Compositae metatrees: the next generation
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Chapter 44
Compositae metatrees: the next generation
Vicki A. Funk, Arne A. Anderberg, Bruce G. Baldwin, Randall J. Bayer,
J. Mauricio Bonifacino, Ilse Breitwieser, Luc Brouillet, Rodrigo Carbajal, Raymund Chan,
Antonio X. P. Coutinho, Daniel J. Crawford, Jorge V. Crisci, Michael O. Dillon,
Susana E. Freire, Mercè Galbany-Casals, Núria Garcia-Jacas, Birgit Gemeinholzer, Michael
Gruenstaeudl, Hans V. Hansen, Sven Himmelreich, Joachim W. Kadereit, Mari Källersjö,
Vesna Karaman-Castro, Per Ola Karis, Liliana Katinas, Sterling C. Keeley, Norbert Kilian,
Rebecca T. Kimball, Timothy K. Lowrey, Johannes Lundberg, Robert J. McKenzie,
Mesfin Tadesse, Mark E. Mort, Bertil Nordenstam, Christoph Oberprieler, Santiago Ortiz,
Pieter B. Pelser, Christopher P. Randle, Harold Robinson, Nádia Roque, Gisela Sancho,
John C. Semple, Miguel Serrano, Tod F. Stuessy, Alfonso Susanna, Matthew Unwin,
Lowell Urbatsch, Estrella Urtubey, Joan Vallès, Robert Vogt, Steve Wagstaff, Josephine Ward
and Linda E. Watson
INTRODUCTION volumes listed the tribes mostly in the order of Bentham
1873a rather than beginning with Heliantheae, which
Constructing a large combined tree of Compositae, a Bentham thought was most primitive (Bentham 1873b).
‘metatree’ (also called ‘meta-supertree’ by Funk and The papers in the 1977 volumes did accept some changes
Specht 2007 and ‘megatree’ by R. Ree, pers. comm.) such as the recognition of Liabeae and the conclusion
allows one to examine the overall phylogenetic and bio- that Helenieae were not a ‘good’ group, both more or
geographic patterns of the family. The fi rst modern at- less accepted by Cronquist in 1977. However, most pro-
tempts to understand the family were by the authors in posed changes such as the new tribe Coreopsideae, etc.
Heywood et al. (1977) plus the paper by Cronquist (1977), were not accepted by the synantherological community.
which was initially intended to be in the Heywood Cronquist (1977) believed that the primitive characters
publication. Literature prior to 1977 has been discussed of the family were as follows (slightly modified): shrubby;
in detail in other chapters (for the early literature, see leaves opposite; inflorescence cymose; heads few, each
Chapter 1). In Cronquist’s 1977 paper he reaffirmed his with many florets; involucre leafy, several-seriate; recep-
agreement with Bentham’s 13-tribe classification of the tacle chaff y; ray florets present and fertile; disk florets
family and the concept that Heliantheae s.l. were the perfect and fertile; lobes of the disk corollas with well
primitive members (Cronquist 1955; Bentham 1973a, b). developed mid-vein; pappus chaff y, of five members; and
Cronquist (1977) pointed out that the Heywood et al. anthers connate, not tailed. Cronquist stated that the
748 Funk et al.
presence of ray florets may have predated the origin of has been some discussion on the pros and cons of the
Compositae, so that even discoid tribes might have had ‘supertree’ and ‘supermatrix’ methods (Steel et al. 2000;
a radiate ancestry. Gatesy et al. 2002; Bininda-Emonds et al. 2003), and
The acceptance of the modified Bentham system was both methods are compared with the metatree approach
not universal. There were at least two papers in the by Funk and Specht (2007). The metatree for this analysis
Heywood et al. volumes ( Jeff rey 1977; Skvarla 1977) was constructed in the following manner:
and two individuals who published elsewhere (Carlquist 1. A ‘base tree’ was formed from the phylogeny of
1966, 1976; Robinson 1981) who had reservations about Panero and Funk (2008) with a few alterations. The
the concept of “13 tribes rooted in the Heliantheae”. All most important change was the addition of taxa
of these dissenting authors observed that the data they from the Heliantheae Alliance. The Heliantheae
were generating did not support all of the above-listed Alliance section of the Panero and Funk tree
characteristics as primitive in the family. However, for (which had only a few taxa) was replaced with the
the most part, the synantherological community contin- branching pattern of the Heliantheae Alliance from
ued to use the Bentham classification. Baldwin (Baldwin et al. 2002; Chapter 41). Also,
Not too long after 1977, opinions began to change with some refi nements were made using the work of
the advent of cladistic methodology and molecular data. Ortiz (Chapters 18 and 19) and Ortiz et al. (Chapter
Jansen and Bremer and their collaborators (Bremer 1987, 17) for Carduoideae, and Funk and Chan (Chapter
1992, 1994; Jansen and Palmer 1987, 1988; Hansen 1991a, 23) for Cichorioideae. The base tree was reduced
b; Jansen et al. 1991a, b; Bremer and Jansen 1992; Jansen to a matrix using Brooks Parsimony Analysis
and Kim 1996; Bremer and Gustafsson 1997) reordered (BPA; Brooks 1982; Brooks and McLennan 2002),
Compositae by placing Barnadesiinae as the sister group of wherein any branching diagram can be reduced to
the family and placing Heliantheae (including Eupatorieae) a series of zeros and ones in a data matrix. We used
highly nested in the phylogeny of the family. MacClade to generate the data matrix (Maddison
Bremer’s cladistic analysis (1994) was the fi rst revi- and Maddison 2001). The data matrix was run in
sion of the whole family based on morphology since a tree program (PAUP 4.0b10; Swofford 2002) to
Bentham, and he recognized many of the problem areas check for errors. All trees have been “ladderized to
in the cladograms of the family and tribes, but the mor- the right” for consistency, although anyone familiar
phology did not generate enough data to resolve many of with cladistics will understand that the tree can be
the issues. Over ten years later Kadereit and Jeffrey (2007) “rotated” at any node. This feature is amply dem-
reordered the genera, tribes, and subfamilies within the onstrated by comparing the rooted tree (Fig. 44.1)
family based on morphology and molecular results, and and the unrooted tree (Fig. 44.2).
this work is now the standard reference for descriptions 2. The most recent (and available) tree for each clade
of the tribes and genera of the family. (see below) was reduced to a matrix (as above) and
This chapter seeks to link the most recent molecular these matrices were added to the original matrix.
trees together in a metatree framework (Funk and Specht Each time a new clade tree was added, the overall
2007) and to use that tree to provide a basis for under- analysis was re-run to insure an accurate replication
standing the systematics, evolution, and biogeography of of the newly added tree, as well as to confi rm that
the family. the addition did not result in topological changes
elsewhere in the metatree. It should be noted that
when a phylogeny for a tribe contained many taxa
MATERIALS AND METHODS from the same area in a monophyletic group or a
grade, these were often pruned to decrease the size
Construction of the metatree of the tree without subtracting any biogeographi-
The metatree for Compositae was developed using a com- cal information. For instance, the phylogeny of
pilation of trees. The name metatree was adopted for this Gnaphalieae contained a clade of 58 terminal taxa
type of tree because it is a “tree of trees”, one that is based all endemic to Australia; this clade was reduced to
on a fi xed ‘base tree’ topology (Funk and Specht 2007). 25 taxa.
