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Phylogeny and evolution of the Astereae (Compositae or Asteraceae)



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Chapter 37
Luc Brouillet, Timothy K. Lowrey, Lowell Urbatsch, Vesna Karaman-Castro,
Gisela Sancho, Steve Wagsta and John C. Semple
Tribe Astereae, with an estimated 222 genera and ca. 3100
species (emended from Nesom and obinson, 2007), is
the second largest tribe of family Compositae. Members
of the tribe range from the Arctic to the tropics, though
they are more numerous in temperate regions. They are
mostly found in open habitats, from salt marshes and bogs
to deserts, and from sea level to the alpine zone. They
are mostly characterized by their ecaudate and ecalcarate
anther bases (though a few genera have caudate anther
bases), and a disc  oret style with two distinct marginal
stigmatic lines and deltate to triangular or lanceolate style
appendages that are glabrous adaxially and with sweeping
hairs abaxially.
The tribe is part of subfamily Asteroideae and belongs
to a clade that includes Calenduleae, Gnaphalieae, and
Anthemideae, the latter often considered its sister tribe
(e.g., Panero and Funk 2002, 2008), though this ques-
tion is still not fully resolved (e.g., Goertzen et al. 2003).
Grau (1977), Bremer (1994), Nesom (1994g), and Nesom
and obinson (2007) summarized the taxonomic history
of the tribe. These authors addressed subtribal classi ca-
tion of Astereae, acknowledging the somewhat arti cial
nature of the traditional divisions. The most signi cant
departure from the classic scheme was that of Nesom
(1994g, 2000a), which culminated in the recent classi ca-
tion by Nesom and obinson (2007). This classi cation
incorporates some, but not all, the molecular phylogenetic
information published between 1994 and 2004. Though
an evolutionary tree was presented by Nesom (1994g), it
was not based on a phylogenetic analysis of characters.
The only cladistic analysis of tribe Astereae (Zhang and
Bremer 1993) was subsequently used by Bremer (1994).
This analysis, however, was based on a few representa-
tives selected from the existing subtribes, using traditional
morphological characters and the sampling did not allow
for the assessment of whether or not the subtribes were
monophyletic. Nesom (1994g) reviewed this analysis and
made extensive comments on the data and conclusions.
Much of the taxonomic literature published since Grau
(1977) has been summarized by Bremer (1994), Nesom
(1994g, i, 2000a), Müller (2006), Flora of North America
Editorial Committee (2006), and in the phylogenetic pa-
pers cited herein. This literature will not be repeated here
unless necessary.
Molecular phylogenies in the tribe initially were based
on cpDNA FLP analyses, mostly of North American
(Suh and Simpson 1990; Zanowiac 1991; Morgan and
Simpson 1992; Morgan 1993, 1997; Lane et al. 1996;
Xiang and Semple 1996; Zhang 1996; see Semple et al.
1999) or Asian genera (Gu et al. 1994; Ito et al. 1995,
1998). Few papers using cpDNA sequence data were pub-
lished for Astereae (Denda et al. 1999; Bayer and Cross
2002; Liu et al. 2002; Watanabe et al. 2006; Forest et
al. 2007); these data are not cumulative since di erent
markers were used in the studies, which further used a
restricted sample of taxa.
Mostly after the publication of the seminal paper by
Noyes and ieseberg (1999), phylogenies of genera or
Brouillet, Lowrey, Urbatsch, Karaman-Castro, Sancho, Wagstaff and Semple590
groups of genera based on the nuclear ribosomal inter-
nal transcribed spacer (ITS) were published (Morgan
1997, 2003; Noyes 2000a; Lowrey et al. 2001; Markos
and Baldwin 2001; Cross et al. 2002; Fiz et al. 2002; Liu
et al. 2002; Wagsta and Breitwieser 2002; oberts and
Urbatsch 2003, 2004; Urbatsch et al. 2003; Beck et al.
2004; Brouillet et al. 2004; Eastwood et al. 2004; Field
et al. 2006; Watanabe et al. 2006; Selliah and Brouillet
2008). Some papers also added the nrDNA 3ETS region
as a phylogenetic tool in Astereae (Markos and Baldwin
2001, 2002; Morgan 2003; oberts and Urbatsch 2003,
2004; Urbatsch et al. 2003; Brouillet et al. 2004; Selliah
and Brouillet 2008).
Given the large number of available ITS sequences
in Astereae and the fact that no phylogeny exists for the
whole tribe, we  rstly present an ITS-based phylogeny of
the entire tribe. The resulting lineages are discussed in re-
lation with the pertinent literature. Secondly, we review
brie y the various characters (morphological, cytologi-
cal, chemical) and their distribution in the tribe. Thirdly,
economic uses and biological data on Astereae are sum-
marized. Fourthly, we examine the biogeography of the
tribe in terms of the ITS phylogeny, contrasting it with
hypotheses based on traditional classi cations. Finally, we
examine the impact of the current molecular phylogeny
on generic delimitation and contrast our Astereae ITS
phylogeny with the recent classi cation of Nesom and
obinson (2007).
The phylogeny was r econstruc ted using Gen Bank-avai lable
nrDNA ITS sequence data for Astereae, as well as repre-
sentat ive s of the sist er t ri be s C al endu le ae, Gn aph al ie ae , a nd
Anthemideae. Details on sequencing methods and phylo-
genetic analyses, as well as discussions of speci c results,
are available in the papers where the data were initially
published (cited above), as well as in papers being con-
currently prepared with the present analysis (Sancho and
Karaman-Castro 2008; Karaman-Castro and Urbatsch,
pers. comm.; Brouillet et al., pers. comm.). To align the
more than 850 available sequences, and more speci cally
the Astereae sequences with those of the outgroups, we
used the 80%-consensus aligned sequences for the tribes of
Asteraceae as determined from the ITS secondary struc-
ture by Goertzen et al. (2003). Small misalignments in
the consensus sequences of Astereae, Gnaphalieae, and
Anthemideae were noted in ITS1 (before helix 1A and in
helix 1B) and in ITS2 (in helices 2A and 2C). These most
likely resulted, for Astereae, from the fact that the sequences
used by Goertzen et al. were mostly from derived genera
with numerous indels, such as Erigeron. The availability of
sequences of primitive Astereae, such as Nannoglottis and
Printzia Cass., and the large number of available ITS se-
quences, allowed us to correct these misaligned portions.
