Bremer 1992

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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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2) 1992)

141

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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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8/15/2019 Bremer 1992


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

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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.

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145

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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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