This type of tree has also been called a meta-supertree or 3. A summary tree (Fig. 44.1) was produced in which
megatree (R. Ree, pers. comm.), and some authors refer each major clade was reduced to a single branch.
to it as a supertree. It is, however, neither a tree produced This tree also shows the phylogenetic position of
by a combined analysis of coded cladograms obtained critically placed taxa and is displayed as an unrooted
from individual datasets (classic ‘supertrees’) nor is it the tree in Fig. 44.2.
result of analyzing a dataset in which data from multiple See the section on optimization for an explanation of
datasets have been combined (‘supermatrix’ trees). There the biogeographic areas and how they were assigned.Chapter 44: Compositae metatrees: the next generation 749
Sources of the trees extending eastwards through Paraguay to Uruguay and
General references for this study were Bremer (1994), southern Brazil and down through Argentina to southern
Heywood (1993), Heywood et al., (1977), Hind (1996), Patagonia (Heywood 1993).
and Kadereit and Jeff rey (2007). Below, the origin of each Cassini, in his famous 1816 diagram (Chapter 41: Fig.
phylogeny on the metatree is discussed. 41.1), showed Calyceraceae and Campanulaceae to be
closely related to Compositae. Even though he did not
Outgroups have it in the diagram, he also thought Goodeniaceae
Lundberg (Chapter 10) examined the relationships among were close (see Chapter 1).
the families now contained in Asterales, including Com-
positae. His work indicated that Calyceraceae were the Compositae
sister group of Compositae (1st outgroup) and that Good- The base tree. — The basic structure of the tree was
eniaceae (2nd outgroup) were the sister group of the taken from Panero and Funk (2002, 2008) and Baldwin
Calyceraceae + Compositae clade. The next most closely (Baldwin et al. 2002; Chapter 41); see above for details.
related family is Menyanthaceae, and it is followed by a The trees in Panero and Funk (2008) contained exten-
clade containing Stylidiaceae, Alseuosmiaceae, Phellinac- sive sampling from the base of the tree, Mutisieae (sensu
eae, and Argophyllaceae. The distribution of these eight fam- Cabrera), three to ten genera representing all other tribes
ilies (Fig. 44.1) shows that the Compositae + Calyceraceae (including the Heliantheae Alliance), and many taxa that
clade is nested in a grade of Australasian taxa (Australia, had been “hard to place” in previous studies (includ-
New Guinea, New Caledonia, and New Zealand). Each ing Hecastocleis, Gymnarrhena, and Corymbium). The Panero
of these families is discussed below (listed in reverse order and Funk phylogeny was based on data from ten chlo-
of relatedness to Compositae). roplast gene regions (ndhF, trnL-trnF, matK, ndhD, rbcL,
Argophyllaceae. — Two genera with ca. twenty spe- rpoB, rpoC1, exon1, 23S-trnI, and ndhI). Relationships
cies that are distributed on Australia, Lord Howe Island, within tribes of the Heliantheae Alliance were taken from
New Caledonia, New Zealand, and Rapa Island. Baldwin et al. (2002) and Chapter 41 and were based
Phellinaceae. — One genus with eleven species, all of on data from the ITS region of rDNA. Modifications
which are found on New Caledonia. were made in Cichorioideae (based on Chapter 23) and
Alseuosmiaceae. — Five genera and ten species all in Carduoideae (based on Ortiz, Chapters 18 and 19; and
located on Australia, New Caledonia, New Guinea, and Ortiz et al. (Chapter 17).
New Zealand. Mutisieae s.l. sensu Cabrera (Chapter 12). — The
Stylidiaceae. — Six genera with 245 species found in tribe Mutisieae (sensu Cabrera) has 84 genera and ca.
Australia and New Zealand with a few species in East 900 species. The paraphyly of Mutisieae (sensu Cabrera)
Asia and South America. was suggested by morphological studies (Cabrera 1977;
Menyanthaceae. — Five genera with sixty species Hansen 1991b) as well as the fi rst molecular studies of
having an almost cosmopolitan distribution; however, the family. The subtribe Barnadesiinae was recognized
four of the five genera are found in Australia, and because as being the sister group to the rest of the family ( Jansen
the closely related taxa are found in the Australia–New and Palmer 1987, 1988; Bremer 1994; Kim and Jansen
Zealand–New Guinea–New Caledonia area, this family is 1995). Kim et al. (2002) showed that the remainder of
treated as having an Australasian distribution at its base. the tribe (sensu Cabrera) could not be supported as a
Goodeniaceae. — The second outgroup of Compos- monophyletic group. Most recently, Panero and Funk
itae is a moderate-sized family of herbs and some shrubs: (2002, 2008) published phylogenies based on molecular
Goodeniaceae (fourteen genera, over 400 species). The data from ten chloroplast regions that (1) confi rmed that
family is largely confi ned to Australia, particularly west- Mutisieae (sensu Cabrera) were paraphyletic, (2) identi-
ern Australia, with only a few species extending else- fied additional clades, and (3) elevated several groups to
where, mostly in the Pacific area (Gustafsson et al. 1996, tribal and subfamily levels. Except for Barnadesieae, the
1997). A recent study (Howarth et al. 2003) has shown phylogeny of Panero and Funk (2008) formed the base
that the base of the phylogeny of Goodeniaceae is in tree for Mutisieae (sensu Cabrera) with a few additions
Australia with dispersals by members of Scaevola into the from Kim et al. (2002) and Katinas et al. (2007).
Pacific area, coastal areas in southern Asia and Africa, and Barnadesieae (Chapter 13). — The subfamily Barn-
the east coast of the Americas. adesioideae (nine genera; 91 species) has one tribe, and
Calyceraceae. — The fi rst outgroup of Compositae, it is the sister group for the rest of Compositae. This
and therefore its sister group, is Calyceraceae, a small has been known since the seminal papers by Jansen and
family (six genera, ca. sixty species) of annual and pe- Palmer (1987, 1988) established the presence of a chlo-
rennial herbs. The family is entirely South American, roplast DNA inversion shared by the rest of the family,
being most abundant in the Andes south from Bolivia, but not by Barnadesieae or other flowering plants. The750 Funk et al.
MUT. WUNDER. CARDU. CICHORIOIDEAE
Outgroups Stifftieae Wun. Cichorieae
Hya.
Gongylolepis clade (29)
Stenopadus clade (30)
Cichorieae C4 (900+)
Cichorieae C5 (200+)
Alseuosmiaceae (10)
Leucomeris clade (3)
Tarchonantheae (13)
Menyanthaceae (60)
A-Gorteriinae (131+)
Goodeniaceae (440)
Warionia C1-3 (313)
Argophyllaceae (20)
A-Arctotidinae (76+)
Hecastocleideae (1)
Eremothamneae (3)
Hyaloseris clade (7)
Vernonieae (1500+)
Gymnarrheneae (1)
Dicomeae (75-100)
Nassauvieae (313)
Barnadesieae (91)
Calyceraceae (60)
Oldenburgieae (4)
Stylidiaceae (245)
Platycarpheae (3)
Onoserideae (52)
Gochnatieae (70)
Phellinaceae (11)
Distephanus (40)
Cardueae (2500)
Wunderlichia (5)
Mutisieae (254)
Hyalis clade (3)
Moquinieae (2)
Heterolepis (3)
Liabeae (174)
Pertyeae (70)
Stifftia (8)
Fig. 44.1. A summary tree based on the metatree (Figs. 44.3–44.7). The tribes or clades have been represented by one to four-
branches. The branches and internodes were colored according to the distribution of the taxon or the optimization of those
distributions. The numbers by the terminal taxa reflect the number of species in that clade. Note that some areas have been
combined (e.g., Mexico and North America) and that the red color in Vernonieae represents Tropical America. Subfamilies
that have more than one tribe are indicated on the summary tree in capital letters (see Chapter 11 for details). A = Arctotideae;
CARDU. = Carduoideae; Hya. = Hyalideae; MUT. = Mutisioideae; S = Senecioneae; Wun. = Wunderlichieae; WUNDER.