Duplicate sequences or sequences of dubious origin were
removed from the matrix. The resulting matrix included
752 taxa (737 Astereae and 15 outgroup species) by 852
characters, of which 468 were parsimony informative.
Gaps were treated as missing characters. Multistate data
were treated as uncertainties. Given that sequences came
from a number of laboratories, there were missing data,
particularly in the outgroups where many lacked the 5.8S
portion; otherwise, data were m issing mostly from the be-
ginning of ITS1 or the end of ITS2; missing data were
usually a few bases only.
Given the size of the matrix, a PAUPat (Sikes and
Lewis 2001) parsimony analysis was carried out using
PAUP*10.6b (Swo ord 2002) (swap = tbr, nrep = 1,
addseq = random, nchuck = 1, chuckscore = 1, wtset = 1).
Subsequently, a parsimony analysis starting from the trees
obtained in the 15 ratchet runs was done in PAUP* (tbr,
maxtree = 20,000) in order to obtain a greater number
of trees as a basis for the strict consensus tree. Base chro-
mosome numbers and distribution (as branch color) were
manually mapped onto the phylogeny.
esulting parsimony trees had a length of 7928, a consis-
tency index (non informative characters removed) of 0.15,
a homoplasy index of 0.84, and a retention index of 0.79;
the rescaled consistency index was 0.12. The strict con-
sensus tree is presented in Fig. 37.1A–E, where subtribes
following Nesom and obinson (2007) are indicated to
the right of the cladograms. Given the number of parsi-
mony informative characters, much lower than the num-
ber of taxa analyzed, and the relatively recent radiations
in many groups of Astereae, homoplasy is high and in
part responsible for the low resolution of the trees. More
localized analyses (see studies cited above), which exclude
taxa from other groups or continents, often produce more
resolved trees. Support for clades is usually non-signi -
cant, in large part due to the low number of characters
de ning each branch, particularly along the spine of the
tree. The addition of phylogenetically signi cant indels
to the analysis may increase support for individual clades
(Brouillet, pers. obs.).
The ITS sequence data, as analyzed here, do not allow
us to determine the relationships between Astereae and
the three tribes to which it is considered closely related,
Calenduleae, Gnaphalieae, and Anthemideae.
Tribe Astereae is monophyletic and resolves into a
number of large clades that will each be described below;
many have been the object of recent phylogenetic analy-
ses. The phylogeny (Fig. 37.1A–E) includes a basal African
Chapter 37: Astereae 591
grade, with disjunct Chinese, South American, and New
Zealand lineages, and a polytomy of crown groups in
Australasia, South America, and North America.
Basal lineages: out of Africa, repeatedly
Printzia (Figs. 37.2A, 37.3A) and Denekia Thunb. (Fig.
37.3B) have been shown by Bayer and Cross (2002) and
Brouillet et al. (pers. comm.) to belong to tribe Astereae
and are sister to the remaining members of the tribe (Fig.
37.1A). Using rbcL data, Forest et al. (2007, Supplement
3) also placed Printzia within Astereae, but not as sister to
the genera studied. Both genera are South African, which
supports the hypothesis of an African origin for the tribe
(Noyes and ieseberg 1999). The tailed anthers of Printzia
and Denekia is a symplesiomorphy shared with related
tribes and tends to con rm the early diverging position
of the taxa in a mostly tail-less tribe. Printzia is placed
in subtribe Hinterhuberinae by Nesom and obinson
(2007), a position not supported here. Denekia has tradi-
tionally been placed in tribe Gnaphalieae (e.g., Bayer et
al. 2007). Placement within Astereae is novel and post-
dates the Nesom and obinson classi cation. The capitu-
lum morphology of Denekia, with its bilabiate peripheral
(rays) and functionally male disc orets, appears odd in
Astereae, but some aspects of its morphology certainly
agree with a placement near Printzia, notably the tomen-
tose abaxial leaf faces. These two genera may deserve a
subtribe of their own.
Liu et al. (2002) showed that the Chinese Nannoglottis
Ma xim. is sister to other Astereae, though they postulated
an African origin followed by dispersal to Asia. Our data
(Fig. 37.1A) con rm that it is one of the earliest diverg-
ing lineages and that it must have d ispersed from Africa to
China at some early stage of Astereae evolution.
Brouillet et al. (pers. comm.) showed that Mairia Nees
is an isolated genus among African Astereae, between
Nannoglottis and the Paleo South American and New
Zealand clades (Fig. 37.1A) (Paleo South American
and New Zealand clades: see below). The study also
con rmed the separation of Mairia into Mairia s.str.,
Polyarrhena Cass., and Zyrphelis Cass. (Nesom 1994a). The
latter are members of subtribe Homochrominae (below).
As with Printzia, Mairia was placed within the Southern
Hemisphere subtribe Hinterhuberinae in the Nesom and
obinson classi cation, a position not supported here.
At the next node in the phylogeny (Fig. 37.1A), Pteronia
L. and the Homochrominae form a polytomy with the re-
maining Astereae. In some trees, Pteronia segregates  rst,
while in others, it is sister to Homochrominae. Pteronia is
a large genus of shrubs with discoid heads. The majority
of genera in Homochrominae are radiate, although Felicia
Cass. (83 species; Fig. 37.2B), Amellus L. (12 spp.), Engleria
O. Ho m. (2 spp.), and Chrysocoma L. (20 spp.) have taxa
with both radiate and discoid heads. Two South African
genera, Heteromma Benth. and Engleria, were not included
in the present analysis. Heteromma is wholly discoid while
Engleria has one discoid and one radiate species. All spe-
cies in both genera are herbaceous perennials with the ex-
ception of one annual species of Engleria. The a nities of
these two taxa are currently unknown. Broader sampling
of the southern African taxa is needed to resolve the rela-
tionships between Pteronia, Homochrominae, and the few
genera that remain to be sampled.
Subtribe Homochromin ae had been ca lled Felici inae by
Nesom (1994g). He had included the African Felicia
group, the Asian Lachnophyllum Bunge group, and the
American-European Monoptilon Torr. and A. Gray
group in this subtribe. Nesom and obinson (2007)
restricted Homochrominae to the Felicia group, thus
making it strictly African, as was done by Grau (1973).
