Tan)
~ COMPOSITAE &
* NEWSLETTER
Number 30 June 1997
Scientific Editor: Bertil Nordenstam
Technical Editor: Gunnel Wirénius Nohlin
Published and distributed by The Swedish Museum of Natural History,
Department of Phanerogamic Botany,
P.O. Box 50007,
S-104 05 Stockholm, Sweden | ] B Re
(Director: Professor Bertil Nordenstam) R vi
ISSN 0284-8422
NEW yor
BOT, K
| ANICAL GARDEN
CONTENTS
C. Pérez Morales, F. Llamas Garcad, C. Acedo Casado & A. Penas Merino:
Typification and definition of Doronicum austriacum Jacq. (Asteraceae) 1
M.S. Ayodele: Studies on the reproductive biology of Vernonia Schreb.
(Asteraceae). IV. Seasonal flowering sequence among plant forms of
Vernonia in Nigeria B)
A. Badr, E. Abdelrazik Kamel & N. Garcia-Jacas: Chromosomal studies
in the Egyptian flora. VI. Karyotype features of some species in subfamily
Asteroideae (Asteraceae) is
J. L. Reveal: Early suprageneric names in Asteraceae 29
Bertil Nordenstam: Nomenclatural notes on Ecuadorian Senecioneae AG
M. Idu & L.S. Gill: Nature of ergastic substances in some West African
Asteraceae seeds - VIII 50
A. Ouyahya: La germination et le pouvoir germinatif de quelques Artemisia
du Maroc a
Comp. Newsl. 30, 1997 1
TYPIFICATION AND DEFINITION OF
DORONICUM AUSTRIACUM JACQ.
(ASTERACEAE)
C. Pérez Morales, F. Llamas Garca, C. Acedo Casado & A. Penas Merino
Department of Plant Biology (Botany)
University of Léon
E-24071 Leon
Spain
Abstract
This study tries to clarify the morphological differential characters of Doronicum
austriacum (Asteraceae). Until now they were not clearly defined, mainly those
referring to the presence or absence of some types of trichomes in its indumentum.
Furthermore, taking into account Jacquin’s protologue as well as materials
attributable to him, we consider as lectotype of Doronicum austriacum, a specimen
from the original materials deposited in the Herbarium LINN.
Introduction
Doronicum austriacum Jacq., iS a conflictive taxon with some identification
problems when using the current keys, because the differential characters usually
employed are not diagnostic enough to make a correct identification.
Various workers have added a series of corrections and interpretations to the
protologue resulting in a lack of knowledge about the taxon considered by different
authors as D. austriacum.
In order to differentiate it from its closest relatives (D. pardalianches L., D.
carpetanum Boiss. & Reuter ex Willk., D. cataractarum Widder) the shape of the
lower leaves and the planindumentum have been used as the most significative
characters. Neither in the consulted floras nor in specific papers, exists total
concordance among the authors relating to the commented characters, and the
biggest contradictions are those related with the indumentum, that on the other
bo
Comp. Newsl. 30, 1997
hand is one of the taxonomic characters most employed now, because of its high
differential value at specific level (Pérez Morales & Penas Merino 1990, Pérez
Morales et al. 1994).
Jacquin (1774), when describing D. austriacum, pointed out the presence of hairs
on both sides of the leaves, stems and bracts, but he did not say they were
glandular. Amo Y Mora (1872), Rouy (1903) and Fiori (1984) agreed with Jacquin
considering this is a plant more or less hairy. Coste (1937) and Guinochet &
Vilmorin (1982) added that the involucral bracts lack glandular hairs. But Cavillier
(1911), Ferguson (1975), Pignatti (1982) and Hegi (1987) said D. austriacum has a
glandular indumentum at least in some parts of the plant.
Results and Discussion
In our former studies on the genus, we have observed specimens identified by
other authors as D. austriacum having glandular hairs all over the plant, and other
specimens lacking this hair type on the involucral bracts, peduncles and stems.
Using the current keys both specimen types can be included in D. austriacum
because of little precision in the use of the indumentum character. But only the
specimens lacking glandular hairs on the involucral bracts, peduncles and stems
agree with the protologue where there is no mention of the presence of this kind of
trichomes.
Because of these incongruences we became interested in the search of Jacquin’s
original material in different European herbaria. In LIV, the few sheets attributable
to Jacquin contain plants from the West Indies. In UPS there is not any sheet of D.
austriacum. In B, neither in the general herbarium nor in the Willdenow collection
exist specimens of D. austriacum attributable to Jacquin. In OXF, in the Sherardian
Collections, there is a specimen of D. austriacum attributable to Jacquin. When we
observed that specimen we saw the presence of glandular hairs on the involucral
bracts and on the peduncle. Jacquin did not mention this kind of trichomes in the
protologue. In BM, now there is no Jacquin material because they have been
moved to LINN. There, there are two sheets numbered 1002.3 and 1002.4 labelled
as D. austriacum. When we cxamined those two sheets we realized they belong to
two different taxa. We observed in one of them (1002.4) characters agreeing with
the protologue and with other material studied by us, lacking glandular hairs. The
specimen of the sheet 1002.3, on the other hand, does not agree with the
protologue, but has great similarity with D. pubescens C. Pérez Morales, A. Penas,
F. Llamas & C. Acedo, because it is glandular-pubescent all over the plant.
Comp. Newsl. 30, 1997 3
So, because Jacquin does not mention in the protologue the presence of glandular
hairs, we designate as lectotype of D. austriacum, the only specimen of the sheet
1002.4. Its label, placed on the left side, reads: n° 175/ D. austriacum, and on the
right side of the specimen is written on the sheet: a Jacquin/austriacum.
Acknowledgements
This work was supported by the Spanish CICYT grant NAT 90-0871-CO3-01.
We are grateful to Dr. J.E. Jarvis for his help in the search of material, and the
Curators of the consultated Herbaria who gave us information.
4 Comp. Newsl. 30, 1997
References
Amo y Mora, M. del 1872. Flora Fanerogdmica de la Peninsula Ibérica. IV:
2554. Granada.
Cavillier, F. 1911. Novuelles études sur le genre Doronicum. Annuaire du Con-
servatoire et du Jardin Botaniques de Genéve 13-14: 195-368.
Coste, J. 1937. Flore descriptive et illustrée de la France, de la Corse et des
contrées limitrophes. 2: 296. Paris.
Ferguson, I.K. 1976. Doronicum L. In: Tutin T.G., Heywood, V.H., Burges,
N.A., Moore, D.M., Valentine, D.H., Walters, S.M. & D.A. Webb (eds.), Flora
Europaea 4: 190-191. Cambridge.
Fiori, A. 1984. Nuova Flora Analitica D’ Italia. Vol. 11: 602. Florencia.
Guinochet, M. 1982. Doronicum L. In: Guinochet, M. & R. Vilmorin, Flore de
France 4: 1465-1466. Paris.
Hegi, G. 1987. /llustrierte Flora von Mittel-Europa. Teil 4: 710-714. Berlin.
Jaqcuin, N.J. von 1774. Flora Austriaca, Vol. 2. Vienna.
Pérez Morales, C. & A. Penas Merino 1990. Sobre algunos Doronicum ibéricos.
Lagascalia 15 (2): 151-160.
Pérez Morales, C. Penas Merino, A., Llamas, F. & C. Acedo 1994. Doronicum
pubescens sp. nov. Lazaroa 14: 5-12.
Pignatti, S. 1982. Flora D' Italia 3: 113-117. Bolonia.
Rouy, M.G. 1903. Le genre Doronicum dans la flore européene et dans la flore
atlantique. Revue de Botanique Systématique et de Géographie Botanique 2:
17-56.
Comp. Newsl. 30, 1997 S
STUDIES ON THE REPRODUCTIVE BIOLOGY OF
VERNONIA SCHREB. (ASTERACEAE)
IV. Seasonal flowering sequence among
plant forms of Vernonia in Nigeria
M.S. Ayodele
Biological Sciences Department
University of Agriculture
PM B 2240
Abeokuta, Nigeria
Abstract
The flowering sequences of 15 species of Vernonia, representing five growth forms
(habits) are reported.
Two major modes of flowering, and their variants, were identified among species
invesugated. These were the synchronised blooming and fruiting phases, common
among the arborescent species; and the intermittent blooming/fruiting and
vegetative growth common among the shrubby and herbaceous species.
Flower production sequence, by the shrubby and herbaceous growth forms, was
influenced by the rainfall regime of their locations (i.e. habitats). Difference in
flowering periods among the species may be one isolation mechanism in the
evolution and in preserving species identity in Nigerian species of Vernonia.
Introduction
Species of the genus Vernonia display different plant forms (i.e. growth habits)
namely: arborescent, shrubby and herbaceous forms. There are annuals,
herbaceous or woody perennials, and scramblers. There are also weedy climbers or
stragglers (Hutchinson & Dalziel 1963, Faust 1972, Olorode 1984). The various
growth forms have different types of inflorescence. However, some types of
inflorescence cut across a number of growth forms (Ayodele 1994),
6 Comp. Newsl. 30, 1997
Inflorescence types are associated with adaptation to habitats and pollinators.
Certain reproductive strategies have also been identified with the different types of
inflorescence (Heywood et al. 1977 and Ayodele 1994).
Flowering is a phase of development in plants, regulated by factors external (and
sometimes internal) to the plant. Of all phenological events in plants, the onset of
flowering is considered the most significant. This is because it marks the transition
of the plant from a vegetative to a reproductive mode (Black & Edelman 1970,
Lawn et al. 1995).
The objective of this paper is to report the observations made on the flowering
phase of the development of different species of Vernonia, as they occurred among
field populations and plants of Vernonia raised in experimental garden and
screenhouse.
Materials and methods
Series of field trips, covering different ecological locations of Vernonia in Nigeria,
were made. Regular observations were recorded on field populations during the
flowering periods for three consecutive growing seasons (1989-1992). The
flowering events in the experimental garden and screenhouse were regularly
monitored and compared with those of the plants in the wild. This was done at the
different developmental stages of the plants during each growing season.
Randomly selected plants (25 per species, where abundant) were marked with
labels for regular entries in the garden. Data collected included: date of
sowing/transplanting; date of appearance of flower buds; flower buds at anthesis;
flower maturation time (i.e. number of days to first bloom of the plants); period of
continuous bloom in the plants. For this study, the flowering period of a species
was taken as the period when S50 % or more in a population were in flower.
The means and standard deviation of all data collected were estimated. The ranges
of flowering initiation and period of continuous flowering were also estimated. The
yearly period of flowering for each species, was graphically represented for
comparison of all the species investigated, at the different locations.
Comp. Newsl. 30, 1997 7
Results and discussion
Growth forms in Vernonia
Five growth forms were recorded, namely: Erect tall woody tree, Erect woody
shrub, Straggling woody shrub, Erect annual herb and Erect perennial herb
(Table 1). The growth habit of V. amygdalina is noteworthy. Plants of this species,
growing around homestead and subjected to regular topping, were observed to be
flowering shrubs. Whereas plants growing at more distant locations, where they
had uninterrupted growth, were observed to be tall trees with stem girth as large as
the arboreous species V. conferta.
Flowering and rainfall regime relationship at different locations
The period of rainfall varied among the locations of Vernonia. There was the
longer, dual peak (April - October) rainfall in locations towards the southern part
of the country; and the shorter, single peak (June - October) up North in the
savanna zone.
The period of availability of large populations of species of Vernonia in the wild,
coincided with the period of abundant rainfall at the different locations. This was
also the stage of luxurious vegetative growth in the native habitats of the species.
The duration of this stage was determined by the duration of the rains. The
beginning of the rainy season signals among others: the production of new shoots
by perennial trees and shrubs; shrubs in burnt locations, e.g. V. tenoreana, develop
new shoots from underground rootstocks; fruits dispersed from the herbaceous
species during the previous season, commence germination, and seedlings develop
to flowering adult plants.
The vegetative phase is longer in plants growing at locations with longer rainfall
period. Commencement and duration of flowering were different, even for the
same species (e.g. V. galamensis) occurring in locations with differing rainfall
regimes. However, accessions of this species from different locations, when raised
in the garden/screen-house, flowered and remained in flower for the same duration.
Mode of flowering among the plants of Vernonia
There were major similarities, differences and overlaps in the mode and duration
of flowering among the different species and plant forms (Table 1, Fig. 1).
8 Comp. Newsl. 30, 1997
The synchronised flowering and fruiting sequence was common among the
arboreous species (e.g. V. conferta and V. colorata) and some herbs (e.g.
V. nestor). Plants of these species were usually found in bloom about the same
period of the year, irrespective of the location. A blooming stage is followed by the
fruiting stage, during which plants are found to carry flower or fruits of about the
same stage of development at all locations. This occurs usually only during the dry
season; between the end of a rainy season and the beginning of another
(November - March, Fig. 1). The strong wind which accompanies the first rains
each year, sweeps clean all plant canopies still carrying undispersed fruits.
