American Journal of Botany 97(5): 856–873. 2010.
PATTERNS AND CAUSES OF INCONGRUENCE BETWEEN PLASTID
AND NUCLEAR SENECIONEAE (ASTERACEAE) PHYLOGENIES1
Pieter B. Pelser2,8, Aaron H. Kennedy3,4, Eric J. Tepe5, Jacob B. Shidler4,
Bertil Nordenstam6, Joachim W. Kadereit7, and Linda E. Watson3
2University
of Canterbury, School of Biological Sciences, Private Bag 4800, Christchurch 8140 New Zealand; 3Oklahoma State
University, Department of Botany, 104 Life Science East, Stillwater, Oklahoma 74078 USA; 4Miami University, Department of
Botany, 316 Pearson Hall, Oxford, Ohio 45056 USA; 5University of Cincinnati, Department of Biological Sciences, 614
Rieveschl Hall, Cincinnati, Ohio 45221 USA; 6Swedish Museum of Natural History, Department of Phanerogamic Botany, P.O.
Box 50007 SE-104 05 Stockholm, Sweden; and 7Johannes Gutenberg-Universität Mainz, Institut für Spezielle Botanik und
Botanischer Garten, 55099 Mainz Germany
One of the longstanding questions in phylogenetic systematics is how to address incongruence among phylogenies obtained
from multiple markers and how to determine the causes. This study presents a detailed analysis of incongruent patterns between
plastid and ITS/ETS phylogenies of Tribe Senecioneae (Asteraceae). This approach revealed widespread and strongly supported
incongruence, which complicates conclusions about evolutionary relationships at all taxonomic levels. The patterns of incongruence that were resolved suggest that incomplete lineage sorting (ILS) and/or ancient hybridization are the most likely explanations. These phenomena are, however, extremely difficult to distinguish because they may result in similar phylogenetic patterns.
We present a novel approach to evaluate whether ILS can be excluded as an explanation for incongruent patterns. This coalescence-based method uses molecular dating estimates of the duration of the putative ILS events to determine if invoking ILS as an
explanation for incongruence would require unrealistically high effective population sizes. For four of the incongruent patterns
identified within the Senecioneae, this approach indicates that ILS cannot be invoked to explain the observed incongruence. Alternatively, these patterns are more realistically explained by ancient hybridization events.
Key words: ancient hybridization; deep coalescence; ETS; incomplete lineage sorting; incongruence; ITS; molecular dating;
plastid sequences.
Since the onset of the use of DNA sequences for phylogeny
reconstruction, molecular systematics has experienced a steady
increase in the number of DNA regions used to resolve evolutionary relationships (Degnan and Rosenberg, 2009). This development has significantly contributed to our understanding
of the evolutionary history of numerous lineages by clarifying
relationships that were previously unresolved in studies using
fewer markers and less data (e.g., Kuzoff and Gasser, 2000;
Pryer et al., 2004; Panero and Funk, 2008). In addition, multigene studies have enabled a more detailed understanding of
macroevolution by revealing congruence or incongruence be-
tween phylogenies obtained from the analysis of different
genes, genic regions, and genomes (Degnan and Rosenberg,
2009). Incongruence may, among others, indicate differences
in the evolutionary histories of the DNA regions employed
(i.e., gene tree–species tree discordance), which could result
from hybridization or incomplete lineage sorting (ILS; Doyle,
1992; Maddison, 1997; Buckley et al., 2006; Liu and Pearl,
2007). ILS is the failure of ancestral polymorphisms to track
speciation events accurately, which may result in incongruence between gene trees and species trees. Unfortunately, ILS
can result in phylogenetic patterns similar to those observed
for hybridization events (Doyle, 1992; Seelanan et al., 1997;
Holder et al., 2001; Buckley et al., 2006; Holland et al., 2008;
Joly et al., 2009). Therefore ILS and hybridization are often
difficult to distinguish. Furthermore, in the absence of an effective methodology to distinguish between them (Joly et al.,
2009), there are few empirical studies that present a detailed
assessment of incongruent patterns and even fewer in which
the specific causes for these patterns were studied (Wiens and
Hollingsworth, 2000; Van der Niet and Linder, 2008; Morgan
et al., 2009). There is therefore an urgent need for research that
explores ways to examine incongruent patterns and to determine the causes (Buckley et al., 2006; Holland et al., 2008;
Degnan and Rosenberg, 2009). In this paper, we aim to contribute to the development of new approaches to the study of
incongruent phylogenetic patterns by documenting strongly
supported topological incongruence in tribe Senecioneae
(Asteraceae) and using a new approach to establish whether
ILS can be excluded as an explanation for incongruent patterns. This coalescent-based method uses estimates of the
1 Manuscript received 21 September 2009; revision accepted 22 February
2010.
The authors thank A. Aksoy, R. J. Bayer, R. Cairns-Wicks, P. Carillo
Reyes, G. V. Cron, V. A. Funk, E. B. Knox, R. R. Kowal, A. Marticorena,
J. L. Panero, T. Sultan Quedensley, T. F. Stuessy, and I. R. Thompson for
providing DNA or tissue samples. They are grateful to the curators of B,
BHCB, CANB, E, F, HYO, L, LPB, M, MEL, MJG, MO, MU, P, PRE,
PREM, S, TENN, TEX, U, UC, US, WAG, WIS, WU, and XAL for
permission to include their Senecioneae specimens in their studies. R. H.
Ree kindly provided his python script for coding indels. H. P. Comes, J. H.
Degnan, M. Fishbein, and A. Varsani are acknowledged for helpful
discussions. P. C. Wood and the Miami University Center for Bioinformatics
and Functional Genomics provided technical support. The authors greatly
appreciate the time and efforts of two anonymous reviewers and their help
to improve an earlier version of this manuscript. Funding for this project
was provided by NSF grant DEB-0542238 to L.E.W. and P.B.P.
8 Author for correspondence (e-mail: pieter.pelser@canterbury.ac.nz)
doi:10.3732/ajb.0900287
American Journal of Botany 97(5): 856–873, 2010; http://www.amjbot.org/ © 2010 Botanical Society of America
856
May 2010]
Pelser et al.—Incongruence between Senecioneae phylogenies
duration of putative ILS events to determine if invoking ILS as
an explanation for incongruence would require unrealistically
high effective population sizes.
The Senecioneae are one of the largest tribes in the Asteraceae (ca. 3100 species and 155 genera) with an almost worldwide distribution, and it exhibits remarkable morphological and
ecological diversity. Senecio is the largest genus in the tribe (ca.
1000 species) and is perceived as taxonomically difficult because of its size and extensive morphological diversity. In a
recent study, ITS sequence data were used to reconstruct a phylogeny for the Senecioneae and to redefine a previously polyphyletic Senecio (Pelser et al., 2007). This study included 186
Senecio species and 114 of the 150 genera recognized by
Nordenstam (2007). Because the molecular phylogeny placed
several lineages outside of core Senecio, a new circumscription
resulted with several taxa being transferred to other genera
within the tribe and others to be described as new genera (Nordenstam et al., 2009c; unpublished data). Furthermore, seven
genera (Aetheolaena, Cadiscus, Culcitium, Hasteola, Iocenes,
Lasiocephalus, and Robinsonia) were deeply embedded within
Senecio and thus, are being transferred into the genus to arrive
at a monophyletic delimitation (Pelser et al., 2007; Nordenstam
et al., 2009b; unpublished data). It is important to note that
while this new delimitation of Senecio is largely based on an
ITS phylogeny, the major clades are also supported by plastid
sequence data composed of ndhF, the trnL intron, and the psbAtrnH, 5′ and 3′ trnK, and trnL-F intergenic spacers (Pelser et al.,
2007). In addition, the plastid data augmented the utility and
efficacy of the ITS for phylogeny reconstruction in the Senecioneae by supporting many of the resolved evolutionary relationships at the species, generic, and subtribal levels in the tribe.
Furthermore, many patterns of relationships in the ITS Senecioneae phylogeny were corroborated by morphological, karyological, and/or biogeographic data and supported the taxonomic
conclusions from previous Senecioneae studies (reviewed in
Pelser et al., 2007; Nordenstam et al., 2009a). Although the
plastid data provided additional support for the ITS phylogeny
with overall congruence between the two topologies observed,
some incongruence was present that prevented taxonomic conclusions for some lineages (Pelser et al., 2007). Because the
plastid phylogeny was only based on a subset of the Senecioneae taxa that were sampled for the ITS analyses (73 vs. 614
species), it was not possible to sufficiently resolve these patterns for a detailed analysis of the extent and understanding of
the underlying causes of the incongruence at the generic level.
In the current study, we therefore have significantly increased
the taxon and marker sampling for the plastid and nuclear data
and conducted extensive phylogenetic analyses. The goals were
(1) to identify strongly supported relationships that are incongruent between trees produced with different data sets and (2) to
explore possible causes of the incongruence.
MATERIALS AND METHODS
Taxon and character sampling—The taxa selected for this study were chosen from the plastid and ITS phylogenies of Pelser et al. (2007) to represent the
Senecioneae genera included in that study. In addition, 31 Senecioneae genera
were added for which material was not previously available to us (Appendix
S1; see Supplemental Data with the online version of this article) resulting in a
sampling of approximately 94% of all Senecioneae genera. A total of 27 genera
from 26 other Asteraceae tribes was included for outgroup comparisons (Appendix S1). DNA samples of representatives of the taxa not included in Pelser
et al. (2007) were obtained from tissue samples taken with permission from
857
herbarium specimens at B, BHCB, F, L, MO, MU, P, S, TENN, TEX, UC, US,
WAG, and XAL, or from field-collected leaf tissue preserved on silica gel.
Phylogenetic analyses were performed with DNA sequences from eight regions: the ITS (ITS1, 5.8S, ITS2) and ETS of the nuclear genome, plus the
ndhF gene, the trnL intron, and psbA-trnH, 5′ and 3′ trnK, and trnL-F intergenic
spacers of the plastid genome. To root tribe Senecioneae in subfamily Asteroideae, partial sequences of the plastid rbcL gene were obtained for representatives of the major Senecioneae lineages and the other Asteraceae tribes, in
addition to the above markers. However, the relationship among outgroups is
not discussed since this is not a goal of this paper.
DNA extraction, PCR amplification, and sequencing—Total genomic
DNA was extracted using the Qiagen DNeasy Plant Mini Kit (Qiagen, Valencia, California, USA). PCR amplification of the ITS, ndhF, psbA-trnH, 5′ and
3′ trnK, trnL, and trnL-F regions followed Pelser et al. (2002, 2003, 2007) or
minor modifications thereof. The ETS region was partially amplified with
primer AST1 (Markos and Baldwin, 2001) or a modification of primer ETS2
(Bayer et al., 2002; 5′-CAA CTT CCA CCT GGC TTA CCT CC-3′) as forward
primers and 18S-ETS (Baldwin and Markos, 1998) as the reverse primer. The
rbcL gene was amplified in two parts using forward primers 1F and a modification of primer 636F (Fay et al., 1997; 5′-GCG TTG GAG AGA CCG TTT
CT-3′), and reverse primers 1460R (Savolainen et al., 2000) and a modification
of primer 724R (Fay et al., 1997; 5′-TCG CAT GTA CCC GCA GTA GC-3′).
PCR products were purified with the Wizard SV Gel and PCR Clean-Up System (Promega, Madison, Wisconsin, USA). Cycle sequencing was carried out
with the BigDye Terminator v.3.1 (Applied Biosystems, Foster City, California, USA) cycle sequencing kit. The fluorescently labeled samples were run on
an ABI 3130xl or 3730 automated sequencer at the Center for Bioinformatics
and Functional Genomics at Miami University (Oxford, Ohio, USA). The program Sequencher v.4.8 (Gene Codes, Ann Arbor, Michigan, USA) was used for
trace file editing.
DNA sequence alignment and phylogeny reconstruction—DNA sequences
were aligned with the program CLUSTALX v.1.83 (Thompson et al., 1997)
using the default penalty settings of the program. This alignment was edited
using the program Se-Al v.2.0a11 (Rambaut, 1996) by excluding portions of
sequences that could not be unambiguously aligned (mostly of ITS and ETS
accessions of tribes distantly related to the Senecioneae) and replacing those
with a missing data symbol. A Python script (Richard Ree, Field Museum, Chicago, Illinois, USA) was used to code indels as binary characters using the
simple indel coding method of Simmons and Ochoterena (2000).
For several species, sequences of multiple accessions were available (Appendix S1). In the first round of heuristic searches performed under maximum
parsimony (MP, see below) for each of the DNA regions individually, all available sequences were included. When multiple accessions of the same species
formed a clade, a consensus sequence was generated in which polymorphisms
were coded as ambiguous nucleotide characters, and this consensus sequence
was used for subsequent phylogenetic analyses (Pelser et al., 2007). This strategy was preferred over selecting one conspecific accession among those available to avoid subjective decisions in the process of selecting a single examplar
and to be able to include all available data that potentially contribute to the
phylogeny reconstruction. In addition, this approach allowed for the inclusion
of longer sequence reads for taxa of which individual accessions only resulted
in partial sequences. Comparisons to results of parsimony analyses did not reveal decreased resolution when multiple accessions per species were included,
and analyses in which consensus sequences also were included placed these
among the individual accessions of the same taxon.
The MP phylogeny was reconstructed with the program TNT 1.0 (Goloboff
et al., 2008) using the Driven Search option with the default settings for Sectorial Searches (RSS, CSS, and XSS), Ratchet, Tree Drifting, and Tree Fusing, 10
initial random addition sequences, terminating the search after finding minimum length trees five times. Bootstrap support (BS; Felsenstein, 1985) was
calculated with Poisson independent reweighting using 1000 replicates. Bayesian inference (BI) analyses were performed using the program MrBayes 3.1.2
(Huelsenbeck and Ronquist, 2001) on the Redhawk Cluster at Miami University (EM64T Cluster with 128 dual nodes with 4 GB of memory and 150 GB of
disk space per node). Prior to the BI analyses, the Akaike information criterion
(AIC) in the program MrModeltest 2.2 (Nylander, 2004) was employed to select nucleotide substitution models for each of the DNA regions. These analyses selected the GTR+I+Γ model for each of the individual DNA regions and
the combinations of DNA regions analyzed, which was therefore used in all BI
analyses. Indel characters were included as “restriction type” data in the BI
analyses. These analyses were performed using two independent simultaneous
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American Journal of Botany
runs. The Markov chain Monte Carlo analyses (MCMC; Geyer, 1991) were run
with up to 40 chains per analysis, temperature settings ranging between 0.0001
and 0.2, and one tree per 1000 generations saved. BI analyses were run until the
average deviation of split frequencies between both simultaneous analyses
reached a value below 0.01. The burn-in values were determined empirically
from the likelihood values. Trees were visualized using the program FigTree
v.1.2.2 (Rambaut, 2009).
Identification and localization of incongruence—The incongruence length
difference test (ILD; Farris et al., 1995) is one of the most widely applied methods for assessing incongruence (Darlu and Lecointre, 2002; Hipp et al., 2004).