The current study con rms this circumscription, at
least insofar as the genera included are concerned (Fig.
37.1A). The Nesom (1994g) evolutionary tree showed
Homochrominae as nested within a polytomy of sub-
tribes mainly with Australasian and South American ele-
ments, but some European and North American as well;
such a nities are not supported here. Homochrominae
constitute one of the major radiations among the early
diverging lineages. A major di erence in the composi-
tion of Homochrominae between Nesom and obinson
(2007) and the current analysis is the inclusion of the
St. Helena endemic trees and shrubs Commidendron Lem.
and Melanodendron DC. In their classi cation, Nesom and
obinson left these genera unplaced. Nesom (1994g) had
placed them within woody Baccharidinae, though not-
ing the lack of comfortable  t in any subtribe. Noyes
and ieseberg (1999) noted the position of Commidendron
near Felicia and Amellus, while Eastwood et al. (2004)
showed a relationship of both to Felicia. Our phylogeny
con rms the relationship between the two groups, but
places Commidendron and Melanodendron as a clade sister
to the Felicia clade. It would appear that these arbores-
cent genera evolved from an ancestor common to the
two groups. Within the Felicia lineage, Felicia does not
appear to be monophyletic. Felicia uliginosa (J.M. Wood
& M.S. Evans) Grau and F. clavipilosa Grau group with
Nolletia Cass., well nested within a lineage that also in-
cludes Amellus, Poecilolepis Grau, and Chrysocoma. There
are two other unresolved Felicia lineages, the F. na-
maquana (Harv.) Merx. and F.  lifolia Burtt Davy lin-
eages. Polyarrhena and Zyrphelis appear related to the F.
lifolia lineage, but it is impossible to determine whether
the genera are sister to this lineage or whether Felicia is
paraphyletic to them. Felicia is most likely polyphyletic as
currently circumscribed and needs more study.
Brouillet, Lowrey, Urbatsch, Karaman-Castro, Sancho, Wagstaff and Semple592
The Madagascan genus Madagaster G.L. Nesom was re-
cently seg regated from Aster by Nesom (1993b), who placed
it in his southern Hemisphere subtribe Hinterhuberinae;
he considered these taxa unrelated to Aster, including
the African asters. As with Printzia and Mairia, our study
does not support a position in this subtribe (Fig. 37.1A).
Instead, it is one of many isolated African lineages along
the basal spine of the Astereae phylogeny. More genera
from Africa need to be included in phylogenetic analyses
before the position of Madagaster can be settled.
Conyza gouani Willd. is the only African Conyza Less.
that does not group with Grangeinae (see below) (Fig.
37.1A). Further study is required before its position can
be ascertained and it can be assigned to a genus of its own.
Nesom and obinson (2007) hinted at the polyphyletic
nature of Conyza. The morphology of this species appears
quite distinct from that of other African conyzas, and its
isolated position is therefore not surprising.
Nesom (1994h) and Nesom and obinson (2007) main-
tained the African asters within the mainly Asian Aster L.
s.str. but our results (Fig. 37.1A) do not support such a
relationship. As with several other African lineages, these
asters appear to be isolated. Additional study is needed to
better resolve their a nities.
Brouillet et al. (pers. comm.) have shown that subtribe
Grangeinae is a monophyletic group of African and south
Asian genera, and does not appear to include Australasian
or American elements (Fig. 37.1A). A major di erence
with the generic composition given by these authors is
the addition of the African conyzas, Psiadia Jacq., and
Welwitschiella O. Ho m. to the subtribe. Nesom and
obinson, however, mentioned the potential relatedness
of African conyzas to the subtribe. Earlier, Nesom (1994g)
had suggested a relationship of the African conyzas to
the Nidorella Cass. group of Grangeinae, as was proposed
by Wild (1969a, b). Our data support this relationship,
but only generally: Nidorella appears more closely related
to Grangea Adans. than to the conyzas in our analysis.
Nesom (1994g) had placed Psiadia in Baccharidinae, a
relation not supported here. It was left unplaced in the
Nesom and obinson classi cation. Finally, Welwitschiella,
a genus traditionally placed within Heliantheae s.l., also
belongs here (Brouillet et al. pers. comm.). This genus has
not been treated within Astereae before.
Fiz et al. (2002) studied the evolution of subtribe
Bellidinae s.str. (Fig. 37.2C) and showed it had relation-
ships with Bellidiastrum Cass. and the Galatella Cass. group
(Fig. 37.1A). The current analysis supports this view and
suggests that Bellidinae should be expanded to include
Bellidiastrum and the Galatella group, placed in Asterinae
by Nesom and obinson (2007) and often treated as Aster
species in the past. Furthermore, an African origin for
Bellidinae is suggested by their placement on the tree, in
a polytomy with mainly African Grangeinae.
As with Nannoglottis, Callistephus Cass. (Brouillet et al.
2001; Semple et al. 2002) occupies an isolated position
within the Astereae phylogeny (Fig. 37.1A–C), usu-
ally within the crown group polytomy. It never groups
with Aster s.str., in contradiction to its classi cation in
subtribe Asterinae (Nesom 1994g; Nesom and obinson
2007). Given that few Asiatic genera have been included
in molecular phylogenetic analyses, it is di cult to reach
further conclusions concerning this taxon.
Paleo South American clade
The Paleo South American clade (Fig. 37.1B) is placed in
the phylogeny between the South African Mairia lineage
(above) and the New Zealand clade (below). In more re-
stricted phylogenies (Brouillet et al., pers. comm.), it is
sometimes sister to the New Zealand clade. The inter-
relationships of these two clades remain to be ascertained.
The clade includes taxa placed within Hinterhuberinae by
Nesom and obinson (2007). Our phylogeny shows that
subtribe to be polyphyletic. The Paleo South American
clade consists of two sister subclades: the Chiliophyllum
Phil. nom. cons. and the Oritrophium (H.B.K.) Cuatrec.
