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Nord. J.
Bot.
- Section of holarctic and general taxonomy
A review of the phylogeny and classification of the Asteraceae
Kire Bremer, Robert
K.
Jansen, Per Ola Karis, Mari Kiillersjo, Sterling
C.
Keeley, Ki-Jwng Kim,
Helen
J.
Michaels, Jeffrey
D.
Palmer and Robert
S.
Wallace
Bremer, K., Jansen, R. K., Karis, P. O., Kallersjo, M., Keeley, S. C., Kim, K.-J.,
Michaels, H. J., Palmer, J . D. Wallace,
R .
S. 1992. A review of the phylogeny and
classification of the Asteraceae. Nord.
J
Bot. 12: 141-148. Copenhagen. ISSN
0107-055X.
The A steraceae ar e commonly divided into two large subfamilies, the Cichorioideae
(syn. Lactucoideae; Mutisieae, Cardueae, Lactuceae, Vernonieae, Liabeae, Arcto-
teae) and the Asteroideae (Inuleae, Astereae, An themideae, Sen ecioneae, Calen-
duleae, Heliantheae, Eupatorieae). Recent phylogenetic analyses based
on
morphological and chloroplast DNA data conclusively show that the Mutisieae-
Barnadesiinae ar e the sister grou p to the rest of the family and that the Asteroideae
tribes form a monophyletic group. The Vernonieae and Liabeae are sister tribes and
the Eupatorieae are nested within a paraphyletic Heliantheae; otherwise tribal in-
terrelationships are still largely uncertain. The M utisieae-Barnadesiinae are excluded
from the Mutisieae and elevated to the new subfamily Barnadesioideae. The two
subfamilies Barnadesioideae and Asteroideae are monophyletic, whereas the status
of the Cichorioideae remains uncertain. Analyses of chloroplast D NA d ata support
the monophyly of the Cichorioideae; however, morphological data indicate that the
subfamily is paraphyletic. Further studies are needed to test the monophyly of the
Cichorioideae, as well as to furth er resolve tribal interrelationships in the two larger
subfamilies.
K .
Bremer, Dept
o
Systematic Botany, Uppsala U niversity, Box 541, S-7.5121 Uppsa-
la, Sweden. R. K . Jansen, Dept o Botany, Univ. o Texas, Austin, Texas 78713,
U.S .A. P .
0
Karis, Dept
of
Phanerogamic Botany, Swedish Museum o Natural
History,
S-104 05
Stockholm, Sweden. M . Kallersjo, Lab .
o
Molecular Systematics,
Smithsonian Institution, Washington, DC 20560, U.S.A.
- S .
C. Keeley, Dept of
Botany, Univ. o Hawaii , Honolulu, Hawaii 96822, U. S. A. K.-J . Kim , Dept o
Botany, Univ. o Texas, Austin, Texas 78713, U. S. A. H. J . Michaels, Dept o
Biology, Bowling Green State University, Bowling Green , Ohio 43402, U .S .A .
.
D .
Palmer, Dept o Biology, Indiana University, Bloomington , Indiana 47405, U . S .A .
R .
S .
Wallace, Dept o Botany, Iowa State University, Am es, Io wa
50011
U . S . A .
Introduction
In this review we compare the recent analyses of
Asteraceae phylogeny published by our two research
groups based
on
morphological (K. Bremer,
P.
0 Ka-
ris,
M.
Kallersjo) and chloroplast D NA (R . Jansen,
S.
Keeley,
K.-J.
Kim, H. M ichaels,
J .
Palmer,
R .
Wallace)
data. We
(1)
recognize three subfamilies, the Asteroi-
deae,
t he
Cichorioideae (syn. Lactucoideae), and the
small South American Barnadesioideae, (2) acknowl-
edg e the monophyly of the large subfamily Ast eroi dea e,
which includes the tribes Astereae, Anthemideae,
Inuleae, Senecioneae, Calenduleae, Heliantheae, and
Eupatorieae, and 3) discuss the uncertain statu s
of
the
subfamily Cichorioideae, which includes the Mutisieae,
Cardu eae, Lactuceae, Vern onieae , Liabe ae, and Arcto-
teae.