Its use has, however, been criticized, because of a high false positive rate
(Cunningham, 1997; Darlu and Lecointre, 2002; Hipp et al., 2004). Because alternative methods for testing incongruence (e.g., the Templeton [Templeton,
1983], Kishino–Hasegawa [Kishino and Hasegawa, 1989], and Shimodaira–
Hasegawa [Shimodaira and Hasegawa, 1999] tests) may suffer from errors as
well (Cunningham, 1997; Shimodaira and Hasegawa, 1999; Buckley et al.,
2001; Shimodaira, 2002; Hipp et al., 2004), incongruence is probably best studied using a combination of methods (Hipp et al., 2004). In this study, we therefore explored patterns of phylogenetic incongruence using two approaches: the
ILD test and an assessment of incongruent patterns that are supported by high
BS or BI values. To reduce the chance of false positives and following the
recommendations of Cunningham (1997), we considered only ILD P-values
below 0.01 as evidence of significant incongruence.
Strongly supported incongruence is here defined as incongruent patterns that
are supported by BS values ≥80% and/or posterior probabilities (PP) ≥0.95 as
well as ILD values of P < 0.01. The ILD tests were conducted in PAUP* with
500 to 10 000 replicates and 1 to 100 random addition sequences, depending on
the size and complexity of the data sets analyzed. Invariant characters were removed from the data sets prior to performing ILD tests (Cunningham, 1997).
This methodology revealed strong incongruence between trees obtained from
the plastid and ITS/ETS data sets, but not among the plastid markers or between
the ITS and ETS. Subsequent phylogenetic analyses were therefore focused on
identifying lineages with strongly supported, but incongruent, topological
placements in trees obtained from the plastid vs. the ITS/ETS data sets.
Lineages with strongly supported, yet incongruent, phylogenetic placements
in the plastid vs. ITS/ETS trees were identified using a novel two-step approach
designed to examine complex patterns involving multiple incongruent lineages
for which some also have internal incongruence. First, plastid and ITS/ETS
trees were visually compared to identify the largest mutually exclusive lineages
that form a clade in all or some consensus trees. In this way, lineages that are
resolved as closely related in both plastid and ITS/ETS trees were identified.
Among these lineages are clades that are retrieved in all trees (e.g., Tussilagininae s.s.; Figs. 1, 2), but also those lineages that form a clade in some, but not all,
trees. The latter category of lineages were only included when their taxa are in
close phylogenetic proximity (e.g., Lachanodes-S. thapsoides group; Figs. 1,
2). Although Senecio s.s. is not resolved as monophyletic in either the plastid or
the ITS/ETS trees, support for its nonmonophyly is low (Figs. 1, 2), and it
comprises a single clade in trees obtained from a combined plastid-ITS/ETS
data set (not shown). Senecio s.s. was therefore also identified as one of the
major lineages present in both plastid and ITS/ETS trees.
Secondly, these identified major lineages were then examined for the presence of strongly supported internal incongruence by evaluating branch support
values and then subjecting them to ILD tests that only included the taxa of the
lineage under investigation. This method allowed for the identification of
strongly incongruent accessions and subclades within each of these major lineages without the confounding effects of unrelated incongruence within or
among other major lineages. In addition, branch support values and ILD tests
were employed in a similar fashion to study the incongruence among the major
lineages. Due to the large size of the data sets, one or two placeholder species
were included in the latter round of ILD tests to eliminate the confounding effects of internal incongruence within the major lineages.
After strongly incongruent lineages were identified, MP and BI analyses of
a combined plastid-ITS/ETS data set were performed in which each taxon in a
strongly incongruent lineage was included twice: once as a plastid-only accession (ITS/ETS characters coded as missing) and once as an ITS/ETS-only accession (plastid characters coded as missing; Pirie et al., 2008). This approach
was used to resolve a generic level tree without the confounding effects of taxa
with strongly supported incongruence between the plastid and ITS/ETS partitions and to examine their alternative phylogenetic placements relative to a
backbone composed of lineages among which relationships are not strongly
incongruent. Because BI analyses of this data set using various temperature
settings and numbers of chains did not reach convergence (40 chains per run,
[Vol. 97
temperature setting of 0.0001), a user-defined starting tree was supplied using
the MP 50% majority rule consensus topology of the combined plastid-ITS/
ETS data set. Each chain was then initiated with a slightly perturbed version of
this starting tree by setting the parameter nperts to 1. This approach resulted in
convergence of the chains.
ITS orthology/paralogy assessment—To determine if specimens possessed
divergent ITS copies, potentially indicating that the incongruence is caused by
inaccurate assessment of orthology, we cloned and sequenced the ITS region
for accessions of strongly incongruent lineages. Because ITS trees were not
strongly incongruent with ETS trees and both regions are adjacent, the ETS
region was not cloned and sequenced. ITS PCR products were cloned using the
TOPO TA Cloning Kit for Sequencing (Invitrogen, Carlsbad, California, USA).
Between 5 and 10 clones per accession were PCR-amplified directly from
plated culture with the manufacturer’s supplied M13 plasmid primers. PCR
products were cycle sequenced with M13 plasmid primers. Sequencing and
alignment followed the protocol outlined above.
Long-branch attraction—Because long-branch attraction (Felsenstein,
1978) is a possible explanation for topological incongruence, MP and BI phylograms were visually inspected for the presence of exceptionally long branches
associated with strongly incongruent lineages. A series of phylogenetic analyses was subsequently carried out in which accessions with long branches were
individually excluded in turn. This was done to determine whether the exclusion of these accessions resulted in changes in topology, which could indicate
the presence of long-branch attraction.
Incomplete lineage sorting—Topological incongruence caused by hybridization and ILS can be difficult to distinguish, because both phenomena may
result in similar topological patterns. Coalescent theory, however, predicts that
ancestral polymorphisms are likely to coalesce within approximately 5Ne generations (Ne being the effective population size; Rosenberg, 2003; Degnan and
Rosenberg, 2009) and that congruence between gene trees and species trees
becomes highly probable. Information about generation times and estimates of
the duration of an ILS event can therefore be used to calculate the minimum Ne
that must be assumed to explain incongruence due to ILS. If these calculations
result in Ne estimates that are much higher than observed in nature, then ILS can
be excluded, and hybridization is favored as the likely explanation for the observed incongruence.
To calculate Ne estimates, we estimated molecular dates using penalized
likelihood analyses performed in the program r8s 1.71 (Sanderson, 2002, 2003)
on the Redhawk Cluster at Miami University. Separate analyses were carried
out for the plastid and the ITS/ETS data sets using the topologies and branch
lengths obtained in the BI analyses. Some accessions were pruned from the input trees to meet the r8s requirement of nonzero branch lengths. Several calibration points outside and within the Senecioneae were used based on previous
age estimates in Asteraceae (Wikström et al., 2001; Kim et al., 2005; Hershkovitz
et al., 2006), fossil evidence (Graham, 1996), and inferred ages for islands
or archipelagos with endemic Senecioneae taxa (Baker et al., 1967; McDougall
and Schmincke, 1976; McDougall et al., 1981; Stuessy et al., 1984; Cronk,
1987; Ancochea et al., 1990; Geldmacher et al., 2000; Table 1). The smoothing
parameters for the penalized likelihood analyses were determined using crossvalidation tests.
In addition to estimating divergence dates using penalized likelihood, molecular dating studies were performed using a Bayesian approach with the program BEAUti/BEAST v.1.4.7 (Drummond and Rambaut, 2007) on the
Redhawk Cluster at Miami University. To compare the dating results of the
BEAST analyses to those estimated by r8s, we also used the input trees from the
r8s analyses as the starting trees in BEAST, and their topology was fixed. The
BEAST analyses were performed with the GTR+I+Γ model, the uncorrelated
relaxed lognormal clock, and the Yule tree prior. The calibration points used in
the r8s analysis were specified using uniform distributions with an upper and
lower bound. Following the instructions in the BEAST manual, the weights of
the operators for the treeModel were modified to improve the efficiency of the
MCMC: the upDownOperator, uniformOperator on internalNodeHeights, narrowExchangeOperator, and subtreeSlideOperator were set at 115 and the wilsonBaldingOperator and the wideExchangeOperator were set at 23. Nine
(plastid data) and 11 (ITS/ETS data) independent BEAST runs totaling
136 687 000 (plastid data) and 318 672 000 (ITS/ETS data) generations were
carried out to ensure that all parameters had an effective sampling size greater
than 200 (after a burnin of 10% was removed). Convergence was examined
using the program Tracer v.1.4 (Rambaut and Drummond, 2007).
Pelser et al.—Incongruence between Senecioneae phylogenies
May 2010]
Table 1.
Calibration points for r8s and BEAST molecular dating
analyses.
Calibration point
Root (fixed)
Age
constraint Age estimates from
used (Myr) literature (Myr)
35
33–36.5
MRCA of
Cichorioideae and
Asteroideae
Anthemideae
24–38
24–38
MRCA of Helianthus
and Tagetes
13–19
Late Oligocene
(fossil pollen)
13–19
MRCA of Senecio
and Blennosperma
Lachanodesa
16–19
16–19
Pladaroxylona
Pericallis auritaa
Pericallisa
Bethencourtiaa
≥23
≤14.5
≤14.5
≤14.3
≤14.3
≤11.6
Lordhoweaa
≤6.9
Robinsoniaa
≤4.2
Robinsonia
masafueraea
aIsland
bOldest
≤2.4
14.5
(age of St. Helena)
14.5
(age of St. Helena)
14.3
(age of Porto Santo)
14.3
(age of Porto Santob)
References
Wikström et al., 2001;
Kim et al., 2005;
Hershkovitz et al., 2006
Wikström et al., 2001;
Kim et al., 2005;
Hershkovitz et al., 2006
Graham, 1996
Wikström et al., 2001;
Kim et al., 2005;
Hershkovitz et al., 2006
Kim et al., 2005;
Hershkovitz et al., 2006
Baker et al., 1967;
Cronk, 1987
Baker et al., 1967;
Cronk, 1987
Geldmacher et al.,
2000
McDougall and
Schmincke, 1976;
Geldmacher et al.,
2000
Ancochea et al., 1990
11.6
(age of Tenerifeb)
6.9
McDougall et al., 1981
(age of Lord Howe
Island)
4.2
Stuessy et al., 1984
(age Juan Fernández
Islands)
2.4
Stuessy et al., 1984
(age of Masafuera)
or archipelago endemics.
island in distribution area.
After obtaining age estimates of divergences, we calculated the duration
of each putative ILS event as follows. First, the estimated time of onset of an
ILS event was determined from both the plastid and ITS/ETS BEAST chronograms. Although the plastid and ITS/ETS BEAST chronograms resulted
in different age estimates for the onset of ILS events, we assume that these
estimates are of the same speciation event in the species tree. To approximate the age of each speciation event that marks the start of an ILS event, we
used the overlap in the 95% highest posterior density (HPD) intervals of the
plastid and ITS/ETS chronograms. For a few of the strongly incongruent
lineages, r8s estimates of the onset of putative ILD events were slightly outside the ranges obtained from BEAST. In these cases, the ranges of the assumed time interval of the ILS onset calculated from the BEAST chronograms
were extended with the r8s ages. Secondly, the ages of the most recent common ancestor (MRCA) of each strongly incongruent lineage and its nonincongruent sister group were recorded from the plastid and ITS/ETS
chronograms. These age estimates signify the end of a putative ILS event in
859
either data set. Again, the 95% HPD intervals of the BEAST analyses were
used to account for some uncertainty in the data, and these age estimate
ranges were extended with the r8s results when the r8s estimates fell outside
of the ranges calculated with BEAST. Finally, the ranges of age estimates for
the onset and the end of the putative ILS event were used to calculate its
minimum and maximum duration.
Using these estimates of putative ILS durations and information about
generation times in the Senecioneae, the effective population sizes needed to
explain incongruent patterns invoking ILS were calculated. Most Senecioneae species flower in their first or second year, and generation times of
one or two years are characteristic for most lineages. To test for robustness
of our conclusions to violations of this assumption, effective population
sizes were also calculated using generation times of 5 and 10 yr. Senecioneae
populations typically range between several dozen to a few thousand individuals (e.g., Widén and Andersson, 1993; Comes and Abbott, 1998; MüllerSchärer and Fischer, 2001). Ne values of 20 000 (plastid data) and 40 000
(ITS/ETS data) were therefore selected as conservative estimates of maximum population sizes. These coalescent calculations were carried out for
each of the strongly incongruent lineages, but were also performed for scenarios in which the conflicting phylogenetic positions of multiple taxa were
assumed to be nonindependent (i.e., a single ILS event resulting in incongruent positions of multiple lineages). Because species-level sampling within
Senecio was limited (48/~1000 species) and accurate estimates of the duration of putative ILS events could not be obtained, these calculations were not
performed for Senecio species.
RESULTS
Identification and localization of incongruence— Although
an ILD test comparing all five plastid markers resulted in P =
0.005, the consensus trees obtained from the individual markers
did not reveal well-supported (BS values ≥ 80% and/or PP ≥
0.95) incongruence. Sequence data from all plastid markers
were therefore combined into a single data set, which was used
for all subsequent analyses. The ITS and ETS trees (not presented) were mostly congruent, and incongruent clades were
not well-supported. Further analyses were carried out with a
combined ITS/ETS data set, even though an ILD test resulted in
P = 0.004. The results of the MP and BI analyses of the combined plastid data and the combined ITS/ETS data sets are summarized in Table 2 and presented in Figs. 1 and 2. The ILD test
for the plastid vs. the ITS/ETS data sets indicated significant
incongruence (P = 0.004), and a visual comparison of their topologies and branch support values revealed well-supported
incongruence (Figs. 1, 2). Therefore, further studies of incongruence within Senecioneae were confined to examining the
strongly conflicting topologies of the plastid vs. the ITS/ETS
data sets. Discussion in this paper is limited to relationships
within the Senecioneae only and does not include outgroup
relationships.
Although all analyses of plastid and ITS/ETS data resolved
the core of Senecioneae (excluding Doronicum and Abrotanella) as monophyletic (Figs. 1, 2; BS 100%, PP 0.99 and 1.00)
and supported Capelio as sister to the well-supported clade
composed of the remaining genera forming the Senecioneae
Table 2.
Character information and clade support data. In the combined plastid-ITS/ETS data set strongly incongruent taxa were included as separate
plastid and ITS/ETS accessions (see text).
Data set
Plastid
ITS/ETS
Combined plastid-ITS/ETS
No. of clades with
No. of
characters
Gap
characters
Variable
characters
Informative
characters
≥50% BS
≥80% BS
PP ≥0.5
PP ≥0.95
12494
3303
15797
1293
914
207
3253 (26.0%)
1605 (48.6%)
4861 (30.7%)
1406 (11.3%)
1096 (33.2%)
2505 (15.9%)
125 (55.1%)
116 (50.5%)
158 (56.8%)
87 (38.3%)
76 (33.0%)
100 (36.0%)
194 (85.5%)
195 (84.8%)
248 (89.2%)
132 (58.2%)
127 (55.2%)
180 (64.7%)
860
American Journal of Botany
[Vol. 97
Fig. 1. Bayesian inference (BI) topology from the plastid data set. Bayesian consensus percentages (posterior probabilities × 100) are placed above
branches, bootstrap support above 50% below branches. Branch color indicates congruence with ITS/ETS BI tree (Fig. 2): green: congruent; yellow: clade
absent in ITS/ETS BI tree, but not contradicted; orange: clade incongruent, but not with posterior probabilities ≥0.95 in both trees; red: incongruent with
posterior probabilities ≥0.95 in both trees.