= Wunderlichioideae.Chapter 44: Compositae metatrees: the next generation 751
ASTEROIDEAE
Senecioneae
(3500)
S-Tussilagininae Grade
Anthemideae (1800)
Neurolaeneae (153)
Gnaphalieae (1240)
Coreopsideae (550)
Chaenactideae (29)
Eupatorieae (2200)
Heliantheae (1461)
Calenduleae (120)
Athroismeae (55)
Astereae (3080)
Abrotanella (20)
Helenieae (120)
S-Senecioninae
Corymbieae (9)
Doronicum (40)
Millerieae (380)
Tageteae (267)
Polymnieae (3)
Perityleae (84)
Madieae (203)
S-Othonninae
Feddeeae (1)
Inuleae (687)
Bahieae (83)
South America
Brazil
Guiana Shield
North & central Andes
Southern Andes, southern South America
General South America
North America
North America, Mexico
Central America, Caribbean
Eurasia
Eurasia, Europe
Eastern & central Asia
Africa
Southern Africa
Madagascar, tropical Africa
Northern Africa, Mediterranean, southern Europe
General Africa
Australia and the Pacific
Australia, N. Guinea, N. Caledonia, N. Zealand
Widespread or ambiguous
first phylogeny of this tribe was done by Gustafsson et al. congeneric Perdicium, which are found in Africa and
(2001), but it was not completely resolved. The phylogeny to a lesser extent in Asia, all Mutisieae (sensu Cabrera)
for the tribe was taken from Gruenstaeudl et al. (2009). It from Africa are no longer part of Mutisioideae (sensu
was based on DNA sequence data of nine chloroplast gene Panero and Funk) and are now in Carduoideae. Using
regions (atpI-atpH IGS, matK, psbA-trnH IGS, rbcL, partial ITS and ndhF sequence data, Ortiz and his collaborators
rpoC1 gene + intron, rps16-trnK IGS, partial trnK intron, (Chapters 17–19) have shown that these segregate African
trnL intron, trnL-trnF IGS), the nuclear ribosomal ITS Mutisieae form three (or four) distinct groups that are
region (ITS1, 5.8S, ITS2), recoded DNA insertions/dele- separated by striking morphological as well as molecular
tions, and selected morphological characters from previous differences. Currently, there are three tribes: Dicomeae,
investigations. In their analysis all genera were monophyl- Oldenburgieae, and Tarchonantheae. However, it is pos-
etic except for Dasyphyllum, which fell into two groups sible, but not yet certain, that the tribe Dicomeae may fall
reflecting the subgenera and their respective distributions into two distinct groups that are not sister taxa. In addi-
“east of the Andes” and “west of the Andes”. There are tion, there is still some ambiguity as to the relationships
two possible positions for Schlechtendalia, one of which is among some of the tribes.
basal for the tribe, and the other is more highly nested. The tribe Dicomeae contains seven African genera (ca.
The ambiguity of the position of Schlechtendalia does not 75–100 species) occurring in tropical and southern Africa
affect the biogeographic hypothesis for this tribe. and Madagascar with a minor presence in the Arabian
African Mutisieae (Chapters 17–19). — With the ex- Peninsula, India, and Pakistan. The tribe Tarchonantheae
ception of Gerbera and the closely related and sometimes contains two African genera (13 species) occurring752 Funk et al.
1. Cichorieae Eupatorieae mainly in tropical and southern Africa, and Madagascar,
2. Eremothamneae but it is also present on the Arabian Peninsula. The tribe
3. Moquineae Oldenburgieae has only the genus Oldenburgia (4 species),
4. Calenduleae
Perityleae which is endemic to the Cape Floristic Region of South
Madieae Africa.
Cardueae (Chapter 20). — Cardueae (thistles; 73 gen-
Millerieae
era, ca. 2500 species) are now known to be nested within
Heliantheae a paraphyletic Mutisieae (sensu Cabrera). This tribe is
Polymnieae
the sister group of the African Mutisieae clades. The
Neurolaeneae tribes Cardueae, Tarchonantheae, Oldenburgieae, and
Bahieae
Dicomeae form a monophyletic group that is now the
subfamily Carduoideae. The Cardueae tree used for the
Chaenactideae
Coreopsideae
metatree is based on matK, trnL-F, and ITS sequence data
Tageteae (Susanna et al. 2006).
Helenieae Cichorieae (Lactuceae; Chapter 24). — The phy-
Feddeeae logeny of the mainly north temperate dandelion tribe
Athroismeae Cichorieae (Lactuceae) has long been problematic. It has
93 genera arranged in eleven subtribes, but the number
Inuleae of species varies depending on one’s species concept. If
one excludes the problematic genera Hieracium, Pilosella,
Senecioneae 4 Astereae and Taraxacum, there are about 1400 species (Kilian et al.,
Chapter 24). The Cichorieae tree used in this study was
Abrotanella Anthemideae
Doronicum
provided by Gemeinholzer and her collaborators based
Gnaphalieae on recent molecular analyses of a large ITS dataset (428
A-Gorteriinae taxa of 83 genera; Gemeinholzer and Bachmann 2003;
Heterolepis
Kilian et al., Chapter 24; Gemeinholzer et al., unpub.).
Corymbieae
The analyses revealed the existence of five major clades,
A-Arctotidineae 2
with a total of eleven subclades, within the tribe.
The position of Gundelia (Gundelieae) as basal within
Cichorieae was suggested by Karis et al. (2001) based
Platycarpheae
Gymnarrheneae
on ndhF data, and this was supported by Panero and
Liabeae
1 Funk (2008), who also found Warionia to be at the base.
Distephanus 3 However, the current studies of Gemeinholzer and her
Pertyeae
Vernonieae collaborators comprising more basally branching taxa
Dicomeae
place the Northern African genus Warionia at the base
Oldenburgieae of Cichorieae with the Mediterranean Gundelia slightly
Tarchonantheae more highly nested. Since both are from the same bio-
Hecastocleideae Cardueae geographic area, the two different placements of Gundelia
and Warionia do not affect the biogeographic analysis.
