoides
Sonchus arvensis L.
Spilanthes ocymifolia (LaM.)
A. H. Moore
Staehelina uniflosculosa
(Stu. & SM.)
Tanacetum cilicium (Botss.)
GRIERSON
Tanacetum parthenium (L.)
Scu. Br.
Tanacetum pseudoachillea
C. WINKL.
Tanacetum vulgare L.
Telekia speciosa (SCHREB.) BAUMG.
Telekia speciosissima (L.) Less.
Tragopogon tommasinii Scu. Br.
*Tridax procumbens L.
Vernonia macrocyanus O. HoFFM.
Xanthium spinosum L.
Xanthium strumarium L. subsp.
italicum (Moretm!) D. Love
Zinnia peruviana (L.) L.

tro tttee tt
t+etettetetest
++et¢¢t¢¢4
++etetet

H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H
H

+
+ +
+ +
+ +

* = Herb L= Liana S = Shrub T= Tree

'- Species earlier investigated by Gi & Asn (1989)
2. Species earlier investigated by EARLE & Jones (1962)
3. Species earlier investigated by Omoicur & Gi (1988)

Comp. Newsl. 32, 1998 31

Allelopathic effect of Calotropis procera on the
germination of Helianthus annuus seeds

MacDonaLp Inu & C. A. OMONHINMIN
Department of Botany
University of Benin
P. M. B. 1154, Benin City, Nigeria
macidu @uniben.edu.ng

Abstract

The effect of 2 %, 10 % w/v aqueous extract of Calotropis procera leaves and flowers
on the germination of nine (9) cultivars of Helianthus annuus seed was investigated.

The interaction resulted in poor germination performance of below 30 % in all the
cultivars, while the control (distilled water) treatment recorded high performance of
above 60 % germination. Only a slight difference in inhibitory effect was recorded
between the different extract concentrations and plant parts solution extract treatments.

Introduction

From De CANDOLLE (1832) to date, a lot of work has been carried out on the allelopathic
effect of plants, both agricultural or non-agricultural, on the growth performance of
other plants. Such plants are believed to affect their neighbour’s performance by
direct releases of the allelochemicals (Rice 1984, GarciA & ANDERSON 1984, GILL et
al. (1993), or indirectly through the effect of the toxins on the characteristics of the
growth media (soils) where they both survive, by altering the physical as well as
chemical characteristics of the soil (IGBoANUGU 1986, INDERMT & DaAKsHINI 1994).

Calotropis procera is a common weed of most farm lands in the northern part of
Nigeria. It has been speculated that it affects the performance of most crop plants in
fields where it exists, possibly due to its high alkaloid and glucoside content (BOUGUENT
1972, DAUBENMRE 1974, GpiLE 1986). However, this remains only a supposition as
no investigation of such claim has been undertaken with respect to Helianthus annuus.

Thus, the aim of the present study is to ascertain the inhibitory effect of Calotropis
procera plant (parts) leachates on the germination of Helianthus annuus seeds.

32 Comp. Newsl. 32, 1998

Materials and methods

Helianthus annuus seeds (9 cultivars: V, - Funtua, V2 - Cakinki, V3; - Saturn, V,-
Record, V; - Perodirk, V, - Cherniank, V7 - Isa-anka, Vz- Vnumik and V,- Smena),
used for this study were collected from AFCOTT (AFCOTT Nig. Ltd, Ngurore South-
East Yola, Nigeria) through a seed exchange programme.

Leaf and flower components of Calotropis procera plants were collected in August
1995 from Yola, Nigera. The fresh materials were subdried and ground. 2g and 10g
weight of each of the leaf and flower parts were dissolved in 100 mls of distilled
water to give 2 % weight/volume (w/v) and 10 % w/v extract solutions of the leaf/
flower components of the C. procera plant. The preparations were filtered and
refrigerated. 150 seeds of each of the 9 Helianthus annuus cultivars were divided
into 5 parts of 30 seeds for each treatment (2 %, 10 % leaves; 2 %, 10 % flower and
control). Each set of 30 seeds were further subdivided into triplicates of 10 seeds.
Petri-dishes were lined with filter paper each with 10 Helianthus annuus seeds, and
moistened daily with the 2 %, 10 % leaves and flowers solution of C. procera. The
control treatment was moistened with distilled water. The experimental set up was
kept at constant temperature (30°+ 3° C) in a growth chamber. Germination was
recorded regularly at 48 hrs interval.