May 2010]
Pelser et al.—Incongruence between Senecioneae phylogenies
861
Clade 1
"
i
,,
c'
00
セ M
'""
55-
Clada2
00
$
'"セ@
セ@
$
Fig. 2. Bayesian inference (BI) topology from ITS/ETS data set. Bayesian consensus percentages (posterior probabilities × 100) are placed above
branches, bootstrap support above 50% below branches. Branch color indicates congruence with plastid BI tree (Fig. 1): green: congruent; yellow: clade
absent in plastid BI tree, but not contradicted; orange: clade incongruent, but not with posterior probabilities ≥0.95 in both trees; red: incongruent with
posterior probabilities ≥0.95 in both trees.
862
American Journal of Botany
(BS 100%, PP 0.99 and 1.00), relationships among many of the
other taxa are obscured by complex patterns of topological incongruence. Among the 29 major lineages of Senecioneae that
were identified (Table 3), only two exhibit strongly supported
internal incongruence: the New World 1 group and Senecio
(Table 3).
Within New World 1 group, incongruence is located in
clade 3 (Figs. 1, 2) in which Lomanthus and the CaxamarcaPseudogynoxys clade take incongruent positions (Table 3).
An ILD test of the data set that was solely composed of the
accessions of clade 3 indicated significant incongruence between the plastid and ITS/ETS data (P = 0.001). This conflict
is not significant (P = 0.256) if Lomanthus is excluded, but
remains statistically significant if only the Caxamarca-Pseudogynoxys clade is excluded (P = 0.001). Similarly, in a series
of MP and BI analyses of a data set in which only the accessions of clade 3 were included, incongruence supported with
high BS and PP values only disappeared when Lomanthus was
excluded (not shown).
Within Senecio, S. flavus, S. cadiscus, S. pinnatifolius, S.
hispidissimus, S. psilocarpus, and S. squarrosus are placed
in strongly supported incongruent positions (Table 3; P =
0.001). Removing all six taxa from the data sets resulted in
trees without high BS and PP values for incongruent patterns
(not shown) and an insignificant ILD value (P = 0.06). When
any five of these six species were excluded, ILD tests indicated significant incongruence (P = 0.001–0.007), and incongruent patterns were supported with high support values (not
shown).
In addition to documenting strongly supported internal
incongruence for two of the 29 major Senecioneae lineages, 13
others exhibit well-supported incongruence with regard to their
phylogenetic positions relative to the 16 remaining lineages
among which well-supported incongruence was not identified
(Table 4). These 13 incongruent lineages are Caputia, New
World 2 group, Jacobaea, Lordhowea, gynuroids, Lamprocephalus-Oresbia clade, Phaneroglossa, Cineraria, Steirodiscus, Packera, Emilia-Bethencourtia group, Senecio otites, and
Io. An ILD test in which placeholders for each of the 29 major
Senecioneae lineages were included indicated significant incongruence between the plastid and ITS/ETS data (Table 5: test
1; P = 0.001). Excluding the placeholders for the 13 lineages
that have well-supported incongruent phylogenetic positions
still resulted in significant incongruence among the remaining
lineages (Table 5: test 2; P = 0.006). Only when Othonninae
(i.e., Othonna) was also excluded, an ILD test did not indicate
significant incongruence (Table 5: test 3; P = 0.052). Adding
placeholders for each of the incongruent major lineages individually to the nonsignificant incongruent data set of test 3 introduced significant incongruence for all of these groups (Table
5: tests 4–13) except for the Emilia-Bethencourtia group, Senecio otites, and Io (tests 14–16). Although adding all three of
these placeholders (test 17), the Emilia-Bethencourtia group
and Senecio otites (test 18), or Senecio otites and Io (test 19),
resulted in significant incongruence (Table 5), incongruence
was insignificant (P = 0.013) in an ILD test in which the EmiliaBethencourtia group and Io were added to the insignificantly
incongruent selection of placeholder species used in test 3 (test
20). Inspection of the BS and PP values in trees obtained from
the data sets used in ILD tests 1–20 generally confirmed these
findings (not shown).
In summary, after exploring incongruence within and
among the major Senecioneae lineages and following the cri-
[Vol. 97
teria outlined in the Materials and Methods, we identified the
following taxa and lineages as having strongly incongruent
placements: Caputia, Cineraria, Steirodiscus, Phaneroglossa,
the Lamprocephalus-Oresbia clade, Lordhowea, the gynuroids, Jacobaea, Packera, Senecio otites, Lomanthus, the
New World 2 group, S. flavus, S. hispidissimus, S. psilocarpus, S. squarrosus, S. cadiscus, and S. pinnatifolius. The MP
and BI analyses of the combined plastid-ITS/ETS data set in
which the strongly incongruent lineages were included as separate plastid and ITS/ETS accessions resulted in trees that
generally have slightly higher support than the individual
plastid and ITS/ETS consensus trees (Fig. 3; Table 2). There
are only a few topological differences between the MP 50%
majority rule consensus tree (not shown) and the BI phylogeny (Fig. 3), and all of these differences are weakly supported.
The phylogenetic positions of the plastid and ITS/ETS accessions of the strongly incongruent lineages in the combined
plastid-ITS/ETS trees (Fig. 3) correspond well to those in the
separate plastid and ITS/ETS trees (Figs. 1, 2).
ITS orthology/paralogy assessment— Cloned ITS sequences
selected from 18 strongly incongruent Senecioneae lineages
(Appendix S1) revealed a relatively high level of polymorphism. Most specimens had between 3 and 24 (0.4–3.2%) polymorphic nucleotide positions; however, the cloned sequences
from Oresbia had higher levels (7.8%) with 59 polymorphic
nucleotide positions. Despite this high level of polymorphism,
sequences of the ITS clones obtained from the same specimen always formed well-supported clades in MP analyses
(results not shown) and with the directly sequenced ITS PCR
products.
Long-branch attraction— Visual inspection of the plastid
and ITS/ETS trees revealed several species in the Tussilagininae s.s. clade with relatively long branches: Blennosperma,
Crocidium, Ischnea, Robinsonecio, and the Arnoglossum-Barkleyanthus-Yermo clade (Appendix S2). Although the relationships among Blennosperma, Crocidium, and Ischnea and
between these and other genera in the Tussilagininae s.s. clade
are not affected by well-supported incongruence, the relationship between Robinsonecio and the Arnoglossum-Barkleyanthus-Yermo clade shows incongruence that is well supported in
the BI trees. In contrast to being more distantly related in the
ITS/ETS phylogenies (Fig. 2), plastid data place Robinsonecio
sister to the Arnoglossum-Barkleyanthus-Yermo clade (Fig. 1).
Reciprocal exclusion of Robinsonecio and the ArnoglossumBarkleyanthus-Yermo clade (results not shown) in both the ITS/
ETS and plastid data sets, however, does not change the relationships in clade 4.
Within Senecioninae, Emilia, Jacobaea, and Packera have
branches that are much longer than other genera in this subtribe
(Appendix S2). These three genera form a clade in the ITS/ETS
MP trees (not shown); however, Packera is placed more distant
to the other two genera in the ITS/ETS BI trees (Fig. 2). Reciprocal exclusion of two of these three genera from MP analyses
(results not presented) resulted in the phylogenetic positions of
Emilia and Jacobaea remaining unchanged when the two other
genera were excluded (resp. Jacobaea and Packera, and Emilia
and Packera). Emilia and Jacobaea were therefore not affected
by long-branch attraction, but when both are excluded, Packera
occupies a different position in the ITS/ETS trees. Because the
results of the BI analyses of the ITS/ETS data agree with the
plastid trees in indicating a close relationship between the New
Identification of the major Senecioneae lineages and analyses of the incongruent patterns within them. Placeholders of the major groups were selected to study incongruent phylogenetic
patterns between them (see Table 5).
Forms clade in
ITS/ETS
trees
1. Tussilagininae s.s.
Y
Y
2. New World 1 groupc
N
Y
3. Quadridentates
Y
Y
4. Seneciod
N
N
Plastid trees (Fig. 1)
Arnoglossum-Barkleyanthus-Yermo clade sister
to Robinsonecio and Psacaliopsis (BS <50%,
PP 0.98)
Robinsonecio sister to the ArnoglossumBarkleyanthus-Yermo clade (BS 54%, PP 1.00)
Roldana sister to Digitacalia, Telanthophora,
Nelsonianthus, Pittocaulon, and Villasenoria
(BS 71%, PP 1.00)
Lomanthus sister to a clade formed by
Charadranaetes, Jessea, and Talamancalia (BS
97%, PP 1.00)
Caxamarca and Pseudogynoxys sister to
Dorobaea and Garcibarrigoa (BS 55%, PP
1.00)
Caucasalia, Dolichorrhiza, and Iranecio form
a clade (BS 91%, PP 1.00)
Senecio flavus is deeply nested within the
core of Senecio s.s. and a member of a clade
composed of S. nevadensis, S. viscosus, and
clade 6 (BS 89%, PP 1.00)
S. cadiscus and S. pinnatifolius are not most
closely related to S. nevadensis, S. viscosus,
and clade 6, which instead are more closely
related to other Senecio species and form
clade 7 with these (BS 96%, PP 1.00)
S. hispidissimus, S. psilocarpus, and S.
squarrosus are members of clade 7 (BS 96%,
PP 1.00)
S. triodon sister to S. gayanus and S.
lastarrianus (BS 100%, PP 1.00)
Senecio lineatus sister to S. hollandii (BS
100%, PP 1.00) and Bethencourtia sister to
Emilia (BS 77%, PP 1.00)
ITS/ETS trees (Fig. 2)
Arnoglossum-Barkleyanthus-Yermo clade sister
to Pippenalia-Psacalium clade (BS 75%, PP
1.00)
Psacaliopsis sister to Robinsonecio (BS 60%,
PP 0.96)
Roldana sister to the ArnoglossumBarkleyanthus-Yermo clade, Pippenalia, and
Psacalium (BS <50%, PP 0.99)
Lomanthus sister to a clade composed of
Misbrookea, Werneria, and Xenophyllum (BS
<50%, PP 0.97)
Caxamarca and Pseudogynoxys sister to the
Charadranaetes-Jessea-Tamancalia clade (BS
77%, PP 0.97)
Dolichorrhiza, Iranecio, and Pojarkovia form a
clade (BS <50%, PP 1.00)
Senecio flavus is placed basal to the core of
Senecio s.s. (BS 100%, PP 1.00)
Support for internal
conflict
BS PP
ILD test
(P)
Placeholder taxa
N
Y
0.114
Tephroseris
Y
Y
0.001
Caxamarca
N
Y
0.022
Caucasalia
Y
Y
0.001
S. viscosus
N
Y
0.026
-
Y
Y
0.015
Emilia
S. cadiscus and S. pinnatifolius are most closely
related to S. nevadensis, S. viscosus, and
members of Clade 6 (BS 99%, PP 1.00)
S. hispidissimus, S. psilocarpus, and S.
squarrosus sister to S. ilicifolius and S. vestitus
(BS 91%, PP 1.00)
S. triodon sister to S. fistulosus, S. gayanus, S.
jarae, and S. lastarrianus (BS 54%, PP 0.98)
Senecio lineatus sister to Bethencourtia
(MP: 93%) or is unresolved on a clade with
Bethencourtia and Jacobaea (BI: PP 0.97)
Y
Y
5. Emilia-Bethencourtia
group
Y
N
6. Senecio segregates
7. New World 2 group
8. Brachyglottidinae
9. Othonninae
10. Bolandia-MesogrammaStilpnogyne clade
11. Gynuroids
12. Synotoids
13. Madagascan group
14. Lachanodes-Senecio thapsoides
group
15. Caputia
Y
Y
Y
Y
Y
N
N
Y
Y
Y
N
N
N
N
N
N
N
N
N
N
0.9998
0.255
0.709
1
1
S. deltoideus
Dendrophorbium
Centropappus
Othonna
Bolandia
Y
Y
N
N
Y
N
Y
Y
N
N
N
N
N
N
N
N
0.97
0.157
1
1
Kleinia
Dauresia, Mikaniopsis
Faujasia
Arrhenechthites
Y
Y
n.a. n.a.
n.a.
Y
Y
n.a.
Y
Y
n.a.
n.a. n.a.
n.a. n.a.
n.a. n.a.
n.a.
n.a.
n.a.
Caputia medleywoodii (ined.)
Lamprocephalus
Crassocephalum
Capelio
16. Lamprocephalus-Oresbia clade
17. Crassocephalum-Erechtites clade
18. Capelio
863
Clade 5 within Senecio (Figs. 1, 2)
Pelser et al.—Incongruence between Senecioneae phylogenies
Plastid
trees
Major Senecioneae groupa
Conflicting relationships within groupb
May 2010]
Table 3.
American Journal of Botany
Notes: BS = incongruent clades supported with maximum parsimony bootstrap support ≥80% in plastid and ITS/ETS trees; PP = incongruent clades supported with Bayesian inference posterior
probabilities ≥0.95 in plastid and ITS/ETS trees; groups in boldface indicate strong internal incongruence according to the criteria used in this study (incongruent patterns supported by bootstrap
values ≥80% and/or PP ≥0.95 and ILD values of P < 0.01).
aSpecies/generic composition outlined in Figs. 1 and 2.
bOnly shown for groups in which incongruent relationships were supported with either MP bootstrap support ≥80% or BI posterior probabilities ≥0.95
cInternal incongruence for the phylogenetic position of Lomanthus.
dInternal incongruence for the phylogenetic positions of Senecio flavus, S. hispidissimus, S. psilocarpus, S. squarrosus, S. cadiscus, and S. pinnatifolius.
Chersodoma
Cineraria
Dendrosenecio
Io
Jacobaea
Lordhowea
Pericallis
Phaneroglossa
Steirodiscus
Senecio otites
Packera
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
19. Chersodoma
20. Cineraria
21. Dendrosenecio
22. Io
23. Jacobaea
24. Lordhowea
25. Pericallis
26. Phaneroglossa
27. Steirodiscus
28. Senecio otites
29. Packera
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
ILD test
(P)
BS PP
ITS/ETS trees (Fig. 2)
Plastid trees (Fig. 1)
ITS/ETS
trees
Plastid
trees
Major Senecioneae groupa
Table 3.
Continued.
Forms clade in
Conflicting relationships within groupb
Support for internal
conflict
Placeholder taxa
864
[Vol. 97
World 2 group and Packera, Packera may very well be a member of the New World 2 group.