Gochnatieae Arctotideae, Eremothamneae, Platycarpheae, and
Stenopadus clade
Heterolepis (Chapters 25, 26, 29, 31). — The tribe Arcto-
Wunderlichia
tideae (African Daisies) is a diverse and interesting group
(18 genera, 215 species). Recent molecular studies are am-
Leucomeris clade biguous as to the monophyly of this tribe, and some former
Hyalis clade members have been moved out of the tribe based on mor-
Gongylolepis clade Onoserideae
phology and/or molecular data. The positions of Heterolepis
Dinoseris clade Mutisieae (Funk and Karis, Chapter 31) and the tribe Eremothamneae
Stifftia Nassauvieae (2 genera, 3 species; Robinson and Funk, Chapter 26) vary
Barnadesieae depending on the data used in the analysis, and the new
tribe Platycarpheae (2 genera and 3 species) is most likely
Calyceraceae closely related to the Liabeae + Vernonieae clade (Funk et
Fig. 44.2. An unrooted representation of the summary tree. al., Chapter 29). Although Arctotideae cannot be unam-
The size of the circle indicates the number of species found biguously diagnosed, the two core subtribes are distinctive
in that clade. Colors are the same as in Fig. 44.1. based on morphology as well as molecular data (Funk etChapter 44: Compositae metatrees: the next generation 753
al. 2004; Karis et al., Chapter 25). Recently published The relationship of Senecioneae to other clades is un-
phylogenies using both chloroplast and nuclear DNA and certain. The tribe is variously positioned as (1) the sister
representing all of the genera (some with many species) group to the rest of Asteroideae, (2) the sister group
provided the structure for the trees (Funk and Chan 2008; to the Calenduleae + Gnaphalieae + Astereae + Anthemi
McKenzie and Barker 2008) and the relationships among deae clade, or, in the least likely scenario, (3) the sis-
the clades was taken from Funk et al. 2004 and Funk and ter group to the Inuleae + Athroismeae + Heliantheae
Chan, Chapter 23). Alliance clade. The support for its inclusion is stron-
Liabeae (Chapter 27). — Liabeae are a monophyletic gest for option 1, but the relatively short branches make
Neotropical tribe containing approximately 174 species its placement there tentative (see Pelser and Watson,
distributed in 17 genera and occupying a wide variety of Chapter 33). This ambiguity will not be resolved until
habitats throughout Mexico, Central America, the West more taxa and characters from both plastid and nuclear
Indies, and the Andes. The greatest diversity in the tribe markers are included in a tribal-level study of the sub-
is found in Peru, where no fewer than 14 genera and over family. At this time we are following the resolution fa-
70 species are represented. After a long history of moving vored by Panero and Funk (2008), which shows the
from tribe to tribe, the current members were brought Senecioneae in a polytomy with the clade formed by
together by Robinson (1983). A previous morphological the Inuleae + Athroismeae + Heliantheae Alliance and the
analysis resolved a northwestern Andean origin (Funk et clade containing Calenduleae + Gnaphalieae + Astereae +
al. 1996). The tree for our study was based on Dillon et Anthemideae. Doronicum and Abrotanella, the two addi-
al. (Chapter 27) and contains all the genera of the tribe tional taxa in this polytomy, are Senecioneae genera that
except the monotypic Bishopanthus, which is only known have been hard to place and may have to be excluded
from the type. Although the type was relatively recently from the tribe (Pelser et al. 2007).
collected, it is just a small piece of the original collection, Calenduleae (Chapter 35). — The placement of Ca len-
most of which was destroyed by one of the collectors. duleae as the sister taxon to the Gnaphalieae + Anthem-
Vernonieae and Moquinieae (Chapters 28, 30). — ideae + Astereae clade is based on the Panero and Funk
The tribe Vernonieae with 126 genera and 1500 spe- (2008) analysis as well as those by Kim and Jansen (1995)
cies has until recently had most of its species placed and Eldenäs et al. (1999). The sister group relation-
in the large and complicated genus Vernonia (ca. 1000; ship of Calenduleae to the other three tribes is strongly
Jones 1977; Keeley and Robinson, Chapter 28). The tribe supported in the Panero and Funk (2008) study, even
is widely distributed with centers of diversity in tropi- though the number of taxa sampled is small. The tribe
cal Africa and Madagascar, Brazil, and North America. Calenduleae has 12 genera with 120 species (Nordenstam
However, it has been the subject of recent revisions that 2007b), and most genera have distinct centers of distribu-
concentrated on recognizing monophyletic genera from tion in southern Africa; most of the species occur in the
within the 1000 species of the core genus Vernonia s.l. Cape Floristic Region. However, one genus, Calendula, is
(e.g., Robinson 1999), fi rst in the Americas and more re- found in northern Africa and the Mediterranean north to
cently in Africa and Asia. Vernonieae have recently been Central Europe and east into Turkey, Iraq, and Iran; but
examined by Keeley et al. (2007; Keeley and Robinson, it is nested in the higher portion of the tree and so does
Chapter 28) based on ndhF, trnL-trnF, and ITS sequence not affect the biogeographic pattern.
data. Their work supports the monophyly of the tribe and Gnaphalieae (Chapter 36). — Gnaphalieae are a mod-
the non-monophyly of Vernonia. However, in the analysis erately large tribe whose members were traditionally in-
of the subfamily Cichorioideae (Funk and Chan, Chapter cluded in the tribe Inuleae. It has only been recently
23), Distephanus had alternative placements: as the sister that the tribe has been shown to be isolated from the
group to the rest of Vernonieae, or unresolved at the base remainder of “old” Inuleae (Anderberg 1989, 1991). The
with the tribe Moquinieae (Robinson, Chapter 30). approximately 180–190 genera and ca. 1240 species of
Senecioneae (Chapter 34). — Senecioneae are the Gnaphalieae are most numerous in the southern hemi-
largest tribe with over 150 genera containing 3500 species sphere, with strong centers of diversity in southern Africa,
(Nordenstam 2007a) and they have a global distribution. Australia, and South America (Anderberg 1991; Bayer et
Pelser et al. (2007), recently published a phylogenetic anal- al. 2007). The tree for this study was provided by Bayer
ysis of the tribe based on ITS data that, while unresolved and his collaborators (Ward et al., Chapter 36) and it is
at the base, showed several well supported clades. The based on chloroplast DNA sequences for matK, the trnL
genus Senecio, which contained the majority of the species intron, and the trnL-trnF intergenic spacer. The principal
of the tribe, was shown to be non-monophyletic, and the improvement of this tree over previously published DNA
authors indicated that revisions of the generic boundaries sequence phylogenies for Gnaphalieae is that it includes
that are needed to achieve monophyletic groups are com- a broad sampling of genera from Africa and Australasia
pleted or in progress. together with some from other continents.754 Funk et al.