The second mode was the intermittent alternation of flowering/fruiting and
vegelative growth; common among the shrubs and majority of the herbaceous
species investigated. Flowers and fruits at different developmental stages, could be
found on the plants at any time of the growing season. Flowering is prolonged in
these species, up to 7 months in V. tenoreana (Fig. 1). The moisture status of a
location enhanced the duration of flowering in the species. The observed
differences in the time and duration of flowering in for instance, V. tenoreana,
V. stenostegia and V. kotschyana, all shrubs occurring at different locations in the
wild (Figs. 1 & 2), were eliminated when the three species were grown in the
garden and screenhouse.
Flowering period among the herbaceous species varied among plant populations at
different locations and was generally shorter in duration than among the other plant
forms (Fig. 1, Table 1).
The case of V. cinerea is pertinent for mentioning. There is hardly any time of the
year that a population of V. cinerea cannot be found flowering in any of its native
habitats (Fig. 1). The species was observed capable of thriving at locations with
minimal moisture status, producing its flowers relatively earlier than the other
species. They were thus readily located on lawns and among omamental plants as
weeds throughout the year. The adaptational success of V. cinerea had earlier
earned it the pantropical species” title (Jones 1974). This success was reported
(Ayodele 1992) as suggesting the potentials for an all purpose genotype” earlier
highlighted by Baker (1965).
Flowering among growth forms cohabiting in similar habitats
Some species of Vernonia of different growth habits coexisted in similar habitats.
Certain species were capable of growing in mosaic habitats. Arboreous species
were found in rainforest area. The majority of the herbaceous forms, and some
shrubby forms, were found in both the derived savanna and Guinea savanna areas.
Comp. Newsl. 30, 1997 9
Certain herbs were restricted to the drier Sahel and Guinea savanna area, e.g.
V. ambigua, V. nestor and V. perrottetii.
The dry habitats in the Sahel and Guinea savanna have a short wet season.
Vernonia species growing in these habitats were characterized by a short period of
vegetative growth, early flowering initiation and short period of flowering
(Table 1, Fig. 1), when compared with species growing in wet habitats further
south in the country.
The plants of these species in the drier areas (especially the herbs) were by various
adaptations able to ensure the rapid completion of their life cycle within the period
of the short growing season, characteristic of savanna grassland.
Flowering in arboreous species was more overlapping with that in the shrubs and
later than within the herbs (Fig. 1, Table 1). There was a greater overlapping of
flowering period among the herbs and the shrubs and between the two growth
forms. The interesting case of V. migeodi (a herb) and V. glaberrima (a shrub) is
noteworthy.
Both are species of dry habitats common in Guinea savanna and derived savanna
locations. The two species were usually found growing in a competitive
community which included tall grasses and tall-growing shrubs. Vernonia migeodi
and V. glaberrima usually concluded their flowering and fruiting processes earlier
in the year than the other species of Vernonia with similar growth habits (Fig. 1).
They were the first colonies of plants, vegetatively regenerated or by seed
germination in the different locations. This is with regard to plant succession in
their communities, especially after the annual grassland fires. These species flower
early enough to produce fruits that are dispersed before the plants are smothered
out or shaded by other tall-growing members of the community.
Similarities and differences in flowering periods for species within the same
habitats are relevant for the production of natural hybrids. The potential for
hybridization in Vernonia is reported high (Jones 1972). Cases of natural and
artificial hybridization are rather common place occurrence between species of
Vernonia in the Americas (Jones 1966, Berry et al. 1970, Faust 1972, Jones 1972
and 1973, Lai & Lessman 1974). Jones (1977) reported that natural hybrids among
species are common in Vernonia and often make classification difficult at species
level. Reports are scanty on the preponderance of natural hybrids among Nigerian
species of Vernonia.
However, in spite of the seemingly easy hybridization in the genus, there are
isolating mechanisms which preserve the essential integrity of the species of
Vernonia. These include ecological, seasonal, geographical isolating mechanisms
10 Comp. Newsl. 30, 1997
and in certain combinations, hybrid inviability or sterility in later generations
(Jones 1976). Of all these, ecological isolation was noted as especially significant
(Jones 1968 and 1972).
It would be of interest to investigate the occurrence of natural hybrids at the
locations where there are overlaps in flowering season among the species of
Vernonia.
There are current efforts to domesticate some species of Vernonia, e.g.
V. galamensis as oilseed crop (Perdue & Dierig 1994). The present investigation
on flowering sequence reveals the economical advantage of growing species like
V. galamensis in the dry locations, where the vegetative growth is reduced and the
flowering initiation and period are shorter than in the wet locations.
References
Aoydele, M.S. 1992. Cytogenetic and reproductive studies on some species of
Vernonia Schreb. (Asteraceae) in Nigeria. Ph.D. Thesis, Obafemi Awolowo
University, Ile-Ife, Nigeria. 290 pp.
Ayodele, M.S. 1994. Studies on the reproductive biology of Vernonia Schreb.
(Asteraceae). I. Types of inflorescence among different growth habits. Comp.
Newsl. 25: 15-23.
Baker, H.G. 1965. Characteristics and modes of origin of weeds. Jn: Baker, H.G.
& G.L. Stebbins (eds.), Genetics of colonising species, pp. 147-168. Academic
Press, New York.
Berry, C.D., Lessman, K.J. & G.A. White 1970. Natural cross-fertilization in
Vernonia anthelmintica (L.) Willd. Crop Science 10: 104-105.
Black, M. & J. Edelman 1970. Plant Growth. Heineman Educational Books Ltd.,
London, 193 pp.
Dierig, D.A. 1994. Domestication of Vernonia galamensis, a new epoxy fatty acid
oilseed crop. Progress reports for AMFRR Program. USDA-ARS US. Water
Conserv. Lab.
Faust, W.Z. 1972. A biosystematic study of the /nteriores species group of genus
Vernonia (Compositae). Brittonia 24: 363-378.
Comp. Newsl. 30, 1997 11
Heywood, W.H., Harborne, J.B. & B.L. Turner 1977. An overture to the
Compositae. /n: Heywood, V.H., Harborme, J.B. & B.L. Turner (eds.), The
Biology and Chemistry of the Compositae 1: 1-20. Academic Press, London &
New York.
Hutchinson, J. & J.M. Dalziel 1963. Flora of West Tropical Africa. 2” ed.
revised by F.N. Hepper Vol. 2: 271-283. Crown Agents, London.
Jones, S.B. Jr 1966. Experimental hybridization in Vernonia (Compositae).
Brittonia 18: 19-44.
Jones, S.B. Jr 1968. An example of a Vernonia hybrid in a disturbed habitat.
Rhodora 70: 486-491.
Jones, S.B. Jr 1972. A systematic study of the Fasciculatae group of Vernonia
(Compositae). Brittonia 24: 28-45.
Jones, S.B. Jr 1973. Revision of Vernonia Section Eremosis (Compositae), in
North America. Brittonia 24: 86-115.
Jones, S.B. Jr 1974. Vernonieae (Compositae). Chromosome numbers. Bull.
Torrey Bot. Club 101: 1-34.
Jones, S.B. Jr 1976. Cytogenetics and affinities of Vernonia (Compositae) from
the Mexican highlands and eastern North America. Evolution 30: 455-462.
Jones, S.B. Jr 1977. Vermonieae - systematic review. /n: Heywood, V.H.,
Harbome, J.B. & B.L. Tumer (cds.), The Biology and Chemistry of the
Compositae 1: 503-521. Academic Press, London & New York.
Lai, W.Y. & K.J. Lessman 1974. Combining ability for eight characters of a
four-parent diallele cross in Vernonia anthelmintica (L.) Willd. Crop Science
14: 569-571.
Lawn, R.J., Summerfield, R.J., Ellis, R.H., Oi, A., Roberts, E.H., Chay, P.M.,
Brouwer, J.B., Rose, J.L. & S.J. Yeates 1995. Towards the reliable
prediction of time to flowering in six annual crops VI. Applications in crop
improvement. Expl. Agric. 31: 89-108.
Olorode, O. 1984. Taxonomy of West African Flowering Plants. Longman Group
Lid., London & New York. 158 pp.
Perdue, R.E. Jr 1989. Preliminary trials of Vernonia galamensis. Update Report
on planting guidelines USDA Germplasm Introduction and Evaluation Lab.
Beltsville, MD. USA.
Comp. News|. 30, 1997
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Fig 2 Map of Nigeria Showing the locational distributic
of Some Species of Vernonia.
(Key to numerals are as in Table | )
REPUBLIC OF
K/m 50
NIGER
wet International boundary
~~ State boundary
State capital
OONONAWNOE
Vernonia
Vernonia
Vernonia
Vernonia
Vernonia
Vernonia
Vernonia
Vernonia
Vernonia
Vernonia.
Vernonia
Vernonia
Vernonia
Vernonia
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Comp. Newsl. 30, 1997 15
CHROMOSOMAL STUDIES IN THE EGYPTIAN
FLORA VI. KARYOTYPE FEATURES OF SOME
SPECIES IN SUBFAMILY ASTEROIDEAE
(ASTERACEAE)
Abdelfattah Badr’
Ehab Abdelrazik Kamel’
Naria Garcia-Jacas’
* Botany Department, Faculty of Science, Tanta University, Tanta, Egypt
* Department of Biological Sciences, Faculty of Education,
Ain Shams University, Cairo, Egypt
“Institut Botanic de Barcelona, Barcelona, Spain
Abstract
In this paper, chromosome numbers of 23 species from the flora of Egypt —
including four new counts —, distributed in six tribes of subfamily Asteroideae
(Asteraceae), are reported. Detailed karyotype features, 1. e. chromosome length
(MCL) and karyotype asymmetry expressed as arm ratio (MAR), total form per-
cent (TF%), intrachromosomal asymmetry index (Al) and interchromosomal
asymmetry index (A2), are described.
Introduction
The Asteraceae are represented in the Egyptian wild flora by 93 genera and 230
species (Tackholm 1974). The family is also represented in the weeds of Egypt by
26 species (Boulos & El-Hadidi 1989).
The Asteraceae show a great array of chromosome numbers. Following Solbrig
(1977), numbers vary from as low as n = 2 in Haplopappus gracilis (Nutt.) Gray
and Brachycome lineariloba (DC.) Druce to as high as n= 103 in Werneria apicu-
lata Sch. Bip., n = 106 in Werneria nubigena Kunth, and n = 110-120 in Monta-
noa guatemalensis Robins. & Greenm. However, the most common basic number
in the family is x = 9.
16 Comp. Newsl. 30, 1997
The members of the family in the Egypuan flora have not been a subject of exten-
sive cytological investigations, but Nordenstam (1972) reported chromosome
counts from 24 species of Egyptian Asteraceae. Also, chromosome numbers of
some species which grow in Egypt are known from chromosome counts in plants
from near floras, particularly that of Europe. In the present study, chromosome
numbers and detailed karyotype features of 23 Egyptian species belonging to
subfamily Asteroideae are reported.
Material and methods
Material of 23 species belonging to six tribes of the subfamily Asteroideae was
collected from their natural habitats. The studied species and the localities from
which they were collected are given in Table 1. Collectors are in all the cases A.
Badr and E. Kamel. Vouchers of the collections are preserved in the herbarium of
the Biological Science and Geology Department, Faculty of Education, Ain Shams
University (Egypt).
Cytological preparations were carried out on root tips obtained from seeds
germinated on sterile moist filter papers in Petri dishes at 20-25°C. Roots were pre-
treated with 0.05% colchicine solution for 3-4 h and fixed in Camoy for 24 h and
stored in 70% ethanol at 4°C. Cytological preparations were made using the
Feulgen squash method and well-spread c-metaphase chromosomes were photo-
graphed from temporary preparations at a magnification of 2000x. Slides of the
original karyotypes are also preserved in the Laboratory of Cytogenetics of the
same department.
A karyogram for each species was constructed by arranging the chromosomes in
homologous pairs by order of their length and arm ratio as measured from the pho-
tographic prints and the number of chromosome types were determined as de-
scribed by Levan & al. (1965). Measurements of chromosome lengths were taken
on the same photographs of the karyogram. Karyograms are based in one plate.
The variation in chromosome length (MCL) and chromosome arm ratio (MAR)
within the karyotype has been estimated by calculating the standard error (SE) of
these parameters. Karyotype asymmetry deduced from the ratio between the short
arms of the chromosomes and their total length was expressed as total form percent
(TF%) as proposed by Huziwara (1962). Karyotype asymmetry expressed by the
ratio between the chromosome arms has been also estimated as the intrachromo-
somal asymmetry index (A1) as suggested by Romero Zarco (1986).
Comp. Newsl. 30, 1997 17
The value of Al is framed as to be close to zero if all chromosomes are meta-
centric and near to one if all chromosomes are telocentric. Karyotype asymmetry
due to the ratio between size of different chromosomes has been also estimated as
the interchromosomal asymmetry index (A2) using Pearson’s dispersion coeffi-
cient, that is the ratio between the standard deviation and the mean chromosome
length (Romero Zarco 1986).
The existence of previous chromosome counts for the studied species has been
verified in the indexes of plant chromosome numbers by Fedorov (1969), Goldblatt
(1981, 1984, 1985, 1988), Goldblatt & Johnson (1990, 1991, 1994, 1996) and
Moore (1971, 1972, 1973, 1974, 1977).