For
brevity an d simplicity we generally use tribal
names in
a
broad sense, for example, Cardueae sens.
Accepted 6-12-91
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OF
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NORD. J. BOT. 12: 141 148
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J . Bot. I
2) 1992)
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lat ., Inuleae sens. lat., and Heliantheae se ns. lat ., al- ily were partly unresolved. Within the subfamily Aste-
though we realize that reclassification into smaller tribes roideae alternative ar ran gem ent s were restricted to the
has been proposed (e.g ., Dittrich 1977; Turner Po- subtribal level, leaving a single patt ern of tribal in-
well 1977; Stro the r 1977; Anderbe rg 1989). Th e nam e terrelationships (Fig. 1B).
Mutisieae is applied sensu stricto, excluding the form er Breme r (1987) dem ons trate d that the subfamily Aste-
subtribe Barnad esiinae. Th e tribal classification is com- roideae is a well-supported mono phyletic gro up. Th e
mented upon
in
the discussion and those tribes currently basal sister group relationship betw een th e Mutisieae-
supported by both morphological and molecular da ta Barnade siinae and the rest of the family (Fig. l A ,C ),
are listed und er classification. revealed by the cpDNA inversion, was also supported
Until the 1970s the tribe Lactuce ae was considered to by several morphological cha racters, such as presence
be distinct from all oth er Asteraceae a nd was classified of the typical Asteraceae twin hairs on the fruits and
in
its own subfamily, the Liguliflorae. Th e oth er tribes spiny pollen.
were placed together in the Tubuliflorae (e.g., Hoff- Althoug h oth er results were less cer tain , the analysis
mann 1890). This classification still persists in some suggested that (1) the tribe Mutisieae and the subfamily
floras and handbooks. D uring the 1970s there was a Cichorioideae are paraphyletic even if the Barnadesii-
growing recognition that all the tribes, including the nae are excluded, (2) the Arcto teae and the Carduea e
Lactuceae, could be arranged in two large groups are related, (3) the Vernonieae and Liabe ae are sister
(Robinson Brettell 1973; Carlquist 1976), fore- groups, (4) the Inuleae sens. lat. are paraphyletic and a
shadowe d by diagrams in Carlquist (1961) and Poljakov basal grade within the A ster oide ae, and
5 )
the Aste-
(1967). Carlquist (1976) formally tre ated these two reae and Eu pat orie ae are sister groups. La ter molecular
groups as the subfamilies Cichorioidea e and Asteroi- and morphological studies have weake ned the sister
dea e. Their precise circumscription was modified by group relationship between the Vernonieae and Lia-
later autho rs (Robinson 1977, 1981, 1983; Tho rne beae and have refuted a close relationship between the
1983). Rece nt subfamilial and tribal classifications of Arctote ae and Ca rduea e, and between the A stereae
the Asteraceae are summarized in Tab. 1. and E upatorie ae (Jansen e t al. 1990, 1991a; Karis 1992,
Karis et al. 1992).
Following their Mutisieae investigations, Jansen and
collaborators performed a large study of cpDNA re-
Recent cladistic analyses
striction site variation in the en tire family (Jansen et al.
1990.1991a. b). Thev sam de d 57 genera from all maior
Since 1985 cladistic analyses of molecular and morph- tribes and identified 328 'phylog etically informative
ological dat a have illuminated the phylogeny and classi- restriction site mutations. Th e dat a matrix was analyzed
fication of the Asteraceae. Jansen Palmer (1987) using both Wagner and Dollo parsimony , as well as the
reported the discovery of a 22 kb cpD NA inversion in bootstrap method (Felsenstein 1985). T he Wagner
Lactuca
and several oth er A stera cea e. Th e inversion analysis resulted in 20 equally parsimoniou s tree s with a
proved to be absent in three genera of the Mutisieae- consistency index of 0.46, and Dollo parsimony gener-
Barnade siinae, suggesting a basal dichotom y between ated 16 tree s with a consistency index of 0.44. Jansen et
the Barnadesiinae and the rest
of
the family. Subse- al. (1990) preferred t he Dollo tree topology (Fig.