Incomplete lineage sorting— Molecular dating analyses
using r8s yielded age estimates that were often older than ages
retrieved with BEAST (Figs. 4 and 5) and sometimes outside
the 95% HPD interval of the BEAST estimates (Appendix
S3). The 95% HPD intervals of the plastid and ITS/ETS analyses were often overlapping. Estimates of the duration of putative ILS events obtained with r8s are within or slightly
outside the ranges suggested by BEAST. Using these estimates and assuming generation times of 1, 2, 5, and 10 yr,
coalescent analyses indicate that for four of the 10 strongly
incongruent lineages that were examined (Caputia, the Lamprocephalus-Oresbia clade, the New World 2 generic group
and Packera, and Jacobaea) effective population sizes must
be assumed that are higher than the selected thresholds of
Ne = 20 000 (plastid data) and 40 000 (ITS/ETS data) to explain
their incongruent phylogenetic positions by ILS (Table 6). If
multiple strongly incongruent phylogenetic positions of lineages are assumed to be the result of single ILS events, even
higher effective population sizes of ancestral lineages are required (not shown).
DISCUSSION
Topological incongruence— One of the longstanding and increasingly prominent questions of phylogenetic systematics is
how to address incongruence between phylogenies obtained
from multiple data sets and how to determine its cause (e.g.,
Sullivan, 1996; Wiens and Hollingsworth, 2000; Rokas et al.,
2003; Holland et al., 2008; Degnan and Rosenberg, 2009). Incongruence between phylogenetic trees inferred using different
molecular markers may result from biological phenomena as
well as analytical artifacts. Among the potential causes are undetected paralogous sequences in one or more data sets (Doyle,
1992; Álvarez and Wendel, 2003); however, topological incongruence may also be found when different genomes or portions
of individual genomes may have different evolutionary histories (e.g., due to hybridization or ILS; Doyle, 1992; Maddison, 1997; Buckley et al., 2006; Liu and Pearl, 2007). Phylogeny
reconstruction methods may also introduce error via longbranch attraction (Graybeal, 1998; Wiens and Hollingsworth,
2000), which can be particularly problematic in data sets with a
relatively sparse taxon sampling and composed of highly divergent sequences. In addition, incongruence may result from errors in phylogeny reconstruction due to low numbers of
informative characters (sampling error; Huson and Bryant,
2006). Bootstrap values and posterior probabilities are often
used as measures of sampling error (Huson and Bryant, 2006).
The presence of well-supported (BS values ≥ 80% and/or PP ≥
0.95) incongruent patterns in the Senecioneae therefore suggests that these patterns are not due to sampling error.
Due to the multicopy nature of ribosomal DNA, differences
among ITS, ETS, and plastid trees may be due to incorrect ITS/
ETS homology assessments, confounding interpretations essentially through undetected paralogous copies (Álvarez and
Wendel, 2003). Cloning of ITS PCR products of incongruent
taxa or representatives of incongruent lineages indeed revealed
base-pair differences among ITS copies within individual samples. Parsimony analyses in which both directly sequenced PCR
products and sequences of ITS clones were included (results
May 2010]
Table 4.
865
Well-supported (BS ≥ 80% and/or PP ≥ 0.95) incongruent patterns among genera and major Senecioneae generic assemblages.
Lineage
Caputia
New World 2 group
Jacobaea
Packera
Lordhowea
Gynuroids
Lamprocephalus-Oresbia clade
Phaneroglossa
Cineraria
Steirodiscus
Emilia-Bethencourtia group
Senecio otites
Io
Pelser et al.—Incongruence between Senecioneae phylogenies
Plastid trees
ITS/ETS trees
Sister to the synotoids (BS 95%, PP 1.00)
Nested among core members of clade 1 (BS 100%, PP 0.92), not
among core members of clade 2 (BS 95%, PP 1.00)
Nested among core members of clade 1 (BS 100%, PP 0.92), not
among core members of clade 2 (BS 95%, PP 1.00)
Nested among core members of clade 1 (BS 100%, PP 0.92), not
among core members of clade 2 (BS 95%, PP 1.00)
Nested among core members of clade 1 (BS 100%, PP 0.92), not
among core members of clade 2 (BS 95%, PP 1.00). Sister to
Phaneroglossa (BS 98%, PP 1.00)
Nested among core members of clade 1 (BS 100%, PP 0.92), not
among core members of clade 2 (BS 95%, PP 1.00)
Nested among core members of clade 1 (BS 100%, PP 0.92), not
among core members of clade 2 (BS 95%, PP 1.00). Sister to the
Gynuroids, Jacobaea, Packera, the New World 2 group and the
core groups of Clade 1 nested between them (BS 100%, PP 0.92)
Nested among core members of clade 1 (BS 100%, PP 0.92), not
among core members of clade 2 (BS 95%, PP 1.00)
Core member of clade 2. Not placed in the clade composed of
Dendrosenecio, the Quadridentates, New World 1 group, the
Madagascan group, Io, the Emilia-Bethencourtia group, the
Senecio segregates group, Pericallis, the Bolandia-MesogrammaStilpnogyne clade, and Senecio otites (BS 95%, PP 1.00)
Core member of clade 2. Not placed in the clade composed of
Dendrosenecio, the Quadridentates,
New World 1 group, the Madagascan group, Io, the EmiliaBethencourtia group, the Senecio segregates group, Pericallis,
the Bolandia-Mesogramma-Stilpnogyne clade, and Senecio
otites (BS 95%, PP 1.00)
Core member of clade 2. Sister to Pericallis, the New World 1
group, Senecio otites, Io, and the Madagascan group (BS <50%,
PP 1.00)
Core member of clade 2. Sister to the New World 1 group (BS
99%, PP 1.00)
Core member of clade 2. Forms a clade with the Madagascan
group (BS 70%, PP 0.99)
not shown), however, showed that all cloned sequences of a
taxon consistently form one clade including the directly sequenced ITS products of the same taxon. Similarly, for species
and genera for which ITS sequences obtained from different
specimens or species were included in our data sets, these different accessions were always resolved as each other’s closest
relatives. Unless copies of greater divergence have been lost or
remained undiscovered (a possibility that cannot be confirmed
nor excluded), it is unlikely that paralogy is the cause of the
widespread ITS/ETS vs. plastid incongruence found in the
Senecioneae.
Long-branch attraction (Felsenstein, 1978) is another possible explanation for the topological incongruence observed. Reciprocal exclusion of accessions with long branches in MP and
BI analyses showed that only the phylogenetic position of
Packera in the ITS/ETS MP trees appears to be affected by
long-branch attraction. Our results indicate that this is caused
by the long branches of Emilia and Jacobaea. Long-branch attraction, however, does not explain the plastid vs. ITS/ETS incongruence regarding the position of Packera, which persists
when Emilia and Jacobaea are excluded from the MP analyses
(not shown). Although more detailed, tree-wide analyses aimed
at identifying branches that may be susceptible to long-branch
Sister to the Brachyglottidinae (BS 100%, PP 1.00)
Nested among core members of clade 2, except
Dendrosenecio (BS 97%, PP 1.00)
Nested among core members of clade 2, except
Dendrosenecio (BS 97%, PP 1.00)
Nested among core members of clade 2, except
Dendrosenecio (BS 97%, PP 1.00)
Nested among core members of clade 2, except
Dendrosenecio (BS 97%, PP 1.00). Sister to the
Gynuroids, Jacobaea, Packera, the New World 2 group
and the core groups of clade 2 in which they are nested
(BS 97%, PP 1.00)
Nested among core members of clade 2, except
Dendrosenecio (BS 97%, PP 1.00)
Sister to Dendrosenecio (core member of clade 2) and
Phaneroglossa (BS 99%, PP 1.00)
Most closely related to Dendrosenecio (core member of
clade 2) and the Lamprocephalus-Oresbia clade (BS 99%,
PP 1.00)
Forms a clade with, among others, New World 1 group,
the Madagascan group, Io, the Emilia-Bethencourtia
group, the Senecio segregates group, Pericallis, the
Bolandia-Mesogramma-Stilpnogyne clade, and Senecio
otites (BS 100%, PP 1.00)
Forms a clade with, among others, New World 1 group,
the Madagascan group, Io, the Emilia-Bethencourtia
group, the Senecio segregates group, Pericallis, the
Bolandia-Mesogramma-Stilpnogyne clade, and Senecio
otites (BS 100%, PP 1.00)
Core member of clade 2. Sister to species of the Senecio
segregates group of which S. deltoideus, S. scandens, and
S. saxatilis form a clade with Emilia (BS <50%, PP 1.00)
Core member of clade 2. Sister to Pericallis and the New
World 2 group (BS <50%, PP 0.95)
Core member of clade 2. Does not form a clade with the
Madagascan group, which instead is sister to the New
World 1 group (BS <50%, PP 1.00)
attraction have not been performed (e.g., Huelsenbeck, 1997;
Wiens and Hollingsworth, 2000; Johnson et al., 2008), visual
comparisons of MP and BI trees (the latter claimed to be less
susceptible to long-branch attraction; e.g., Bergsten, 2005) did
not reveal well-supported incongruence related to strongly supported incongruent patterns between plastid and ITS/ETS
trees.
Hybridization and ILS are important biological explanations
for incongruence between data sets and are often difficult to
distinguish from each other (Doyle, 1992; Seelanan et al., 1997;
Holder et al., 2001; Buckley et al., 2006; Holland et al., 2008;
Joly et al., 2009). The detection of hybrids can for instance be
obscured by backcrossing, introgression, extinction of parental
species, and secondary hybridization (Doyle, 1992). Hybridization events may also be difficult to recognize when they result
in homoploid hybrids, are ancient, or were followed by speciation or dispersal in combination with extinction in the parental
distribution area. Hybridization has been demonstrated within
many Senecioneae genera (e.g., Bedfordia, Blennosperma,
Brachyglottis, Cineraria, Crassocephalum, Dendrosenecio,
Dolichoglottis, Emilia, Euryops, Farfugium, Hubertia, Jacobaea, Jessea, Ligularia, Packera, Petasites, Senecio, Traversia; Nordenstam, 1963, 1968, 1978, 1996; Ornduff, 1964;
American Journal of Botany
866
[Vol. 97
Table 5.
Incongruence length difference (ILD) tests performed to study incongruent patterns among placeholders of the major Senecioneae groups (see
Table 3).
ILD test
Test 1
Test 2
Test 3
Test 4
Test 5
Test 6
Test 7
Test 8
Test 9
Test 10
Test 11
Test 12
Test 13
Test 14
Test 15
Test 16
Test 17
Test 18
Test 19
Test 20
Accessions included
Placeholders of all major groups
Placeholders of all major groups, except for those in well-supported incongruent phylogenetic positions (BS
≥80% and/or PP ≥0.95; Caputia, the New World 2 group, Jacobaea, Lordhowea, gynuroids, LamprocephalusOresbia clade, Phaneroglossa, Cineraria, Steirodiscus, Emilia-Bethencourtia group, Senecio otites, Io, and
Packera)
Placeholders of test 2 minus placeholder of Othonninae
Placeholders of test 3 plus placeholder of Caputia
Placeholders of test 3 plus placeholder of New World 2 group
Placeholders of test 3 plus Jacobaea
Placeholders of test 3 plus Lordhowea
Placeholders of test 3 plus placeholder of the Gynuroids
Placeholders of test 3 plus placeholder of the Lamprocephalus-Oresbia clade
Placeholders of test 3 plus Phaneroglossa
Placeholders of test 3 plus Cineraria
Placeholders of test 3 plus Steirodiscus
Placeholders of test 3 plus Packera
Placeholders of test 3 plus placeholder of the Emilia-Bethencourtia group
Placeholders of test 3 plus Senecio otites
Placeholders of test 3 plus Io
Placeholders of test 3 plus placeholder of the Emilia-Bethencourtia group, Io, and Senecio otites
Placeholders of test 3 plus placeholder of the Emilia-Bethencourtia group and Senecio otites
Placeholders of test 3 plus Io and Senecio otites
Placeholders of test 3 plus Io and placeholder of the Emilia-Bethencourtia group
Chapman and Jones, 1971; Drury, 1973; Olorode and Olorunfemi,
1973; Jeffrey, 1986; Yamaguchi and Yahara, 1989; Beck et al.,
1992; Lowe and Abbott, 2000; Kirk et al., 2004; Kadereit et al.,
2006; Cron et al., 2008; Pan et al., 2008; Vanijajiva and
Kadereit, 2009) and is therefore a likely hypothesis for explaining incongruence between plastid and ITS/ETS phylogenies.
ILS is especially likely when species rapidly radiate and
population sizes are large (Maddison, 1997). Because of the
stochastic nature of the coalescence process, ILS may yield
gene trees with random patterns of relationships among taxa
(Buckley et al., 2006), which may result in gene tree–species
tree incongruence. Among other methods, gene tree parsimony (Page and Charleston, 1997) or Bayesian hierarchical
model approaches based on coalescent theory (Liu and Pearl,
2007) can be used to reconstruct species trees from gene trees
that are incongruent due to lineage sorting. However, these
approaches do not account for hybridization (Liu and Pearl,
2007), which can result in similar phylogenetic patterns
(Buckley et al., 2006). Using coalescent-based approaches,
we can distinguish ILS from hybridization by testing whether
patterns of incongruence are random (ILS) or nonrandom (hybridization; Buckley et al., 2006). These studies, however,
require more than two unlinked genomic data sets to distinguish between both hypotheses (Buckley et al., 2006). Because our Senecioneae data were composed of only two
unlinked data sets (ITS/ETS region and the plastid markers),
this approach could not be used to test whether patterns of
incongruence in the Senecioneae are random or not. Instead,
we used the assumption that ancestral polymorphisms coalesce within approximately 5Ne generations (Rosenberg,
2003; Degnan and Rosenberg, 2009) to assess whether ILS
may be regarded as a plausible explanation for the observed
incongruence. This approach requires information about generation times and the duration of the ILS events that could be
invoked to explain incongruent patterns, as well as effective
population sizes in the Senecioneae. Because of the relatively
ILD test
results (P)
0.001
0.006
0.052
0.001
0.001
0.001
0.001
0.001
0.001
0.001
0.002
0.001
0.001
0.084
0.024
0.020
0.001
0.001
0.006
0.013
ancient nature of many of the historical events resulting in the
incongruence and the diversity and enormity of the tribe, making assumptions about effective population sizes and generation times of ancestral lineages is somewhat precarious.