Results and discussion
The cytological features of the 23 investigated species are summarised in Table 1.
Tribe Anthemideae
The chromosome numbers and karyotype description are shown for six species of
this tribe. One of them (Achillea fragrantissima) is reported for the first time
(Table 1). Somatic number of 2n = 2x = 18 is found in Achillea fragrantissima,
Chamomilla recutita and Chrysanthemum coronarium. 2n = 4x = 36 1s recorded in
Achillea santolina and Artemisia monosperma. In Artemisia judaica, 2n = 2x = 16
is recorded. The same chromosome numbers have been reported previously for
Achillea santolina, Artemisia judaica, Chamomilla recutita and Chrysanthemum
coronarium. In this last species both diploid (2n = 18) and tetraploid (2n = 36)
numbers were recorded. Our report of 2n = 36 in Artemisia monosperma differs
from that of Nordenstam (1972), who found 2n = 34 in materials from Egypt.
In the tribe Anthemideae, x = 9 seems to be the dominant basic number: it is re-
corded in five of the six species examined here. Only in Artemisia judaica (2n =
16) a basic number of x = 8 is reported. In the genus Artemisia, Fedorov (1969)
listed this chromosome number of x = 8 in 21 species, whereas x = 9 was listed
in 123 species of the genus. Vallés (1987) and Oliva & Vallés (1994) confirm that
x = 9 is dominant in the genus.
The highest MCL (4.20¢0.16*) is recorded in Chrysanthemum coronarium,
whereas the shortest MCL (1.91 *0.11+) in Achillea fragrantissima. The chromo-
somes in this tribe are clearly longer than those of other tribes (Table 1).
18 Comp. Newsl. 30, 1997
Karyotypes of the six species include only metacentric and submetacentric
chromosomes (Fig. 1, A-F) with close similarity between them in the MAR values:
the highest (1.50* 0.09) is recorded in Artemisia monosperma and the lowest
(1.39 0.09) in both Artemisia judaica and Chrysanthemum coronarium. Similar
high values of TF% are also found in the examined species of this tribe. The high
degree of karyotype symmetry in these species is also indicated by similar Al and
A2 values (Table 1).
Tribe Astereae
The Astereae are represented in this study by Aster squamatus and Conyza
linifolia. Chromosome number of Aster squamatus is 2n = 2x = 20, while in
Conyza linifolia 2n = 6x = 54 is scored. Both numbers were previously recorded.
The MCL is 1.48+0.12+* in Aster squamatus and 1.02+0.06¢ in C. linifolia. The
latter species has the shortest chromosomes among the species studied (Fig. 1, G
and H). Aster squamatus has a higher MAR and a lower TF%, as compared to
Conyza linifolia (Table 1). Both species have similar Al values, but Aster
squamatus has considerably higher A2 value (Table 1).
Tribe Calenduleae
In this tribe the chromosome number of Calendula arvensis is examined: 2n = 4x =
36. The same number was recorded by other authors. Nevertheless, a very different
number of 2n = 44 is reported in other counts. The MCL in Calendula arvensis is
1.17* 0.08» and its karyotype is the most symmetric of the studied species: MAR=
1.08* 0.03 and TF%= 48.10. The symmetry of the karyotype of this species (Fig. 2,
A) is also indicated by the Al value (0.07) which is the lowest among the species
investigated (Table 1).
Tribe Heliantheae
The karyotypes of five species in this tribe are studied. In the two species of Xan-
thium, i.e. X. spinosum and X. strumarium, 2n = 4x = 36 is recorded. In Bidens pi-
losa, 2n = 2x = 24 is observed, whereas, in both Helianthus annuus and Verbesina
encelioides the number recorded is 2n = 2x = 34. All our results coincide with
previous counts. However, in B. pilosa both tetraploid (2n = 48) and hexaploid (2n
= 72) numbers were also reported.
Comp. Newsl. 30, 1997 19
The longest chromosomes among the five species of Heliantheae are found in X.
strumarium (MCL= 2.0¢ 0.17*), while the shortest (MCL= 1.19+ 0.08+) are recor-
ded in V. encelioides. The karyotypes of the five species are symmetric (Fig. 2, B-
F), being composed of metacentric chromosomes with small variation among them
in the MAR. The highest MAR (1.45 0.05) is recorded in V. encelioides, whereas
the lowest MAR (1.33¢ 0.05) is found in H. annuus. The low MAR values recorded
in the species of Heliantheae are correlated with high values of the TF%. The
similarity among the studied species of this tribe in karyotype symmetry is also
reflected by similar Al and A2 values (Table 1).
Tribe Inuleae
The tribe Inuleae is represented here by seven species. Chromosome counts and
karyotype descriptions of three of them (Pulicaria undulata, Phagnalon barbey-
anum and Pluchea dioscoridis) are presented here for the first time (Table 1).
Somatic numbers vary between 2n = 2x = 8 in /phiona mucronata to 2n = 4x = 40
in Pluchea dioscoridis. In Pallenis spinosa the somatic number is 2n = 2x = 10,
whereas in Pulicaria undulata 2n = 2x = 12 is recorded. Both in /nula crithmoides
and Phagnalon barbeyanum a diploid number of 2n = 18 is found, while the re-
corded number for Filago desertorum is 2n = 2x = 28. Numbers of /nula crithmoi-
des, Pallenis spinosa and Filago desertorum have been previously scored by other
authors. Our result of 2n = 8 in /phiona mucronata, however, differs from a previ-
ous count of 2n = 18 by Amin (1972), also on Egyptian material.
The highest MCL among the seven species of Inuleae is found in [phiona mucro-
nata (2.0*0.14*), whereas the shortest were observed in Filago desertorum
(1.17* 0.07). The highest MAR value (1.80* 0.15) is recorded in Pulicaria undu-
lata, whereas the lowest (1.11* 0.05) was found in Phagnalon barbeyanum. The
low MAR recorded for the species of this tribe is correlated with high values of the
TF% (Table 1), indicating a high degree of karyotype symmetry. The karyotype
symmetry in the seven species of Inuleae is also illustrated by the presence of only
metacentric and submetacentric chromosomes in the karyotypes of these species
(Fig. 3, A-G). However, the Al and A2 values indicate some degree of karyotype
asymmetry in some species. The Al value ranges between ().10 in P. barbeyanum
to 0.43 in Pulicaria undulata, whereas the highest A2 (0.24) value is found in P.
spinosa and the lowest (0.11) in /nula crithmoides (Table 1).
20 Comp. Newsl. 30, 1997
Tribe Senecioneae
In the two species of this tribe, i.e. Senecio aegyptius and S. vulgaris, 2n = 4x = 40
is recorded. The same number has been reported in other previous counts for both
species. The chromosomes of the two species are similar in length, MCL is
1.27¢0.09¢ in S. aegyptius and 1.19*0.04¢ in S. vulgaris. The chromosomes of
both species are all metacentric (Fig. 3, H and I) with some differences between
them in the MAR, being 1.36¢ 0.03 for S. aegyptius and 1.07+ 0.02 for S. vulgaris.
Differences between these two species in karyotype asymmetry are also reflected
in the values of the TF% and are more clearly manifested in the values of Al and
A2, being 0.26 & 0.23 for S. aegyptius and 0.16 & 0.12 for S. vulgaris respectively
(Table 1).
Conclusions
Of the 23 species studied from the Egyptian flora, polyploid numbers are recorded
in nine species, distributed in the six tribes (Table 1). It is notable that poly-
ploidization occurs only in species with x = 9 or 10.
With regard to the evolution of the basic chromosome number in Asteraceae,
Solbrig (1977) suggested that x = 9 is the ancestral basic chromosome number for
all the family. Basic numbers higher than x = 9 should be the result of cycles of
polyploidy and successive aneuploid reduction; chromosome numbers lower than x
= 9 should be the result of aneuploid reductions. Descending aneuploidy is a gen-
eral trend in the whole family, as it has been repeteadly pointed out by Stebbins,
1950: 449 & 456, tab. 89, and 1971: 93-96.
As to karyotype symmetry, the calculated TF% of the karyotypes of the examined
species ranges between TF%= 35.37 in Pulicaria undulata (Inuleae) to TF%=
48.36 in Senecio vulgaris (Senecioneac). The values of the TF% for the studied
species thus support previous observations (Huziwara 1962, Mehra 1977) that the
karyotype in the Asteraceae is symmetric. The intrachromosomal asymmetry index
(A1), on the other hand, defines some clear differences between the studied species
in the tribes Senecioneae, Inuleae, Heliantheae and Anthemideae. The interchro-
mosomal asymmetry index (A2), however, shows little differences between the
studied species.
Measurements of chromosome length indicate that species in tribe Anthemideae
have substantially longer chromosomes than those in other tribes. The longest
chromosomes are found in Chrysanthemum coronarium (MCL= 4.20+0.16s),
whereas the shortest chromosomes are observed in Conyza linifolia of tribe
Comp. Newsl. 30, 1997 PI
Astereae (MCL= 1.02+ 0.06¢). In all the studied species, however, small differ-
ences in length are recorded among the chromosomes in the karyotype.
References
Amin, A. 1972. Seven chromosome numbers of Egyptian plants. Bot. Notiser 125:
537-538.
Boulos, L. & M.N. El-Hadidi 1989. The weed flora of Egypt. The AUC Press,
Cairo.
Fedorov, A. A. 1969. Chromosome numbers of flowering plants. Academy of
Sciences of the USSR, Leningrad.
Goldblatt, P. 1981. Index to plant chromosome numbers 1975-1978. Monogr.
Syst. Bot. Missouri Bot. Gard. 5.
Goldblatt, P. 1984. Index to plant chromosome numbers 1979-1981. Monogr.
Syst. Bot. Missouri Bot. Gard. 8.
soldblatt, P. 1985. Index to plant chromosome numbers 1982-1983. Monogr.
Syst. Bot. Missouri Bot. Gard. 13.
Goldblatt, P. 1988. Index to plant chromosome numbers 1984-1985. Monogr.
Syst. Bot. Missouri Bot. Gard. 23.
Goldblatt, P. & D.E. Johnson 1990. Index to plant chromosome numbers 1986-
1987. Monoegr.Syst. Bot. Missouri Bot. Gard. 30.
Goldblatt, P. & D.E. Johnson 1991. Index to plant chromosome numbers 1988-
1989. Monogr. Syst. Bot. Missouri Bot. Gard. 40.
Goldblatt, P. & D.E. Johnson1994. Index to plant chromosome numbers 1990-
1991. Monogr. Syst. Bot. Missouri Bot. Gard. 51.
Goldblatt, P. & D.E. Johnson1996. Index to plant chromosome numbers 1992-
1993. Monogr. Syst. Bot. Missouri Bot. Gard. 58.
Huziwara, Y. 1962. Karyotype analysis in some genera of Compositae. VIII - Fur-
ther studies on the chromosomes of Aster. Amer. J. Bot. 49: 116-119.
Leyan, A., Fredga, K. & A.A. Sandberg 1965. Nomenclature for centromeric
position on chromosomes. Hereditas 52: 201-220.
th
i)
Comp. Newsl. 30, 1997
Mehra, P. N. 1977. Cytological invesugations on the Indian Compositae VI.
Chromosomes and evolutionary phylogeny. Cytologia 42: 347-356.
Moore, D. M. 1971. Index to plant chromosome numbers for 1969. Regnum
Vegetabile 77.
Moore, D. M. 1972. Index to plant chromosome numbers for 1970. Regnum
Vegetabile 84.
Moore, D. M. 1973. Index to plant chromosome numbers 1967-1971. Regnum
Vegetabile 90.
Moore, D. M. 1974. Index to plant chromosome numbers for 1972. Regnum
Vegetabile 91.
Moore, D. M. 1977. Index to plant chromosome numbers for 1973/74. Regnum
Vegetabile 96.
Nordenstam, B. 1972. Chromosome numbers in some Compositae from Egypt.
Bot. Notiser 125: 393-396.
Oliva, M., & J. Vallés 1994. Karyological studies in some taxa of the genus
Artemisia (Asteraceae). Can. J. Bot. 72: 1126-1135.
Romero Zarco, C. R. 1986. A new method for estimating karyotype asymmetry.
Taxon 35: 526-530.
Solbrig, O. T. 1977. Chromosomal cytology and evolution in the family
Compositae. Jn: Heywood, V.H., Harborne, J.B. & B. L. Turner (eds.). The
Biology and Chemistry of the Compositae 1: 267-281. Academic Press, London
& New York.
Stebbins, G. L. 1950. Variation and evolution in plants. Columbia University
Press, New York.
Stebbins, G. L. 1971. Chromosomal Evolution in Higher Plants. Amold Press,
London.
Tackholm, V. 1974. Students’ Flora of Egypt. Cairo University Press, Cairo.
Valles, J. 1987. Aportaci6n al conocimiento citotaxonémico de ocho taxones
ibéricos del género Artemisia L. (Asteraceae, Anthemideac). Anales Jard. Bot.
Madrid 44: 79-96.