Results

Table 1 shows the germination percentages recorded for the nine (9) cultivars of
Helianthus annuus. The results obtained show a marked response of germination
inhibition in all the cultivars used. The control treatment recorded a high performance
for all the cultivars, with the lowest percentage germination of 62 %, and the highest
of 78 % in the cultivars Funtua (V,) and Vnumik (V3) respectively. The maximum
recorded for any of the extract solutions treatment was 21 % for the cv. Cherniank
(V.) with the cv. Funtua (V,), Cakinki (V2) and Record (V,) recording the minimum
germination of 10 %. The variance level using Duncan’s multiple range test (P =
0.05) further shows the significance level between the control (distilled water)
treatment and the extract solutions treatments. However, only slight significance
difference existed between the different concentrations (2 %, 10 %) of the leaves or
flowers solutions and between the different plant component (parts) - leaves and
flowers solution treatments (Table 1).

Comp. Newsl. 32, 1998 33

Discussion

The phenomenon of allelopathy is not only an important one, but also plays a
significant role in the distribution of plants in fields. From the present study, it is
quite apparent that Calotropis procera extract solutions greatly hindered the
germination of H. annuus seeds. This suppressive effect may be due to the presence
of noxious compounds in the species C. procera and the genus as a whole. The inci-
dence of such toxins was first elucidated by Watr & BREYER-BRANDWUK (1962).
They showed that species of Calotropis contained a strong cardiac poison in the
latex exudate known as ”Calotropin”. BouGuent (1972) and DauBENMIRE (1974)
identified 7 glucosides as well as calotropin in the latex of some members of the
genus Calotropis. Later, GpiLe (1986) extracted the following alkaloids; Benzoyth-
colone, Benzylusolineolone, Calotropin, Calotoxin, Uscharin, Uscharuchin, Calactin
vorisherine and mudarin from this group of plants.

The presence of an extract solution of a whole plant or plant parts with an array of
toxic compounds such as above in the growth medium of another plant, can be directly
linked to the inhibition of growth of the effector plant (EVANaRI 1949, Rice 1984), or
indirectly by altering the characteristics (physical and chemical) of the growth media
(soils), and/or the availability of nutrient, pH, total phenolic levels (TPL), and microbial
population (BLUM & SHAFER 1988, INDERMIT & DAKSHINI 1992, 1994).

The toxicity of the bioassay 2 % or 10 % w/v plant part was undoubtedly pronounced.
Table 1 shows the percentage germination recorded for the five different treatments
on the 9 cultivars of H. annuus. The control treatment (distilled water) recorded high
germination of above 60 % for all the cultivars, with highest of 78 % for the cv.
Vnumik (V,) and the lowest of 62 % for the cv. Isa-anka (V,), whereas, the highest
recorded for any of the 4 extract solutions was 21 % in the 2 % w/v leaf extract
solution treatment for the cv. Chemiank (V,,). Similarly, ApaMs & Azim1(1991) showed
the suppressive effect of Cyperus rotundus leaf extract on the germination of wheat
grains. Also, Gi et al. (1993) reported on the allelopathic effect of Chromolaena
odorata extract on growth of cowpea.

A high degree of variance in treatment effect between the control and extract solutions
was recorded. This further supports the view that the toxins present in the effector
plant’s extract solutions inhibited germination growth in seeds of H. annuus.

A significant level of treatment effect amongst the leaf and flower extract solutions
and at different concentrations (2 % & 10 % w/v) was not recorded, as only the cv.
Funtua (V,) showed some level of significance of treatment effect for the leaf extract
treatments, while the cv. Funtua, Cakinki and Record recorded identical results for
the flower extract treatments.

34 Comp. Newsl. 32, 1998

Similar results from OasE et al. (1994), showed that the leaf extract of Chromolaena
odorata inhibited germination seedling growth in Zea mays grain to a high degree
irrespective of the duration of extraction. Unlike in most allelopathic studies, the
different plant component (parts) extract solutions, in the present study, show no
significant level of difference in their effect on the germination of seed of H. annuus.
This is because identical degrees of inhibition were exhibited by both leaf and flower
extract solutions.