Furthermore, calculations of the duration of putative ILS
events rest on our molecular dating analyses, which resulted
in large 95% HPD intervals of age estimates and substantial
differences between the dating results of the BEAST and r8s
analyses and between estimates obtained from plastid and
ITS/ETS sequences. We therefore used a conservative approach in estimating these parameters. Coalescence calculations were performed with generation times of 1 and 2 yr,
which are assumed to be typical for most Senecioneae species. In some species, however, longer generation times are
expected, particularly in species with a tree-like (e.g., Aequatorium, Brachyglottis , Dendrosenecio, Lachanodes , Nordenstamia, Pladaroxylon) or succulent (e.g., Curio, Kleinia ,
Othonna) habit or those forming seed banks (e.g., Euryops
annuus, Jacobaea vulgaris). Calculations were therefore
also carried out for generation times of 5 and 10 yr. Senecioneae populations are generally small, with individuals
countable in dozens or hundreds, or containing up to 5000
plants (e.g., Wid é n and Andersson, 1993 ; Comes and Abbott, 1998 ; Golden, 1999 ; Panero et al., 1999 ; M ü llerSch ä rer and Fischer, 2001 ). More rarely, larger populations
are observed, which may comprise several hundreds of
thousands of plants (e.g., Euryops annuus ). Although few
estimates of N e of plant populations based on genetic data
have been published ( Siol et al., 2007 ), N e/ N ( N being the
census number of reproductive individuals) ratios of approximately 5–10% have been suggested as typical (Siol et al.,
2007). The selected values of Ne = 40 000 for the ITS/ETS
data and N e = 20 000 for the plastid data (due to the 1/2
reduction in effective population size necessary for the
plastid markers relative to the nuclear markers in hermaphroditic individuals) are therefore conservative estimates of
Pelser et al.—Incongruence between Senecioneae phylogenies
May 2010]
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Fig. 3. Bayesian inference (BI) topology from the combined plastid-ITS/ETS data set in which each strongly incongruent taxon (in bold) is included
twice: once as a plastid-only accession (ITS/ETS sequences coded as missing data) and once as an ITS/ETS-only accession (plastid sequences coded as
missing data). Bayesian consensus percentages (posterior probabilities × 100) are placed above branches, bootstrap support above 50% below branches.
Relationships between the outgroup species are not shown.
American Journal of Botany
868
H
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maximum effective population sizes. To accommodate for
uncertainty in the calculations of the duration of ILS events,
the coalescent analyses were performed with both minimum
and maximum durations using the 95% HPD intervals of the
BEAST estimates extended with the occasional outliers
from the r8s analyses.
Although ILS could not be eliminated as a possible explanation for the incongruent phylogenetic positions for six of the 10
strongly incongruent lineages that were examined, our coalescent analyses were able to demonstrate that ILS cannot be invoked to explain incongruence regarding the relationships of
Caputia, the Lamprocephalus-Oresbia clade, the New World 2
Pelser et al.—Incongruence between Senecioneae phylogenies
May 2010]
869
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epidosparlum burgessii
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schnea elachoglossa
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Robinsonia evenia
Robinsonia masafuerae
Robinsonia berleroi
Senecio acanthifolius
Senecio hieracium
Senecio amicoides
Senecio triodon
Senecio lastarrianus
Senecio fistulosus
Senecio jarae
Oauresia alliariifolia
Phaneroglossa bolusii
Oendrosenecio kilimanjari
L amprocephalus montanus
Oresbia heterocarpa
Austrosynotis reclirama
Humberlacalia sp.
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Brachyglottis repanda
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Senecio lauricola
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Senecio nemorensis
Senecio elegans
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Senecio pinnatifolius
Senecio oreophyton
Senecio flaccidus
Senecio mairetianus
Senecio niveo-aureus
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Delairea odorata
Senecio meuselii
Gynura Formosana
Kleinia neriifolia
Solanecio biafrae
Stefrodfscus capillaceus
Cineraria abyssinica
Mesogramma apiifolium
o/andia peduncu/osa
tilpnogyne bellidioides
$ enecio ho/landii
$ enecio letifolius
$ enecio retrorsus
$ enecio triqueter
$ enecio lineatus
Bethencourlia palmensis
J acobaea vulgaris
Emilia coccinea
$ enecio saxatilis
$ enecio delloideus
$ enecio scandens
o ambondrombeensis
Huberlia riperia
aujasia squamosa
aujasiopsis flexuosa
riothrix Iycopodioides
arafaujasia fontinalis
Monticalia arbutifo/ia
abrerielle oppositicOrdia
entacalia arborea
Monticalia abietina
crobicaria ilicifolia
L omanthus fosbergll
enophyl/um poposum
Misbrookea strigosissima
Wemeria caespitosa
0 orobaea pimpincllifolia
Garcibarrigoa telembina
Caxamarca sanchezii
seudogynoxys haenkei
alamancalia boquelensis
haradranaetes durandii
essea mullivenia
Monticalia apiculata
Senecio otites
Graphistylis dichroa
H oehnephytum trlxoldes
rbelaezaster ellsworthii
I
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セ@
ヲM
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Fig. 5.
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Pericallis aurita
セ@
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/.nurbia constanzae
L eonis trineurus
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,
Ekmaniopappus mikanioides
Nesampelos lucens
H erreranthus rivalis
A ntillanthus almironcillo
S hafera platyphylla
Lundinia plumbea
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ITS/ETS BEAST chronogram. Strongly incongruent taxa and groups of taxa in boldface.
Omya
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American Journal of Botany
870
[Vol. 97
Table 6.
Results of the coalescent analyses in which effective population (Ne) sizes were calculated for a selection of the strongly incongruent lineages
assuming that the incongruent patterns were caused by incomplete lineage sorting (ILS). Coalescence was assumed to occur within 5Ne generations,
and calculations were performed for generation times of 1, 2, 5, and 10 yr. Estimates of the duration of putative ILS events were calculated with
BEAST and r8s (Figs. 4, 5).
Ne (×1000) for assumed generation times
Strongly incongruent lineage
Data set
Duration of putative ILS
(Myr)
1 yr
2 yr
5 yr
10 yr
Caputia
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
Plastid
ITS/ETS
2.26–11.97
3.73–14.28
4.22–9.3
1.7–8.23
0–5.41
6.67–11.43
0–5.41
5.98–13.13
1.03–6.97
5.95–13.62
4.7–9.79
2.84–7.77
8.08–13.38
5.29–12.41
0–3.45
0.06–8.37
1.66–5.64
0–1.39
1.05–6.3
0–2.98
452–2394
746–2856
844–1860
340–1646
up to 1082
1334–2286
up to 1082
1196–2626
206–1394
1190–2724
940–1958
568–1554
1616–2676
1058–1482
up to 690
12–1647
332–1128
up to 278
210–1260
up to 596
226–1197
373–1428
422–930
170 to 823
up to 541
667–1143
up to 541
598–1313
103–697
595–1362
470–979
284–777
808–1338
529–1241
up to 345
6–837
166–564
up to 139
105–630
up to 298
90.4–478.8
149.2–571.2
168.8–372
68–329.2
up to 216.4
266.8–457.2
up to 216.4
239.2–525.2
41.2–278.8
238–544.8
188–391.6
113.6–310.8
323.2–535.2
211.6–496.4
up to 138
2.4–334.8
66.4–225.6
up to 55.6
42–252
up to 119.2
45.2–239.4
74.6–285.6
84.4–186
34–164.6
up to 108.2
133.4–228.6
up to 108.2
119.6–262.6
20.6–139.4
119–272.4
94–195.8
56.8–155.4
161.6–267.6
105.8–148.2
up to 69.0
1.2–164.7
33.2–112.8
up to 27.8
21.0–126.0
up to 59.6
Gynuroids
Lordhowea
Phaneroglossa
Lamprocephalus and Oresbia
New World 2 group and Packera
Jacobaea
Cineraria and Steirodiscus
Lomanthus
Senecio otites
generic group and Packera, and Jacobaea (Table 6). To result
in the incongruent patterns, ILS events must have continued for
at least 1.03 (Lamprocephalus-Oresbia clade) to 5.29 (Jacobaea) Myr with effective population sizes never falling below
20 600 (Lamprocephalus-Oresbia clade) to 105 800 (Jacobaea)
individuals. This is extremely improbable considering the present day population sizes of Senecioneae species and the likely
occurrence of bottlenecks relative to speciation events throughout the evolutionary history of these lineages. For these four
lineages, ancient hybridization is a much more likely explanation, although additional morphological, karyological, and molecular studies will need to be performed to identify direct
evidence for hybrid origins.
Conclusions— This study reveals new insights into the evolutionary history of Senecioneae by demonstrating strongly
supported incongruence between plastid and ITS/ETS phylogenies. This incongruence is found at various taxonomic levels
and affects the phylogenetic positions of six Senecio species,
nine genera (Caputia, Cineraria, Steirodiscus, Phaneroglossa,
Lordhowea, Jacobaea, Packera, Senecio otites, and Lomanthus), and three generic assemblages (the LamprocephalusOresbia clade, the gynuroids, and the New World 2 group).
Although the emphasis of this study has been on understanding
the incongruence between plastid and ITS/ETS trees, it is important to note that our analyses also support many phylogenetic patterns that are congruent or that are only weakly
incongruent in the Senecioneae. In fact, the plastid data confirms many of the taxonomic and phylogenetic conclusions that
were drawn from the ITS/ETS trees presented in this study and
the ITS trees in Pelser et al. (2007). For example, except for
Caputia, the subtribal delimitation of Senecioneae identified
with ITS sequences (Pelser et al., 2007) remains largely unaffected by the incongruence, well-supported generic-level incongruence was not found within subtribes Brachyglottidinae,
Tussilagininae s.s., and Othonninae, and plastid data provided
additional support for the new delimitation of Senecio proposed
by Pelser et al. (2007). Therefore, the congruent patterns found
in this comprehensive study of the large tribe Senecioneae provide a framework (Fig. 3) for future research focused on the
incongruent phylogenetic positions of its lineages.
In addition to a better understanding of the patterns of phylogenetic incongruence and the taxa involved, our studies also
provided information about the potential causes of topological
incongruence in the Senecioneae. The dense generic-level sampling used in our analyses, the lack of evidence for ITS/ETS
paralogy issues, and the paucity of long-branch attraction associated with the incongruent taxa suggest that ILS and/or hybridization are the most likely causes of the plastid vs. ITS/ETS
incongruence. Using a novel approach based on coalescenttheory, we were able to show that ILS is unlikely to be responsible for four of the 10 strongly incongruent patterns that were
observed. Our study therefore indicates that this approach may
be a valuable tool for testing whether ILS can be invoked as an
explanation of phylogenetic incongruence.
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Appendix S1. Sequences of multiple accessions.
3’trnK. Abrotanella emarginata (Gaudich.) Cass., EF537909; Acrisione cymosa
(J.Rémy) B.Nord., Lammers, Rodriques & Baeza 6447 (MU), GU817416; Adenostyles
alpina (L.) Bluff & Fingerh., Schwerdtfeger 12251 (B), GU817417; Aequatorium
asterotrichum B.Nord., Asplund 18263 (S), GU817418; Anacyclus valentinus L., Tepe
1186 & Moreno-Paez (MU), GU817419; Antillanthus almironcillo (M.Gómez)
B.Nord., El-Ghazaby et al. 23.608 (S), GU817420; Arbelaezaster ellsworthii (Cuatrec.)
Cuatrec., Kilip & Smith 19413 (US), GU817421; Arnoglossum atriplicifolium (L.)
H.Rob., EF537911; Arrhenechthites mixta (A.Rich.) Belcher, EF537912; Austrosynotis
rectirama (Baker) C.Jeffrey, LaCroix 4001 (WAG), GU817423; Bahia dissecta (Gray)
Britton, Tuhy 3868 (MU), GU817424; Barkleyanthus salicifolius (Kunth) H.Rob. &
Brettell, Genelle & Fleming 861 (B), GU817425; Barnadesia sp., Hunziker & Gamerro
12561 (MU), GU817426; Bedfordia arborescens Hochr., Greuter 21319 (B),
GU817427; Bethencourtia palmensis (Nees) Choisy, Nordenstam 9326 (S), GU817428;
Blennosperma nanum S.F.Blake, AF460029; Blepharispermum zanguebaricum Oliv.
& Hiern., Medley 570 (MU), GU817429; Bolandia pedunculosa (DC.) Cron, Cron &
Goodman 681 (J), GU817430; Brachyglottis repanda J.R.Forst. & G.Forst.,
Schwerdtfeger 17407 (B), GU817431; Cabreriella oppositicordia (Cuatrec.) Cuatrec.,
Romero-Castañeda 7428 (US), GU817432; Cacaliopsis nardosmia (A.Gray) A.Gray,
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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Ertter 6502 (UC), GU817433; Calendula arvensis L., Watson 95-13C (MU),
GU817435; Capelio caledonica B.Nord., Nordenstam 9644 (S), GU817436; Caucasalia
parviflora (M.Bieb.) B.Nord., EF537913; Caxamarca sanchezii M.O.Dillon & Sagást.,
Sagástegui et al. 15548 (F), GU817437; Centropappus brunonis Hook.f., Wapstra MW2
(MEL), GU817438; Chaenactis douglasii (Hook.) Hook. & Arn., Vincent 8582 (MU),
GU817439; Charadranaetes durandii (Klatt) Janovec & H.Rob., Gomez P. 2242 (S),
GU817440; Chersodoma jodopappa (Sch.Bip.) Cabr., EF537914; Cichorium intybus
L., Tepe 1673 (MU), GU817441; Cineraria abyssinica Sch.Bip. ex A.Rich., AF460028;
Cirsium discolor (Muhl.) Spreng., Tepe 1670 (MU), GU817442; Cissampelopsis
volubilis (Blume) Miq., Carvalho 3175 (MU), GU817443; Coreopsis sp., Vincent 13226
(MU), GU817444; Corymbium enerve Markötter, Trinder-Smith 124 (US), GU817445;
Cotula coronopifolia L., Watson & Panero 94-26 (MU), GU817446; Crassocephalum
crepidioides (Benth.) S.Moore, AF459991; Cremanthodium humile Maxim.,
EF537915; Crocidium multicaule Hook., Bartholomew 5749 (MO), GU817447; Curio
rowleyanus (H.Jacobsen) P.V.Heath, AF460002; Dauresia alliariifolia (O.Hoffm.)
B.Nord & Pelser, Müller & Tilson 907 (M), GU817448; Delairea odorata Lem., Tepe
2180, Pelser & Marticorena (MU), GU817449; Dendrophorbium bomanii (R.E.Fr.)