23
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Figure captions
Figure 1. Karyograms of A) Achillea fragrantissima; B) Achillea santolina; C)
Artemisia judaica, D) Artemisia monosperma; E) Chamomilla recutita; F)
Chrysanthemum coronarium; G) Aster squamatus; H) Conyza linifolia.
Figure 2. Karyograms of A) Calendula arvensis; B) Bidens pilosa; C) Helianthus
annuus, D) Verbesina encelioides; E) Xanthium spinosum; F) Xanthium
strumarium.
Figure 3. Karyograms of A) Filago desertorum; B) Inula crithmoides; C) Iphiona
mucronata; D) Pallenis spinosa; E) Pulicaria undulata; F) Phagnalon
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Comp. Newsl. 30, 1997
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EARLY SUPRAGENERIC NAMES IN
ASTERACEAE
James L. Reveal
Department of Plant Biology
University of Maryland
College Park, Maryland 20742-5815, U.S.A.
jr19@umail.umd.edu
Abstract
Suprageneric names of Asteraceae found to date as part of the Indices Nominum
Supragenericorum Plantarum Vascularium project, and published from 1753
through 1850 as part of a survey of the botanical literature for such names, are
listed with full bibliographic citation. No taxonomic judgment about the
significance (either taxonomically or nomenclaturally) of any name is present,
leaving that to those who use such names in their research.
Introduction
Scientific names of plants above the rank of genus, except for those at the rank of
family, are poorly known and generally unevaluated as to their validity or even
place of publication. An ongoing effort, the Indices Nominum Supragenericorum
Plantarum Vascularium project, under the sponsorship of the International
Association for Plant Taxonomy and the Norton-Brown Herbarium at the
University of Maryland is attempting to prepare a database of all validly published
names above the rank of genus based on a generic name. The results to date are
available online on the world wide web at
http://matrix.nal.usda.gov:8080/star/supragenericname.html
thanks to the cooperation of the U.S. Department of Agriculture’s National
Agricultural Library in Beltsville, Maryland.
The purpose of this published report is to provide users without access to the
database with a preliminary summary of the findings for Asteraceae. This is a large
and important family with numerous, active workers, and a literature base that is
30 Comp. Newsl. 30, 1997
enormous. My goal is to encourage workers to provide me with information on
names not yet found by sending copies of works that I have missed in my survey of
the literature published through 1850. In this way, information on the suprageneric
nomenclature of Asteraceae will be more widely distributed and hopefully more
correctly applied. My email address is
jr19@umail.umd.edu and my fax number is (301) 314-9082.
Some minor facts about suprageneric nomenclature are important to note. First,
only names at the rank of family may be conserved. Second, names above the rank
of family are not subjected to the concept of priority. Third, while descriptive
names are permitted above the rank of family, such names can not be typified and
may be used in any sense one may wish, the only requirement being that the name
be validly published at that rank. For example, the name Centrospermae has been
used for a group of families related to Chenopodiaceae as well as for a group that
includes Lemnaceae. There is nothing to prevent one from using this name for any
other combination of families, at the rank of order, as long as one or more
representatives had seeds that are centrospermous. For this reason, such names are
not included. Finally, it should be noted that many authors use suprageneric names
without ascertaining their validity. It is strongly urged that all such names be
checked thoroughly before used in print.
Listing of Suprageneric Names for Asteraceae
Acarna Boehm. in C.G. Ludwig, Def. Gen. PI., ed. 3: 195. 1760.
Fam. Acarnaceae Link, Handbuch 1: 684. Jan-Aug 1829.
Adenostyles Cass. in G.F. Cuvier, Dict. Sci. Nat. 1, Suppl.: 59. 12 Oct 1816.
Subfam. Adenostyloideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1073.
1829 (Adenostyleae), based on Tribe Adenostyleae A.H.G. de Cassini, J. Phys.
Chim. Hist. Nat. 88: 201. Mar 1819.
Tribe Adenostyleae Cass., J. Phys. Chim. Hist. Nat. 88: 201. Mar 1819.
Subtribe Adenostylinae Cass. ex Dumort., Anal. Fam. PL: 31. 1829
(Adenostyleae).
Ageratum L., Sp. Pl.: 839. 1 Mai 1753.
Tribe Agerateae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829.
Subtribe Ageratinae Less., Syn. Gen. Compos.: 155. Jul-Aug 1832
(Ageratcac).
Comp. Newsl. 30, 1997 3]
Alomia Kunth in F.W.H.A. von Humboldt, A.J.A. Bonpland & C.S. Kunth, Nov.
Gen. Sp. 4, ed. f*: 118. 26 Oct 1818.
Subtribe Alomiinae Less., Syn. Gen. Compos.: 154. Jul-Aug 1832
(Alomieae).
Ambrosia L., Sp. Pl.: 987. 1 Mai 1753
Order Ambrosiales Dumort., Anal. Fam. Pl.: 15. 1829 (Ambrosarieae).
Fam. Ambrosiaceae Link, Handbuch 1: 816. Jan-Aug 1829, nom. cons.
Subfam. Ambrosioideae Raf., Ann. Gen. Sci. Phys. Bruxelles 6: 88. 1820
(Ambrosidia).
Tribe Ambrosieae Cass., J. Phys. Chim. Hist. Nat. 88: 191. Mar 1819.
Subtribe Ambrosiinae Less., Linnaea 5: 151. Jan 1830 (Ambrosieae).
Anthemis L., Sp. Pl.: 893. 1 Mai 1753.
Fam. Anthemidaceae Martinov, Tekhno-Bot. Slovar: 33. 1820.
Subfam. Anthemidoideae (Cass.) Lindl., Encycl. Pl.: 1073. 1829
(Anthemideae), based on Tribe Anthemideae A.H.G. de Cassini, J. Phys.
Chim. Hist. Nat. 88: 192. Mar 1819.
Tribe Anthemideae Cass., J. Phys. Chim. Hist. Nat. 88: 192. Mar 1819.
Subtribe Anthemidinae (Cass.) Dumort., Fl. Belg.: 69. 1827 (Anthemideae),
based on Tribe Anthemideae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88:
192. Mar 1819.
Aposeris Neck. ex Cass. in J.B.G.M. Bory de Saint-Vincent, Dict. Class. Hist. Nat.
48: 427. Jun 1827.
Fam. Aposeridaceae Raf., New Fl. N. Amer. 4: 106. late Sep 1838
(Aposerides).
Arctotis L., Sp. Pl.: 922. 1 Mai 1753.
Subfam. Arctotidoideae (Cass.) Lindl., Encycl. Pl.: 1073. 1829 (Arctotideae),
based on Tribe Arctotideae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88:
159. Feb 1819.
Tribe Arctotideae Cass., J. Phys. Chim. Hist. Nat. 88: 159. Feb 1819.
Subtribe Arctotidinae (Cass.) Dumort., Anal. Fam. Pl.: 32. 1829
(Arctotideae), based on Tribe Arctotideae A.H.G. de Cassini, J. Phys. Chim.
Hist. Nat. 88: 159. Feb 1819.
Artemisia L., Sp. Pl.: 845. 1 Mai 1753.
Fam. Artemisiaceae Martinov, Tekhno-Bot. Slovar: 48. 1820 (Artemisiae).
Subfam. Artemisioideae Burmeist., Handb. Naturgesch.: 291. 1837
(Artemisieae).
32 Comp. Newsl. 30, 1997
Tribe Artemisieae Kostel., Allg. Med.-Pharm. Fl. 2: 693. Jan-Jun 1833.
Subtribe Artemisiinae Less., Linnaea 5: 163. Jan 1830 (Artemisieae).
Aster L., Sp. Pl.: 872. 1 Mai 1753.
Class Asteropsida Brongn., Enum. Pl. Mus. Paris: xvii, 32. 12 Aug 1843
(Asteroideae).
Order Asterales Lindl., Nix. Pl.: 20. 17 Sep 1833.
Suborder Asterineae Burmett, Outul. Bot.: 901, 1111. Jun 1835 (Asterosae).
Fam. Asteraceae Dumort., Comment. Bot.: 55. Nov-Dec 1822 (Astereae),
nom. cons.
Fam. Compositae Giseke, Prael. Ord. Nat. Pl.: 538. Apr 1792, nom. alt. et
nom. cons.
Subfam. Asteroideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829
(Astereae), based on Tribe Astereae A.H.G. de Cassini, J. Phys. Chim. Hist.
Nat. 88: 195. Mar 1819.
Tribe Astereae Cass., J. Phys. Chim. Hist. Nat. 88: 195. Mar 1819.
Subtribe Asterinae (Cass.) Dumort., Fl. Belg.: 66. 1827 (Astereae), based on
Tribe Astereac A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88: 195. Mar 1819.
Athanasia L., Sp. Pl., ed. 2: 1180. Jul-Aug 1763.
Fam. Athanasiaceae Martinov, Tekhno-Bot. Slovar: 56. 1820 (Athanasiae).
Baccharis L., Sp. Pl.: 860. 1 Mai 1753, nom. cons.
Subfam. Baccharidoideae Burmeist., Handb. Naturgesch.: 294. 1837
(Baccharideae).
Tribe Baccharideae Kostel., Allg. Med.-Pharm. Fl. 2: 665. Jan-Jun 1833.
Subtribe Baccharidinae Less., Linnaea 5: 145. Jan 1830 (Baccharideae).
Barnadesia Mutis ex L.f., Suppl. Pl.: 55, 348. Apr 1782.
Tribe Barnadesieae D. Don, Trans. Linn. Soc. London 16: 273. 27 Mai 1830
(Barnadeseac).
Bellis L., Sp. Pl.: 886. 1 Mai 1753.
Tribe Bellideae Cass. ex D. Don in R. Sweet, Brit. Fl. Gard. 1: ad t. 38. Mar
1830.
Bellium L., Mant. Pl.: 157, 285. Oct 1771.
Tribe Bellieae DC. ex Godr. in J.C.M. Grenier & D.A. Godron, Fl. France 2:
83, 104. Nov 1850.
Comp. Newsl. 30, 1997 33
Bidens L., Sp. Pl.: 831. 1 Mai 1753.
Tribe Bidentideae (Less.) Godr. in J.C.M. Grenier & D.A. Godron, Fl. France
2: 84, 168. Nov 1850, based on [Rankless] Bidentideae C.F. Lessing, Syn. Gen.
Compos.: 229. Jul-Aug 1832.
Subtribe Bidentidinae Griseb., Spic. Fl. Rumel. 2: 226. Jan 1846
(Bidentideae), based on [Rankless] Bidentideae C.F. Lessing, Syn. Gen.
Compos.: 229. Jul-Aug 1832.
Buphthalmum L., Sp. Pl.: 903. 1 Mai 1753.
Subfam. Buphthalmoideae Burmeist., Handb. Naturgesch.: 293. 1837
(Buphthalmeae).
Tribe Buphthalmeae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829.
Subtribe Buphthalminae Less., Linnaea 6: 153. post Mar 1831
(Buphthalmeae).
Calendula L., Sp. Pl.: 921. 1 Mai 1753.
Fam. Calendulaceae Link, Handbuch 1: 776. Jan-Aug 1829.
Subfam. Calenduloideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074.
1829 (Calenduleae), based on Tribe Calenduleae A.H.G. de Cassini, J. Phys.
Chim. Hist. Nat. 88: 161. Feb 1819.
Tribe Calenduleae Cass., J. Phys. Chim. Hist. Nat. 88: 161. Feb 1819.
Subtribe Calendulinae (Cass.) Dumort., Anal. Fam. Pl.: 32. 1829
(Calenduleae), based on Tribe Calenduleae A.H.G. de Cassini, J. Phys. Chim.
Hist. Nat. 88: 161. Feb 1819.
Cardopatium Juss., Ann. Mus. Natl. Hist. Nat. 6: 324. 1805.
Tribe Cardopatieae Kostel., Allg. Med.-Pharm. FI. 2: 620. Jan-Jun 1833
(Cardopateae).
Subtribe Cardopatiinae Less., Syn. Gen. Compos.: 14. Jul-Aug 1832
(Cardopateac).
Carduus L., Sp. Pl.: 820. 1 Mai 1753.
Fam. Carduaceae Dumort., Comment. Bot.: 56. Nov-Dec 1822.
Subfam. Carduoideae Cass. ex Sweet, Hort. Brit.: 213. Jul-Aug 1826
(Carduaceae).
Tribe Cardueae Cass., J. Phys. Chim. Hist. Nat. 88: 155. Feb 1819
(Carduineae).
Subtribe Carduinae (Cass.) Dumort., Fl. Belg.: 73. 1827 (Carduineae), based
on Tribe Cardueae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88: 155. Feb
1819 (Carduineae).
34 Comp. Newsl. 30, 1997
Carlina L., Sp. Pl.: 828. 1 Mai 1753.
Subfam. Carlinoideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829
(Carlineae), based on Tribe Carlineae A.H.G. de Cassini, J. Phys. Chim. Hist.
Nat. 88: 152. Feb 1819.
Tribe Carlineae Cass., J. Phys. Chim. Hist. Nat. 88: 152. Feb 1819.
Subtribe Carlininae (Cass.) Dumort., Fl. Belg.: 72. 1827 (Carlineae), based
on Tribe Carlineae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88: 152. Feb
1819.