In conclusion, the authors are of the view that regardless of the component part
extracted, the concentration of the extract solution of C. procera or the cultivar of H.
annuus seeds employed, the degree of inhibition is similar. A comparable degree of
the allelopathic effect of C. procera on other plants is apparent from the present
study. This suppressive ability might account for the spatial distribution and reduced
growth of plants and notably H. annuus in fields where it occurs.

References

Apams, S. M. & R. Azmi 1991. Effect of purple nutsedge (Cyperus rotundus) leaf
extract on germination and seedling growth of wheat (W. Pavon). Pakistan
Journal of Weed Science 4: 59-61.

Bium, U. & S. R. SHAFER 1988. Microbial populations and phenolic acid in soil. Soil
Biology and Biochemistry 20: 793-800.

BoucuEnt, A. 1972. Plantes Medicinales du Congo Brazzaville. B. Traveaux orstorn.
Paris No. 13, 15.

DAUBENMIRE, R. F. 1974. Plant and environment. 3" ed. John Wiley and Sons, New
York.

De CanDoOLLE, A. P. 1832. Physiologie Végétale 3. Bechet Jeune, Paris.
Evanart, M. 1949. Germination Inhibitors. Botanical Review 15: 159-194.

Garcia, A. G. & I. C. ANDERSON 1984. Monthly variation in allelopathic effects of
corn residues on corn seedlings growth under three tillage practices. Philippine
Journal of Crop Science 9: 61-64.

Gre, Z. O. 1986. Ethnobotany, taxonomy and conservation of medicinal plants.
In: SoroworA, A. (ed.), The State of medicinal plants research in Nigeria.
University Press, Ie-Ife.

Gut, L. S. , ANOLIEFO, G. O. & U. V. Inuoze 1993. Allelopathic effect of aqueous
extract of Siam weed on growth of Cowpea. Chromolaena Newsletter 8: 1-4.

Comp. Newsl. 32, 1998 35

IcBoanucu, A. B. I. 1986. Phytotoxic effect of some Eucalypts on food crops,
particularly on germination and radicle extension. Tropical Science 26:
19-24.

Inpergit, A. & K. M. M. Daksuint 1992. Interference potential of Pluchea lanceolata
(Asteraceae): growth and physiological response of asparagus bean, Vigna
unguiculata var. sesquipedalis. American Journal of Botany 79: 977-981.

Inpergit, A. & K. M. M. Dasuini 1994. Allelopathic effect of Pluchea lanceolata

(Asteraceae) on characteristics of four soils and tomato and mustard growth.
American Journal of Botany 81: 799-804.

OcsE, F. M. O., Git, L. S. & E. O. O. Iseruien 1994. Effect of aqueous extracts of
Chromolaena odorata L. on radicle and plumule growth and seedling height
of maize, Zea mays L. Compositae Newsletter 25: 31-38.

Rice, E. L. 1984. Allelopathy. 2" ed. Academy Press, New York.

Wart, J. M. & M. G. BREYER-BRANDWIJK 1962. Medicinal and poisonous plants of
Southern and Eastern Africa. E. & S. Livingstone, Edinburgh and London.

36 Comp. Newsl. 32, 1998

Table 1. Allelopathic effect of Calotropis procera leaves and flowers extract
solutions on germination of Helianthus annuus seeds.

Treatment

Control

Leaves 2 % w/v
Leaves 10 % w/v
Flower 2 % w/v

Flower 10 % wiv

* Within each variable (cultivars), means followed by the same subscript are not
significantly different at 5 % level.

** w/w = weight/volume

Key to names of cultivars:

Vi — Funtua Vs — Cherniank
V, — Cakinki V; — Isa-anka
V; — Saturn Vs — Vnumik
V;, — Record Vo — Smena

V; — Perodirk

Comp. Newsl. 32, 1998 37

Germination control of Helianthus annuus L.
using two growth regulators - thiourea and
coumarin

MacDonaLp Inu & C. A. OMONHINMIN
Department of Botany
University of Benin
P. M. B. 1154, Benin City, Nigeria
macidu @ unibe.edu.ng

Abstract

Thiourea effectively promoted germination in 9 varieties of Helianthus annuus seeds.
Conversely, coumarin retarded germination growth in the seeds. A significant level
of difference was observed (at 5 % level) between the treatments, with thiourea
recording as high as 85 %, 70 % germination under light and dark condition, and a
minimum degree of 43 % and 38 % germination in the cultivars, ’Funtua” ( V,) and
”’Vnumik” ( Vs), respectively. Coumarin recorded low percentage germination (39
%,27 %) in the cv.”’Saturn” (V3) with the lowest percentage germination of 10 % in
cv. Record” under the light condition. The cultivars *Funtua” (V,) and ’’Cakinki”
(V2) had better germination energies than the others, with cv.’ Vnumik” (V3) exhibiting
the least. The light condition enhanced germination in H. annuus.