quently, Jansen Palm er (1988) undertook an analysis lC , D ) as the best estimate of phylogenetic relationships
of restriction site variation in the Mutisieae. They sam- in the Ast erac eae arguing that (1) the Dollo algorithm is
pled 13 genera of the Mutisieae (including the Barnade- more ap pro pria te for restriction site mutations because
siinae) and three representatives each from the Lactu- convergent gains occur at least an orde r
of
magnitude
ceae, Cardue ae, and Heliantheae. This study corrobor- less than convergent losses (Templeton 1983; DeBry
ated the sister grou p relationship between the Slade 1985), (2) eight of the 20 equally parsimoniou s
Barnadesiinae and the rest of the family. Furtherm ore, Wagner trees have the Dollo tree topology, and (3) a
the Mutisieae sens. str. (excluding the Barnadesiinae) posteriori character analysis, including successive ap-
came out as a monophyletic group.
proximation (Farris 1969), followed by Wagner analysis
Simultaneously with Jansen .Palmer's cpD NA in- always gave trees with the Dollo topology. Further-
vestigations Bremer (1987) performed a cladistic analy-
sis of tribal interrelationships based on mainly morph-
ological data. The study included some 27 tribes and
subtribes and 47 phylogenetically informative charac-
ters, one of them being the cpDNA inversion. Several
hundred equally parsimonious Wagner trees were
foun d, with a consistency index of
0.54
(excluding auta-
pomorphies). The strict consensus tree contained a
number
of
polychotomies especially within the Cicho-
rioideae. Hence tribal interrelationships in that subfam-
more, phylogenetic analyses of the restriction site data
employing character state weighting of sensu Holsinger
Jansen (1992) also produced the Dollo topology (Jan-
sen et al. 1990). We mention the differences between
Wagner and Dollo analyses because they are critical in
comparing the results with those from morphological
data; th e D ollo topologies a re largely incongruent with
the morphological trees whereas some of the Wagner
topologies are similar to the morphological trees.
Jansen et a l.3 analyses indicated tha t (1) the subfam-
142
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ily Asteroideae is strongly supported as a monophyletic
group, (2) the subfamily Cichorioideae likewise is
monophyletic, although this result is less well sup-
ported , (3) 13 of th e 15 investigated tribes are mono-
phyletic, with the Heliantheae and Tageteae being para-
phyletic, (4) the Vernonieae and Liabeae are sister
tribes, and that (5) the Coreopsideae, Tageteae, Helian-
theae, and Eupatorieae are closely related, and the
Eupatorieae are nested within a paraphyletic Helian-
theae. Interrelationships within the Heliantheae-Eupa-
torieae complex have been further clarified by Kim
et
al. (1990) (Fig. 1D).
Using the Barnadesiinae as an outgroup, tribal in-
terrelationships within the Cichorioideae were explored
in more detail by Karis et al. (1992). They perform ed a
much more detailed morphological analysis than the
preliminary o ne by Brerner (1987). They focused on th e
Cichorioideae, sampling 53 gener a, many from the Mu-
tisieae, and used 72 phylogenetically informative char-
acters. Many of the characters were multistate, and the
dat a matrix was equivalent to a da ta set of 110 binary
characters. The matrix yielded two very similar, equally
parsimonious cladograms, differing in a generic rear-
rangement within one of the tribes only, and with
a
consistency index of 0.24.
Th e study by Karis et al. (1992) suggested tha t (1) the
tribe Mutisieae as well as the subfamily Cichorioideae
are paraphyletic, (2) a large part of Mutisieae, exclud-
ing several genera of the subtribe Gochnatiinae, still
form a monophyletic group,
3)
the Mutisieae and the
Card ueae form successive sister groups to the remaining
parts of the family (Fig. lA ), and (4) the sister group to
the Aster oideae can be found within a complex of tribes
consisting
of
the Lactuceae, Vernonieae, Liabeae, and
Arctoteae. A number of alternative topologies were
explored with respect to the number of extra steps
needed. Thus, monophyly of the Mutisieae and the
subfamily Cichorioideae required a considerable num-
ber of extra steps, 9 and 12, respectively. Karis et al.
concluded that the paraphyletic natur e of the Mutisieae
and the Cichorioideae are strongly supported by m orph-
ological data .