Carthamus L., Sp. Pl.: 830. 1 Mai 1753.
Tribe Carthameae Kitt., Taschenb. Fl. Deutschl., 2, 2: 557. 1844, based on
Subtribe Carthaminae A.P. de Candolle, Prodr. 6: 450, 609. early Jan 1838
(Carthameae).
Subtribe Carthaminae DC., Prodr. 6: 450, 609. early Jan 1838 (Carthameae).
Catananche L., Sp. Pl.: 812. 1 Mai 1753.
Tribe Catanancheae D. Don, Edinburgh New Philos. J. 6: 307. Jan-Mar 1829.
Centaurea L., Sp. Pl.: 909. 1 Mai 1753.
Fam. Centaureaceae Martinov, Tekhno-Bot. Slovar: 117. 1820 (Centaureae).
Subfam. Centaureoideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074.
1829 (Centaurieae), based on Tribe Centaureeae A.H.G. de Cassini, J. Phys.
Chim. Hist. Nat. 88: 154. Feb 1819 (Centaurieae).
Tribe Centaureeae Cass., J. Phys. Chim. Hist. Nat. 88: 154. Feb 1819
(Centaurieac).
Subtribe Centaureinae (Cass.) Dumort., Fl. Belg.: 72. 1827 (Centaurieae),
based on Tribe Centaureeae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88:
154. Feb 1819 (Centaurieae).
Chaetanthera Ruiz & Pav., Fl. Peruv. Prodr.: 106. Oct (prim.) 1794.
Tribe Chaetanthereae D. Don, Trans. Linn. Soc. London 16: 232. 27 Mai
1830.
Chondrilla L., Sp. Pl.: 796. 1 Mai 1753.
Tribe Chondrilleae W.D.J. Koch, Syn. Fl. Germ. Helv.: 427. Jan-Oct 1837.
Subtribe Chondrillinae (W.D.J. Koch) M. Lamotte, Cat. Pl. Eur. Centr.: 56.
Jul-Dec 1847 (Chondrilleae), based on Tribe Chondrilleae W.D.J. Koch, Syn.
Fl. Germ. Helv.: 427. Jan-Oct 1837.
Comp. Newsl. 30, 1997 35
Chrysanthemum L., Sp. Pl.: 887. 1 Mai 1753.
Subfam. Chrysanthoideae Burmeist., Handb. Naturgesch.: 292. 1837
(Chrysanthemeae).
Tribe Chrysanthemeae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829.
Subtribe Chrysantheminae Less., Linnaea 6: 167. post Mar 1830
(Chrysanthemeae).
Cichorium L., Sp. Pl.: 813. 1 Mai 1753.
Fam. Cichoriaceae Juss., Gen. Pl.: 168. 4 Aug 1789 (Cichoraceae), nom.
cons.
Subfam. Cichorioideae (Juss.) Chev., Fl. Gen. Env. Paris 2: 531. 5 Jan 1828
(Cichoraceae), based on Fam. Cichoriaceae A.L. de Jussieu, Gen. Pl.: 168. 4
Aug 1789, nom. cons.
Tribe Cichorieae Lam. & DC., Syn. Pl. Fl. Gall.: 255. 30 Jun 1806
(Cichoraceae).
Subtribe Cichoriinae Cass. ex Dumort., Anal. Fam. Pl.: 30. 1829 (Cichoreae).
Cnicus L., Sp. Pl.: 826. 1 Mai 1753.
Fam. Cnicaceae Vest, Anleit. Stud. Bot.: 273, 297. 1818 (Cnicoideae).
Conyza Less., Syn. Gen. Compos.: 203. Jul-Aug 1832, nom. cons.
Subtribe Conyzinae Horan., Char. Ess. Fam.: 93. 1847 (Conyzinae).
Coreopsis L., Sp. Pl.: 907. 1 Mai 1753.
Fam. Coreopsidaceae Link, Handbuch 1: 768. Jan-Aug 1829 (Coreopsideae).
Tribe Coreopsideae Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829.
Subtribe Coreopsidinae Cass. ex Dumort., Anal. Fam. Pl.: 31. 1829
(Coreopsideae).
Cotula L., Sp. Pl.: 891. 1 Mai 1753.
Subtribe Cotulinae (Less.) Kitt., Taschenb. Fl. Deutschl., 2, 2: 609. 1844
(Cotuleae), based on [Rankless] Cotuleae C.F. Lessing, Syn. Gen. Compos.:
260. Jul-Aug 1832.
Crepis L., Sp. Pl.: 805. 1 Mai 1753, nom. cons..
Tribe Crepideae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829 (Crepideae).
Subtribe Crepidinae Cass. ex Dumort., Fl. Belg.: 60. 1827 (Crepideae).
Crupina (Pers.) DC., Ann. Mus. Natl. Hist. Nat. 16: 157. Jul-Dec 1810.
Tribe Crupineae Godr. in J.C.M. Grenier & D.A. Godron, Fl. France 2: 200,
266. Nov 1850.
36 Comp. Newsl. 30, 1997
Cynara L., Sp. Pl.: 827. 1 Mai 1753.
Suborder Cynarineae Raf., Anal. Nat.: 191. Apr-Jul 1815 (Cynarea).
Fam. Cynaraceae Durande, Notions Elém. Bot.: 273. 1782 (Cinarocephalae).
Subfam. Cynaroideae (Durande) Chev., Fl. Gen. Env. Paris 2: 557. 5 Jan
1828 (Cynarocephalae), based on Fam. Cynaraceae J.F. Durande, Notions
Elém. Bot.: 273. 1782 (Cinarocephalae).
Tribe Cynareae Lam. & DC., Syn. PI. Fl. Gall.: 267. 30 Jun 1806
(Cynarocephalae).
Diazeuxis D. Don, Trans. Linn. Soc. London 16: 251. 27 Mai 1830.
Tribe Diazeuxideae D. Don, Trans. Linn. Soc. London 16: 251. 27 Mai 1830
(Diazeuxeae).
Echinops L., Sp. Pl.: 814. 1 Mai 1753.
Fam. Echinopaceae Dumort., Comment. Bot.: 56. Nov-Dec 1822
(Echinopsidea).
Subfam. Echinopsoideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074.
1829 (Echinopseae), based on Tribe Echinopseae A.H.G. de Cassini in G.F.
Cuvier, Dict. Sci. Nat. 14: 200. 14 Aug 1819.
Tribe Echinopseae Cass., J. Phys. Chim. Hist. Nat. 88: 157. Feb 1819.
Subtribe Echinopsinae (Cass.) Dumort., Anal. Fam. Pl.: 32. 1829
(Echinopsideae), based on Tribe Echinopseae A.H.G. de Cassini in G.F.
Cuvier, Dict. Sci. Nat. 14: 200. 14 Aug 1819.
Eclipta L., Mant. Pl.: 157, 286. Oct 1771, nom. cons..
Subfam. Ecliptoideae Burmeist., Handb. Naturgesch.: 293. 1837 (Eclipteae).
Tribe Eclipteae Kostel., Allg. Med.-Pharm. FI. 2: 670. Jan-Jun 1833.
Subtribe Ecliptinae Less., Linnaea 6: 153. post Mar 1831 (Ecliptae).
Elephantopus L., Sp. Pl.: 814. 1 Mai 1753.
Subtribe Elephantopinae Less., Linnaea 5: 135. Jan 1830 (Elephantopeae).
Erigeron L., Sp. Pl.: 863. 1 Mai 1753, nom. cons..
Tribe Erigeroneae Gren. & Godr., Fl. France 2: 83, 92. Nov 1850
(Erigerineac).
Comp. Newsl. 30, 1997 37
Eupatorium L., Sp. Pl.: 836. 1 Mai 1753.
Fam. Eupatoriaceae Martinov, Tekhno-Bot. Slovar: 239. 1820.
Subfam. Eupatorioideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1073.
1829 (Eupatorieae), based on Tribe Eupatorieae A.H.G. de Cassini in G.F.
Cuvier, Dict. Sci. Nat. 16: 9. 8 Apr 1820 (Eupatoriae).
Tribe Eupatorieae Cass., J. Phys. Chim. Hist. Nat. 88: 202. Mar 1819.
Subtribe Eupatoriinae Dumort., Anal. Fam. Pl.: 31. 1829 (Eupatorieae).
Facelis Cass., Bull. Sci. Soc. Philom. Paris 1819: 94. Jun 1819.
Tribe Facelideae Kostel., Allg. Med.-Pharm. FI. 2: 623. Jan-Jun 1833.
Subtribe Facelidinae Less., Linnaca 5: 361. Jul 1830 (Facelideae).
Fidelia Schultz-Bip., Flora 17: 478. 14 Aug 1834.
Subtribe Fideliinae Schultz-Bip., Flora 17: 478. 14 Aug 1834 (Fidelieae).
Flaveria Juss., Gen. Pl.: 186. 4 Aug 1789.
Subtribe Flaveriinae Less., Syn. Gen. Compos.: 235. Jul-Aug 1832.
Gaillardia Foug., Observ. Phys. 29: 55. Jul 1786.
Tribe Gaillardieae (Nutt.) Lecog & Juillet, Dict. Rais. Term. Bot.: 286. 1831
(Galardieae), based on [Rankless] Galardiae T. Nuttall, Gen. N. Amer. PI. 2:
177. 14 Jul 1818.
Subtribe Gaillardinae (Nutt.) Less., Linnaea 6: 516. Jul-Dec 1831
(Galardieae), based on [Rankless] Galardiae T. Nuttall, Gen. N. Amer. PI. 2:
177. 14 Jul 1818.
Gerbera L., Opera Var.: 247. 1758, nom. cons.
Tribe Gerbereae Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829.
Gnaphalium L., Sp. Pl.: 850. 1 Mai 1753.
Fam. Gnaphaliaceae Link ex Rudolphi, Syst. Orb. Veg.: 46. 1830
(Gnaphalieae).
Subfam. Gnaphalioideae Burmeist., Handb. Naturgesch.: 291. 1837
(Gnaphalieae).
Tribe Gnaphalieae (Cass.) Lecoq & Juillet, Dict. Rais. Term. Bot.: 296. 1831,
based on [Rankless] Gnaphalicac A.H.G. de Cassini in G.F. Cuvier, Dict. Sci.
Nat. 19: 122. 26 Jan 1821.
Subtribe Gnaphaliinae (Cass.) Dumort., Anal. Fam. Pl.: 31. 1829
(Gnaphalieae), based on [Rankless] Gnaphalieae A.H.G. de Cassini in G.F.
Cuvier, Dict. Sci. Nat. 19: 122. 26 Jan 1821.
38 Comp. Newsl. 30, 1997
Gorteria L., Syst. Nat., ed. 10: 1189, 1229, 1358, 1377. 7 Jun 1759.
Tribe Gorterieae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829.
Gundelia L., Sp. Pl.: 814. 1 Mai 1753.
Tribe Gundelieae DC. ex Lecog & Juillet, Dict. Rais. Term. Bot.: 306. 1831
(Gundeliaceae).
Helenium L., Sp. Pl.: 886. 1 Mai 1753.
Fam. Heleniaceae Raf., Cincinnati Lit. Gaz. 2: 28. 24 Jul 1824 (Helenidia).
Subfam. Helenioideae Burmeist., Handb. Naturgesch.: 292. 1837 (Helenieae).
Tribe Helenieae Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829.
Subtribe Heleniinae Cass. ex Dumort., Anal. Fam. Pl.: 31. 1829 (Helenieae).
Heltanthus L., Sp. Pl.: 904. 1 Mai 1753.
Fam. Helianthaceae Dumort., Comment. Bot.: 56. Nov-Dec 1822
(Heliantheae).
Subfam. Helianthoideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074.
1829 (Heliantheae), based on Tribe Heliantheae A.H.G. de Cassini, J. Phys.
Chim. Hist. Nat. 88: 189. Mar 1819.
Tribe Heliantheae Cass., J. Phys. Chim. Hist. Nat. 88: 189. Mar 1819.
Subtribe Helianthinae (Cass.) Dumort., Fl. Belg.: 71. 1827 (Heliantheae),
based on Tribe Heliantheae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88:
189. Mar 1819.
Helichrysum Mill., Gard. Dict. Abr., cd. 4: unpaged [462]. 28 Jan 1754, nom. cons.
Fam. Helichrysaceae Link, Handbuch 1: 712. Jan-Aug 1829 (Elichryseae).
Hieracium L., Sp. Pl.: 799. 1 Mai 1753.
Subfam. Hieracioideae Burmeist., Handb. Naturgesch.: 296. 1837
(Hieracieae).
Tribe Hieracieae D. Don, Edinburgh New Philos. J. 6: 306. Jan-Mar 1829
(Hieraceae).
Subtribe Hieraciinae Cass. ex Dumort., Fl. Belg.: 62. 1827 (Hieracieae).
Comp. Newsl. 30, 1997 39
Hyoseris L., Sp. Pl.: 808. 1 Mai 1753.
Subfam. Hyoseridoideae Burmeist., Handb. Naturgesch.: 298. 1837
(Hyoserideae).
