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 Asteraceae are commonly divided into two large subfamilies, the Cichorioideae
(syn. Lactucoideae; Mutisieae, Cardueae, Lactuceae, Vernonieae, Liabeae, Arctoteae) and the Asteroideae (Inuleae, Astereae, Anthemideae, Senecioneae, Calenduleae, Heliantheae, Eupatorieae). Recent phylogenetic analyses based on
morphological and chloroplast DNA data conclusively show that the MutisieaeBarnadesiinae are the sister group 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 interrelationships are still largely uncertain. The Mutisieae-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 DNA data 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 further resolve tribal interrelationships in the two larger
subfamilies.

K . Bremer, Dept of Systematic Botany, Uppsala University, Box 541, S-7.5121 Uppsala, Sweden. - R. K . Jansen, Dept of Botany, Univ. of Texas, Austin, Texas 78713,
U.S.A. - P. 0 . Karis, Dept of Phanerogamic Botany, Swedish Museum of Natural
History, S-104 05 Stockholm, Sweden. - M . Kallersjo, Lab. of Molecular Systematics,
Smithsonian Institution, Washington, DC 20560, U.S.A. - S. C. Keeley, Dept of
Botany, Univ. of Hawaii, Honolulu, Hawaii 96822, U.S.A. - K.-J. Kim, Dept of
Botany, Univ. of Texas, Austin, Texas 78713, U.S.A. - H. J . Michaels, Dept of
Biology, Bowling Green State University, Bowling Green, Ohio 43402, U.S.A. - J . D.
Palmer, Dept of Biology, Indiana University, Bloomington, Indiana 47405, U.S.A . R . S . Wallace, Dept of Botany, Iowa State University, Ames, Iowa 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. Karis, M. Kallersjo) and chloroplast DNA (R. Jansen, S.
Keeley, K.-J. Kim, H. Michaels, J. Palmer, R. Wallace)
data. We (1) recognize three subfamilies, the Asteroideae, the Cichorioideae (syn. Lactucoideae), and the

small South American Barnadesioideae, (2) acknowledge the monophyly of the large subfamily Asteroideae,
which includes the tribes Astereae, Anthemideae,
Inuleae, Senecioneae, Calenduleae, Heliantheae, and
Eupatorieae, and (3) discuss the uncertain status of the
subfamily Cichorioideae, which includes the Mutisieae,
Cardueae, Lactuceae, Vernonieae, Liabeae, and Arctoteae. For brevity and simplicity we generally use tribal
names in a broad sense, for example, Cardueae sens.

Accepted 6-12-91

NORDIC JOURNAL OF BOTANY

NORD. J. BOT. 12: 141-148

10 Nord J . Bot. I?(2) (1992)

141

lat., Inuleae sens. lat., and Heliantheae sens. lat., although we realize that reclassification into smaller tribes
has been proposed (e.g., Dittrich 1977; Turner & Powell 1977; Strother 1977; Anderberg 1989). The name
Mutisieae is applied sensu stricto, excluding the former
subtribe Barnadesiinae. The tribal classification is commented upon in the discussion and those tribes currently
supported by both morphological and molecular data
are listed under classification.
Until the 1970s the tribe Lactuceae was considered to
be distinct from all other Asteraceae and was classified
in its own subfamily, the Liguliflorae. The other tribes
were placed together in the Tubuliflorae (e.g., Hoffmann 1890). This classification still persists in some
floras and handbooks. During the 1970s there was a
growing recognition that all the tribes, including the
Lactuceae, could be arranged in two large groups
(Robinson & Brettell 1973; Carlquist 1976), foreshadowed by diagrams in Carlquist (1961) and Poljakov
(1967). Carlquist (1976) formally treated these two
groups as the subfamilies Cichorioideae and Asteroideae. Their precise circumscription was modified by
later authors (Robinson 1977, 1981, 1983; Thorne
1983). Recent subfamilial and tribal classifications of
the Asteraceae are summarized in Tab. 1.