The Oritrophium clade (Fig. 37.1B) includes rosette
herbs that are scapose (Oritrophium) or acaulescent (Novenia
S.E. Freire). They are adapted to the climatic and edaphic
conditions of the puna and páramo ecosystems, respectively
(Freire 1986; Torres et al. 1996). Oritrophium is distributed
in South America and Mexico. Because of its herbaceous
habit, it was considered a highly derived member of subtribe
Hinterhuberinae (Nesom 1993a). Its rosulate habit, scapose
in orescences, and staminate disk  orets de ne the genus.
ecently two new species of Oritrophium were described
from Mexico (Nesom 1992, 1998b). This is a remarkable
Fig. 37.1. Strict consensus tree of tribe Astereae based on a parsimony phylogenetic analysis of ITS sequence data. A (opposite
page) basal grade of mostly African lineages; B (p. 594) Paleo South American and New Zealand clades; C (p. 596) Australasian
lineages; D (p. 597) South American lineages; E (pp. 598–599) North American clade. Branches are colored according to geo-
graphic distribution, following the color code developed for this volume; Madagascar and St. Helena are color-coded distinctly
from tropical Africa (Fig. 37.1A). Numbers along branches are basic chromosome number (x) for all taxa subtended by the
branch, unless otherwise indicated; × on a branch indicates taxa for which chromosome numbers are lacking. Thin red lines to
the right of trees indicate groups found in the unpublished ML analysis (Brouillet et al., pers. comm.). Black lines and subtribal
names to the right of trees indicate subtribes according to Nesom and obinson (2007). Circled numbers to the right refer to
the groups discussed in text. For a complete biogeography tree of the entire Compositae see chapter 44.
Chapter 37: Astereae 593
Fig. 37.1A. Basal grade of
mostly African lineages.
Denekia capensis
Printzia polifolia
Nannoglottis ravida
Nannoglottis macrocarpa
Nannoglottis hieraciphylla
Nannoglottis latisquama
Nannoglottis delavayi
Nannoglottis carpesioides
Nannoglottis yuennanensis
Nannoglottis gynura
Mairia hirsuta
Pteronia camphorata
Pteronia uncinata
Pteronia glomerata
Pteronia incana
Melanodendron integrifolium
Commidendron rotundifolium
Commidendron spurium
Commidendron robustum
Commidendrum rugosum
Felicia minima
Felicia namaquana
Felicia aethiopica
Felicia aethiopica ecklonis
Polyarrhena reflexa
Felicia filifolia
Felicia echinata
Felicia linifolia
Felicia fruticosa
Zyrphelis microphylla
Zyrphelis decumbens
Aster capensi
Amellus tridactylu
Amellus microglossus
Amellus strigosus
Poeciliopsis ficoidea
Amellus? sp.
Felicia uliginosa
Felicia clavipilosa
Nolletia chrysocomoides
Chrysocoma ciliata
Chrysocoma sp. 1
Chrysocoma sp. 2
Madagaster madagascariensis
Madagaster mandrarensis
Conyza gouanii
Aster comptonii
Aster perfoliatu
Aster harveyanu
Conyza pyrrhopappa
Psiadia punctulata
Welwitschiella neriifolia
Conyza scabrida
Conyza sp.
Grangea maderaspatana
Nidorella polycephala
Nidorella resedifolia
Aster glehni
Galatella sedifolia
Tripolium vulgare
Galatella coriacea
Crinitaria linosyris
Bellidiastum michelii
Bellium crassifolium
Bellium bellidioides
Bellium minutum
Bellis atlantica
Bellis rotundifolia
Bellis pappulosa
Bellis longifolia
Bellis caerulescens
Bellis sylvestris
Bellis microcephala
Bellis azorica
Bellis bernardii
Bellis perennis
Bellis margaritaefolia
Bellis annua
Callistephus chinensis
ustralasian lineages
Paleo South American clade
New Zealand clade
South American lineages
North American clade
St. Helena
10, 9, 7
to 37.1B
to 37.1C–E
Southern Andes,
southern South America
Guiana Shield
North & central Andes
General South America
Eastern & central Asia
Southern &
southeastern Asia
Southern Africa
Tropical Africa
Northern Africa, Mediterranean,
Southern Europe
General Africa
North America
Central America, Caribbean
Australia, New Guinea,
New Caledonia
New Zealand
Pacific Islands
Widespread or ambiguous
Brouillet, Lowrey, Urbatsch, Karaman-Castro, Sancho, Wagstaff and Semple594
Fig. 37.1B. Paleo South
American and New
Zealand clades. For color
chart see Fig. 37.1A.
Oritrophium hieracioides
Oritrophium peruvianum
Novenia acaulis
Llerasia sp.
Llerasia lindeni
Llerasia macrocephala
Nardophyllum chiliotrichoides
Nardophyllum bryoides
Chiliophyllum andinum
Chiliotrichiopsis keideli
Lepidophyllum cupressiforme
Chiliophyllum fuegianum
Chiliotrichum diffusum
Chiliotrichum rosmarinifolium
Damnamenia vernicosa
Olearia colensoi
Olearia chathamica
Pleurophyllum criniferum
Pleurophyllum speciosum
Pleurophyllum hookeri
Olearia lyallii
Olearia semidentata
Olearia angustifolia
Olearia operina
Olearia furfuracea
Celmisia mackaui
Celmisia asteliifolia
Celmisia tomentella
Olearia velutina
Celmisia bellidioides
Celmisia lateralis
Olearia nummularifolia
Olearia albida
Olearia paniculata
Olearia cheesmanii
Olearia arborescens
Olearia rani
Pachystegia insignis
Olearia solandri
Olearia traversii
Olearia virgata
Olearia ledifolia
Olearia pannosa
Olearia argophylla
Olearia megalophylla
Olearia myrsinoides
Olearia chrysophylla
Olearia rosmarinifolia
Olearia oppositifolia
Olearia covenyi
Conyza gouanii
African Aster
Outgroup tribes
Callistephus chinensis
Australasian lineages
South American lineages
North American clade
to 37.1C–E
Paleo South American clade
New Zealand clade
Chapter 37: Astereae 595
nding because the genus has not been found in Central
America. Such a disjunct distribution may be attributed
to transport by birds, possibly from northern Venezuela
(Cuatrecasas 1997). Novenia, a monotypic genus, is distrib-
uted at high elevations in the Andes from Boliv ia to north-
ern Peru and Argentina. Novenia is highly reduced, with
rigid leaves with axillary clumps of tomentum, numerous
sessile, disciform capitula in the center of the rosette, with
reduced pistillate corollas and functionally staminate disk
orets, and multinerved achenes with a multiseriate pappus
of persistent barbellate bristles. These characters and the
number of chromosomes typical of Astereae (n = 9) justi-
ed its transfer from Inuleae (Freire 1986; Anderberg 1991)
to Hinterhuberinae (Nesom 1994g). The genus appears to
be derived from within Oritrophium; more data are needed
to evaluate the relationship of these two genera.