Astereae (Chapter 37). — With 170 genera, ca. 3000 The tree for this clade in the metatree was formed by
species, and a worldwide distribution, Astereae are the using the Baldwin treatment of the Heliantheae Alliance
second largest tribe after Senecioneae. It has centers of (Chapter 41), the Coreopsideae treatment of Crawford et
diversity in southwestern North America, the Andes, al. (Chapter 42), and the Funk et al. paper (2005). The
South Africa, Australia, and New Zealand. The tree branching within Eupatorieae was taken from Robinson
presented in this book (Brouillet et al., Chapter 37) is the et al. (Chapter 43).
fi rst global, molecular phylogenetic analysis of the tribe. The tree for the Heliantheae Alliance section of the
It is based on ITS sequence data and shows that interrela- family contains 160 out of ca. 460 genera and so rep-
tionships among genera are better reflected by geographic resents about 35% of the generic diversity of this clade.
origin than by the current classification. This is the lowest percentage for any clade on the meta-
Anthemideae (Chapter 38). — The tribe Anthemideae tree, however the poor representation is found primar-
is composed of 111 genera and ca. 1800 species with ily in three tribes, Eupatorieae (the tree has 25 genera
main concentrations of species in southern Africa, the represented out of a total of 182; there are 2200 species),
Mediterranean region, and Central Asia. The phylog- Heliantheae (6 out of 113 genera were represented; there
eny for the metatree was generated using data from are 1461 species), and Millerieae (3 genera out of 36
two recent publications that used ndhF (Watson et al. were represented; there are 380 species). When totaled
2000; Himmelreich et al. 2008) and one that used ITS together, these three tribes are represented by only about
(Oberprieler et al. 2007). 10% of the generic diversity within them. The other ten
Inuleae and Plucheeae (Chapter 39). — Plucheeae are tribes in the Alliance are much better represented, some
now known to be nested within Inuleae, and so they are at or close to 100% (see below). Because the members of
recognized as a single tribe with about 66 genera and ca. former Helenieae form the basal grade, the under-repre-
700 species (Anderberg and Eldenäs 2007). The tree for sentation of three of the more highly nested groups does
this study was provided by Anderberg and his collabora- not present an obstacle to the biogeographic analysis, al-
tors (Anderberg et al. 2005) based on ndhF data. Inuleae though it does give an under-estimate of the importance
are a mainly Eurasian and east and southern African of the northern and central Andes.
tribe, but some genera (e.g., Pluchea) have a worldwide The tribe Heliantheae s.l. was broken up by Baldwin
distribution. et al. (2002) and by Panero and Funk (2002) when
Athroismeae (Chapter 40). — The tribe Athroismeae Eupatorieae were found to be nested within what is now
is the sister group to the rest of the large and diverse referred to as the Heliantheae Alliance (Fig. 44.1). Most
clade that is the Heliantheae Alliance. The five genera of the new tribes, however, were actually not new and
(only two were included in Panero and Funk 2008) and had been described previously by others but not picked
55 species in Athroismeae are centered in eastern tropical up by the synantherological community. In fact, only
Africa and were in Inuleae until moved to Heliantheae three of the tribes recognized by Baldwin needed to be
s.l. (Eriksson 1991). described as new (Baldwin et al. 2002): Bahieae (17 out
Heliantheae Alliance (including Eupatorieae) (Chap- of 20 genera were represented in the analysis; there are
ters 41–43). — The tribe Eupatorieae is nested in the 83 species), Chaenactideae (all 3 genera were represented;
Heliantheae Alliance, and former Heliantheae s.l. have 29 species), and Perityleae (4 out of 7 genera were rep-
been reorganized into twelve tribes (Baldwin et al. 2002; resented; 84 species) (see Funk et al., Chapter 11). Other
Panero and Funk 2002; Cariaga et al. 2008). Bremer tribes in the Heliantheae Alliance (not mentioned above)
(1994) divided this part of the family into three groups, include: Coreopsideae (21 genera out of 30 were repre-
Helenieae (including Athroismeae), Heliantheae, and sented; 550 species), Helenieae (all 13 genera were repre-
Eupatorieae, but recognized that the groups would need sented; 120 species), Madieae (35 genera were represented
to be re-arranged once additional information was avail- out of 36; 203 species), Neurolaeneae (1 out of 5 genera
able. The studies of both Baldwin et al. (2002) and Panero was represented; 153 species), Polymnieae (the only genus
and Funk (2002) showed Helenieae and Heliantheae of was represented; 3 species), and Tageteae (17 out of 32
Bremer to be non-monophyletic, and they described ad- genera were represented; 267 species).
ditional tribes where needed. More recently, Cariaga et
al. (2008) published a treatment of the problem genus Area optimization analysis using parsimony
Feddea based on ndhF sequence data. As part of their study The terminal branches of the metatree were colored
the genus was placed in a new tribe by itself, Feddeeae, based on the distribution of each terminal taxon; taxa
located as the sister group of the “rest” of the Heliantheae that span more than one area have multiple colors (Figs.
Alliance (minus Athroismeae). The inclusion of the tribes 44.1–44.7). The internode distributions were mapped
Feddeeae and Eupatorieae in the Heliantheae Alliance onto the metatree using the Farris double pass method
brings the total number of tribes in the Alliance to 13. (1970). The results of the mapping were checked usingChapter 44: Compositae metatrees: the next generation 755
the PAUP ‘Acctran’ option (Swofford 2002). These tech- family in montane areas. More recently, Bremer (1992,
niques provided the hypothesized distributions at deep 1994) developed a method he called ‘Ancestral Areas
branches and nodes. Analysis’ and came to the conclusion that the family
Following the theory that bold hypotheses are better originated in “South America and the Pacific”. DeVore
than weak ones (courtesy of Popper), equivocal situa- and Stuessy (1995) suggested that the family originated
tions were resolved when possible to present the most in southern South America, which was re-emphasized
predictive estimate of the biogeographic history. In a by Bremer and Gustafsson (1997). Graham (1996) sum-
few instances there were equivocal resolutions which marized the fossils for the family but had wide estimates
were left black, or if the two areas were contained in a of the age of some of the pollen. Other than these efforts,
single continent, they were coded for that continent (e.g., little attention has been paid to this topic. Perhaps the
general Africa). In essence, we created an ‘area metatree’ size of the family, its global distribution, the lack of mac-
as opposed to an ‘area cladogram’. In the summary tree rofossils and paucity of discriminating characters in fossil
and unrooted tree (Figs. 44.1, 44.2), some of the biogeo- pollen, and the lack of an agreed upon phylogeny have
graphic areas were combined (e.g., North America was restricted attempts to understand its history.
combined with Mexico). The meta showing the overall phylogeny of Compositae
allows us to use information from the most recent avail-
able molecular phylogenies to look at the family as a
RESULTS AND DISCUSSION whole and to try to discern its origin and history. It is also
an excellent method for determining critical areas of the
The first supertree (= metatree) for Compositae was pub- tree for future work (Funk and Specht 2007).
lished in 2005 (Funk et al.), and since then there has been
considerable progress in the reconstruction of evolution- The metatree and its sections
ary relationships in many clades. In fact, we now have In order to more easily discuss the tree it has been bro-
robust phylogenies for most of the clades in the family. ken into sections. Section 1 (Fig. 44.3) covers the Basal
Descriptions and diagnostic characters for all of the tribes Grade, from the outgroups through monotypic Gym-
and critical clades are found in Chapter 11. Without a narrheneae. Section 2 (Fig. 44.4) covers the large subfamily
doubt the most substantial progress has been made in the Cichorioideae. Section 3 (Fig. 44.5) covers Corymbieae,
large and complicated Astereae, Cichorieae, Senecioneae, Senecioneae, Calenduleae, and Gnaphalieae; Section 4
and Vernonieae tribes, all of which were problematic in (Fig. 44.6) Anthemideae and Astereae; and fi nally, Section
the 2005 publication (Funk et al. 2005) but now have 5 (Fig. 44.7) Inuleae, Athroismeae, and the Heliantheae
their first comprehensive molecular phylogenetic hypoth- Alliance (including the Eupatorieae). Figure 44.8 has
eses (Keeley et al. 2007; Pelser et al. 2007; Brouillet et al., some of the proposed ages of the clades and Figs. 44.9
Chapter 37; Kilian et al., Chapter 24; and other references and 44.10 show some of the morphological variation.