Tribe Hyoserideae Kostel., Allg. Med.-Pharm. Fl. 2: 625. Jan-Jun 1833.
Subtribe Hyoseridinae Less., Syn. Gen. Compos.: 127. Jul-Aug 1832
(Hyoserideae).
Hypochaeris L., Sp. Pl.: 810. 1 Mai 1753.
Subfam. Hypochaeridoideae Burmeist., Handb. Naturgesch.: 298. 1837
(Hypochaerideae).
Tribe Hypochaerideae D. Don, Edinburgh New Philos. J. 6: 307. Jan-Mar
1829 (Hypochoerideae).
Subtribe Hypochaeridinae Less., Syn. Gen. Compos.: 130. Jul-Aug 1832
(Hypochoerideae).
Inula L., Sp. Pl.: 881. 1 Mai 1753.
Subfam. Inuloideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829
(Inuleae), based on Tribe Inuleae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat.
88: 193. Mar 1819.
Tribe Inuleae Cass., J. Phys. Chim. Hist. Nat. 88: 193. Mar 1819.
Subtribe Inulinae (Cass.) Dumort., Fl. Belg.: 67. 1827 (Inuleae), based on
Tribe Inuleae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88: 193. Mar 1819.
Iva L., Sp. Pl.: 988. 1 Mai 1753.
Tribe Iveae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829.
Jacobaea Mill., Gard. Dict. Abr., ed. 4: unpaged. 28 Jan 1754.
Tribe Jacobaeae Dumort., Fl. Belg.: 65. 1827 (Jacobaceae).
Jungia L.f., Suppl. Pl.: 58, 380. Apr 1782.
Tribe Jungieae D. Don, Trans. Linn. Soc. London 16: 224. 27 Mai 1830
(Jungeae).
Eactuca. Sp; Pi:: 795. 1 Mai 1753.
Subfam. Lactucoideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829,
based on Tribe Lactuceae A.H.G. de Cassini in G.F. Cuvier, Dict. Sci. Nat. 20:
355. 29 Jun 1821.
Tribe Lactuceae Cass., J. Phys. Chim. Hist. Nat. 88: 151. Feb 1819.
Subtribe Lactucinae Cass. ex Dumort., Fl. Belg.: 59. 1827.
40 Comp. Newsil. 30, 1997
Lapsana L., Sp. Pl.: 811. 1 Mai 1753.
Tribe Lapsaneae Kostel., Allg. Med.-Pharm. Fl. 2: 624. Jan-Jun 1833.
Subtribe Lapsaninae Cass. ex Dumort., Anal. Fam. Pl.: 30. 1829 (Lapsaneae).
Leontodon L., Sp. Pl.: 798. 1 Mai 1753, nom. cons.
Tribe Leontodonteae (Schultz-Bip.) W.D.J. Koch, Syn. Fl. Germ. Helv.: 417.
Jan-Oct 1837, based on Subtribe Leontodontinae C.H. Schultz-Bipontinus,
Flora 17: 478. 14 Aug 1834 (Leontodonteae verae).
Subtribe Leontodontinae Schultz-Bip., Flora 17: 478. 14 Aug 1834
(Leontodonteae verae).
Leysera L., Sp. Pl., ed. 2: 1249. Aug 1763.
Subtribe Leyseriinae Less., Syn. Gen. Compos.: 363. Jul-Aug 1832
(Leysserieae).
Liabum Adans., Fam. Pl. 2: 131. Jul-Aug 1763.
Subtribe Liabinae Cass. ex Dumort., Anal. Fam. PI.: 31. 1829 (Liabeae).
Liatris Gaertn. ex Schreb., Gen. Pl. 2: 542. Mai 1791, nom. cons.
Tribe Liatrideae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829.
Subtribe Liatridinae Dumort., Anal. Fam. Pl.: 31. 1829 (Liatrideae).
Matricaria L., Sp. Pl.: 890. 1 Mai 1753, nom. cons.
Fam. Matricariaceae Voigt, Hort. Suburb. Calcutt.: 400. Jul-Dec 1845.
Melampodium L., Sp. Pl.: 921. 1 Mai 1753.
Subfam. Melampodioideae Burmeist., Handb. Naturgesch.: 293. 1837
(Melampodieae).
Tribe Melampodieae D. Don, Edwards’s Bot. Reg. 17: ad t. 1458. 1 Dec
1831.
Subtribe Melampodiinae Less., Linnaea 5: 149. Jan 1830 (Melampodieae).
Milleria L., Sp. Pl.: 919. 1 Mai 1753.
Tribe Millerieae Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829.
Subtribe Milleriinae Cass. ex Dumort., Anal. Fam. Pl.: 31. 1829 (Millerieae).
Comp. Newsl. 30, 1997 4]
Mutisia L.f., Suppl. Pl.: 57, 373. Apr 1782.
Fam. Mutisiaceae Burnett, Outl. Bot.: 934, 935, 1094, 1111. Jun 1835.
Subfam. Mutisioideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829
(Mutisieae), based on Tribe Mutisieae A.H.G. de Cassini in G.F. Cuvier, Dict.
Sci. Nat. 20: 379. 29 Jun 1821.
Tribe Mutisieae Cass., J. Phys. Chim. Hist. Nat. 88: 199. Mar 1819.
Subtribe Mutisiinae (Cass.) Dumort., Anal. Fam. Pl.: 31. 1829 (Mutisieae),
based on Tribe Mutisieae A.H.G. de Cassini in G.F. Cuvier, Dict. Sci. Nat. 20:
379. 29 Jun 1821.
Nassauvia Comm. ex Juss., Gen. Pl.: 175. 4 Aug 1789.
Fam. Nassauviaceae Burmeist., Handb. Naturgesch. 1: 290. 1837.
Subfam. Nassauvioideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074.
1829 (Nassauvieae), based on Tribe Nassauvieae A.H.G. de Cassini in G.F.
Cuvier, Dict. Sci. Nat. 20: 378. 29 Jun 1821.
Tribe Nassauvieae Cass., J. Phys. Chim. Hist. Nat. 88: 198. Mar 1819.
Subtribe Nassauviinae (Cass.) Dumort., Anal. Fam. Pl.: 31. 1829
(Nassauvieae), based on Tribe Nassauvieae A.H.G. de Cassini in G.F. Cuvier,
Dict. Sci. Nat. 20: 378. 29 Jun 1821.
Osteospermum L., Sp. Pl.: 923. 1 Mai 1753, nom. cons.
Tribe Osteospermeae Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829.
Othonna L., Sp. Pl.: 924. 1 Mai 1753.
Subfam. Othonnoideae Burmeist., Handb. Naturgesch.: 299. 1837
(Othonneae).
Tribe Othonneae Kostel., Allg. Med.-Pharm. Fl. 2: 621. Jan-Jun 1833.
Subtribe Othonninae Less., Linnaea 6: 93. post Mar 1831 (Othonneae).
Parthenium L., Sp. Pl.: 988. 1 Mai 1753.
Fam. Partheniaceae Link, Handbuch 1: 816. Jan-Aug 1829.
Subfam. Parthenioideae Raf., Ann. Gen. Sci. Phys. Bruxelles 6: 89. 1820
(Parthenidia).
Pectis L., Syst. Nat., ed. 10: 1189, 1221, 1376. 7 Jun 1759.
Subtribe Pectidinae Less., Linnaea 5: 134. Jan 1830 (Pectideae).
Perdicium L., Pl. Rar. Afr.: 22. 20 Dec 1760.
Fam. Perdiciaceae Link, Handbuch 1: 728. Jan-Aug 1829 (Perdicieae).
Tribe Perdicieae D. Don, Trans. Linn. Soc. London 16: 239. 27 Mai 1830
(Perdiceae).
42 Comp. Newsl. 30, 1997
Picris L., Sp. Pl.: 792. 1 Mai 1753, nom. cons.
Fam. Picridaceae Martinov, Tekhno-Bot. Slovar: 482. 1820 (Picrides).
Tribe Picrideae Schultz-Bip., Flora 17: 476, 478. 14 Aug 1834.
Subtribe Picridinae Schultz-Bip., Flora 17: 478. 14 Aug 1834 (Picrideae
verae).
Pluchea Cass., Bull. Sci. Soc. Philom. Paris 1817: 31. Feb 1817.
Subtribe Plucheinae Cass. ex Dumort., Anal. Fam. Pl.: 31. 1829 (Plucheae).
Polyachyrus Lag., Amen. Nat. Espafi. 1(1): 37. post 19 Apr 1811.
Tribe Polyachyreae D. Don, Trans. Linn. Soc. London 16: 229. 27 Mai 1830.
Subtribe Polyachyrinae Endl., Gen. Pl.: 489. Jun 1838.
Pyrethrum Zinn, Cat. Pl. Hort. Gott.: 414. 20 Apr-21 Mai 1757.
Tribe Pyrethreae Horan., Char. Ess. Fam.: 90. 1847 (Pyrethrariae s.
Senecionideae).
Subtribe Pyrethrinae Horan., Char. Ess. Fam.: 90. 1847 (Pyrethrinae).
Relhania L’ Hér., Sert. Angl.: 22. Jan (prim.) 1789, nom. cons.
Tribe Relhanieae Kostel., Allg. Med.-Pharm. FI. 2: 710. Jan-Jun 1833.
Subtribe Relhaniinae Less., Linnaea 6: 232. Jul-Dec 1831 (Relhanieae).
Rodigia Spreng., Neue Entdeck. Pflanzenk. 1: 275. Jan-Mai 1820.
Subtribe Rodigiinae DC., Prodr. 7: 74, 98. late Apr 1838.
Rolandra Rottb., Soc. Med. Havn. Collect. 2: 256. 1775.
Subtribe Rolandrinae Cass. ex Dumort., Anal. Fam. Pl.: 313. 1829
(Rolandreae).
Rudbeckia L., Sp. Pl.: 906. 1 Mai 1753.
Tribe Rudbeckieae Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829.
Subtribe Rudbeckiinae Cass. ex Dumort., Anal. Fam. Pl.: 31. 1829
(Rudbeckieae).
Santolina L., Sp. Pl.: 842. 1 Mai 1753.
Fam. Santolinaceae Martinov, Tekhno-Bot. Slovar: 560. 1820 (Santolinae).
Tribe Santolineae Lindl. in J.C. Loudon, Encycl. Pl.: 1073. 1829.
Scolymus L., Sp. Pl.: 813. 1 Mai 1753.
Tribe Scolymeae Kostel., Allg. Med.-Pharm. Fl. 2: 624. Jan-Jun 1833.
Subtribe Scolyminae Less., Syn. Gen. Compos.: 126. Jul-Aug 1832
(Scolymeae).
Comp. Newsil. 30, 1997 43
Scorzonera L., Sp. Pl.: 790. 1 Mai 1753.
Subfam. Scorzoneroideae Burmeist., Handb. Naturgesch.: 297. 1837
(Scorzonereae).
Tribe Scorzonereae D. Don, Edinburgh New Philos. J. 6: 307. Jan-Mar1829.
Subtribe Scorzonerinae Cass. ex Dumort., Fl. Belg.: 63. 1827 (Scorzonereae).
Senecio L., Sp. Pl.: 866. 1 Mai 1753.
Fam. Senecionaceae Spenn., Handb. Angew. Bot. 1: 339. 1834
(Senecionideae).
Subfam. Senecionoideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074.
1829 (Seneciones, sphalm.), based on Tribe Senecioneae A.H.G. de Cassini, J.
Phys. Chim. Hist. Nat. 88: 195. Mar 1819.
Tribe Senecioneae Cass., J. Phys. Chim. Hist. Nat. 88: 195. Mar 1819.
Subtribe Senecioninae (Cass.) Dumort., Fl. Belg.: 65. 1827 (Senecioneae),
based on Tribe Senecioneae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88:
195. Mar 1819.
Serratula L., Sp. Pl.: 816. 1 Mai 1753.
Fam. Serratulaceae Martinov, Tekhno-Bot. Slovar: 577. 1820 (Serratulae).
Tribe Serratuleae (Less.) W.D.J. Koch, Syn. Fl. Germ. Helv.: 406. Jan-Oct
1837, based on [Rankless] Serratuleae C.F. Lessing, Syn. Gen. Compos.: 4.
Jul-Aug 1832.
Subtribe Serratulinae (Less.) DC. in J. Lindley, Intr. Nat. Syst. Bot., ed. 2:
262. Jul 1836 (Serratuleae), based on [Rankless] Serratuleae C.F. Lessing, Syn.
Gen. Compos.: 4. Jul-Aug 1832.
Silybum Adans., Fam. Pl. 2: 116, 605. Jul-Aug 1763, nom. cons.
Tribe Silybeae Kitt., Taschenb. Fl. Deutschl., 2, 2: 557. 1844, based on
[Rankless] Silybeae C.F. Lessing, Syn. Gen. Compos.: 10. Jul-Aug 1832.
Subtribe Silybinae (Less.) DC. in J. Lindley, Intr. Nat. Syst. Bot., ed. 2: 262.
Jul 1836 (Silybeae), based on [Rankless] Silybeae C.F. Lessing, Syn. Gen.
Compos.: 10. Jul-Aug 1832.
