Biochemical Systematicsand Ecology, Vol. 18, No. 7/8, pp. 529-532, 1990. 0305-1978/90$3.004-0.00
Printed in GreatBritain. © 1991 PergamonPressplc.
Flavonoid Systematics of Ten Sections of Ludwigia (Onagraceae)
JOHN E. AVERETT,* ELSA M. ZARDINIt and PETER C. HOCHT~
*National Wildflower Research Center, Austin, TX 78725, U.S.A.;
1Missouri Botanical Garden, P.O. Box 299, St Louis, MO 63166, U.S.A.
Key Word Index--Ludwigia, Onagraceae; flavonoids; glycoflavones; flavonols; phytochamical evolution.
Abstract--Data for the flavonoids of 19 species in 10 sections of Ludwigia are presented. Eight flavonoids, comprising four
glycoflavones, of which vitexin and isovitexin are reported for the first time in Ludwigia, and four flavonol glycosides, based on
quercetin, are present in these species. Each section treated here has either glycoflavones or flavonols; presence of only one
class is considered to be advanced in the genus as a whole, compared with the presence of both glycoflavones and flavonols
in the more generalized sects Myrtocarpus, Cinerascentes, and Pterocaulon, which were examined earlier. Only glycoflavones
are present in sects Macrocarpon (four species), Serninuda (five species), the ditypic African sect. Africana, the monotypic
African sects Brenania, Cryptosperma, and Prieurea, the monotypic east Asian sect. Nipponia, and the monotypic pantropical
section Fissendocarpa. Only flavonols are present in the monotypic Old World section Caryophylloidea and sect. Oligo-
spermum, which comprises nine species widespread in the Old and New Worlds.
Introduction
As a continuation of a comprehensive study of
the flavonoids of Onagraceae [1], we here report
results for 10 sections of Ludwigia. We previously
reported data from 30 species in seven other
sections of the genus [2], which included those
groups considered to be most generalized [3-5].
Ludwigia is one of the largest and most
diverse genera in the family Onagraceae, with
some 82 species grouped in 23 sections [6, 7].
Recent data indicate that Ludwigia, which is very
distinctive within the family and comprises the
only genus of the tribe Jussiaeeae, constitutes
the sister group of all other Onagraceae [4, 8-10].
The 10 well-delimited sections investigated
here are considered to be specialized in many
features [6]. Six of these sections are monotypic
and comprise autogamous herbs; three of them
(Brenania, Cryptosperma, and Prieurea) are
restricted to tropical Africa, one to east Asia
(Nipponia), one is widely distributed in the Old
World (Caryophylloidea), and one is a pantropical
weed (Fissendocarpa). The ditypic sect. Africana,
is also restricted to Africa. The remaining three
sections are larger, with broad distributions.
Section Macrocarpon consists of four species of
tTo whom correspondence should be addressed.
(Received 8 June 1990)
herbs or low shrubs, three of which grow pri-
marily in southern South America, whereas the
fourth, L. octovalvis (Jacq.) Raven, is a highly
polymorphic, pantropical, autogamous weed.
Section Seminuda includes five species of robust
herbs, two of which occur only in the New
World, one [L. africana (Brenan) Hara] only in
West Africa, and two common to Latin America
and Africa [6]. Finally, sect. Oligospermum is a
widespread polyploid complex comprising nine
species, of which two are endemic to the Old
World, five to the New World, and two occur in
both regions (Zardini and Raven, unpublished
data).
Results
Eight flavonoids were found to be present
among the species sampled (Table 1): vitexin (1),
isovitexin (2), orientin (3), isoorientin (4),
quercetin 3-rhamnoside (5), quercetin 3-arabino-
side (6), quercetin 3-glucoside (7), and quercetin
3-rutinoside (8). Compounds 1-4 are glyco-
flavones and 5-8 are flavonol glycosides. Com-
pounds 1 and 2 have not been reported pre-
viously in Ludwigia [2]. On the other hand, the
glycoflavone orientin-O-acylate and the flavonol
quercetin 3-diglucoside, which were detected in
sects Cinerascentes, Heterophylla, and Ptero-
caulon, and in sects Myrtocarpus and Ptero-
caulon, respectively [2], were not detected in the
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