ily were partly unresolved. Within the subfamily Asteroideae alternative arrangements were restricted to the
subtribal level, leaving a single pattern of tribal interrelationships (Fig. 1B).
Bremer (1987) demonstrated that the subfamily Asteroideae is a well-supported monophyletic group. The
basal sister group relationship between the MutisieaeBarnadesiinae and the rest of the family (Fig. lA,C),
revealed by the cpDNA inversion, was also supported
by several morphological characters, such as presence
of the typical Asteraceae twin hairs on the fruits and
spiny pollen.
Although other results were less certain, the analysis
suggested that (1) the tribe Mutisieae and the subfamily
Cichorioideae are paraphyletic even if the Barnadesiinae are excluded, (2) the Arctoteae and the Cardueae
are related, (3) the Vernonieae and Liabeae are sister
groups, (4) the Inuleae sens. lat. are paraphyletic and a
basal grade within the Asteroideae, and ( 5 ) the Astereae and Eupatorieae are sister groups. Later molecular
and morphological studies have weakened the sister
group relationship between the Vernonieae and Liabeae and have refuted a close relationship between the
Arctoteae and Cardueae, and between the Astereae
and Eupatorieae (Jansen et al. 1990, 1991a; Karis 1992,
Karis et al. 1992).
Following their Mutisieae investigations, Jansen and
collaborators performed a large study of cpDNA restriction site variation in the entire family (Jansen et al.
1990.1991a. b). Thev samded 57 genera from all maior
tribes and identified 328 'phylog&etically informative
restriction site mutations. The data matrix was analyzed
using both Wagner and Dollo parsimony, as well as the
bootstrap method (Felsenstein 1985). The Wagner
analysis resulted in 20 equally parsimonious trees with a
consistency index of 0.46, and Dollo parsimony generated 16 trees with a consistency index of 0.44. Jansen et
al. (1990) preferred the Dollo tree topology (Fig.
lC, D) as the best estimate of phylogenetic relationships
in the Asteraceae arguing that (1) the Dollo algorithm is
more appropriate for restriction site mutations because
convergent gains occur at least an order of magnitude
less than convergent losses (Templeton 1983; DeBry &
Slade 1985), (2) eight of the 20 equally parsimonious
Wagner trees have the Dollo tree topology, and (3) a
posteriori character analysis, including successive approximation (Farris 1969), followed by Wagner analysis
always gave trees with the Dollo topology. Furthermore, phylogenetic analyses of the restriction site data
employing character state weighting of sensu Holsinger
& Jansen (1992) also produced the Dollo topology (Jansen 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; the Dollo topologies are largely incongruent with
the morphological trees whereas some of the Wagner
topologies are similar to the morphological trees.
Jansen et al.3 analyses indicated that (1) the subfamI

Recent cladistic analyses

Since 1985 cladistic analyses of molecular and morphological data have illuminated the phylogeny and classification of the Asteraceae. Jansen & Palmer (1987)
reported the discovery of a 22 kb cpDNA inversion in
Lactuca and several other Asteraceae. The inversion
proved to be absent in three genera of the MutisieaeBarnadesiinae, suggesting a basal dichotomy between
the Barnadesiinae and the rest of the family. Subsequently, Jansen & Palmer (1988) undertook an analysis
of restriction site variation in the Mutisieae. They sampled 13 genera of the Mutisieae (including the Barnadesiinae) and three representatives each from the Lactuceae, Cardueae, and Heliantheae. This study corroborated the sister group relationship between the
Barnadesiinae and the rest of the family. Furthermore,
the Mutisieae sens. str. (excluding the Barnadesiinae)
came out as a monophyletic group.
Simultaneously with Jansen & .Palmer's cpDNA investigations Bremer (1987) performed a cladistic analysis of tribal interrelationships based on mainly morphological data. The study included some 27 tribes and
subtribes and 47 phylogenetically informative characters, one of them being the cpDNA inversion. Several
hundred equally parsimonious Wagner trees were
found, with a consistency index of 0.54 (excluding autapomorphies). The strict consensus tree contained a
number of polychotomies especially within the Cichorioideae. Hence tribal interrelationships in that subfam142

Nord. J . Bot. 12 (2) (1992)