The Chiliophyllum clade (Fig. 37.1B) includes resin-
ous shrubby genera with usually white-tomentose leaves,
radiate capitula, paleate receptacles, and bisexual disk
orets (functionally staminate in Lepidophyllum Cass.):
Chi lio trichum Cass. (Fig. 37.2D), Chiliotrichiopsis Cabrera,
Nardophyllum Hook. & Arn., Lepidophyllum (Fig. 37.3C),
and Llerasia Triana. These are mainly restricted to the arid
puna region of southern Chile and Argentina (Patagonia
and Tierra del Fuego). Llerasia is restricted to the upper-
most montane forest of the Andes, below the páramo
zone, in Peru, Ecuador, and Colombia. The group is char-
acterized by a pappus of  attened bristles in one to several
series. In Nardophyllum, pappus bristles vary from  attened
to terete and widened apically, whereas pappus bristles in
Llerasia are terete and scabrous. The presence of receptacu-
lar paleae is another de ning feature for this group, but
receptacles vary from epaleate in Lepidophyllum and three
species of Nardophyllum, to few-paleate in Chiliotrichum, and
Llerasia, to fully paleate in Chiliotrichiopsis and Chiliophyllum
(Bonifacino and Sancho 2001). Variation in the number of
receptacular paleae was recorded at the population level in
Nardophyllum (Bonifacino 2005). Lepidophyllum is the only
genus in the group with opposite leaves. Chiliophyllum,
a genus of three species from Argentina and Chile, was
traditionally distinguished from Chiliotrichum based on
the color of ray  orets, and from Chiliotrichiopsis based on
the morphology of pappus bristles. In the present analy-
sis, Chiliophyllum appeared polyphyletic. Chiliophyllum
fuegianum O. Ho m. in Dusén groups with Chiliotrichum
and Lepidophyllum, and Chiliophyllum andinum Cabrera
with Nardophyllum and Chiliotrichiopsis, suggesting that
the de ning characteristics of the genus are homoplasious
(Karaman-Castro and Urbatsch, pers. comm.).
New Zealand clade
In this phylogeny (Fig. 37.1B), the New Zealand clade is
located between the Paleo South American clade and a
grade of African lineages (see above). A similar clade was
retrieved by Wagsta and Breitwieser (2002), who noted
that Astereae are the most diverse tribe of Asteraceae in
New Zealand, and by Cross et al. (2002) as their primary
clade II (but including Chiliotrichum); it also coincides with
the Olearia/Celmisia complex of Given and Gray (1986)
and the Celmisia group of Nesom (1994g). The New
Zealand clade is comprised of over 100 endemic species
that trad itionally have been placed in  ve genera: Celmisia
Cass. (Figs. 37.2E, 37.3D, E), Damnamenia Given, Olearia
Moench (Figs. 37.2F, 37.3F), Pachystegia Cheeseman (Figs.
37.2G, 37.3G), and Pleurophyllum Hook. f. (Fig. 37.3H)
(Wagsta and Breitwieser 2002). All have been classi-
ed within Hinterhuberinae by Nesom and obinson
(2007); this taxonomy is not supported here. Olearia
and Pachystegia are woody shrubs, whereas Celmisia,
Damnamenia, and Pleurophyllum are subshrubs or herbs that
arise from a woody base. These  ve genera are character-
ized by tailed anthers, tomentose leaves, and usually large
heads, features they share with the South American genus
Chiliotrichum (Cross et al. 2002). Many of the species are
showy ornamental shrubs that are widely cultivated in
New Zealand. A chromosome number of 2n = 108 is
the most commonly reported for members of the New
Zealand clade, but numbers as high as ca. 432 have been
reported for Olearia albida Hook. f. (Dawson 2000). The
New Zealand clade is not restricted to New Zealand since
members of Celmisia and Olearia s.l. independently dis-
persed to Australasia. Within the New Zealand clade, two
subclades can be recognized: the megaherb clade, includ-
ing Damnamenia, Pleurophyllum, and the macrocephalous
Olearia species (Drury 1968; Given 1973), and the Celmisia
clade, including Celmisia, Pachystegia, and the remaining
New Zealand Olearia species (including the divaricating
tree daisies; Heads 1998). Olearia is clearly polyphyletic
(Cross et al. 2002), hence well-supported clades in New
Zealand will likely be recognized as distinct genera.
Members of the megaherb clade (Fig. 37.1B), Dam-
na menia, Pleurophyllum (the megaherbs), and macrocepha-
lous olearias (Olearia spp.), were considered related by
Drury (1968) and Given (1973). All were included in
the current analysis (Fig. 37.1B). In Given’s taxonomic
scheme, Damnamenia and Pleurophyllum are closely re-
lated to Olearia colensoi Hook. f. and O. lyallii Hook. f.,
while the other olearias of the group (O. oporina Hook. f.,
O. chathamica Kirk, O. angustifolia Hook. f., and O. semi-
dentata Decne. ex Hook.) were related only through Cel-
misia; the Australian Pappochroma palucidola (S.J. Forbes)
G.L. Nesom (as Erigeron pappocroma Labill.) (Nesom
1994f, h, 1998a) was also considered tentatively related to
Celmisia but was not included in molecular analyses and
therefore cannot be evaluated (but see Pappochroma af. in
Australasian lineages below). These taxa are concentrated
in the subantarctic islands of New Zealand and Australia,
and in the highlands of New Zealand (Drury 1968; Given
Brouillet, Lowrey, Urbatsch, Karaman-Castro, Sancho, Wagstaff and Semple596
1973 provides a map of the taxa). Our phylogeny does not
support membership of Celmisia in this clade but all other
taxa identi ed by Given belong here, as was also noted by
Cross et al. (2002). Members of the clade are trees, shrubs
or woody-based herbs, with abaxially tomentose leaves,
racemose capitulescences or solitary capitula, large capit-
ula with usually purple disc  orets (yellow in O. oporina)
and tailed anthers with prominent apical appendages, and
long-villose achenes. Damnamenia is monotypic and based
upon the distinctive subantarctic D. vernicosa (Hook. f.)