in the corresponding chapters). For the fi rst time within Since we have no macrofossil data, the following dis-
these tribes we have a fairly good idea of what the basal cussion is based on extant taxa.
groups are and where different clades are found, and we Section 1, Basal Grade (Figs. 44.3, 44.9A–D). —
know that the large genera with global distributions are Except for Calyceraceae (the sister group of Compositae),
not monophyletic. the most closely related families to Compositae are
Considering the entire metatree, most of the tradi- found in Australia, New Zealand, New Guinea, and
tional thirteen tribes were found to be monophyletic New Caledonia (purple lines; Fig. 44.3). The members of
or could easily be made monophyletic with only a few Calyceraceae are from southern South America.
rearrangements. The big exceptions to this are Mutisieae The fi rst split within Compositae is between the sub-
(sensu Cabrera) and the Heliantheae Alliance, both were family Barnadesioideae and the remainder of the family
broken up into many groups. The genera that were once (Fig. 44.3). Gustafsson et al. (2001) and Stuessy et al.
placed in Mutisieae by Cabrera or others are now in four- (Chapter 13) examined the biogeography and concluded
teen tribes, Helenieae are in seven, and Heliantheae are that the Barnadesioideae clade has its origin in southern
in six (including Feddeeae). South America; this is confi rmed by our analysis. In the
For such a large and interesting family, relatively little sister group of Barnadesioideae the relationships among
has been published on its geographic origin and diversi- the basal groups are largely unresolved and are shown
fication since Bentham (1873b). Bentham (1873b), Small as a trichotomy (Fig. 44.3). However, this part of the
(1919), Raven and Axelrod (1974), and Turner (1977) all tree could have been shown as a polytomy containing
believed that Compositae had their origin in the north- four or even five clades because support for monophyly
west portion of South America, in the Andes. Rzedowski of the subfamily Wunderlichioideae is not consistently
(1972) and Hu (1958) pointed out the high diversity of the strong, nor is its phylogenetic position; this ambiguity is756 Funk et al.
Mutisieae Wund.
Barnadesieae
Outgroups Hyalideae Gochnatieae
Onoser. Nassauvieae
Schlechtendalia-1
Schlechtendalia-2
Stifftieae
Adenocaulon bic
Adenocaulon chi
Alseuosmiaceae
Menyanthaceae
Argophyllaceae
Dasyphyllum-D
Stomatochaeta
Dasyphyllum-A
Goodeniaceae
Richterago am
Cnicothamnus
Richterago an
Aphyllocladus
Calyceraceae
Chaetanthera
Lophopappus
Ianthopappus
Brachyclados
Gochnatia hy
Gongylolepis
Wunderlichia
Phellinaceae
Urmenenetia
Doniophyton
Gochnatia hi
Polyachyrus
Stylidiaceae
Chimantaea
Stenopadus
Pachylaena
Chuquiraga
Leucomeris
Barnadesia
Gerbera se
Dolicasium
Trichocline
Panphalea
Gerbera pi
Duseniella
Hyaloseris
Nassauvia
Moscharia
Cyclolepis
Leucheria
Fulcaldea
Lycoseris
Chaptalia
Leibnitzia
Dinoseris
Arnaldoa
Onoseris
Huarpea
Triptilion
Duidaea
Proustia
Acourtia
Perezia
Nouelia
Mutisia
Jungia
Stifftia
Plazia
Hyalis
Trixis
Fig. 44.3. The metatree of Compositae has been broken up into five figures with two to three parts for each figure. The original
trees are from the various chapters in this volume, but some taxa with redundant distributions have been pruned from the tree to
save space. Figure 44.3 covers the Basal Grade of the family and includes the outgroups through Gymnarrheneae, including this-
tles (some of the internodes have been compressed). All outgroups except for the sister group are Australasian. The extant taxa
from the sister group of the family, Calyceraceae, along with those from the basal grade of Compositae have a southern South
American origin. For subfamily groups see Chapter 11, for color chart see Fig. 44.7. Gy. = Gymnarrheneae; H. = Hecastocleideae;
O. = Oldenburgieae; Onoser. = Onoserideae; Perty. = Pertyeae; Tar. = Tarchonantheae; Wund. = Wunderlichieae.Chapter 44: Compositae metatrees: the next generation 757
O. Dicomeae
H. Cardueae
Tar. Gy.
Centaurothamnus
Perty.
Xeranthemum an
Erythrocephalum
Hecastocleideae
Schmalhausenia
Xeranthemum in
Gymnarrheneae
Rhaponticoides
Amphoricarpos
Tarchonanthus
Lamyropappus
Cheirolophus
Rhaponticum
Pasaccardoa
Cardopatium
Ainsliaea ma
Cousiniopsis
Brachylaena
Schischkinia
Centaurea-1
Centaurea-2
Crocodylium
Echinops pe
Atractylodes
Ainsliaea ap
Oldenburgia
Oligochaeta
Echinops or
Onopordum
Tugarinovia
Ptilostemon
Stizolophus
Psephellus
Carthamus
Macledium
Saussurea
Staehelina
Carlina gu
Acroptilon
Galactites
Chardinia
Pleiotaxis
Myripnois
Carlina la
Tyrimnus
Atractylis
Serratula
Cousinia
Cloiselia
Berardia
Carduus
Synurus
Silybum
Siebera
Dicoma
Alfredia
Arctium
Cirsium
Zoegea
Jurinea
Olgaea
Cynara
Pertya
p. 758
General Africa
indicated in Fig. 44.3 by a dotted line. However, many of in Cuba. It is clear from the optimization that the extant
the main clades basal to the clade formed by Hecastocleis taxa at the base of the Compositae metatree have their
and its sister group are consistently resolved as having a origin in southern South America.
southern South American origin, with the exception of The internode between the southern South American
the tribe Wunderlichieae whose members are found in grade and the beginning of the African radiation (labeled
the Guiana Shield and Brazil. The large Mutisioideae “General Africa” in Fig. 44.3) is left unresolved as to
clade (composed of the tribes Mutisieae, Nassauvieae, and origin because there are no areas shared among the three
Onoserideae) contains mostly southern South American (South American base, African radiation, and the North
taxa, but it also contains Gerbera from tropical and south- American genus Hecastocleis). A species level analysis of
ern Africa and Asia, North America taxa (e.g., Acourtia), the tribe Gochnatieae (4–5 genera) is underway (Sancho
and Leibnitzia from Asia and Mexico. Hyalideae have et al., pers. comm.) and its relationships to Hecastocleis
two clades, one from Asia and one from southern South may provide some insight into the problem, because one
America. Gochnatieae contain genera mainly from south- of the genera (Gochnatia) is found in South America, the
ern South America and Brazil, but there is also a radiation West Indies, and North America.758 Funk et al.
Cichorieae
Arctotideae-Arct
Er. Arctotidieae-Gort
Marshalljohnstonia
He.
Haplocarpha s.str.