----

5
5

Anthemideae
Astereae
Senecioneae
Inuleae
Heliantheae
Filagineae
Arctotideae
Mutisieae
Cynareae
Vernonieae
Cichorieae
Fi tchineae

Poljakov
(1967)

Eupatorieae
Vernonieae
Astereae
Inuleae
Heliantheae
Tageteae
Senecioneae
Liabeae
Anthemideae
Arctoteae
Calenduleae
Cynareae
Mutisieae
Lactuceae

Heywood
et al. (1977)

Eupatorieae
Heliantheae
Helenieae
Senecioneae
Calenduleae
Astereae
Inuleae
Anthemideae

Group 2

Vernonieae
Liabeae
Mutisieae
Cardueae
Echinopeae
Arctotideae

Group I

Asteroideae

Cichorioideae

Wagenitz
(1976)

Senecioneae
Tageteae
Heliantheae
Inuleae
Anthemideae
Ursinieae
Calenduleae
Cotuleae
Astereae

Asreroideae

Helian theae
Astereae
Inuleae
Anthemideae
Senecioneae
Calenduleae

Asteroideae

Lactuceae
Mutisieae
Eremothamneae
Arctotideae
Cardueae
Vernonieae
Liabeae
Eupatorieae

Eupatorieae
Heliantheae
Calenduleae
Inuleae
Senecioneae
Astereae
Anthemideae

Asteroideae

Heliantheae
Tageteae
Eupatorieae
Astereae
Inuleae
Anthemideae
Senecioneae
Calenduleae

Asteroideae

Mutisieae
Vernonieae
Liabeae
Cichorieae
Cardueae
Arctoteae

Cichorioideae

Thorne
(1983)

Inuleae
Gnaphalieae
Astereae
Eupatorieae
Calenduleae
Senecioneae
Anthemideae
Helenieae
Madieae
Heliantheae
Tageteae
Coreopsideae

Asteroideae

Mutisieae
Arctoteae
Carlineae
Echinopsideae
Cardueae
Lactuceae
Eremothamneae
Vernonieae
Liabeae

Cichorioideae

Heliantheae
Astereae
Cichorioideae Anthemideae
Arctotideae
Mutisieae
Inuleae
Cardueae
Senecioneae
Lactuceae
Calenduleae
Vernonieae
Eupatorieae
Liabeae
Vernonieae
Arctoteae
Liabeae
Cardueae
Asreroideae
Mutisieae
Cichorieae
Inuleae
Astereae
Anthemideae
Senecioneae
Calenduleae
Helenieae
Coreopsideae
Tageteae
Heliantheae
Eupatorieae

Barnadesieae

Cronquist
(1955, 1977)

Cardueae).

Barnadesioideae

This paper
(1992)

Arctoteae, Cynareae

Bremer
(1987)

Lactuceae, Arctotideae

Lactuceae
Vernonieae
Liabeae
Mutisieae
Cardueae
Echinopsideae
Gundelieae
Eremothamneae
Arctoteae

Cichorioideae

Robinson
(1981,1983)

Cichorioideae, Cichorieae

Lactucoideae

Jeffrey
(1978)

Mutisieae
Vernonieae
Cardueae
Arctoteae
Cichorieae
Eupatorieae

Cichorioideae

Carlquist
(1976)