C.Jeffrey, EF537917; Dendrosenecio kilimanjari (Mildbr.) E.B.Knox ssp. cottonii
(Hutch. & G.Taylor) E.B.Knox, AF460027; Digitacalia jatrophoides (Kunth) Pippen,
Panero 2330 & Salinas (TEX), GU817450; Dolichoglottis lyallii (Hook.f.) B.Nord.,
Strid 22172 (MO), GU817451; Dolichorrhiza caucasica (M.Bieb.) Galushko,
Schneeweiss 44 (WU), GU817452; Dorobaea pimpinellifolia (Kunth) B.Nord., Tepe
1467 (MU), GU817453; Doronicum pardalianches L., EF537918; Ekmaniopappus
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mikanioides (Urb. & Ekman) Borhidi, Nordenstam & Lundin 500 (S), GU817454;
Elekmania picardae (Krug & Urb.) B.Nord., EF537919; Emilia coccinea (Sims)
G.Don., AF460026; Endocellion sibiricum (J.F.Gmel.) J.Toman, Schönswetter &
Tribsch T480 (WU), GU817455; Erato polymnoides DC., Tepe 1456 (MU), GU817456;
Erechtites valerianifolius (Link ex Spreng.) Less. ex DC., EF537920; Eupatorium
serotinum Michx., Tepe 1667 (MU), GU817457; Euryops pectinatus (L.) Cass.,
AF460025; Farfugium japonicum (L.) Kitam., Wu 1439 (MO), GU817458; Faujasia
squamosa (Bory) C.Jeffrey, Nordenstam 9210 (S), GU817459; Faujasiopsis flexuosa
(Lam.) C.Jeffrey ssp. bourbonensis C.Jeffrey, Nordenstam 9202 (S), GU817460;
Galinsoga quadriradiata Cav., Vincent 13227 (MU), GU817461; Garcibarrigoa
telembina (Cuatrec.) Cuatrec., Holm-Nielsen et al. 6211 (S), GU817462; Gerbera sp.,
Vincent 13223 (MU), GU817463; Graphistylis dichroa (Bong.) D.J.N.Hind, Silva 140
& Zelma (S), GU817464; Gymnodiscus capillaris (L.f.) Less., Coppejans EC1116 (U),
GU817465; Cron & Goodman 650 (J), GU817466; Gynoxys soukupii Cuatrec.,
AF459989; Gynura divaricata (L.) DC. ssp. formosana (Kitam.) F.G.Davies,
AF459988; Helenium autumnale L., Tepe 1664 (MU), GU817467; Helianthus
tuberosus L., Tepe 1672 (MU), GU817468; Helichrysum stoechas DC., Tepe 1092
(MU), GU817469; Herodotia haitiensis Urb. & Ekman, EF537922; Herreranthus
rivalis (Greenm.) B.Nord., Nordenstam s.n. (2/12/2007) (S), GU817470; Hertia pallens
(DC.) Kuntze, Koekemoer & Funk 1963 (US), GU817471; Hoehnephytum trixoides
(Gard.) Cabr., Carmo 131 (BHCB), GU817472; Homogyne alpina (L.) Cass., Uhink
98-56 (MJG), GU817473; Hubertia riparia (DC.) C.Jeffrey, Labat, Phillipson & Lowry
II 2022 (WAG), GU817474; Humbertacalia sp., Phillipson et al. 5641 (P), GU817475;
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Inula viscosa (L.) Aiton, Tepe 1142 (MU), GU817476; Io ambondrombeensis
(Humbert) B.Nord., Malcomber et al. s.n. (P), GU817477; Iranecio cariensis (Boiss.)
C.Jeffrey, Budak 1724, Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu),
GU817478; Ischnea elachoglossa F.Muell., Beaman 12129 (MO), GU817479; Jacmaia
incana (Sw.) B.Nord., Nordenstam 7914 (S), GU817480; Jacobaea vulgaris Gaertn.,
AF460007; AY156987; Jessea multivenia (Benth.) H.Rob. & Cuatrec., Nordenstam
9161 (S), GU817481; Kleinia neriifolia Haw., AF460024; Lachanodes arborea (Roxb.)
B.Nord., Cairns-Wicks s.n., GU817482; Lactuca canadensis L., Vincent 13128 (MU),
GU817483; Lamprocephalus montanus B.Nord., Nordenstam 9542 (S), GU817484;
Leonis trineurus (Griseb.) B.Nord., Smith & al. 3238 (S), GU817485; Lepidospartum
burgessii B.L.Turner, EF537925; Ligularia stenocephala (Maxim.) Matsum. &
Koidz., AF460023; Lomanthus fosbergii (Cuatrec.) B. Nord. & Pelser, Øllgaard et al.
90697 (AAU), GU817561; Lopholaena coriifolia (Sond.) Phillips & C.A.Sm., Van der
Westhuizen & Liamé s.n. (PREM), GU817486; Lordhowea insularis (Benth.) B.Nord.,
AF459987; Luina hypoleuca Benth., Greuter 17706 (B), GU817487; Lundinia plumbea
(Griseb.) B.Nord., Zanoni 45816 (JBSD), GU817488; Madia sp., Vincent 12912 (MU),
GU817489; Mesogramma apiifolium DC., Giess 16074 (M), GU817490; Mikaniopsis
clematoides (Sch.Bip. ex A.Rich.) Milne-Redh., De Wilde & De Wilde-Duyfjes 9006
(B), GU817491; Monticalia apiculata (Sch.Bip. ex Wedd.) C.Jeffrey, Alston 6935 (S),
GU817492; Monticalia arbutifolia (Kunth.) C.Jeffrey, EF537926; Nelsonianthus
tapianus (B.L.Turner) C.Jeffrey, Pérez & Kendizabal 457 (XAL), GU817493;
Nemosenecio nikoensis (Miq.) B.Nord., Koyama 4079 (L), GU817494; Nesampelos
lucens (Poir. in Lam.) B.Nord., EF537927; Neurolaena lobata (L.) R.Br., Acevedo
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9316 (US), GU817495; Nordenstamia kingii (H.Rob. & Cuatrec.) B.Nord., Ståhl
5572A (S), GU817496; Oldfeltia polyphlebia (Griseb.) B.Nord. & Lundin, EF537929;
Oresbia heterocarpa Cron & B.Nord., Nordenstam 9628 (S), GU817497; Othonna
capensis Bailey, AF460022; Packera eurycephala (Torr. & A.Gray) W.A.Weber &
Á.Löve, EF537930; Papuacalia dindondl (P.Royen) Veldkamp, EF537931; Paracalia
jungioides (Hook. & Arn.) Cuatrec., King & Collins 9017 (UC), GU817498;
Parafaujasia fontinalis (Cordem.) C.Jeffrey, Lorence 2602 (MO), GU817499;
Paragynoxys neodendroides (Cuatrec.) Cuatrec., Cleef 9852a (U), GU817500;
Parasenecio adenostyloides (Franch. et Sav. ex Maxim.) H.Koyama, AF459992;
Pentacalia arborea (Kunth) H.Rob. & Cuatrec., Øllgaard & Balslev 8298 (U),
GU817501; Pericallis murrayi (Bornm.) B.Nord., Royl 446 (B), GU817502; Perityle
emoryi Torr., Atwood 26483 (MU), GU817503; Petasites albus (L.) Gaertn., Cubr
40398 (B), GU817504; Phaneroglossa bolusii (Oliv.) B.Nord., AF459984; Pippenalia
delphinifolia (Rydb.) MacVaugh, Spellenberg & Bacon 11048 (TEX), GU817505;
Pittocaulon praecox (Cav.) H.Rob. & Brettell, EF537932; Pladaroxylon leucadendron
(G.Forst.) Hook.f., Cairns-Wicks s.n., GU817506; Pojarkovia pojarkovae (Schischk.)
Greuter, Pelser cult. 191 (L), GU817507; Polymnia canadensis L., Tepe 1665 (MU),
GU817508; Psacaliopsis purpusii (Greenm. ex Brandegee) H.Rob. & Brettell, Panero
2607, Davila & Tenorio (TEX), GU817509; Psacalium cirsiifolium (Zucc.) H.Rob. &
Brettell, EF537928; Pseudogynoxys haenkei (DC.) Cabr., Molina R. 18431 (U),
GU817510; Rainiera stricta (Greene) Greene, Dennis 2317 (U), GU817511;
Robinsonecio gerberifolius (Sch.Bip. ex Hemsl.) T.M.Barkley & Janovec, Garcia P.
171 (MO), GU817512; Robinsonia berteroi (DC.) R.W.Sanders, Stuessy & Martic.,
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AF459982; R. evenia Phil., Stuessy et al. 11308 (CONC), GU817513; R. gracilis
Decne., EF537933; Stuessy et al. 11282 (CONC), GU817514; Stuessy et al. 11312
(CONC), GU817515; R. masafuerae Skottsb., Stuessy et al. s.n. (CONC), GU817516;
Roldana suffulta (Greenm.) H.Rob. & Brettell, Rzedowski 36569 (UC), GU817517;
Rugelia nudicaulis Shuttlew. ex Chapm., Feist, Phillippe, Molano-Flores, Busemeyer
& Carroll 714 (TENN), GU817518; Senecio acanthifolius Hombr. & Jacq., ex Decne.,
EF537923; S. adamantinus Bong., EF537934; S. algens Wedd., Beck 2879 (S),
GU817519; S. arnicoides Hook. & Arn., Zöllner 3474 (L), GU817520; S. boyacensis
(Cuatrec.) Cuatrec., Gonzalez 180 (S), GU817521; S. brasiliensis (Spreng.) Less.,
UEC 50.171 (UEC), GU817522; S. cadiscus B.Nord. & Pelser, Rourke 1118 (S),
GU817434; S. chilensis Less., EF537936; S. ctenophyllus Phil., Zöllner 3959 (L),
GU817524; S. culcitioides Wedd., Øllgaard & Balslev 8822 (U), GU817523; S.
deltoideus Less., EF537937; S. elegans L., Cron & Goodman 687 (J), GU817525; S.
fistulosus Poepp. ex Less., Beck & Liberman 9672 (S), GU817526; S. flaccidus Less.
var. flaccidus, Jardin Thuret cult. s.n. (MJG), GU817527; S. flavus (Decne.) Sch.Bip.,
DQ208168; S. gayanus (Colla) Cabr., Rosas 2157 (INIA), GU817528; S. gramineus
Harv., Hoener 2104 (WAG), GU817529; S. gregorii F.Muell., Albrecht 7091 (NT),
GU817530; S. hemmendorffii Malme, EF537938; S. hieracium Remy, Baeza & Finot
3695 (CONC), GU817531; S. hispidissimus I.Thomps., Thompson 927 (MEL),
GU817532; S. hypsobates Wedd., Øllgaard & Balslev 9863 (U), GU817533; S.
ilicifolius Thunb., Cron & Goodman 686 (J), GU817534; S. integerrimus Nutt. var.
exaltatus (Nutt.) Cronquist, Crockett 437 (MU), GU817535; S. involucratus (Kunth)
DC., EF537910; S. jarae Phil., EF537939; S. lastarrianus Remy, Ricardi 3230 (B),
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GU817536; S. mairetianus DC., EF537942; S. medley-woodii Hutch. (Caputia medleywoodii ined.), AF460005; S. meuselii Rauh, EF537943; S. nemorensis L., AF460004;
S. nevadensis Boiss. & Reut., Vogt 4172 (B), GU817537; S. niveo-aureus Cuatrec.,
Cleef 6665 (S), GU817538; S. oerstedianus Benth. ex Oerst., Nordenstam 9160 (S),
GU817539; S. oreophyton Remy, Beck 21589 (S), GU817540; S. patens (Kunth) DC.,
Zak & Jaramillo 3427 (L), GU817541; S. pflanzii (Perkins) Cuatrec., EF537916; S.
pinifolius (L.) Lam., EF537945; S. pinnatifolius A.Rich. var. lanceolatus (Benth.)
I.Thomps., Thompson 904 (MEL), GU817542; S. polygaloides Phil., Zöllner 5442 (L),
GU817543; S. prenanthoides A.Rich., Thompson 915 (MEL), GU817544; S.
psilocarpus R.O. Belcher & D.E. Albrecht, Thompson 937 (MEL), GU817545; S.
roseus Sch.Bip., Garcia P. 250 (L), GU817546; S. scaposus DC. (Caputia scaposa
ined.), AF460001; S. squarrosus A.Rich., Thompson 907 (MEL), GU817547; S.
stigophlebius Baker, EF537946; S. suaveolens (L.) Ell., EF537921; S. superandinus
Cuatrec., EF537924; S. szyszylowiczii Hiern., Sagastegui A. et al. 15797 (S),
GU817548; S. tauricola V.A.Matthews, Budak 1735, Aksoy & Hamazaoǧlu (Yozgat
Türkiye Florası Herbaryumu), GU817549; S. thapsoides DC., EF537948; S. triodon
Phil. var. triodon, Luebert & Teillier 2266 (CONC), GU817550; S. vestitus P.J.Bergius,
Greuter 21766 (B), GU817551; S. viscosus L., AF459996; Shafera platyphylla
Greenm., Nordenstam s.n. (S), GU817552; Sinacalia tangutica (Maxim.) B.Nord., Lian
et al 93-54 (MO), GU817553; Sinosenecio euosmus (Hand.-Mazz.) B.Nord., Bouford,
Donoghue & Ree 27625 (MO), GU817554; Solanecio biafrae (Oliv. & Hiern)
C.Jeffrey, Brunel 6220 (B), GU817555; Steirodiscus capillaceus (Thunb.) Less., Cron
& Goodman 653 (J), GU817556; Stilpnogyne bellidioides DC., Goldblatt & Porter
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11729 (MO), GU817557; Symphyotrichum novae-angliae (L.) G.L.Nesom, Tepe 1675
(MU), GU817422; Syneilesis palmata (Thunb.) Maxim., Hata s.n. (4-Aug-1979)
(CHR), GU817558; Synotis nagensium (C.B.Clarke) C.Jeffrey & Y.L.Chen,
AF459973; Tagetes sp., Vincent 13224 (MU), GU817559; Talamancalia boquetensis
(Standl.) H.Rob. & Cuatrec., Wilbur 15397 (NY), GU817560; Telanthophora
grandifolia (Less.) H.Rob. & Brettell, EF537949; Tephroseris integrifolia (L.) Holub
ssp. aucheri (DC.) B.Nord., Budak 1688 & Hamazaoǧlu (Yozgat Türkiye Florası
Herbaryumu), GU817562; Tetradymia filifolia Greene, EF537950; Tussilago farfara
L., EF537951; Urostemon kirkii (Hook.f. ex Kirk) B.Nord., Cooper & Nickerson s.n.
(US), GU817563; Vernonia altissima Nutt., Tepe 1668 (MU), GU817564; Villasenoria
orcuttii (Greenm.) B.L.Clark, Robles 389 (XAL), GU817565; Werneria caespitosa
Wedd., Funk 11324 (US), GU817566; Xenophyllum poposum (Phil.) V.A.Funk, Funk
11351 (US), GU817567; Yermo xanthocephalus Dorn, Anderson 13691 (MO),
GU817568; Zemisia discolor (Sw.) B.Nord., Webster et al. 8420 (S), GU817569.