Based on the results from Karis et al. (1992), Karis
(1992) subsequently performed a more restricted analy-
sis involving the Lactuceae, Vernonieae, Liabeae, and
Arcto teae, in search for the sister group of th e Asteroi-
deae. This analysis indicated that the Asteroideae sister
group is limited to the Vernonieae, Liab eae, and Arcto-
teae, and that the Vernonieae and Liabeae are sister
tribes (Fig. 1 A). The suggested sister group of the Aste-
roideae was not strongly supported by the data, how-
ever, and the result must be regarded as preliminary.
Within the subfamily Asteroideae (Fig. lB ), the sta-
tus
of
the Inuleae
in
relation to the rest of the Asteroi-
deae has been assessed by Anderberg (1989). He ana-
lyzed a large data set of many Inuleae genera and in-
cluded the rest of the Asteroideae as one taxon.
Anderberg established three monophyletic groups
within the Inulea e; however, together they did not form
a monophyletic group. Consequently he divided the
Inuleae into three tribes, Plucheeae, Gnaphalieae, and
Inuleae sens. str . (A nderb erg 1991a, b, c).
In addition to the studies based on m orphological and
cpDNA restriction site data, Kim et al. (1992) have
sequenced the rbcL gene from 25 Asteraceae genera
represen ting 16 tribes. W agner analysis of the sequence
data gen erated eight equally parsimonious tr ees with a
consistency index of 0.47. Their results suggest that
both the Asteroideae and the Cichorioideae are mono-
phyletic (Fig. l E , F), in agreement with most trees from
restriction site data. Only limited comparisons of tribal
relationships in trees genera ted by the two cp DN A data
sets can be made because of the lack of resolution of
tribal groups in strict consensus trees (Wagner or Dollo)
for the restriction site data (Fig. lC , D ) . Only two
groups of tribes consistently group together: the Lia-
beae and Vernonieae, and the tribes Core opside ae, Ta-
geteae, Heliantheae, and Eupatorieae. The rbcL tree
does not indicate a close relationship between the Lia-
beae and Vernonieae, but it does provide strong sup-
port for the clade including the other four tribes. An
expande d restriction site comp arison (Keeley Jansen
1991), which included ad ditional ge nera of Liabeae and
Vernonieae, has indicated that the sister group relation-
ship between the two tribes is less strongly supported
than was previously suggested by Jansen et al. (1990,
1991 ).
Discussion
Although the morphological and the cpDNA trees are
largely incongruent, all data sets strongly support the
sister group relationship between the Mutisieae-Barna-
desiinae and the rest of the family. The subtribe Barna-
desiinae has now received formal subfarnilial status, as
Barnadesioideae (Bre mer Jansen 1992). It is a small
subfamily with nine genera a nd nearly 90 species, occur-
ring in South America mainly along the Andes. The
Barnadesioideae genera share a number of synapomor-
phies, both morphological and molecular. T he subfam-
ily is thus a strongly supported monophyletic group. Its
members are characterized primarily by unique axillary
spines and by a unique indum entum of long, unicellular,
barnadesioid hairs on the corollas, cypselas, and pappus
(Cab rera 1959, 1961, 1977; Bremer 1987).
The large subfamily Asteroideae is also monophy-
letic, with synapomorphies both in morphology and
cpDNA . Members
of
the subfamily are characterized by
shallowly lobed corollas, style branches with stigmatic
areas sepa rated in two parallel lines, and caveate pollen
(Bremer 1987). Among the molecular synapomorphies
there is a length mutation at the 3’ end of the rbcL gene
involving a six bp repeat (which is repeated four times)
in all examined Asteroideae taxa (Kim et al. 1992).