Tab. 1 . Recent major classifications of the Asteraceae (Lactucoideae

ily Asteroideae is strongly supported as a monophyletic

group, (2) the subfamily Cichorioideae likewise is
monophyletic, although this result is less well supported, (3) 13 of the 15 investigated tribes are monophyletic, with the Heliantheae and Tageteae being paraphyletic, (4) the Vernonieae and Liabeae are sister
tribes, and that (5) the Coreopsideae, Tageteae, Heliantheae, and Eupatorieae are closely related, and the
Eupatorieae are nested within a paraphyletic Heliantheae. Interrelationships within the Heliantheae-Eupatorieae complex have been further clarified by Kim et
al. (1990) (Fig. 1D).
Using the Barnadesiinae as an outgroup, tribal interrelationships within the Cichorioideae were explored
in more detail by Karis et al. (1992). They performed a
much more detailed morphological analysis than the
preliminary one by Brerner (1987). They focused on the
Cichorioideae, sampling 53 genera, many from the Mutisieae, and used 72 phylogenetically informative characters. Many of the characters were multistate, and the
data matrix was equivalent to a data set of 110 binary
characters. The matrix yielded two very similar, equally
parsimonious cladograms, differing in a generic rearrangement within one of the tribes only, and with a
consistency index of 0.24.
The study by Karis et al. (1992) suggested that (1) the
tribe Mutisieae as well as the subfamily Cichorioideae
are paraphyletic, (2) a large part of Mutisieae, excluding several genera of the subtribe Gochnatiinae, still
form a monophyletic group, (3) the Mutisieae and the
Cardueae form successive sister groups to the remaining
parts of the family (Fig. lA), and (4) the sister group to
the Asteroideae 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 number of extra steps, 9 and 12, respectively. Karis et al.
concluded that the paraphyletic nature of the Mutisieae
and the Cichorioideae are strongly supported by morphological data.
Based on the results from Karis et al. (1992), Karis
(1992) subsequently performed a more restricted analysis involving the Lactuceae, Vernonieae, Liabeae, and
Arctoteae, in search for the sister group of the Asteroideae. This analysis indicated that the Asteroideae sister
group is limited to the Vernonieae, Liabeae, and Arctoteae, and that the Vernonieae and Liabeae are sister
tribes (Fig. 1A). The suggested sister group of the Asteroideae was not strongly supported by the data, however, and the result must be regarded as preliminary.
Within the subfamily Asteroideae (Fig. lB), the status of the Inuleae in relation to the rest of the Asteroideae has been assessed by Anderberg (1989). He analyzed a large data set of many Inuleae genera and included the rest of the Asteroideae as one taxon.
Anderberg established three monophyletic groups

144

within the Inuleae; however, together they did not form

a monophyletic group. Consequently he divided the
Inuleae into three tribes, Plucheeae, Gnaphalieae, and
Inuleae sens. str. (Anderberg 1991a, b, c).
In addition to the studies based on morphological and
cpDNA restriction site data, Kim et al. (1992) have
sequenced the rbcL gene from 25 Asteraceae genera
representing 16 tribes. Wagner analysis of the sequence
data generated eight equally parsimonious trees with a
consistency index of 0.47. Their results suggest that
both the Asteroideae and the Cichorioideae are monophyletic (Fig. l E , F), in agreement with most trees from
restriction site data. Only limited comparisons of tribal
relationships in trees generated by the two cpDNA 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. l C , D ) . Only two
groups of tribes consistently group together: the Liabeae and Vernonieae, and the tribes Coreopsideae, Tageteae, Heliantheae, and Eupatorieae. The rbcL tree
does not indicate a close relationship between the Liabeae and Vernonieae, but it does provide strong support for the clade including the other four tribes. An
expanded restriction site comparison (Keeley & Jansen
1991), which included additional genera of Liabeae and
Vernonieae, has indicated that the sister group relationship between the two tribes is less strongly supported
than was previously suggested by Jansen et al. (1990,
1991a).

Discussion
Although the morphological and the cpDNA trees are
largely incongruent, all data sets strongly support the
sister group relationship between the Mutisieae-Barnadesiinae and the rest of the family. The subtribe Barnadesiinae has now received formal subfarnilial status, as
Barnadesioideae (Bremer & Jansen 1992). It is a small
subfamily with nine genera and nearly 90 species, occurring in South America mainly along the Andes. The
Barnadesioideae genera share a number of synapomorphies, both morphological and molecular. The subfamily is thus a strongly supported monophyletic group. Its
members are characterized primarily by unique axillary
spines and by a unique indumentum of long, unicellular,
barnadesioid hairs on the corollas, cypselas, and pappus
(Cabrera 1959, 1961, 1977; Bremer 1987).
The large subfamily Asteroideae is also monophyletic, with synapomorphies both in morphology and
cpDNA. Members of the subfamily are characterized by
shallowly lobed corollas, style branches with stigmatic
areas separated 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).
Nord. J . Bot. I2 ( 2 ) (1992)