Stevia Cav., Icon. 4: 32. Sep-Dec 1797.
Tribe Stevieae Horan., Char. Ess. Fam.: 93. 1847 (Steviariae s Eupatoriaceae).
Stifftia J.C. Mikan, Del. Fl. Faun. Bras.: ad t. 1. 1820, nom. cons.
Tribe Stifftieae D. Don, Trans. Linn. Soc. London 16: 291. 27 Mai 1830.
44 Comp. Newsl. 30, 1997
Tagetes L., Sp. Pl.: 887. 1 Mai 1753.
Subfam. Tagetoideae (Cass.) Lindl., Encycl. Pl.: 1074. 1829 (Tagetineae),
based on Tribe Tageteae A.H.G. de Cassini in G.F. Cuvier, Dict. Sci. Nat. 20:
367. 29 Jun 1821 (Tagetineae).
Tribe Tageteae Cass., J. Phys. Chim. Hist. Nat. 88: 162. Feb 1819
(Tagetneae).
Subtribe Tagetinae (Cass.) Dumort., Anal. Fam. Pl.: 31. 1829 (Tagetineae),
based on Tribe Tageteae A.H.G. de Cassini in G.F. Cuvier, Dict. Sci. Nat. 20:
367. 29 Jun 1821 (Tagetineae).
Tanacetum L., Sp. Pl.: 843. 1 Mai 1753.
Fam. Tanacetaceae Vest, Anleit. Stud. Bot.: 273, 298. 1818 (Tanacetoideae).
Taraxacum F.H. Wigg., Prim. Fl. Holsat.: 56. 29 Mar 1780, nom. cons.
Tribe Taraxaceae D. Don, Edinburgh New Philos. J. 6: 307. Jan-Mar 1829.
Tarchonanthus L., Sp. Pl.: 842. 1 Mai 1753.
Tribe Tarchonantheae Kostel., Allg. Med.-Pharm. FI. 2: 668. Jan-Jun 1833.
Subtribe Tarchonanthinae Cass. ex Dumort., Anal. Fam. Pl.: 31. 1829.
Tragopogon L., Sp. Pl.: 789. 1 Mai 1753.
Tribe Tragopogoneae Schultz-Bip., Flora 17: 476. 14 Aug 1834.
Trichospira Kunth in F.W.H.A. von Humboldt, A.J.A. Bonpland & C.S. Kunth,
Nov. Gen. Sp. 4, ed. f*: 21. 26 Oct 1818.
Subtribe Trichospirinae Less., Linnaea 6: 690. Jul-Dec 1831 (Trichospireae).
Trixis P. Browne, Civ. Nat. Hist. Jamaica: 312. 10 Mar 1756.
Tribe Trixideae Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829.
Subtribe Trixidinae Less., Linnaea 5: 6. Jan 1830 (Trixidea).
Tussilago L., Sp. Pl.: 865. 1 Mai 1753.
Subfam. Tussilaginoideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074.
1829 (Tussilagineac), based on Tribe Tussilagineae A.H.G. de Cassini, J. Phys.
Chim. Hist. Nat. 88: 200. Mar 1819.
Tribe Tussilagineae Cass., J. Phys. Chim. Hist. Nat. 88: 200. Mar 1819.
Subtribe Tussilagininae (Cass.) Dumort., Fl. Belg.: 64. 1827 (Tussilagineae),
based on Tribe Tussilagineae A.H.G. de Cassini, J. Phys. Chim. Hist. Nat. 88:
200. Mar 1819.
Comp. Newsl. 30, 1997 45
Urospermum Scop., Intr. Hist. Nat.: 122. Jan-Apr 1777.
Tribe Urospermeae Cass. ex Schultz-Bip., Flora 17: 475. 14 Aug 1834.
Vernonia Schreb., Gen. Pl. 2: 541. Mai 1791, nom. cons.
Fam. Vernoniaceae Burmeist., Handb. Naturgesch. 1: 296. 1837.
Subfam. Vernonioideae (Cass.) Lindl. in J.C. Loudon, Encycl. Pl.: 1074. 1829
(Vernonieae), based on Tribe Vernonieae A.H.G. de Cassini, J. Phys. Chim.
Hist. Nat. 88: 203. Mar 1819.
Tribe Vernonieae Cass., J. Phys. Chim. Hist. Nat. 88: 203. Mar 1819.
Subtribe Vernoniinae Cass. ex Dumort., Anal. Fam. Pl.: 31. 1829
(Vernonieae).
Xanthium L., Sp. Pl.: 987. 1 Mai 1753.
Fam. Xanthiaceae Vest, Anleit. Stud. Bot.: 273, 298. 1818 (Xanthoideae).
Xeranthemum L., Sp. Pl.: 857. 1 Mai 1753.
Subfam. Xeranthemoideae Burmeist., Handb. Naturgesch.: 300. 1837
(Xeranthemeae).
Tribe Xeranthemeae Kostel., Allg. Med.-Pharm. FI. 2: 621. Jan-Jun 1833.
Subtribe Xerantheminae Cass. cx Dumort., Anal. Fam. PIl.: 32. 1829
(Xexanthemeae [sphalm.]).
46 Comp. Newsl. 30, 1997
NOMENCLATURAL NOTES ON ECUADORIAN
SENECIONEAE
Bertil Nordenstam
Department of Phanerogamic Botany
Swedish Museum of Natural History
P. O. Box 50007, S-104 05 Stockholm, Sweden
bertil.nordenstam@nrm.se
Abstract
Seven new combinations for Ecuadorian taxa in the genera Aetheolaena,
Dendrophorbium and Monticalia are presented. Senecio hypomallus Benoist is
synonymized with Aetheolaena heterophylla (Turcz.) B. Nord.
Introduction
Continuing work on Ecuadorian Senecioneae for the ‘Catalogue of the Vascular
Plants of Ecuador’ and the ‘Flora of Ecuador’ necessitates some further
combinations in addition to those already recently published in this journal
(Nordenstam 1996). One of the reasons for these new combinations is my
standpoint that Aetheolaena should be kept distinct from Lasiocephalus, which is a
small Andean genus typified by L. ovatus Schltdl. The species of Aetheolaena are
herbs or half-shrubs, often scandent, with petiolate leaves and corymbose capitula,
whereas Lasiocephalus comprises shrubs with imbricate, sessile or subsessile
leaves and larger solitary flower-heads, somewhat resembling those of Culcitium.
Comp. Newsl. 30, 1997 47
Nomenclature
1. The type of Senecio hypomallus Benoist (Ecuador, valley of Lloa, 3. VII.1930,
Benoist 2719, P holotype!) was examined and found to be conspecific with
Aetheolaena heterophylla (Turcz.) B. Nord. The synonymy of this species is as
follows.
Aetheolaena heterophylla (Turcz.) B. Nord.
Syn.: Gynoxys heterophylla Turcz., Lasiocephalus heterophyllus (Turcz.) Cuatrec.;
Senecio pindilicensis Hieron.; S. hypomallus Benoist.
2. Aetheolaena decipiens (Benoist) B. Nord., comb. nov.
Basionym: Senecio decipiens Benoist, Bull. Soc. Bot. France 83: 807 (1936). Syn.:
Lasiocephalus decipiens (Benoist) Cuatrec. - Orig. coll.: Ecuador, Loja, 2200
m, IV. 1905, Rivet 971 (P holotype!).
This is close to Aetheolaena campanulata (Sch. Bip. ex Klatt) B. Nord., which was
described from Bolivia and appears to be distinct from the Ecuadorian species.
3. Aetheolaena ledifolia (Kunth) B. Nord., comb. nov.
Basionym: Culcitium ledifolium Kunth in H.B.K., Nov. Gen. Sp. Pl. ed. fol. 4: 133
(1818). Syn.: Lasiocephalus ledifolius (Kunth) C. Jeffrey. - Orig. coll.:
Ecuador, Humboldt & Bonpland s.n. (P, non vidi).
4. Aetheolaena otophora (Wedd.) B. Nord. var. christophori (Cuatrec.) B.
Nord., comb. nov.
Basionym: Senecio otophorus Wedd. var. christophori Cuatrec., Fieldiana, Bot.
27(2): 22 (1951). Syn.: Lasiocephalus otophorus (Wedd.) Cuatrec. var.
christophori (Cuatrec.) Cuatrec. - Orig. coll.: Colombia, Dep. Cundinamarca,
Bogotd, San Cristobal, 3000-3300 m alt., [X.1917, F. W. Pennell 2047 (NY
holotype, non vidi; F, US, isotypes, non vidi).
This is a felty-lanate variety from high elevations in Colombia and Ecuador.
48 Comp. Newsl. 30, 1997
5. Aetheolaena otophora (Wedd.) B. Nord. var. microcephala (Hieron.) B.
Nord., comb. nov.
Basionym: Senecio otophorus Wedd. var. microcephalus (microcephala’’)
Hieron., Engl. Bot. Jahrb. 19(1): 66 (1894). - Orig. coll.: Ecuador, Prov. Azuay,
Cuenca, Paramo de Huarijacaja, Chaning et Pilzhum, X.1880, Lehmann 4912
(K?, non vidi).
This variety is rather characteristic with small and sometimes elongate capitula. I
have seen three collections, all from Azuay Prov. in Ecuador.
6. Aetheolaena pichinchensis (Cuatrec.) B. Nord., comb. nov.
Basionym: Culcitium pichinchense Cuatrec., An. Univ. Madrid 4(2): 215 (1935).
Syn.: Senecio pichinchensis (Cuatrec.) Cuatrec., nom. illeg. (non S.
pichinchensis Greenm.); Lasiocephalus pichinchensis (Cuatrec.) Cuatrec.;
Senecio quitensis Cuatrec. - Orig. coll.: Isern 308 (MA holotype, non vidi; F
isolype, non vidi).
7. Dendrophorbium kleinioides (Kunth) B. Nord., comb. nov.
Basionym: Cacalia kleinioides Kunth in H.B.K., Nov. Gen. Sp. Pl. ed. fol. 4: 128-
129 (1818). Syn.: Psacalium kleinioides (Kunth) DC.; Pentacalia kleinioides
(Kunth) Cuatrec. - Orig. coll.: Colombia, prope Guaduas, Humboldt &
Bonpland s.n. (P holotype, non vidi) . - Further syn.: Senecio karstenii Hieron.;
Dendrophorbium karstenii (Hieron.) C. Jeffrey; Senecio pennellii Greenm.
8. Monticalia angustifolia (Kunth) B. Nord., comb. nov.
Basionym: Cacalia angustifolia Kunth in H.B.K., Nov. Gen. Sp. PI. ed. fol. 4: c.
124-125 (1818) ed. quarto 4: 159 (1820). Syn.: Senecio humboldtianus DC. -
Orig. coll.: Ecuador, Quito, prope Mulalo, Humboldt & Bonpland s.n. (P, non
vidi).
De Candolle (1838) changed the specific name when he transferred the species to
Senecio, because the epithet angustifolia was pre-occupied in that genus by S.
angustifolius (Thunb.) Willd., a South African species. Monticalia angustifolia is
an erect small shrub with closely set, sessile, narrow leaves, and several, more or
Comp. Newsl. 30, 1997 49
less erect, discoid capitula, and styles with truncate tips lacking a central hair
pencil, all features characteristic of the genus Monticalia.
References
De Candolle, A. P. 1838. Prodromus systematis naturalis regni vegetabilis 6.
Paris.
Nordenstam, B. 1996. New combinations in Ecuadorean Senecioneae. Comp.
Newsl. 29: 47-50.
50 Comp. Newsl. 30, 1997
NATURE OF ERGASTIC SUBSTANCES IN SOME
WEST AFRICAN ASTERACEAE SEEDS - VIII
M. Idu & LS. Gill
Department of Botany
University of Benin
P.M.B. 1154
Benin City
Nigeria
Abstract
The ergastic substances of seeds from 46 species of Asteraceae were examined for
alkaloids, fats and oils, inulin, protein, starch and tannin. All species examined
were herbs excepts five. Fat and oils were found to be present in all the
investigated taxa, 14 species had alkaloid, 22 had protein. Starch grains and tannin
were found to be absent in all the investigated species.
Introduction
The ergastic substances are secondary products of plant metabolism which might
have been formed at certain stages of metabolic process and are retained when the
taxon in question underwent further evolution (Erdtman 1956). According to Gill
& Ayodele (1986) the present food crisis where the number of cultivated crops is
radically insufficient to provide for the world food supply, a knowledge of the
stored products of the seeds of wild plants cannot be overemphazied. This can be
done with the view to harness the resources of wild plants.
Furthermore, the future energy needs of mankind will depend on renewable plant
resources to replace the present decreasing fossil fuel reserves (Abelson 1978).
The present paper is part of ongoing project on the nature of ergastic substances in
angiospermic seeds. The earlier contributions on this project are Gill & Ayodele
(1986), Omoigui & Gill (1988) and Gill et al. (1991).
Comp. Newsl. 30, 1997 51
The results of the survey for ergastic substances of 46 Asteraceae seeds distributed
in 7 tribes are reported here.