(Given 1973). It was formerly considered as the sole mem-
ber of Celmisia subgenus Ionopsis, but was recently recog-
nized as a distinct genus. The distinctness of this taxon
from Celmisia is con rmed here. Pleurophyllum includes
three subantarctic island species, P. speciosum Hook. f.,
P. crini ferum Hook. f., and P. hookeri Buchanan, which
Fig. 37.1C. Australasian lineages.
For color chart see Fig. 37.1A.
Callistephus chinensis
to 37.1D, E
to 37.1A, B
Dichromochlamys dentatifolia
Ixiochlamys filicifolia
Ixiochlamys cuneifolia
Dimorphocoma minutula
Kippistia suaedifolia
Isoetopsis graminifolia 17
Elachanthus pusillus
Chondropyxis halophila
Minuria cunninghamii
Minuria sp.
Olearia elliptica elliptica
Olearia stuartii
Olearia glandulosa
Achnophora tatei
Tetramolopium vagan
Camptacra barbata
Camptacra gracilis
Olearia arguta
Lagenophora pumila
Pappochroma nitidum
Peripleura diffusa
Minuria macrorhiza
Tetramolopium pumilum
Tetramolopium alinae
Tetramolopium macrum
Tetramolopium mitiaroense
Tetramolopium arenarium
Tetramolopium remyi
Tetramolopium consanguineum
Tetramolopium rockii
Tetramolopium humile humile
Tetramolopium filiforme
Tetramolopium lepidum
Vittadinia pleurochaeta
Vittadinia pustulata
Vittadinia cerviculari
Vittadinia condyloide
Vittadinia dissecta
Vittadinia sulcata
Vittadinia eremaea
Vittadinia australis
Vittadinia humerata
Vittadinia megacephala
Vittadinia australasiaca
Vittadinia gracilis
Vittadinia cuneata
Vittadinia blackii
Peripleura obovata
Peripleura bicolor
Peripleura sericea
Peripleura hispidula
Tetramolopium sylvae
Ixiochlamys integerrimae
Remya kauaiensis
Olearia ferresii
Olearia flocktoniae
Olearia muelleri
Olearia calcarea
Olearia magniflora
8, 7,
6, 5,
Erodiophyllum acanthocephalum
Erodiophyllum elderi
Calotis inermis
Calotis xanthosoidea
Calotis dentex
Calotis cuneifolia
Calotis squamigera
Calotis hispidula
Calotis multicaulis
Calotis plumulifera
Calotis porphyroglossa
Calotis cuneata cuneata
Calotis scabiosifolia scabiosifolia
Calotis anthemoidea
Calotis scapigera
Calotis erinacea
Calotis cymbacantha
Calotis lappulacea
Calotis latiuscula
Brachyscome radicans
Brachyscome rigidula
Brachyscome dichromosomatica A1
Brachyscome breviscapis
Brachyscome lineariloba race B
Brachyscome tesquorum
Brachyscome ciliocarpa
Brachyscome angustifolia
Brachyscome exilis
Brachyscome parvula
Brachyscome leptocarpa
Brachyscome iberidifolia
Brachyscome decipiens
Brachyscome procumbens
Brachyscome muelleri
Brachyscome melanocarpa
Brachyscome cardiocarpa
Brachyscome humilis
Brachyscome smithwhitei
Brachyscome ciliaris
Brachyscome multifida multifida
Brachyscome campylocarpa
Brachyscome eriogona
Brachyscome goniocarpa
Brachyscome gracilis
Brachyscome tetrapterocarpa
Brachyscome curvicarpa
Brachyscome heterodonta
Brachyscome dentata
Brachyscome papillosa
Myriactis humilis
Keysseria maviensis
Olearia rudis
Olearia picrifolia
Rhynchospermum verticillatum
Aster taiwanensis
Aster savatieri
Aster diplostephioides
Aster alpinus alpinus
Aster alpinus vierhapperi
Aster tataricu
Aster viscidulus
Aster spathulifolius
Aster kantoensis
Aster hispidus
Aster yunnanensis
Aster ageratoides
Aster pinnatifidus
Aster iinumae
Aster holophyllus
Aster yakushensis
Aster amellus
Aster scaber
Aster miquelianus
Aster rugulosus
Aster dimorphophyllus
Aster komonoensis
Olearia ramosissima
Olearia floribunda
Olearia elliptica praeternissima
Olearia microphylla
Olearia passerinoides passerinoides
Olearia pimeleoides
Olearia ramulosa
Olearia teretifolia
Olearia ballii
Olearia astroloba
Olearia lasiophylla
Olearia nernstii
Olearia stellulata stellulata
Olearia stellulata lirata
Olearia phlogopappa
Olearia phlogopappa subrepanda
Olearia tomentosa
Olearia cordata
Olearia ciliata
9, 8, 7,
6, 5, 4,
3, 2
South American lineages
NorthAmerican clade
Chapter 37: Astereae 597
have purple or whitish-purple rays (Drury 1968). Drury
(1968) noted the similarity of the macrocephalous olearias
to Pleurophyllum, suggesting the inclusion of the former
into the latter; Given (1973) proposed to include O. colen-
soi and O. lyallii in Pleurophyllum and to create a new genus
for the others.
The genus Celmisia (Fig. 37.1B) is not fully re-
solved in our analysis. This may be due to the fact that
only about 20% of the taxa have been included so far
(S. Wagsta , pers. comm.). The clade shows a polytomy
comprised of an apparently isolated O. furfuracea Hook. f.,
a large Pachystegia clade, and two Celmisia lineages, one of
which is associated with the Olearia albida subclade (clade
F of Cross et al. 2002) and the New Guinean O. velutina
Koster (O. sp. WNG in Cross et al., 2002). The Olearia
groups recognized here coincide with those of the Cross
Fig. 37.1D. South
American lineages.
