Monophyly of the Asteridae and Identification of Their Major Lineages
Inferred From DNA Sequences of rbeL
Richard G. Olmstead; Helen J. Michaels; Kathy M. Scott; Jeffrey D. Palmer
Annals of the Missouri Boranical Garden, Vol. 79, No. 2 (1992), 249-265.
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Mon Feb 9 14:26:22 2004MONOPHYLY OF THE
ASTERIDAE AND
Richard G. Olmstead,’
Helen J. Michaels, Kathy M. Scott
IDENTIFICATION OF THEIR. “4/4”? Palmer
MAJOR LINEAGES INFERRED
FROM DNA SEQUENCES OF
rbcLt
‘Ansreact
A pasiony analysis of 57 angionperm rcL sequences was conducted to test the monophyly af the Astrdae and
to tently major Ineages within the heeise, Tove major eladen, the Caryophlida, the Rosiae pls Dilerisae,
tnd the Aste senu lato, emerge from an unrelvedraiaton inthe ghee” dvats. The Amerie sna. at
{nce the Eas, Coral, and APisles in adtion tothe Asterida ses. sr. Two tao lineages within the
‘tera som at are estifd: the Dipscales, piles, Asterles, ac Capel in one, andthe Centanales
‘Srophularley amily, Beagle, at Slanals in the oer. Thr analyse demonstrates the wt of molecu
plnlogenice to help place problematic taxa, such asthe Menyanthaces, eacea, and Caliichacese, within the
‘trae. Inplcaone from th pylogencic analysis and erence fom the fowl record lad to the sugpestion tat
the org and dveriiaton of the mae higher dat neags occured during eelavely short period of ime abot
80-95 millon years ago
‘The modern concept of the Asteridee, sensu
‘Takhtajan (1980) and Cronquist (1981), is derived
from the ancestral Monopetalae (dle Jussieu, 1789)
and Gamopetalae (de Candolle, 1813) by the elim:
ination of many groups of plants bearing the org
inal defining feature of fused corollas (Wagenitz,
1992), Cronquist (1981: 852) stated that “the
Asterdae are the most advanced subclass of di
cotyledons.” This statement puts into words @ gen-
erally held perception, based on traditional as-
sumptions regarding trends in character evolution
inthe angiosperms, that the subclass is of relatively
recent origin compared to other major groups of
dicots Spome, 1969, 1975; Stebbins, 1974).
‘There is no consen:us of opinion concerning the
monophyly of the Asteridae. Whereas a combi
nation of floral and embryological characters seems
ta define @ natural group, portrayed as monophy-
letic in the treatments of Cronquist (1981) and
‘Takhtajan (1980, but not 1987), chemical char-
acters suggest two separate asterid lineages, each
derived independently from ancestors in the Ros
dao (Dahlgren, 1980). As Wagenitz(1977) pointed
‘out, no division of the Asteridae into separate lin
ceages can be constructed without having to pos
tulate parallel evolution in morphology, embryol-
‘ogy, and phytocheristry. Parsimony-hased methods
‘of phylogeny reconstruction offer a means of as
sessing phylogenetic information in which paral:
lelisms exist, by establishing objective criteria for
accepting one hypothesis of relationships (Le. tree)
cover another hypothesis. Parsimony-hased phylog-
eny reconstructions among major groups in the
dicots are few. Donoghue & Doyle (1989), in their
analysis of basal angiosperm lineages, identified a
“higher”-icot clade (ie, derived relative to the
bhasal dicots). This clade, to which all Asteridae,
Rosidae, Dillenidee, Caryophylidae, and Hama-
rmelidae, as well as certain members of the Mag-
nolidae, belong is characterized by the presence
of tricolpate pollen. Hamby & Zimmer (1991)
conducted @ parsimony analysis of nuclear ribo:
somal RNA sequences in angiosperms and other
seed plant groups, bu found litle resolution among
"We thank S. Dewi
M. Donoghue, M. Chase, K. Kron, B. Zimmer, and M. Kills for eritial reading of
the manuscript and M, Chase, G. Furie, D. Giana, E. Golenberg, D. Lex J. Nogent, J. Retig, and K. Sytsma
fo proving paid spines, Ths rsa bs en suport by AS grant BLT 760
“Department of Biology, Indiana University, Bloomington, Indiana 47405, US.A.
® Preset sve: Department of P.O. Biology, University of Cslorado, Boulder, Colorado 80309, U.S.A
“Department of Biology, Bowling Green State University, Bowling Green, Ohio 43403, US.A,
ANN. Missourt Bor. Garb. 79: 249-265. 1992.‘Annals of the
Missouri Botanical Garden
Burmanniacese
Lilisceae
Orchidaceae
Commelridae
Poaceae
Dicots
Magnotidae
Lauraceae
‘Magnolacece
[Nelumbonacese
Nymphacscese
Hamamelidae
Cereiphyllaceae
Platanaceae
Caryophylidae
Amaranthacese
yophyllaceee
Chenopodiaceae
Phytlacaceae
Plumboginacese
Polygonaceae
Diese
Beastcncese
Ericacese
Fouguiriacese
Malvaceae
Vielacene
Rosidae
Apiaceae
Araliacese
CCornacese
Fabaceae
Grosulrineae
Hydrangescese
Linacese
Onagraceae
Polygalacese
Serifagscese
Voehysincese
Asteridae
Apoeynacene
Asteraceae
Bignoniaceae
Boraginaceee
Calltrichacene
Calyeeraceae
Burmannia biflora
Lilium superbum
Oncidium exeavatum
Cenchrus setigeras
Puccinellia ditans
Persea americana
‘M. macrophylla
Nelumbo lute
Nuphar variegata
Nymphaea odorata
Cercidiphyllum japonica!
Platans racemosa
Amaranthus hypochondriacus
Stellaria media
Spinacia oleracea
Phytolacea americana
Plumbogo cepensis
Rheum >cultorum
Brassica campestris
Rhododendron hippophacoides
Fouguleriasplendent
Gossypium hirtun
Viola sonar
Coriandram saioune
Hedera heli
Comus mas
Medicago sativa
Bresia madagascarenss
Carpentria californica
Linum perenne
Clarkia vaniana
Seeuridaca diversiflia.
Heuchera micrantha
Parnassia finbriata
Penthorum sedoides
Quatea sp.
Apocynum cannabinune
Barnadesia earyophylia.
Catalpa sp
Borage ofteinalie
Calltriche heterophylla
Boopis anthemoides
MIC (unpublished)
MIC (unpublished)
MIC (unpublised)
Dobley et al. (1990)
Doabley et al. (1990)
Goenberg eta. (1990)
Goleberg eta. (1990)
Tes et a. (1991)
Tes et al. (1991)
Tes et a. (1991)
RGO sn
EMG (unpublished)
Michalowski et a (1990)
JHR, JRM & HDW (unpublished)
araweki otal. (1981)
SHR, JME & HDW (unpublished)
DEG eta. (onpublbed)
DEG etal. (unpublished)
IMN (unpublished)
MWC & KK (unpublished)
Matthaei BG 860162
Galo eta. (1990)
[RCO (ao voucher)
IDP (no voucher)
RKY sn
Donoghue tal (1992)
Aldrich etal (1986)
Solis eral. (1990)
Solis etal. (1990)
MWC (unpublished)
KIS & BC (unpublished)
MWC (unpublished)
Solis etal. (1990)
Solis etal (1990)
Solis etal. (1990)
MIC (unpublished)
GO (no voucher)
Michaels etal (in prep.)
CHD sn
RCO (oo voucher)
cP 2152
Michael etal (in prep.)