Acanthocephalus
Haplocarpha sca
Munzothamnus
Eremothamnus
Schlagintweitia
Chaetadelpha
Crepidiastrum
Heterorhachis
Lagoseriopsis
Hoplophyllum
Syncalathium
Pterachaenia
Arctotis s.str.
Urospermum
Pleiacanthus
Glyptopleura
Prenanthella
Cymbonotus
Hypochaeris
Rhagadiolus
Tragopogon
Catananche
Rothmaleria
Lygodesmia
Garhadiolus
Rafinesquia
Arctotis ano
Nothocalais
Berkheya-4
Berkheya-2
Berkheya-3
Scorzonera
Prenanthes
Heterolepis
Tourneuxia
Geropogon
Taraxacum
Arctotis lan
Lasiospora
Uropappus
Arctotheca
Leontodon
Hedypnois
Microseris
Hieracium
Cichorium
Dymondia
Scolymus
Hispidella
Notoseris
Hololeion
Hirpicium
Gundelia
Warionia
Andryala
Cicerbita
Aposeris
Launaea
Sonchus
Hyoseris
Cullumia
Cuspidia
Gazania
Nabalus
Youngia
Gorteria
Picrosia
Phitosia
Lactuca
Didelta
Crepis
Tolpis
Krigia
Picris
p. 760
p. 757
Fig. 44.4. Monotypic Cichorioideae (internodes have been compressed). For subfamily groups see Chapter 11, for color chart see
Fig. 44.7. Di. = Distephanus; Er. = Eremothamneae; He. = Hecastocleis; Mo. = Moquinieae; Pl. = Platycarpheae.
The largest clade of the basal grade contains the sub- for the sister group relationship between Oldenburgieae
family Carduoideae (Tarchonantheae, Oldenburgieae, and Tarchonantheae. The thistles (Cardueae) are mono-
Dicomeae, Cardueae; Fig. 44.3); this is followed on the phyletic and show a Mediterranean–northern African ra-
metatree by the Pertyeae (Asia) and Gymnarrheneae diation with numerous incursions into Eurasia and Asia.
(northern Africa). At the base of Carduoideae are several The combination of the Mediterranean–northern African
former members of Mutisieae from southern and tropi- base of the thistles and the tropical and southern African
cal Africa (African Mutisieae). The relationships of these Tarchonantheae, Oldenburgieae, and Dicomeae give a
clades to one another are unresolved at this time, except ‘general Africa’ base to this clade.Chapter 44: Compositae metatrees: the next generation 759
Vernonieae
Liabeae
Di.
Mo.
Stramentopappus
Gymnanthemum
Pl.
Hesperomannia
Chiantopappus
Lessingianthus
Pseudonoseris
Vernonanthura
Munnozia cam
Vernoniastrum
Paranephelius
Heterocypsela
Chrysactinium
Ferreyranthus
Elephantopus
Centratherum
Eirmocephala
Baccharoides
Baccharoides
Pseudostifftia
Cabobanthus
Vernonia ang
Munnozia gig
Parapolydora
Sinclairiopsis
Lepidaploa-1
Lepidaploa-2
Munnozia wil
Hilliardiella-2
Hilliardiella-1
Vernonia pro
Munnozia lyr
Chrysolaena
Vernonia tax
Vernonia bal
Vernonia fas
Distephanus
Vernonia ab
Vernonia bu
Centrapalus
Eremanthus
Microliabum
Vernonia su
Platycarpha
Bothriocline
Strobocalyx
Critoniopsis
Moquinieae
Vernonia br
Philoglossa
Brachythrix
Tarlmounia
Muschleria
Orbivestus
Cacosmia
Gorceixia
Eremosis
Albertinia
Sampera
Sinclairia
Sipolesia
Leiboldia
Stokesia
Dillandia
Oligactis
Chresta
Liabum
Ethulia
Linzia
Erato
Tropical
America
The sister group of Carduoideae is the remainder of that paper, it was thought that the biogeographic pathways
the family (Pertyeae, Gymnarrheneae, Cichorioideae, of Cichorioideae would lead from the Mediterranean via
Corymbieae, and Asteroideae) all of which, except for Eurasia to Asia and across to North America but it seems
Pertyeae (Asia), presumably originated in Africa. The fi rst that the Asian and North American taxa are separately
group to split off is Pertyeae followed by Gymnarrheneae derived from Mediterranean clades.
(Northern Africa) followed by Cichorioideae. At the base of the rest of the subfamily Cichorioideae
Section 2, subfamily Cichorioideae (Figs. 44.4, 44.9E, there are five clades containing members of the former
F). — This large clade contains six tribes: Cichorieae Arctotideae: two are subtribes of that tribe (Arctotidinae
(Fig. 44.4; also referred to as Lactuceae) is the sister and Gorteriinae), two are now recognized at the tribal
group to the remainder. This tribe has a Mediterranean– level (Eremothamneae and Platycarpheae), and one is an
northern African base with independent radiations in unplaced genus (Heterolepis). All are from southern Africa
North America and Asia. Interestingly, the main North (Fig. 44.4) and are prominent members of the Cape Floral
American clade of Cichorieae is not nested within the Region, which is the subject of intense conservation in-
Asian radiation as was predicted (Funk et al. 2005). In terest. Because all of the basal taxa in each subtribe are in760 Funk et al.
Senecioneae
Co.
Senecio m-w grp
Crassocephalum
Senecio saxatilis
Dendrophorbium
Senecio mel grp
Senecio stig grp
Charadranaetes
Ekmaniopappus
Cremanthodium
Senecio arnaldii
Cissampelopsis
Senecio gla grp
Senecio pin grp
Dendrosenecio
Lepidospartum
Blennosperma
Barkleyanthus
Senecio otites
Nordenstamia
Senecio s.str.
Robinsonecio
Gymnodiscus
Austrosynotis
Dolichoglottis
Arnoglossum
Paragynoxys
Chersodoma
Brachyglottis
Parasenecio
Adenostyles
Sinosenecio
Steirodiscus
Lopholaena
Antillanthus
Endocellion
Corymbium
Abrotanella
Papuacalia
Urostemon
Doronicum
Homogyne
Othonna-1
Othonna-2
Pentacalia
Elekmania
Crocidium
Monticalia
Dorobaea
Jacobaea
Mattfeldia
Tussilago
Acrisione
Petasites
Sinacalia
Cineraria
Werneria
Dauresia
Pericallis
Gynoxys
Bolandia
Faujasia
Hubertia
Rainiera
Roldana
Euryops
Packera
Oldfeltia
Iranecio
Ischnea
Gynura
Kleinia
Emilia
Hertia
Luina
Australasia
& Pacific
Sub-Saharan
Africa
p. 758
Fig. 44.5. Corymbieae, Senecioneae, Calenduleae, and Gnaphalieae. For subfamily groups see Chapter 11, for color chart see
Fig. 44.7. Co. = Corymbieae.