For color chart see
Fig. 37.1A.
ustralasian lineage
Podocoma hirsuta
Podocoma notobellidiastrum
Heterothalamus spartioides
Baccharis boliviensis
Myriactis panamensis
Diplostephium rupestre
Guynesomia scoparia
Callistephus chinensis
Baccharis sp. 1
Baccharis dracunculifolia
Baccharis neglecta
Baccharis sp. 2
Microgyne trifurcata
Sommerfeltia spinulosa
steropsis macrocephala
Inulopsis scaposa
rchibaccharis androgyna
rchibaccharis schiedeana
rchibaccharis asperifolia
Plagiocheilus bogotensis
Plagiocheilus solivaeformis
Laennecia sophiifolia
Laennecia schiedeana
Westoniella chirripoensis
Westoniella triunguifolia
Westoniella eriocephala
Westoniella kohkemperi
North American clade
Blakiella bartsiaefolia
Laestadia pinifolia
Laestadia costaricensis
Laestadia muscicola
Hinterhubera ericoides
Hinterhubera imbricata
Parastrephia teretiuscula
Parastrephia lepidophylla
Parastrephia quadrangularis
Parastrephia lucida
Parastrephia phylicaeformis
Hinterhubera adenopetala
Hinterhubera laseguei
Hinterhubera lanuginosa
Hinterhubera columbica
Floscaldasia hypsophila
Diplostephium ericoides
ztecaster matudae
ztecaster pyramidatus
to 37.1A
to 37.1E
to 37.1C
Brouillet, Lowrey, Urbatsch, Karaman-Castro, Sancho, Wagstaff and Semple598
Callistephus chinensis
Australasian lineages
South American lineages
Eucephalus elegans
Eucephalus paucicapitatus
Eucephalus glaucescens
Eucephalus gormanii
Eucephalus breweri
Eucephalus engelmanii
Eucephalus ledophyllus
Eucephalus brickellioides
Eucephalus vialis
Doellingeria infirma
Doellingeria umbellata
Doellingeria sericocarpoides
Egletes liebmannii
Ionactis stenomeres
Ionactis elegans
Ionactis caelestis
Ionactis linariifolia
Chaetopappa ericoides
Chaetopappa effusa
Chaetopappa bellioides
Monoptilon bellioides
Pentachaeta exilis
Pentachaeta aurea
Rigiopappus leptocladus
Tracyina rostrata
Ericameria zionis
Ericameria albida
Ericameria resinosa
Ericameria parryi
Ericameria palmeri
Ericameria ophidis
Ericameria nauseosa
Ericameria naus. consimilis
Ericameria martirensis
Ericameria linearifolia
Ericameria laricifolia
Ericameria juerezensis
Ericameria gilmanii
Ericameria discoidea linearis
Ericameria discoidea disc.
Ericameria cuneata
Ericameria crispa
Ericameria cooperi
Ericameria compacta
Ericameria brachylepis
Ericameria paniculata
Ericameria teretifolia
Ericameria arborescens
Ericameria parishii
Ericameria ericoides
Ericameria pinifolia
Ericameria fasciculata
Ericameria bloomeri
Ericameria greenei
Ericameria suffruticosa
Ericameria cervina
Ericameria lignumviridis
Ericameria watsonii
Ericameria obovata
to 37.1C, D
Geissolepis suaedifolia
P. asperifolia adenolepis
Pityopsis flexuosa
Pityopsis graminifolia
Pityopsis falcata
Pityopsis pinifolia
Bradburia pilosa
Bradburia hirtella
C. gossypina cruiseana
Chrysopsis mariana
Chrysopsis gossypina trichophylla
Chrysopsis gossypina hyssopifolia
Chrysopsis gossypina gossypina
Chrysopsis subulata
Chrysopsis dressei
Chrysopsis linearifolia
Chrysopsis godfreyi
Chrysopsis lanuginosa
Chrysopsis scabrella
Chrysopsis delaneyi
Chrysopsis highlandsensis
Noticastrum marginatum
Croptilon divaricatum
Croptilon rigidifolium
Heterotheca viscida
Heterotheca oregana compacta
Heterotheca oregana rudis
Heterotheca fulcrata subaxillaris
Heterotheca villosa
Heterotheca sessilifolia echioides
Heterotheca rutteri
Heterotheca horrida
Heterotheca camporum camporum
Heterotheca camp. glandulissiumum
Heterotheca zionensis
Heterotheca jonesii
Heterotheca shevockii
Erigeron lepidopodus
Erigeron rhizomatus
Erigeron neomexicanus
Erigeron eruptens
Erigeron podophyllus
Erigeron wislizeni
Erigeron inornatus
Erigeron compositus
Erigeron vagus
Erigeron barbellatus
Erigeron tener
Erigeron linearis
Erigeron ochroleucus
Erigeron aphanactis
Erigeron pumilus
Erigeron arenarioides
Erigeron tweedyi
Erigeron pygmaeus
Erigeron aliceae
Erigeron eatonii
Erigeron scoparioides
Aphanostephus ramosissimus
Aphanostephus skirrhobasis
Erigeron bellioides
Erigeron procumbens
Erigeron dryophyllus
Erigeron veracruzensis
Erigeron philadelphicus
Erigeron quercifolius
Erigeron argentatus
Erigeron utahensis
Erigeron speciosus
Erigeron pulchellus
Erigeron pinnatus
Erigeron pinnatisectus
Erigeron kachinensis
Erigeron coronarius
Erigeron caespitosus
Erigeron glabellus
Erigeron glaucus
Erigeron fraternus
Erigeron galotti
Erigeron formosissimus
Erigeron subtrinervis
Erigeron lonchophyllus
Erigeron cinereus
Erigeron humilis
Conyza apurensis
Erigeron leptorhizon
Conyza canadensis
Conyza ramosissima
Erigeron miyabeanus
Erigeron grandiflorus
Erigeron simplex
Erigeron uniflorus
Erigeron acris
Erigeron thunbergii
Erigeron borealis
Erigeron scopulinus
Erigeron leiomerus
Erigeron ursinus
Erigeron annuus
Erigeron tenuis
Erigeron strigosus
Erigeron karvinskianus
Erigeron bellidiastrum
Erigeron divergens
Erigeron religiosus
Apopyros warmingii
Erigeron maximus
Hysterionica jasionoides
Neja filiformis
Erigeron fernandezianus
Erigeron rosulatus
Erigeron flettii
Conyza chilensis
Conyza floribunda
Conyza bonariensis
Gundlachia riskindii
Gundlachia truncata
Gundlachia triantha
Gundlachia corymbosa
Gundlachia diffusa
Gymnosperma glutinosum
Amphiachyris dracunculoides
Xylovirgata pseudobaccharis
Medranoa parrasana
Neonesomia johnstonii
Neonesomia palmeri
Thurovia triflora
Chihuahuana purpusii
Bigelowia nudata
Bigelowia nutallii
Gutierrezia sarothrae
Gutierrezia texana
Euthamia leptocephala
Euthamia occidentalis
Euthamia graminifolia
Euthamia tenuifolia
Oclemena acuminata
Oclemena nemoralis
Oclemena reticulata
Ericameria nana
Fig. 37.1E. North American clade. Abbreviations: A. =
Astranthiinae; B. = Boltoniinae; C. = Chaetopappinae;