Materials and Methods
Seeds used in this investigation were randomly collected in nature from different
localities in southern Nigeria. Seed materials was stored at 27 + 2° C in the
laboratory. The method used for the determination of ergastic substances was
outlined by Gill & Abili (1989). Vouchers of the specimens studied are deposited
in the herbarium of the University of Benin, Nigeria.
Results
Table 1 summarizes the taxa studied along with their life form and nature of
ergastic substances. Starch grains and tannin were absent in all the investigated
taxa whereas fats and oils and inulin were observed in all the taxa, alkaloid and
protein in 14 and 22 taxa respectively.
Discussion
For more than three decades now, much attention has been focused on the
comparative studies of basic molecules in rclauion to taxonomic problems.
De Wet & Scott (1965) are of the opinion that essential oil can be used as a
taxonomic criterion and that chemical characters are often found to more reliable
than the gross morphology in determining the taxonomic affinities.
Gill & Ayodele (1986) and Gill et al. (1991) have established a relationship
between life form and nature of ergastic substances and suggested that starch
granules are generally associated with herbaceous habits, but this view does not
hold in the present study with seeds of Asteraceae, where all the surveyed taxa are
predominantly herbaceous and devoid of starch granules. However, Gill & Abili
(1989) are of the opinion that an absolute reliance on the presence or absence of a
particular or a group of ergastic substances will be a gross mistake in making any
taxonomic decisions. Pirie (1955) suggests that various types of information
regarding a taxon should be taken into consideration for any taxonomic
classification.
Further investigation is needed on these taxa to know the nature of protein,
alkaloid, fats and oils present in them for commercial exploitation.
52 Comp. Newsl. 30, 1997
References
Abelson, P.H. 1978. Bioenergy. Science 204: 1164.
De Wet, J.M.J. & B.D. Scott 1965. Essential oils as taxonomic criteria in
Bothriochloa. Bot. Gaz. 126: 209-214.
Erdtman, H. 1956. Flavonoid Heartwood Constituents of Conifers. Sci. Proc.
Roy. Soc. Dublin 24: 129-138.
Gill, L.S. & M.A. Abili 1989. Nature of ergastic substances in some angiospermic
seeds. V. Feddes Repert. 100: 71-79.
Gill, L.S. & J.R. Ayodele 1986. On the nature of ergastic substances in the seeds
of some tropical and temperate angiosperms. III. J. Plant. Anat. and Morphol.
3: 35-49.
Gill, L.S., Nyawuame, H.G.K., Aibangbe, M.I. & D.A. Agho 1991. Nature of
ergastic substances in some Mediterranean angiospermous seeds. VI. Feddes
Repert. 102: 613-628.
Omoigui, I.D. & L.S. Gill 1988. Nature of ergastic substances in some West
African Compositae. Feddes Repert. 99: 143-145.
Pirie, N.W. 1955. The principles of microbial classification. Summing Up. J. Gen.
Microbiol. 121: 382-386.
53
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LA GERMINATION ET LE POUVOIR
GERMINATIF DE QUELQUES ARTEMISIA
DU MAROC
Mme Aicha Ouyahya
Département de Botanique et Ecologie végétale
Institut Scientifique
B.P. 703
Rabat, Maroc
Résumé
Le présent article traite de la germination et du pouvoir germinatif des armoises
marocaines réputées par leurs valeurs pastorale, thérapeutique et écologique. Les
résultats obtenus suggérent que la germination des Artemisia étudiés dépend en
grande partie des conditions écologiques du biotope et de la maturité physiologi-
que des diaspores.
Abstract
The present paper describes the germination and the germinal power of Moroccan
species of the genus Artemisia. These species have a pastoral and a therapeutical
value. The results show that the germination of Artemisia depends of climatic
conditions in the biotope and physiological maturity of achenes.
Introduction
Dans une note antérieure (Ouyahya 1983), on a étudie la germination et le pouvoir
germinatif de cing armoises endémiques marocaines. Dans le présent travail, a titre
de comparaison, on a repris la méme étude sur d’autres populations et sur d’autres
taxons afin de formuler des hypotheses relatives aux aptitudes germinatives des
diaspores, aux facteurs externes et internes propices a l’activité germinative et au
role de cette dermiére dans la régénération des populations des steppes d’armoise.
58 Comp. Newsl. 30, 1997
Matériel et méthode
Les diaspores étudiées proviennent soit des échantillons récoltés sur le terrain, soit
des échantillons envoyés par des jardins botaniques.
Pour chaque taxon étudié et pour chaque mois (durant six mois) un lot de
semences de 100 akénes pris au hasard, aprés traitement a hypochlorite 4 50 %
(bain de 5 minutes), a été placé sur des papiers filtres imbibés d’eau distillée, dans
des boites de Pétri, 4 l’obscurité dans un tiroir, 4 la température de la salle (T°C).
Des observations réguli¢res faites tous les jours permettent d’une part de suivre le
développement de la plantule, d’autres part de connaitre la progression du
pourcentage de germination des akénes.
Résultats et discussion
Le tableau 1 réunit les valeurs de quelques paramétres: temps de latence (TL);
temps moyen de germination (TMG), c’est-a-dire temps nécessaire pour atteindre
50 % de germination; durée de la période de germination (DPG); capacité de
germination (CG), c’est-a-dire taux maximal atteint dans les conditions notées.
A partir du tableau 1 et des courbes de germination de figures 1 et 2, plusieurs
constatations s’imposent; dans un premier temps il appait possible de séparer les
taxons en trois groupes bien distincts.
Le premier groupe rassemble les taxons qui germent abondamment et conservent
un pouvoir germinatif encore assez élevé: A. absinthium, A. caerulescens, A.
verlotiorum, A. vulgaris, et ce sont sans doute des espéces qui croissent dans des
biotopes humides. Cependant, rappelons que les akénes de ces derniéres espéces
ont un an de plus. Clor et al. (1974) ont mentionné que les diaspores d’A. herba-
alba ne peuvent germer qu’apres une dormance obligatoire de presqu’une année.
Les especes du second groupe sont caractérisées par une capacité de germination
moyenne, elles germent aussi bien que les précédentes. Ce sont des orophytes
endémiques marocaines a répartition géographique assez large qui se localisent
surtout dans des régions semi-aride: A. mesatlantica et A. negrei.
Le dernier groupe renferme A. atlantica var. maroccana et A. herba-alba. Ces
deux taxons marocains possedent un pouvoir germinatif faible et germent plus
lentement que les autres taxons étudiés. A. herba-alba croit dans des régions de
préférence arides, depuis le littoral atlantique jusqu’a 2000 m d’altitude; par contre
Comp. Newsl. 30, 1997 59
A. atlantica var. maroccana ne se rencontre qu’a partir de 1600 m d’altitude dans
des localités semi-arides.
Notons également que la capacité de germination varie beaucoup d’un taxon a
autre et vraisemblablement en fonction de la maturité physiologique des
diaspores et de la température de la salle (facteur limitant de la germination, Come
1970).
Au cours de la germination, on a pu constater que pour certains échantillons
(A. absinthium, éch. 0102; A. herba-alba, éch. 2044, 2043, 2047, 2048; A. mes-
atlantica, éch. 2411, 2413, 2414 et A. negrei, éch. 2511, 2512) le taux maximal de
la germination n’a été atteint qu’a une température élevée (25 <T’C<28), ce qui
laisserait supposer que ces plantes ont besoin a la fois d’humidité et de chaleur
pour germer. Le r6le de l’humidité n’appelle aucun commentaire particulier. Par
contre, l’action des températures élevées laisse supposer que la dormance exige
que, pour étre levée, la graine ait franchi une thermophase précise: les stations
concemées sont ordinairement dans les étages bioclimatiques semi-aride, aride et
saharien, mais a hiver frais ou froid. Pendant cette période hivernale froide les
graines sont généralement dormantes, et ne peuvent satisfaire cette thermophase
que durant le printemps (les Artemisia étudiés ne fructifient qu’au mois de
novembre; cette fructification d’automne constitue une stratégie adaptative adoptée
par beaucoup d’espéces des régions arides), comme cela pourrait étre le cas pour
d’autres stations situées dans des sous-étages a hiver tempéré ou doux, ou méme
pour les espéces cultivées et acclimatisées depuis longtemps en jardin botanique
(échantillons n° 1201, 2811, 2907).
On peut s’approcher de cette hypothése les observations de Fernandez & Caldwell
(1975), observations portant sur A. tridentata. Ils ont révélé que le taux de
croissance des racines est maximum aux mois d’avril et juin (observations réalisées
dans les chambres implantées sur le terrain de récolte). Ceci est en rapport, non
avec la levée de dormance, mais seulement avec les phénomenes_habituels de la
croissance.
A partir de la figure 2, on a pu déceler les faits suivants:
- pour les localités ou cohabitent les trois armoises marocaines: A. herba-alba,
A. mesatlantica et A. negrei, la derni¢re germe plus rapidement et possede un
pouvoir germinatif encore plus élevé que les deux autres.
- les courbes de germination de ces trois armoises révélent une succession
frappante. En fait, cette succession pourrait étre corrélée a |’étagement altitudinal
d’une part, et aux conditions écologiques d’autre part et sans omettre la maturité
physiologique des diaspores.
60 Comp. Newsl. 30, 1997
En effet, A. negrei est une espéce endémique marocaine, polyploide (2n=56 a 65)
et orophile. Elle se rencontre surtout dans le Haut Atlas a partir de 2200 m
d’altitude avec des xérophytes €pineux (conditions micro- et mésoclimatiques
dures, surtout froides et humides).
A. mesatlantica est aussi une espece endémique marocaine, diploide (2n=18),
orophile et a répartition assez large (Moyen Atlas, Haut Atlas et Anti Atlas). Elle
croit en-dessous de A. negrei dans le Haut Atlas, a partir de 1800 m d’ altitude.
A. herba-alba est une espece a répartition écologique et géographique plus large;
elle se rencontre aussi bien sur silice que sur calcaire; elle s’étend depuis le Maroc
jusqu’en Egypte, Sud de l’Espagne et de la France, Proche-Orient, Iran, Irak et
Afghanistan. Elle est, pour le moment, connue comme diploide au Maroc et
tétraploide en Espagne et en Tunisie. Elle croit en-dessous d’A.mesatlantica dans
le Moyen Atlas, le Haut Atlas et l’Anti Atlas.
Ces résultats mettent en évidence que le phénoméne de germination est lié
directement aux conditions écologiques de la plante d’une part, et a la maturité
physiologique des diaspores d’autre part.
Bibliographie
Clor, M.A., Al Ani, T.A. & F. Charchafchi 1974. Range resources of Iraq, XV.
Germination, storage conditions and after-ripening of the seeds of Artemisia
herba-alba. Tech. Bull. no. 76. Inst. for Appl. Research on Natural Resources,
Abu Ghraib, Iraq. 17 pp.
Come, D. 1970. Les obstacles a la germination. Masson et Cie, Paris.
Fernandez, O.A. & M.M. Caldwell 1975. Phénologie et dynamique de la
croissance des racines de trois arbustes de climat semi-désertique frais (en
culture en champ). J.Ecol. G.B. 63 (2): 703-714.
Ouyahya, A. 1983. Etude sur la germination et le pouvoir germinatif de cing
armoises endémiques du Maroc. Bull. Inst. Sct. Rabat 7: 75-82.
Ouyahya, A. 1987. Systématique du genre Artemisia L. au Maroc. These doc. és-
sciences Univ. Aix-Marseille III, 433 pp.
61
Comp. Newsl. 30, 1997
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Comp. Newsl. 30, 1997 63
Légende
Tableau 1. Tableau résumant les résultats obtenus relatifs aux caractéristiques de
germination pour six mois de tests de germination (nombre de diaspores germées
par jour sur 100 akénes) sur huir armoises: cing marocaines (A. absinthium, A.
atlantica var. maroccana, A. herba-alba, A. mesatlantica et A. negrei) et les trois
autres provenant du jardin botanique de Nantes (A. verlotiorum) et du jardin
botanique de Sienne (A. vulgaris et A. caerulescens).
Figure 1. Courbes de germination des akénes d’Artemisia. Chaque courbe
correspond a 100 akénes. C.G.: Capacité de germination (température de la salle,
16 <T°C<19, mois de mars).
Figure 2. Courbes de germination des akénes d’Artemisia agés de 4 mois
(température de la salle, 16<T°C<19, mois de mars). 2045, 2046: A. herba-alba;
2410, 2411, 2413, 2414: A. mesatlantica; 2508, 2509, 2511, 2512: A. negrei.
64
ca’
60
40
80
40
é0
40
80
40
40
40
Comp. Newsl. 30, 1997
2814 A. ver/oforum
2907 A. vu/garis
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1201 A. coeru/escens
2024
. 2045
2048
2045
2049
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2445
2444
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2410
2443
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2508
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D 2509
2312
E
_-———- 0808
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maroecana
35 6 9 12 15 18 2t 24 27 30 Jours
Figure 1}
Comp. Newsl. 30, 1997 65
GG. 2509
2444
80 A
40
Tierhist: (2700m)
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80 B
2443
40
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2440
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2542
60 D EEE , oda
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Figure 2
66 Comp. Newsl. 30, 1997
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