SS NN f\ N d SS. Pp. * Be o¢ “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 euouls — IN ya eyue-ess — ‘A yuemeyg = SA =e (HOM OIGS) yupoeg — ‘SA sUeoO JsouUn SsnnuuSe 4 Jo vVelevitie. pioooy = —~Stst«SA Doos UO SSeuClwwieul 1@ j2eler 7 ce umes — ‘A Ssitoilen pure) SAA GA SA ZA GA Sh FAL EX Sen A emuny — TA , - ulLeLUNOD fF] eBeino!Iyt [| JO11U0D Fy UONCUILLSS %