U. = Unplaced. For color chart see Fig. 37.1A.
Chapter 37: Astereae 599
Cuniculotinus gramineus
Sericocarpus asteroides
Sericocarpus tortifolius
Sericocarpus oregonensis
Sericocarpus linifolius
Stenotus pulvinatus
Stenotus lanuginosus
Stenotus acaulis
Stenotus armerioides
Nestotus macleanii
Nestotus stenophyllus
Columbiadoria hallii
Petradoria pumila
Toiyabea alpina
Chrysoma pauciflosculosa
Brintonia discoidea
Tonestus eximius
Tonestus peirsonii
Tonestus graniticus
Tonestus pygmaeus
Tonestus lyallii
Oreochrysum parryi
Lorandersonia microcephala
Lorandersonia pulchella
Lorandersonia spathulata
Lorandersonia baileyi
Lorandersonia salicina
Lorandersonia linifollia
Eastwoodia elegans
Solidago patula
Solidago bicolor
Solidago nitida
Solidago rigida
Solidago shortii
Solidago sempervirens
Solidago petiolaris
Solidago fistulosa
Solidago canadensis
Amphipappus fremontii spinos.
Acamptopappus shockleyi
Acamptopap. sphaerocephala
Hesperadoria scopulorum
Chrysothamnus molestus
Chrysothamnus stylosus
Chrysothamnus humilis
Chrysothamnus eremobius
Chrysothamnus greenei
Chrysothamnus depressus
Chrysothamnus vaseyi
Chrysothamnus visc. axillaris
Chrysothamnus visc. viscidiflorus
Oreostemma alpigenum haydenii
Oreostemma elatum
Oreostemma alpigenum andersonii
Herrickia glauca pulchra
Herrickia horrida
Herrickia glauca glauca
Herrickia wasatchensis
Herrickia kingii barnebyana
Herrickia kingii kingii
Eurybia paludosa
Eurybia radulina
Eurybia radula
Eurybia avita
Eurybia gracilis
Eurybia furcata
Eurybia chlorolepis
Eurybia divaricata
Eurybia sibirica
Eurybia eryngiifolia
Eurybia integrifolia
Triniteurybia aberrans
Machaeranthera tanacetifolia
Xylorhiza wrightii
Xylorhiza tortifolia
Oonopsis wardii
Oonopsis engelmannii
Arida turneri
Arida parviflora
Arida blepharophylla
Arida riparia
Dieteria canescens
Dieteria bigelovii
Xanthisma texanum drummondii
Xanthisma gracile
Xanthisma viscidum
Xanthisma spinulosum
Xanthisma stenolobum
Xanthisma blephariphyllum
Xanthisma gymnocephalum
Xanthisma coloradoense
Xanthisma crutchfieldii
Xanthisma rhizomatum
Haplopappus foliosus
Haplopappus macrocephalus
Haplopappus marginalis
Haplopappus glutinosus
Haplopappus paucidentatus
Xanthocephalum gymnospermoides
Isocoma menziesii vernonioides
Isocoma wrightii
Isocoma acradenia
Isocoma tenuisecta
Isocoma rusbyi
Grindelia lanceolata
Isocoma veneta
Grindelia ciliata
Grindelia nana
Rayjacksonia phyllocephala
Pyrrocoma lanceolata
Pyrrocoma clementis
Pyrrocoma crocea
Hazardia detonsa
Hazardia whytnei
Hazardia cana
Hazardia squarrosa
Hazardia squarrosa grindelioides
Benitoa occidentalis
Corethrogyne filaginifolia californica
Corethrogyne filaginifolia filaginifolia
Lessingia virgata
Lessingia arachnoidea
Lessingia ramulosa
Lessingia nemaclada
Lessingia nana
Lessingia micradenia micradenia
Lessingia micradenia glabrata
Lessingia leptoclada
Lessingia hololeuca
Lessingia glandulifera
Lessingia germanorum
Lessingia tenuis
Lessingia glandulifera peirsonii
Lessingia glandulifera tomentosa
Lessingia glandulifera glandulifera
Lessingia pectinata
6, 5, 4
4, 3, 2
ninae Solidagininae
Townsendia florifer
Astranthium integrifolium
Dichaetophora campestris
Batopilasia byei
Chloracantha spinosa
Boltonia asteroides
Boltonia diffusa
Canadanthus modestus
Ampelaster carolinianus
Almutaster pauciflorus
Psilactis brevilingulata
Psilactis odysseus
Psilactis asteroides
Psilactis boltoniae
Psilactis tenuis
Symphyotrichum trilineatum
Symphyotrichum chapmanii
Symphyotrichum urophyllum
Symphyotrichum texanum
Symphyotrichum drummondii
Symphyotrichum cordifolium
Symphyotrichum depauperatum
Symphyotrichum porteri
Symphyotrichum puniceum
Symphyotrichum parviceps
Symphyotrichum elliottii
Symphyotrichum dumosum
Symphyotrichum tradescantii
Symphyotrichum racemosum
Symphyotrichum attenuatum
Symphyotrichum foliaceum
Symphyotrichum subspicatum
Symphyotrichum turbinellum
Symphyotrichum boreale
Symphyotrichum tenuifolium
Symphyotrichum spathulatum
Symphyotrichum eulae
Symphyotrichum bracteolatum
Symphyotrichum hallii
Symphyotrichum eatonii
Symphyotrichum cusickii
Symphyotrichum frondosum
Symphyotrichum ciliatum
Symphyotrichum laurentianum
Symphyotrichum ascendens
Symphyotrichum pygmaeum
Symphyotrichum oblongifolium
Symphyotrichum novaeangliae
Symphyotrichum georgianum
Symphyotrichum sericeum
Symphyotrichum concolor
Symphyotrichum plumosum
Symphyotrichum ericoides
Symphyotrichum falcatum
Symphyotrichum vahlii