Inuleae
Astereae
Eupatorieae
Calenduleae
Senecioneae
Anthemideae
Helenieae
Heliantheae s. str.
Tageteae
Coreopsideae

BARNADESIOIDEAE
Mutisieae
Cardueae
Lactuceae
Vemonieae
Liabeae
Arctoteae
ASTEROIDEAE

Karis et al. 1992

Karis 1992
morphology

Astereae
Anthemideae
Inuleae
Senecioneae
Calenduleae
Eupatorieae
Heliantheae s. str.
Tageteae.
Coreopsideae
Helenieae

BARNADESIOIDEAE
Mutisieae
Cardueae
Vemonieae
Liabeae
1-Lactuceae
Lactuceae
Arctoteae
ASTEROIDEAE

_
I

IJ

c
r

Jansen et al. 1990,199la

cpDNA restriction sites

BARNADESIOIDEAE
Cardueae
Lactuceae
Liabeae
7
Arctoteae
Mutisieae
Vemonieae
+
ASTEROIDEAE

Kim et al. 1992

rbcL sequences

Bremer 1987
morphology

Jansen et al. 1990,199la

Kim et al. 1990
cpDNA restriction sites

I
Senecioneae

Calenduleae
Anthemideae
Astereae
Coreopsideae
Tageteae
Heliantheae s. str.
LEuDatorieae

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 combined from Karis et al. (1992) and Karis (1992). - B. Single Wagner tree from Bremer (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 et al. (1990). - E. Strict consensus Wagner tree from Kim et al. (1992). - F. Strict consensus Wagner tree from Kim et al.
(1992); sequence data for Inuleae and Helenieae are not available.

Nord. 1. Bot. 12 ( 2 ) (1992)

145

The status of the subfamily Cichorioideae (excluding

Barnadesioideae) remains unresolved, and the interrelationships among its tribes are not clearly resolved by
our hitherto published analyses. The morphological and
the cpDNA trees are incongruent in several groupings,
especially if Bremers (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 others persist. Karis et al. suggest that a large
part of the Mutisieae form a monophyletic group, excluding 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. Jansen 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, Vernonieae, Liabeae, Arctoteae, and subfamily Asteroideae
form a monophyletic group, excluding Mutisieae and
Cardueae (Fig. 1A). In contrast to this hypothesis, Jansen et al.3 (1990, 1991a) cpDNA analyses support the
monophyly of the Cichorioideae (excluding Barnadesioideae) . 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 cpDNA data there is overwhelming support for a position of the Eupatorieae as
firmly nested within a paraphyletic Heliantheae sens.
lat. In the morphological tree, Eupatorieae and Astereae are sister tribes, although the position of the Eupatorieae together with the Heliantheae is only one step
longer (Bremer 1987; see also Sandersons (1989) bootstrap analysis of Bremers data). Morphological data
could support the arrangement of the Eupatorieae together with the Heliantheae, although this requires acceptance of slightly less parsimonious trees (Bremer
1987). We conclude that the Astereae-Eupatorieae
grouping in Bremers (1987) analysis was erroneous and
due to parallelisms. The other tribal arrangements suggested by Fig. 1B, D, and F are only weakly supported
in both data sets.
With respect to tribal classification, we propose reinstatement of three tribes, which since Robinsons (1981)
treatment have been included in Heliantheae sens. lat.
The Helenieae and Coreopsideae form groups outside
the Heliantheae sens. str., which are more closely related to the Eupatorieae than to either of these three
tribes. The Tageteae (Strother 1977) are nested within
the basal part of the Heliantheae-Eupatorieae complex
and are tentatively reinstated in our classification. The
Tageteae are paraphyletic in the cpDNA trees of Jansen
et al. (1990, 1991a), but the extended data set (Kim et
al. 1990) supports the monophyly of this tribe.
The Inuleae were split into three tribes by Anderberg
(1989, 1991a, b, c). Four genera from Anderbergs three
tribes were included in the cpDNA analysis (Jansen et
146