Introduction

Seeds have been shown to differ in their response to growth regulatory compounds;
this applies to both hormonal and non-hormonal compounds. Likewise, these growth
chemicals whether applied exogenously or naturally occurring exert either stimulatory
or inhibitory effects under different conditions and concentrations, thus either as
germination stimulators, or germination inhibitors when they suppress or nullify
germination growth (Mayer & PoLJAKOFF-MAYBER 1989).

Growth regulators (stimulators and inhibitors) commonly occur naturally and interact
in seeds to effect dormany/germination together with other dormancy/germination
mechanisms present in the seed or its environment (KELLy et al. 1992). They can be
synthesized and exogenously applied to achieve the same effect as the endogenously
present ones. Most often these interactive regulators act antagonistically; a commonly

38 Comp. Newsl. 32, 1998

known interactive pair is the GA/ABA Complex (Ketrinc 1977, EGLey 1972) which
occurs in growth regions of plant components. Similarly, the pair thiourea and cou-
marin have been reported for seeds, where they control the germination growth of
such seeds. In the case of lettuce seeds, while thiourea effectively stimulated
germination to as high as 100 %, coumarin reduced germination from 50 % level
(control) to zero. Such interactions are light, temperature and respiration linked (MAYER
& POLJAKOFF-MAYBER 1989).

The present study sets to determine the effect of exogenously applied thiourea and
coumarin on the germination growth of Helianthus annuus seeds.

Materials and methods

Seeds for the study were obtained from AFCOTT (AFCOTT Nigeria Plc, Ngurore,
25km South-East Yola, Nigeria). Seeds of 9 cultivars: V, - Funtua, V2 - Cakinki, V;
- Saturn, V,- Record, V;- Perodirk, V,- Cherniank, V;- Isa-anka, V;- Vnumik and
V.- Smena, whose healthy state have been determined to be 100 % with tetrazolium
salt, were used. 5g/L thiourea solution and 0.04g/100m1 coumarin solution were em-
ployed for the experiment.

The three treatments, viz. control (A), thiourea (B) and coumarin (C) were set up for
light and dark condition treatments (A1 and A2, B1 and B2, C1 and C2). Each illumi-
nation regime treatment was replicated in triplicates for each cultivar. Following a
randomized design 180 seeds of each cultivar were selected for the triplicated treat-
ments under the light and dark conditions. Treated seeds were placed in filter-paper
lined petri-dishes. The Al, B1, C1 sets were placed under continuous light, while the
A2, B2, C2 were placed under continuous dark condition. Germination was recorded
at 2 day intervals for 28 days.

Statistical analysis was carried out using Duncan’s multiple range test at 5 % level
(P—O03):

Results and discussion

Figs. 1 and 2 show the percentage germination of treatments for the 9 cultivars of H.
annuus under light and dark conditions. Thiourea recorded the highest germination
of 85 % under the light condition for the Vj, this is higher than that recorded under
the dark condition (70 %) for the same cultivar. The smallest percentage recorded for
the thiourea treatment was 43 % and 38 % under both illumination regimes respectively
for the cv. Vs(Vnumik). The control treatment recorded a maximum germination of

Comp. Newsl. 32, 1998 39

70 % for V; (Fig. 1, Table 1) and the minimum levels were above 30 %. However,
germination percentages recorded for the coumarin treatment were low, with the
highest of above 53 % and the lowest 10 % for the same cultivar, V, (Figs. 1 and 2).

From the foregoing, it is apparent that thiourea irrespective of the illuminaton
conditions or kinds of cultivars, stimulated germination to a high level, while coumarin
exhibited inhibitory effect. These stimulatory/inhibitory effects of thiourea and
coumarin may have been due to the action on the storage materials of the seeds, the
oxidative phosphorylation (phosphate/oxygen ratio) and the coupling action in ger-
minating seeds either directly or indirectly; this view is supported by Mayer & Pot-
JAKOFF-MAYBER (1989). Germination was higher for all the cultivars for control,
thiourea and coumarin treatment under the light condition, suggesting light
dependency (stimulatory effect) of the seeds of H. annuus. Hsiao et al. (1988) showed
the stimulatory effect of growth stimulators on germination of the witchweed (Striga
asiatica). BASKIN & BASKIN (1974) also reported up to 100 % germination in seeds of
Isanthus brachiatus treated with gibberellic acid. UGBoroGHO & AGomo (1989) show-
ed the germination retarding effect of colchicine on seeds of Vigna unguiculata. The
germination enhancing effect of light was demonstrated by BASKIN & BasKIN (1975)
in Helenium amarum seeds.