144
Nord.
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I
Inuleae
I
BARNADESIOIDEAE
Mutisieae
Cardueae
Lactuceae
Vemonieae
Liabeae
Arctoteae
ASTEROIDEAE
A
Karis et al. 1992
Karis 1992
morphology
BARNADESIOIDEAE
Mutisieae
Cardueae
_I
Vemonieae
Liabeae
J 1-Lactuceae
BARNADESIOIDEAE
Mutisieae
Vemonieae
Lactuceae
Arctoteae
ASTEROIDEAE
c
Jansen
et al. 1990,199la
cpDNA restriction sites
r
BARNADESIOIDEAE
Cardueae
Lactuceae
Liabeae
rctoteae
Mutisieae
Vemonieae
STEROIDEAE
E Kim et al. 1992
rbcL
sequences
Astereae
Eupatorieae
Calenduleae
Senecioneae
Anthemideae
Helenieae
Heliantheae
s.
str.
Tageteae
Coreopsideae
B Bremer 1987
morphology
stereae
Anthemideae
Inuleae
Senecioneae
Calenduleae
Eupatorieae
Heliantheaes.
str.
Tageteae.
Coreopsideae
Helenieae
D Jansen et al. 1990,199la
Kim et al. 1990
cpDNA
restriction sites
enecioneae
Calenduleae
Anthemideae
Astereae
Coreopsideae
Tageteae
Heliantheaes. str.
L uDatorieae
F
Kim et al. 1992
rbcL
sequences
Fig.
1.
Phylogenetic hypotheses in the Asteraceae derived from morphology, cpDNA restriction sites, and
rbcL
sequences. A.
Single Wagner tree comb ined from Karis et
al.
(1992) and Karis (1992). B. Single Wagner tree from Brem er (1987). C. Strict
consensus
Dollo
tree from Jansen
et
al. (1990,1991a).
D.
Strict consensus
Dollo
tree combined from Jansen
et
al. (1990, 1991a)
and Kim e t al. (1990). E. Strict consensus Wagner tree from Kim et a l. (1992). F. Strict consensus Wagner tre e from K im et al.
(1992); sequence data for Inuleae and Helenieae are not available.
Nord.
1 Bot.
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2 ) 1992)
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The status
of
the subfamily Cichorioideae (excluding
Barnadesioideae) remains unresolved, and the interre-
lationships among its tribes are not clearly resolved by
our h itherto published analyses. The morphological and
the cpD NA trees are incongruent in several groupings,
especially
if
Bremer’s (1987) tree is compared to those
based on cpDNA. The more detailed analysis of Karis
et al. (1992) has removed some of these conflicts,
whereas othe rs persist. Karis et al. suggest that a large
part of the Mutisieae form a monophyletic group, ex-
cluding an uncertain number of Mutisieae-Gochnatiinae
genera (such as the Stenopadus generic group from the
Guayana Highlands, or the East Asian Ainsliaea ,
which form a more
or
less unresolved paraphyletic
grade at the base. Janse n et al. (1990, 1991a) suggested
that the Mutisieae are monophyletic; all taxa sampled
(including, for example, Ainsliaea consistently group
together.
Karis et al. (1992) suggest that the Lactuceae, Verno-
nieae, Liabeae, Arctoteae, and subfamily Asteroideae
form a monophyletic group, excluding Mutisieae and
Card ueae (Fig. 1 A). In contrast to this hypothesis, Jan-
sen et al.3 (1990, 1991a) cpDNA analyses support the
monophyly of the Cichorioideae (excluding Barnade-
sioideae) . Thus Cichorioideae and Asteroideae should
be sister groups, as shown
in
Fig. 1C and E .