al. 1990, 1991a), but they consistently formed a monophyletic group. Since the sample of Inuleae genera was
rather small, the cpDNA analysis may not be very reliable regarding the Inuleae problem, and Anderbergs
proposed reclassification should be seriously considered. However, pending results from Kariss current
morphological analysis of a large set of Asteroideae
genera and from Jansens extended restriction site
analysis of the family, we retain Inuleae sens. lat. in the
list of Asteroideae tribes.
The Cardueae were split into three tribes by Dittrich
(1977). Dittrichs three tribes were all included in both
the morphological (Bremer 1987; Karis et al. 1992) and
molecular (Jansen et a]. 1990, 1991a) studies, and since
they form a monophyletic group we see no reason to
split the Cardueae sens. lat. and we retain it as a single
tribe.
A large part of the Mutisieae still forms a monophyletic group in the most recent morphological study
(Karis et al. 1992), and all genera sampled in the
cpDNA analyses (Jansen & Palmer 1988; Jansen et al.
1990, 1991a) consistently group together. Hence the
Mutisieae may be retained after exclusion of the Barnadesiinae. Yet morphological data as well as an extended
cpDNA 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 Guyana Highlands (Karis et al. 1992) or Brachylaena and Tarchonanthus (Keeley & Jansen 1991).
Thus Keeley & Jansen (1991) proposed a new tribe,
Tarchonantheae, for the latter two genera. However we
have provisionally retained these genera within the Mutisieae 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 understanding of Asteraceae phylogeny. We are reasonably
confident that (1) the Barnadesioideae and the rest of
the family are monophyletic sister groups, (2) the Asteroideae are monophyletic, (3) the Vernonieae and Liabeae are sister groups, and (4) the Eupatorieae are
nested within the Heliantheae. With increased efforts
we may succeed in resolving the phylogeny in considerable detail and provide a basis for a more accurate and
phylogenetically informative classification. Further
studies are needed to evaluate the status of the subfamily Cichorioideae and to resolve tribal interrelationships
within the two subfamilies Cichorioideae and Asteroideae .
Nard. 1. Bat. 12 (2) (1992)

Classification

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Ann. Missouri Bot. Gard. 79 (in press), 1992.

Type: Barnadesia Mutis ex L. f.
Axillary spines generally present. Corollas, cypselas,
and pappus villous with long unicellular barnadesioid
hairs. Corolla usually actinomorphic or pseudobilabiate
(1 adaxial and 4 abaxial lobes). Anthers calcarate or
ecalcarate, caudate or ecaudate. Styles shortly bilobed,
glabrous o r papillose, with continuous stigmatic areas.
Pollen smooth to granular, spineless, sometimes with
depressions or honey-combed with smooth ridges.

Cichorioideae Kitamura
Mem. Coll. Sci. Kyoto Imp. Univ. Ser. B, Biol. 13: 4,
1937.
Type: Cichorium L.
Syn.: Lactucoideae Solbrig, Taxon 12: 230, 1963.
Axillary spines and barnadesioid hairs absent. Corolla
actinomorphic and usually deeply lobed, bilabiate (2
adaxial and 3 abaxial lobes), ligulate (Lactuceae), or
sometimes radiate (Arctoteae, Liabeae). Anthers
mostly calcarate and caudate. Styles often pilose on the
shaft and along the branches, with continuous stigmatic
areas. Pollen mostly spiny, sometimes honey-combed,
usually ecaveate.

Asteroideae
Type: Aster L.
Axillary spines and barnadesioid hairs absent. Corolla
actinomorphic and usually shallowly lobed, o r radiate.
Anthers mostly ecalcarate and often ecaudate. Styles
generally pilose mainly at the tips of the branches,
mostly with stigmatic areas in two separate lines. Pollen
mostly spiny and caveate, never honey-combed.

Acknowledgements - This study was supported by NFR grant

2401-108 to K. Bremer, NSF grants BSR-8415934 to J . Palmer

and R. Jansen and BSR-8708246 to R. Jansen, and a University of Connecticut Research Foundation grant to R. Jansen. K. Bremers participation in the symposium Phylogeny of
the Asteridae was covered by NSF grant BSR-9005921 to
Larry Hufford. We thank Meredith A. Lane for useful comments.
Nord. J . BoC. I2 (2) (1992)

147

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