Analysis of variance (Table 1) showed a marked and significant level of difference
between thiourea and coumarin treatments. Significant differences were recorded in
some of the cultivars between the control and thiourea treatments. Also a major level
of difference was apparent between the control and coumarin treatments. Within the
same treatment, significant difference was recorded between the light and dark regi-
mes with the light treatment recording a higher degree of germination than the dark
in all cases. From the comparison, the thiourea treatment showed a higher degree of
germination in all the cultivars, and the coumarin treatment recorded the lowest degree.
Also, the cultivars V, and V2 recorded higher level of germination, and cv. V; obtained
the lowest germination for all the treatments.

From the above it is quite conclusive that thiourea stimulates germination while
coumarin retards germination in H. annuus seeds. The cv. Funtua and Cakinki are
the most readily germinating cultivars while cv. Vnumik shows the lowest initial
germination energy.

The stimulatory effect of light on germination of H. annuus seeds is an area of further
investigation.

The performance of the cultivars in the present study can be useful in the delimitation
of infraspecific taxa in a complex species such as Helianthus annuus.

40 Comp. Newsl. 32, 1998

References

Baskin, J. M. & C. C. Baskin 1974. Breaking dormancy in seeds of Jsanthus
brachiatus (Labiatae) with gibberellic acid. Phyton 32: 159-165.

Baskin, J. M & C. C. Baskin 1975. Do seeds of Helenium amarum have a light
requirement for germination? Bull. Torrey Bot. Club 102: 73-75.

Ectey, G. H. 1972. Influence of seed envelope and growth regulators upon seed
dormancy in witchweed (Striga lutea Lour.). Annals of Botany 36: 755-770.

Hsiao, I. A., WorsuaM, A. D. & D. E. MorELAND 1988. Effects of chemicals often
regarded as germination stimulants on seed conditioning and germination of
witchweed (Striga asiatica). Annals of Botany 62: 17-24.

KE ty, K. M., vAN STADEN, J. & W. E. BELL 1992. Seed coat structure and dormancy.
Plant Growth Regulation 11: 201-209.

Ketrinc, D. L. 1977. The Physiology and Biochemistry of Seed Dormancy and
Germination. North-Holland, Amsterdam.

Mayer, A. M. & A. PotjakorF-MayBeErR 1989. The Germination of Seeds. 4th Ed.
Pergamon Press, Oxford.

Ucsorocuo, R. E. & C.O. Acomo 1989. Seed germination and economic importance
of Vigna unguiculata (L.) WavpERS (Papilionaceae) in Nigeria. Feddes
Repertorium 100: 147-155.

Comp. Newsl. 32, 1998 4]

Table 1. Germination percentages (P = 0.05) of Helianthus annuus seeds treated
with thiourea and coumarin and of untreated (control) seeds under
light and dark condition.

Treatments

light dark | light dark | light dark

85a 70b
65a 62a
53a) 52a

46a 38b
69a 53b
52a 52a
53a 52a
43a 38a
59a 43b

Percentages followed by the same letter are not significantly different at 5 % level.

Key to cultivars:

V, — Funtua V;— Cherniank
V,. — Cakinki V; — Isa-anka
V; — Saturn V,— Vnumik
V, — Record V,. — Smena

V; — Perodirk

Comp. Newsl. 32, 1998

42

95 CSacrminatiam

100

LO ace ee it

Varieties

Aig) saieetror hemmonesrteonbeseea
Germination of H.annuus Umder ligmt
Gonaeitian

__] Control
[|] Thiourea

[7] Coumarin

V; — Funtua

V, — Cakinki
V3 — Saturn

Vi, — Record

Vs — Perodirk
Vs as Cherniank
V; — _Isa-anka
V3 — Vnumik
Vo — Smena

43

Comp. Newsl. 32, 1998

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