As for the Asteroideae, tribal interrelationships are
also largely obscure. In the cpD NA data th ere is over-
whelming support for a position of the Eupatorieae as
firmly nested within a paraphyletic Heliantheae sens.
lat. In the morphological tree, Eupatorieae and Aste-
reae are sister tribes, although the position of the Eu-
patorieae together with the He lianthea e is only one ste p
longer (Bremer 1987; see also Sanderson’s (1989) boot-
strap analysis of Bremer’s data). Morphological data
could support the arrangement of the Eupatorieae to-
gether with the Heliantheae, although this requires ac-
ceptance of slightly less parsimonious trees (Bremer
1987). We conclude that the Astereae-Eupatorieae
grouping in Bremer’s (1987) analysis was err oneou s and
due to parallelisms. T he o ther tribal arrangements sug-
gested by Fig. 1B, D , and F are only weakly supported
in both data sets.
With respect to tribal classification, we propose rein-
stat em ent of thr ee tribes, which since Robinson’s (1981)
treatment have been included in Helianth eae sens. lat.
The Helenieae and Coreopsideae form groups outside
the Heliantheae sens. str., which are more closely re-
lated to the Eupatorieae than
to
either of these three
tribes. The Tageteae (S trother 1977) are nested within
the basal part of the Heliantheae-Eupatorieae complex
and are tentatively reinstated
in
our classification. T he
Tageteae are paraphyletic in the cpDN A trees
of
Jansen
et al. (1990, 1991a), but the extended data set (Kim et
al. 1990) supports the monophyly of this tribe.
The I nuleae were split into three tribes by Anderb erg
(1989, 1991a, b, c). Four genera from Anderberg’s th ree
tribes were included in the cpDNA analysis (Jansen et
al. 1990, 1991a), but they consistently formed a mono-
phyletic grou p. Since the sample of Inulea e genera was
rather small, the cpDNA analysis may not be very re-
liable regarding the Inuleae problem, and Anderberg’s
proposed reclassification should be seriously con-
sidered. However, pending results from Karis’s current
morphological analysis of a large set
of
Asteroideae
genera and from Jansen’s extended restriction site
analysis of the family, we retain In ulea e sens. lat. in the
list
of
Asteroideae tribes.
Th e Cardu eae were split into three trib es by Dittrich
(1977). Dittrich’s three tribes were all included in both
the morphological ( Brem er 1987; Karis et al. 1992) and
molecular (Ja nsen e t a]. 1990, 1991a) studies, a nd since
they form a monophyletic group we see no reason to
split the Car duea e sens. lat. and we retain it as a single
tribe.
A large part of the Mutisieae still forms a mono-
phyletic group in the most recent morphological study
(Karis et al. 1992), and all genera sampled in the
cpD NA analyses (Jansen Palmer 1988; Jansen et al.
1990, 1991a) consistently group together. Hence the
Mutisieae may be retained after exclusion of the Barna-
desiinae. Yet morphological data as well as an extended
cpDN A restriction site data set (Keeley Jansen 1991)
suggest that several other genera could be excluded
from the Mutisieae and classified in new tribes of the
Cichorioideae. Examples include the Stenopadus group
from the G uyana Highlands (Karis et al. 1992) or
Bra-
chylaena and Tarchonanthus (Keeley Jansen 1991).
Thu s Keeley Jansen (1991) proposed a new tribe,
Tarchonantheae, for the latter two gene ra. However we
have provisionally retained these genera within the Mu-
tisieae in our joint classification (Tab. l), pending a
more detailed resolution of the basal phylogeny of the
Cichorioideae.
Conclusions
Cladistic analysis of morphological and cpDNA data
has in a few years substantially improved our under-
standing of Asteraceae phylogeny. We are reasonably
confident that (1) the Barnadesioideae and the rest of
the family are monophyletic sister groups, 2) the Aste-
roideae are monophyletic, (3) the Vernonieae and Lia-
beae are sister groups, and (4) the Eupatorieae are
nested within the Heliantheae. With increased efforts
we may succeed in resolving the phylogeny in consider-
able detail and provide a basis for a more accurate and
phylogenetically informative classification. Further
studies are needed to evaluate the status of th e subfam-
ily Cichorioideae and to resolve tribal interrelationships
within the two subfamilies Cichorioideae and Asteroi-
deae
146
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