5’trnK. Abrotanella emarginata (Gaudich.) Cass., EF042141; AY929860; Acrisione
cymosa (J.Rémy) B.Nord., Lammers, Rodriques & Baeza 6447 (MU), GU817582;
Adenostyles alpina (L.) Bluff & Fingerh., Schwerdtfeger 12251 (B), GU817583;
Aequatorium asterotrichum B.Nord., Asplund 18263 (S), GU817584; Anacyclus
valentinus L., Tepe 1186 & Moreno-Paez (MU), GU817585; Antillanthus almironcillo
(M.Gómez) B.Nord., El-Ghazaby et al. 23.608 (S), GU817586; Arbelaezaster
ellsworthii (Cuatrec.) Cuatrec., Kilip & Smith 19413 (US), GU817587; Arnoglossum
atriplicifolium (L.) H.Rob., EF042143; Arrhenechthites mixta (A.Rich.) Belcher,
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EF042144; Austrosynotis rectirama (Baker) C.Jeffrey, LaCroix 4001 (WAG),
GU817589; Bahia dissecta (Gray) Britton, Tuhy 3868 (MU), GU817590;
Barkleyanthus salicifolius (Kunth) H.Rob. & Brettell, Genelle & Fleming 861 (B),
GU817591; Barnadesia sp., Hunziker & Gamerro 12561 (MU), GU817592; Bedfordia
arborescens Hochr., Greuter 21319 (B), GU817593; Bethencourtia palmensis (Nees)
Choisy, Nordenstam 9326 (S), GU817594; Blennosperma nanum S.F.Blake,
AF460089; Blepharispermum zanguebaricum Oliv. & Hiern., Medley 570 (MU),
GU817595; Bolandia pedunculosa (DC.) Cron, Cron & Goodman 681 (J), GU817596;
Brachyglottis repanda J.R.Forst. & G.Forst., AY554067; Cabreriella oppositicordia
(Cuatrec.) Cuatrec., Romero-Castañeda 7428 (US), GU817597; Cacaliopsis nardosmia
(A.Gray) A.Gray, Ertter 6502 (UC), GU817598; Calendula arvensis L., Watson 9513C (MU), GU817600; Capelio caledonica B.Nord., Nordenstam 9644 (S), GU817601;
Caucasalia parviflora (M.Bieb.) B.Nord., EF042145; Caxamarca sanchezii
M.O.Dillon & Sagást., Sagástegui et al. 15548 (F), GU817602; Centropappus brunonis
Hook.f., AY554058; Chaenactis douglasii (Hook.) Hook. & Arn., Vincent 8582 (MU),
GU817603; Charadranaetes durandii (Klatt) Janovec & H.Rob., Gomez P. 2242 (S),
GU817604; Chersodoma jodopappa (Sch.Bip.) Cabr., EF042146; Cichorium intybus
L., Tepe 1673 (MU), GU817605; Cineraria abyssinica Sch.Bip. ex A.Rich., AF460088;
Cirsium discolor (Muhl.) Spreng., Tepe 1670 (MU), GU817606; Cissampelopsis
volubilis (Blume) Miq., Carvalho 3175 (MU), GU817607; Coreopsis sp., Vincent 13226
(MU), GU817608; Corymbium enerve Markötter, Trinder-Smith 124 (US), GU817609;
Cotula coronopifolia L., Watson & Panero 94-26 (MU), GU817610; AY554071;
Crassocephalum crepidioides (Benth.) S.Moore, AF460050; Cremanthodium humile
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Maxim., EF042147; Crocidium multicaule Hook., Bartholomew 5749 (MO),
GU817611; AY929879; Curio rowleyanus (H.Jacobsen) P.V.Heath, AF460062;
Dauresia alliariifolia (O.Hoffm.) B.Nord & Pelser, Müller & Tilson 907 (M),
GU817612; Delairea odorata Lem., Tepe 2180, Pelser & Marticorena (MU),
GU817613; Dendrophorbium bomanii (R.E.Fr.) C.Jeffrey, EF042149; Dendrosenecio
kilimanjari (Mildbr.) E.B.Knox ssp. cottonii (Hutch. & G.Taylor) E.B.Knox,
AF460087; Digitacalia jatrophoides (Kunth) Pippen, Panero 2330 & Salinas (TEX),
GU817614; Dolichoglottis lyallii (Hook.f.) B.Nord., Strid 22172 (MO), GU817615;
Dolichorrhiza caucasica (M.Bieb.) Galushko, Schneeweiss 44 (WU), GU817616;
Dorobaea pimpinellifolia (Kunth) B.Nord., Tepe 1467 (MU), GU817617; Doronicum
pardalianches L., Cubr 40487 (B), GU817618; Ekmaniopappus mikanioides (Urb. &
Ekman) Borhidi, Nordenstam & Lundin 500 (S), GU817619; Elekmania picardae
(Krug & Urb.) B.Nord., EF042150; Emilia coccinea (Sims) G.Don., AF460086;
Endocellion sibiricum (J.F.Gmel.) J.Toman, Schönswetter & Tribsch T480 (WU),
GU817620; Erato polymnoides DC., Tepe 1456 (MU), GU817621; Erechtites
valerianifolius (Link ex Spreng.) Less. ex DC., EF042151; Eriothrix lycopodioides
DC., Nordenstam 9208 (S), GU817622; Eupatorium serotinum Michx., Tepe 1667
(MU), GU817623; Euryops pectinatus (L.) Cass., AF460030; Farfugium japonicum
(L.) Kitam., Wu 1439 (MO), GU817624; Faujasia squamosa (Bory) C.Jeffrey,
Nordenstam 9210 (S), GU817625; Faujasiopsis flexuosa (Lam.) C.Jeffrey ssp.
bourbonensis C.Jeffrey, Nordenstam 9202 (S), GU817626; Galinsoga quadriradiata
Cav., Vincent 13227 (MU), GU817627; Garcibarrigoa telembina (Cuatrec.) Cuatrec.,
Holm-Nielsen et al. 6211 (S), GU817628; Gerbera sp., Vincent 13223 (MU), GU817629;
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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Graphistylis dichroa (Bong.) D.J.N.Hind, Silva 140 & Zelma (S), GU817630;
Gymnodiscus capillaris (L.f.) Less., Coppejans EC1116 (U), GU817631; Cron &
Goodman 650 (J), GU817632; Gynoxys soukupii Cuatrec., AF460048; Gynura
divaricata (L.) DC. ssp. formosana (Kitam.) F.G.Davies, AF460047; Haastia
pulvinaris Hook.f., AY554075; Helenium autumnale L., Tepe 1664 (MU), GU817633;
Helianthus tuberosus L., Tepe 1672 (MU), GU817634; Helichrysum stoechas DC.,
Tepe 1092 (MU), GU817635; Herodotia haitiensis Urb. & Ekman, EF042153;
Herreranthus rivalis (Greenm.) B.Nord., Nordenstam s.n. (2/12/2007) (S), GU817636;
Hertia pallens (DC.) Kuntze, Koekemoer & Funk 1963 (US), GU817637;
Hoehnephytum trixoides (Gard.) Cabr., Carmo 131 (BHCB), GU817638; Homogyne
alpina (L.) Cass., Uhink 98-56 (MJG), GU817639; Hubertia riparia (DC.) C.Jeffrey,
Labat, Phillipson & Lowry II 2022 (WAG), GU817640; Humbertacalia sp., Phillipson et
al. 5641 (P), GU817641; Inula viscosa (L.) Aiton, Tepe 1142 (MU), GU817642; Io
ambondrombeensis (Humbert) B.Nord., Malcomber et al. s.n. (P), GU817643; Iranecio
cariensis (Boiss.) C.Jeffrey, Budak 1724, Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası
Herbaryumu), GU817644; Ischnea elachoglossa F.Muell., Beaman 12129 (MO),
GU817645; Jacmaia incana (Sw.) B.Nord., Nordenstam 7914 (S), GU817646;
Jacobaea vulgaris Gaertn., AF460068; AF548776; AY554084; Jessea multivenia
(Benth.) H.Rob. & Cuatrec., Nordenstam 9161 (S), GU817647; Kleinia neriifolia
Haw., AF460085; Lachanodes arborea (Roxb.) B.Nord., Cairns-Wicks s.n., GU817648;
Lactuca canadensis L., Vincent 13128 (MU), GU817649; Lamprocephalus montanus
B.Nord., Nordenstam 9542 (S), GU817650; Leonis trineurus (Griseb.) B.Nord., Smith
& al. 3238 (S), GU817651; Lepidospartum burgessii B.L.Turner, EF042156; Ligularia
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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stenocephala (Maxim.) Matsum. & Koidz., AF460084; Lomanthus fosbergii
(Cuatrec.) B. Nord. & Pelser, Øllgaard et al. 90697 (AAU), GU817729; Lopholaena
coriifolia (Sond.) Phillips & C.A.Sm., Van der Westhuizen & Liamé; s.n. (PREM),
GU817652; Lordhowea insularis (Benth.) B.Nord., AF460046; Luina hypoleuca
Benth., Greuter 17706 (B), GU817653; Lundinia plumbea (Griseb.) B.Nord., Zanoni
45816 (JBSD), GU817654; Madia sp., Vincent 12912 (MU), GU817655; Mesogramma
apiifolium DC., Giess 16074 (M), GU817656; Mikaniopsis clematoides (Sch.Bip. ex
A.Rich.) Milne-Redh., Friis, Hounde & Jacobsen 499 (WAG), GU817657; Miricacalia
makineana (Yatabe) Kitam., Kobayashi 33484 (HYO), GU817658; Misbrookea
strigosissima (A.Gray) V.A.Funk, EF042157; Monticalia abietina (Willd. ex Wedd.)
C.Jeffrey, Cleef 8411 (U), GU817659; Monticalia apiculata (Sch.Bip. ex Wedd.)
C.Jeffrey, Alston 6935 (S), GU817660; Monticalia arbutifolia (Kunth.) C.Jeffrey,
EF042158; Nelsonianthus tapianus (B.L.Turner) C.Jeffrey, Pérez & Kendizabal 457
(XAL), GU817661; Nemosenecio nikoensis (Miq.) B.Nord., Koyama 4079 (L),
GU817662; Nesampelos lucens (Poir. in Lam.) B.Nord., EF042159; Neurolaena lobata
(L.) R.Br., Acevedo 9316 (US), GU817663; Nordenstamia kingii (H.Rob. & Cuatrec.)
B.Nord., Ståhl 5572A (S), GU817664; Oldfeltia polyphlebia (Griseb.) B.Nord. &
Lundin, EF042161; Oresbia heterocarpa Cron & B.Nord., Nordenstam 9628 (S),
GU817665; Othonna capensis Bailey, AF460083; Packera eurycephala (Torr. &
A.Gray) W.A.Weber & Á.Löve, EF042162; Papuacalia dindondl (P.Royen)
Veldkamp, EF042163; Paracalia jungioides (Hook. & Arn.) Cuatrec., King & Collins
9017 (UC), GU817666; Parafaujasia fontinalis (Cordem.) C.Jeffrey, Lorence 2602
(MO), GU817667; Paragynoxys neodendroides (Cuatrec.) Cuatrec., Cleef 9852a (U),
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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GU817668; Parasenecio adenostyloides (Franch. et Sav. ex Maxim.) H.Koyama,
AF460051; Pentacalia arborea (Kunth) H.Rob. & Cuatrec., Øllgaard & Balslev 8298
(U), GU817669; Pericallis murrayi (Bornm.) B.Nord., EF042164; Perityle emoryi
Torr., Atwood 26483 (MU), GU817670; Petasites albus (L.) Gaertn., OPTIMA Iter VIII
1084 (RNG), GU817671; Phaneroglossa bolusii (Oliv.) B.Nord., AF460042;
Pippenalia delphinifolia (Rydb.) MacVaugh, Spellenberg & Bacon 11048 (TEX),
GU817672; Pittocaulon praecox (Cav.) H.Rob. & Brettell, EF042165; Pladaroxylon
leucadendron (G.Forst.) Hook.f., Cairns-Wicks s.n., GU817673; Pojarkovia pojarkovae
(Schischk.) Greuter, Pelser cult. 191 (L), GU817674; Polymnia canadensis L., Tepe
1665 (MU), GU817675; Psacaliopsis purpusii (Greenm. ex Brandegee) H.Rob. &
Brettell, Panero 2607, Davila & Tenorio (TEX), GU817676; Psacalium cirsiifolium
(Zucc.) H.Rob. & Brettell, EF042160; Pseudogynoxys haenkei (DC.) Cabr., Molina R.
18431 (U), GU817677; Rainiera stricta (Greene) Greene, Dennis 2317 (U), GU817678;
Robinsonecio gerberifolius (Sch.Bip. ex Hemsl.) T.M.Barkley & Janovec, Garcia P.
171 (MO), GU817679; Robinsonia berteroi (DC.) R.W.Sanders, Stuessy & Martic.,
AF460040; R. evenia Phil., Stuessy et al. 11308 (CONC), GU817680; R. gracilis
Decne., EF042166; Stuessy et al. 11312 (CONC), GU817681; R. masafuerae Skottsb.,
Stuessy et al. s.n. (CONC), GU817682; Roldana suffulta (Greenm.) H.Rob. & Brettell,
Rzedowski 36569 (UC), GU817683; Rugelia nudicaulis Shuttlew. ex Chapm., Feist,
Phillippe, Molano-Flores, Busemeyer & Carroll 714 (TENN), GU817684; Scrobicaria
ilicifolia (L.f.) B.Nord., Cleef, Garcia-B. & Jaramillo-M. 3528 (U), GU817719; Senecio
acanthifolius Hombr. & Jacq., ex Decne., EF042154; S. adamantinus Bong.,
EF042167; S. algens Wedd., Beck 2879 (S), GU817685; S. arnicoides Hook. & Arn.,
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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Zöllner 3474 (L), GU817686; S. boyacensis (Cuatrec.) Cuatrec., Gonzalez 180 (S),
GU817687; S. brasiliensis (Spreng.) Less., UEC 50.171 (UEC), GU817688; S. cadiscus
B.Nord. & Pelser, Rourke 1118 (S), GU817599; S. chilensis Less., EF042170; S.
ctenophyllus Phil., Zöllner 3959 (L), GU817690; S. culcitioides Wedd., Øllgaard &
Balslev 8822 (U), GU817689; S. deltoideus Less., EF042171; S. elegans L., Cron &
Goodman 687 (J), GU817691; S. fistulosus Poepp. ex Less., Beck & Liberman 9672 (S),
GU817692; S. flaccidus Less. var. flaccidus, Jardin Thuret cult. s.n. (MJG), GU817693;
S. flavus (Decne.) Sch.Bip., EF042172; S. gayanus (Colla) Cabr., Rosas 2157 (INIA),
GU817694; S. gramineus Harv., Hoener 2104 (WAG), GU817695; S. gregorii
F.Muell., Albrecht 7091 (NT), GU817696; S. hemmendorffii Malme, EF042174; S.
hieracium Remy, Baeza & Finot 3695 (CONC), GU817697; S. hispidissimus
I.Thomps., Thompson 927 (MEL), GU817698; S. hollandii Compton, Germishuizen
6586 (WAG), GU817699; S. hypsobates Wedd., Øllgaard & Balslev 9863 (U),
GU817700; S. ilicifolius Thunb., Cron & Goodman 686 (J), GU817701; S. integerrimus
Nutt. var. exaltatus (Nutt.) Cronquist, Crockett 437 (MU), GU817702; S. involucratus
(Kunth) DC., EF042142; S. jarae Phil., EF042175; S. lastarrianus Remy, Ricardi 3230
(B), GU817703; S. latifolius DC., EF042176; S. lineatus DC., AF460066; S.
mairetianus DC., EF042178; S. medley-woodii Hutch. (Caputia medley-woodii ined.),
AF460065; S. meuselii Rauh, EF042179; S. nemorensis L., AF460064; S. nevadensis
Boiss. & Reut., Vogt 4172 (B), GU817704; S. niveo-aureus Cuatrec., Cleef 6665 (S),
GU817705; S. oerstedianus Benth. ex Oerst., Nordenstam 9160 (S), GU817706; S.
oreophyton Remy, Beck 21589 (S), GU817707; S. otites Kunze ex DC., EF042180; S.