southern Africa, the lack of evidence for the monophyly African clades (Fig. 44.4). Liabeae are predominantly
of Arctotideae does not affect the biogeographic hypoth- central Andean and the tribe is believed to have origi-
eses produced in this study. nated in northern Peru and southern Ecuador with small
The tribe Platycarpheae (southern Africa) is the sis- incursions into Central America and radiations in Mexico
ter taxon of the Liabeae + Vernonieae clade (including (Sinclairia) and the Caribbean (Liabum). The basal branches
Distephanus and Moquinieae) but without strong support. of Vernonieae are from the area we have designated as
This clade is nested in a grade formed by the southern ‘tropical Africa and Madagascar’. New to the analysis isChapter 44: Compositae metatrees: the next generation 761
Calenduleae Gnaphalieae
Pseudognaphalium
Chrysanthemoides
Osteospermum-3
Osteospermum-1
Rhynchopsidium
Ozothamnus dio
Helichrysum pat
Acanthocladium
Helichrysum leu
Trichanthodium
Dimorphotheca
Pterygopappus
Myriocephalus
Argentipallium
Leontopodium
Hyalochlamys
Cephalosorus
Xerochrysum
Phaenocoma
Plecostachys
Hyalosperma
Triptilodiscus
Arrowsmithia
Nephrotheca
Gamochaeta
Ozothamnus
Langebergia
Bryomorphe
Oligocarpus
Dolichothrix
Antennaria
Monoculus
Pithocarpa
Gnephosis
Feldstonia
Craspedia
Calendula
Tripteris-1
Norlindhia
Tripteris-2
Jalcophila
Anaphalis
Garuleum
Podolepis
Anaxeton
Asteridea
Petalacte
Lemooria
Inuloides
Relhania
Cassinia
Gibbaria
Mniodes
Quinetia
Rosenia
Raoulia
Athrixia
Oedera
Ewartia
Waitzia
Stoebe
Millotia
p. 762
p. 764
the small Brazilian tribe Moquinieae (Pseudostifftia and southern, in effect covering the whole continent. As a
Moquinia). In some of the analyses the inclusion of this result the fi nal biogeographic resolution of the subfamily
tribe results in Distephanus changing position from being is listed as ‘General Africa’.
the sister group of the rest of the tribe to being ambiguous Sections 3–5 (Figs. 44.5–44.7) cover Corymbieae
at the base of the Vernonieae-Moquinieae clade (Keeley (Cor ymbioideae) and its sister group Asteroideae.
and Robinson, Chapter 28; Funk and Chan, Chapter 23). Section 3, tribes Corymbieae, Senecioneae, Calen-
More highly nested members of Vernonieae are from duleae and Gnaphalieae (Figs. 44.5, 44.10). — The tribe
Brazil and North America. In Vernonieae, the unusual Corymbieae (Corymbioideae) consists of only one genus,
North American genera Stokesia and Elephantopus are not Corymbium, and this distinctive group is restricted to
in the main North American clade but rather represent southern South Africa (Nordenstam 2007c; Fig. 44.5).
two independent lineages (Fig. 44.4). With the excep- Asteroideae encompass the remainder of the family
tion of Liabeae, every tribe or subtribe in Cichorioideae phylogeny, and it is the largest subfamily. It was recog-
s.str. has its origins in Africa, either north, tropical or nized by Cassini (1816) and Bentham (1873a) due to the762
Osmitopsis
Hippia
Soliva
Cotula
Adenanthellum
Lidbeckia
Thaminophyllum
Hilliardia
Inezia
Schistostephium
Inulanthera
Ursinia
p. 761
Leucoptera
Adenoglossa
Eriocephalus
Athanasia
Hymenolepis
Lasiospermum
Eumorphia
Gymnopentzia
Phymasperma
Brocchia
Leucocyclus
Heliocauta
Gonospermum
Cota
Anthemis
Nananthea
Phalacrocarpum
Hymenostemma
Daveaua
Anthemideae
Otospermum
Coleostephus
Plagius
Lepidophorum
Santolina
Endopappus
Argyranthemum
Trichanthemis
Handelia
Microcephala
Sclerorhachis
Lepidolopsis
Funk et al.
Xylanthemum
Myxopappus
Oncosiphon
Cymbopappus
Rennera
Hippolytia
Leucanthemella
Artemisiella
Phaeostigma
Arctanthemum
Chrysanthemum
Neopallasia
Sphaeromeria
Artemisia
Picrothamnus
Denekia
Printzia
Nannoglottis
Mairia
Oritrophium
Lepidophyllum
Chiliotrichum dif
Llerasia
Nardophyllum
Chiliophyllum and
Damnamenia
Olearia col
Astereae
Pleurophyllum
Olearia lya
Celmisia
Olearia che
Fig. 44.6. Anthemideae and Astereae. All taxa are in the subfamily Asteroideae; see Fig. 44.7 for the color chart.
Pachystegia
Olearia covChapter 44: Compositae metatrees: the next generation 763
Astereae
Tetramolopium alp
Tetramolopium lep
Rhynchospermum
Symphyotrichum
Commidendrum
Chrysothamnus
Melanodendron
Archibaccharis
Diplostephium
Erodiophyllum
Welwitschiella
Brachyscome
Pteronia cam
Plagiocheilus
Sommerfeltia
Haplopappus
Sericocarpus
Hinterhubera
Vittadinia ple
Vittadinia bla
Bellidiastrum
Parastrephia
Chrysocoma
Callistephus
Heterotheca
Polyarrhena
Conyza gou
Pteronia inc
Poeciliopsis
Madagaster
Doellingeria
Eucephalus
Ixiochlamys
Olearia ram
Westoniella
Conyza pyr
Gundlachia
Chrysopsis
Camptacra
Olearia stu
Podocoma
Felicia min
Ericameria
Xanthisma
Olearia fer
Microgyne
Laennecia
Felicia aet
Peripleura
Aster com
Isoetopsis
Baccharis
Keysseria
Felicia fru
Aster cap
Aster dim
Euthamia
Lessingia
Tripolium
Aster har
Zyrphelis
Aster gle
Galatella
Nidorella
Herrickia
Felicia fil
Solidago
Erigeron
Aster tai
Amellus
Conyza
Egletes
Nolletia
Psiadia
Bellium
Remya
Calotis
Bellis
combination of its capillary pappus, true rays, and a recep- among Senecioneae and its potential sister groups,
tive area in two lines on the inside of the style branches. there are not enough data to determine whether or not
Senecioneae have long been one of the largest and these two genera should stay in the tribe as subtribes
most difficult groups to understand; they are truly a or be moved to tribes of their own. The authors of the
global tribe with major radiations in sub-Saharan Africa, Senecioneae Chapter (Nordenstam et al., Chapter 34)
West and East Asia, Andean South America, and Mexico. have reserved fi nal judgment on this matter until they
Because of uncertainty about the phylogenetic positions have more information.
of the core of Senecioneae and two of the genera usually Within the core Senecioneae clade there are four clades
assigned to this tribe, relationships among these taxa and that form a polytomy. One represents the bulk of the
the clade formed by the other Asteroideae tribes are pres- species, which are found in two monophyletic subtribes
ently unresolved. that are sister taxa (Othonninae + Senecioninae). This
Two genera of Senecioneae, Doronicum (Eurasia and major clade has a sub-Saharan African base with highly
northern Africa) and Abrotanella (Australasia and southern nested groups of species from South America and Central
South America), “fall out” of monophyletic Senecioneae. America–Caribbean basin. The second clade, the core of
Since there is a general problem of tribal relationships subtribe Tussilagininae, has clades in Asia, Eurasia, NorthYou can also read