patens (Kunth) DC., Zak & Jaramillo 3427 (L), GU817708; S. pflanzii (Perkins)
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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Cuatrec., EF042148; S. pinifolius (L.) Lam., EF042181; S. pinnatifolius A.Rich. var.
lanceolatus (Benth.) I.Thomps., Thompson 904 (MEL), GU817709; S. polygaloides
Phil., Zöllner 5442 (L), GU817710; S. prenanthoides A.Rich., Thompson 915 (MEL),
GU817711; S. psilocarpus R.O. Belcher & D.E. Albrecht, Thompson 937 (MEL),
GU817712; S. retrorsus DC., EF042182; S. roseus Sch.Bip., Garcia P. 250 (L),
GU817713; S. saxatilis Wall. ex DC., EF042183; S. scandens Buch.-Ham. ex D.Don,
AF460061; S. scaposus DC. (Caputia scaposa ined.), AF460060; S. squarrosus
A.Rich., Thompson 907 (MEL), GU817714; S. suaveolens (L.) Ell., EF042152; S.
superandinus Cuatrec., EF042155; S. szyszylowiczii Hiern., Sagastegui A. et al. 15797
(S), GU817715; S. tauricola V.A.Matthews, Budak 1735, Aksoy & Hamazaoǧlu (Yozgat
Türkiye Florası Herbaryumu), GU817716; S. thapsoides DC., EF042185; S. triodon
Phil. var. triodon, Luebert & Teillier 2266 (CONC), GU817717; S. triqueter Less.,
EF042186; S. vestitus P.J.Bergius, Greuter 21766 (B), GU817718; S. viscosus L.,
AF460055; Shafera platyphylla Greenm., Nordenstam s.n. (S), GU817720; Sinacalia
tangutica (Maxim.) B.Nord., Lian et al 93-54 (MO), GU817721; Sinosenecio euosmus
(Hand.-Mazz.) B.Nord., Bouford, Donoghue & Ree 27625 (MO), GU817722; Solanecio
biafrae (Oliv. & Hiern) C.Jeffrey, Brunel 6220 (B), GU817723; Steirodiscus
capillaceus (Thunb.) Less., Cron & Goodman 653 (J), GU817724; Stilpnogyne
bellidioides DC., Goldblatt & Porter 11729 (MO), GU817725; Symphyotrichum novaeangliae (L.) G.L.Nesom, Tepe 1675 (MU), GU817588; Syneilesis palmata (Thunb.)
Maxim., Hata s.n. (4-Aug-1979) (CHR), GU817726; Synotis nagensium (C.B.Clarke)
C.Jeffrey & Y.L.Chen, AF460031; Tagetes sp., Vincent 13224 (MU), GU817727;
Talamancalia boquetensis (Standl.) H.Rob. & Cuatrec., Wilbur 15397 (NY),
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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GU817728; Telanthophora grandifolia (Less.) H.Rob. & Brettell, EF042187;
Tephroseris integrifolia (L.) Holub ssp. aucheri (DC.) B.Nord., Budak 1688 &
Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu), GU817730; Tetradymia filifolia
Greene, EF042188; Traversia baccharoides Hook.f., AY554090; Tussilago farfara L.,
EF042189; Urostemon kirkii (Hook.f. ex Kirk) B.Nord., AY554065; Vernonia
altissima Nutt., Tepe 1668 (MU), GU817731; Villasenoria orcuttii (Greenm.)
B.L.Clark, Robles 389 (XAL), GU817732; Werneria caespitosa Wedd., Funk 11324
(US), GU817733; Xenophyllum poposum (Phil.) V.A.Funk, Funk 11351 (US),
GU817734; Yermo xanthocephalus Dorn, Anderson 13691 (MO), GU817735; Zemisia
discolor (Sw.) B.Nord., Webster et al. 8420 (S), GU817736.
ndhF. Abrotanella emarginata (Gaudich.) Cass., AJ012679; Acrisione cymosa
(J.Rémy) B.Nord., Lammers, Rodriques & Baeza 6447 (MU), GU817808; Adenostyles
alpina (L.) Bluff & Fingerh., Schwerdtfeger 12251 (B), GU817826; Aequatorium
asterotrichum B.Nord., Asplund 18263 (S), GU817827; Anacyclus valentinus L., Tepe
1186 & Moreno-Paez (MU), GU817828; Antillanthus almironcillo (M.Gómez)
B.Nord., Ståhl, Cejas, Rova & Urquiola 1339 (MO), GU817829; Arbelaezaster
ellsworthii (Cuatrec.) Cuatrec., Kilip & Smith 19413 (US), GU817830; Arnoglossum
atriplicifolium (L.) H.Rob., EF537953; Arrhenechthites mixta (A.Rich.) Belcher,
EF537954; Austrosynotis rectirama (Baker) C.Jeffrey, LaCroix 4001 (WAG),
GU817831; Bahia dissecta (Gray) Britton, Tuhy 3868 (MU), GU817832;
Barkleyanthus salicifolius (Kunth) H.Rob. & Brettell, Ramos Marchena 575 (B),
GU817833; Barnadesia sp., Hunziker & Gamerro 12561 (MU), GU817789; Bedfordia
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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arborescens Hochr., Greuter 21319 (B), GU817834; Bethencourtia palmensis (Nees)
Choisy, Nordenstam 9326 (S), GU817835; Blennosperma nanum S.F.Blake, L39433;
Blepharispermum zanguebaricum Oliv. & Hiern., Medley 570 (MU), GU817809;
L39456; Bolandia pedunculosa (DC.) Cron, Cron & Goodman 681 (J), GU817836;
Brachyglottis repanda J.R.Forst. & G.Forst., Schwerdtfeger 17407 (B), GU817796;
Cabreriella oppositicordia (Cuatrec.) Cuatrec., Romero-Castañeda 7428 (US),
GU817810; Cacaliopsis nardosmia (A.Gray) A.Gray, Ertter 6502 (UC), GU817837;
Calendula arvensis L., Watson 95-13C (MU), GU817838; Capelio caledonica B.Nord.,
Nordenstam 9644 (S), GU817839; Caucasalia parviflora (M.Bieb.) B.Nord., EF537955;
Caxamarca sanchezii M.O.Dillon & Sagást., Sagástegui et al. 15548 (F), GU817840;
Centropappus brunonis Hook.f., Wapstra MW2 (MEL), GU817841; Chaenactis
douglasii (Hook.) Hook. & Arn., Vincent 8582 (MU), GU817842; Charadranaetes
durandii (Klatt) Janovec & H.Rob., Gomez P. 2242 (S), GU817843; Chersodoma
jodopappa (Sch.Bip.) Cabr., EF537956; Cichorium intybus L., Tepe 1673 (MU),
GU817844; Cineraria abyssinica Sch.Bip. ex A.Rich., EF537957; Cirsium discolor
(Muhl.) Spreng., Tepe 1670 (MU), GU817845; Cissampelopsis volubilis (Blume) Miq.,
Carvalho 3175 (MU), GU817846; Coreopsis sp., Vincent 13226 (MU), GU817847;
Corymbium enerve Markötter, Trinder-Smith 124 (US), GU817848; Cotula
coronopifolia L., Watson & Panero 94-26 (MU), GU817849; Crassocephalum
crepidioides (Benth.) S.Moore, EF537958; Cremanthodium humile Maxim.,
EF537959; Cremanthodium humile Maxim., AY723234; Crocidium multicaule Hook.,
Bartholomew 5749 (MO), GU817805; Curio rowleyanus (H.Jacobsen) P.V.Heath,
EF538008; Dauresia alliariifolia (O.Hoffm.) B.Nord & Pelser, Müller & Tilson 907
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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(M), GU817850; Delairea odorata Lem., L39435; Dendrophorbium bomanii (R.E.Fr.)
C.Jeffrey, EF537961; Dendrosenecio kilimanjari (Mildbr.) E.B.Knox ssp. cottonii
(Hutch. & G.Taylor) E.B.Knox, Knox 50 (MICH), GU817851; Digitacalia
jatrophoides (Kunth) Pippen, Panero 2330 & Salinas (TEX), GU817852;
Dolichoglottis lyallii (Hook.f.) B.Nord., Strid 22172 (MO), GU817793; Dolichorrhiza
caucasica (M.Bieb.) Galushko, Schneeweiss 44 (WU), GU817853; Dorobaea
pimpinellifolia (Kunth) B.Nord., Tepe 1467 (MU), GU817854; Doronicum
pardalianches L., EF537962; Ekmaniopappus mikanioides (Urb. & Ekman) Borhidi,
Nordenstam & Lundin 500 (S), GU817817; Elekmania picardae (Krug & Urb.)
B.Nord., EF537963; Emilia coccinea (Sims) G.Don., EF537964; Endocellion sibiricum
(J.F.Gmel.) J.Toman, Schönswetter & Tribsch T480 (WU), GU817855; Erato
polymnoides DC., Tepe 1456 (MU), GU817790; Erechtites valerianifolius (Link ex
Spreng.) Less. ex DC., EF537965; Eriothrix lycopodioides DC., Nordenstam 9208 (S),
GU817856; Eupatorium serotinum Michx., Tepe 1667 (MU), GU817857; Euryops
pectinatus (L.) Cass., EF537966; Farfugium japonicum (L.) Kitam., AY723242;
Faujasia squamosa (Bory) C.Jeffrey, Nordenstam 9210 (S), GU817858; Faujasiopsis
flexuosa (Lam.) C.Jeffrey ssp. bourbonensis C.Jeffrey, Nordenstam 9202 (S),
GU817859; Galinsoga quadriradiata Cav., Vincent 13227 (MU), GU817860;
AF384727; Garcibarrigoa telembina (Cuatrec.) Cuatrec., Holm-Nielsen et al. 6211 (S),
GU817861; Gerbera sp., Vincent 13223 (MU), GU817862; Graphistylis dichroa (Bong.)
D.J.N.Hind, Silva 140 & Zelma (S), GU817863; Gymnodiscus capillaris (L.f.) Less.,
Coppejans EC1116 (U), GU817864; Gynoxys soukupii Cuatrec., EF537967; Gynura
divaricata (L.) DC. ssp. formosana (Kitam.) F.G.Davies, Panero & Hsiao 6457 (TEX),
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GU817865; Helenium autumnale L., Tepe 1664 (MU), GU817866; Helianthus
tuberosus L., Tepe 1672 (MU), GU817867; Helichrysum stoechas DC., Tepe 1092
(MU), GU817868; Herodotia haitiensis Urb. & Ekman, EF537969; Herreranthus
rivalis (Greenm.) B.Nord., Nordenstam s.n. (2/12/2007) (S), GU817869; Hertia pallens
(DC.) Kuntze, Koekemoer & Funk 1963 (US), GU817870; Hoehnephytum trixoides
(Gard.) Cabr., Carmo 131 (BHCB), GU817871; Homogyne alpina (L.) Cass., Uhink
98-56 (MJG), GU817811; Hubertia riparia (DC.) C.Jeffrey, Labat, Phillipson & Lowry
II 2022 (WAG), GU817872; Humbertacalia sp., Phillipson et al. 5641 (P), GU817873;
Inula viscosa (L.) Aiton, Tepe 1142 (MU), GU817874; Io ambondrombeensis
(Humbert) B.Nord., Malcomber et al. 1380 (MO), GU817875; Iranecio cariensis
(Boiss.) C.Jeffrey, Budak 1724, Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası
Herbaryumu), GU817876; Ischnea elachoglossa F.Muell., Beaman 12129 (MO),
GU817877; Jacmaia incana (Sw.) B.Nord., Nordenstam 7914 (S), GU817798;
Jacobaea vulgaris Gaertn., EF537971; Jessea multivenia (Benth.) H.Rob. & Cuatrec.,
Nordenstam 9161 (S), GU817878; Kleinia neriifolia Haw., EF537972; Lachanodes
arborea (Roxb.) B.Nord., Cairns-Wicks s.n., GU817879; Lactuca canadensis L.,
Vincent 13128 (MU), GU817880; Lamprocephalus montanus B.Nord., Nordenstam
9542 (S), GU817881; Leonis trineurus (Griseb.) B.Nord., Smith & al. 3238 (S),
GU817882; Lepidospartum burgessii B.L.Turner, EF537974; Ligularia stenocephala
(Maxim.) Matsum. & Koidz., EF537975; Lomanthus fosbergii (Cuatrec.) B. Nord. &
Pelser, Øllgaard et al. 90697 (AAU), GU817935; Lopholaena coriifolia (Sond.)
Phillips & C.A.Sm., L39434; Lordhowea insularis (Benth.) B.Nord., Trodd & Thorne
3703 (L), GU817799; Luina hypoleuca Benth., Greuter 17706 (B), GU817883;
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Lundinia plumbea (Griseb.) B.Nord., Zanoni 45816 (JBSD), GU817806; Madia sp.,
Vincent 12912 (MU), GU817807; Mesogramma apiifolium DC., Giess 16074 (M),
GU817792; Mikaniopsis clematoides (Sch.Bip. ex A.Rich.) Milne-Redh., De Wilde &
De Wilde-Duyfjes 9006 (B), GU817884; Miricacalia makineana (Yatabe) Kitam.,
Kobayashi 33484 (HYO), GU817812; Misbrookea strigosissima (A.Gray) V.A.Funk,
EF537976; Monticalia abietina (Willd. ex Wedd.) C.Jeffrey, Cleef 8411 (U),
GU817885; Monticalia apiculata (Sch.Bip. ex Wedd.) C.Jeffrey, Alston 6935 (S),
GU817886; Monticalia arbutifolia (Kunth.) C.Jeffrey, EF537977; Nemosenecio
nikoensis (Miq.) B.Nord., Koyama 4079 (L), GU817887; Nesampelos lucens (Poir. in
Lam.) B.Nord., EF537978; Neurolaena lobata (L.) R.Br., AF384754; Nordenstamia
kingii (H.Rob. & Cuatrec.) B.Nord., Ståhl 5572A (S), GU817888; Oldfeltia
polyphlebia (Griseb.) B.Nord. & Lundin, EF537980; Oresbia heterocarpa Cron &
B.Nord., Nordenstam 9628 (S), GU817889; Othonna capensis Bailey, EF537981;
Packera eurycephala (Torr. & A.Gray) W.A.Weber & Á.Löve, EF537982;
Papuacalia dindondl (P.Royen) Veldkamp, EF537983; Parafaujasia fontinalis
(Cordem.) C.Jeffrey, Lorence 2602 (MO), GU817890; Paragynoxys neodendroides
(Cuatrec.) Cuatrec., Cleef 9852a (U), GU817813; Parasenecio adenostyloides
(Franch. et Sav. ex Maxim.) H.Koyama, EF537984; Pentacalia arborea (Kunth)
H.Rob. & Cuatrec., Øllgaard & Balslev 8298 (U), GU817891; Pericallis murrayi
(Bornm.) B.Nord., EF537985; Perityle emoryi Torr., Atwood 26483 (MU), GU817892;
Petasites albus (L.) Gaertn., Cubr 40398 (B), GU817893; OPTIMA Iter VIII 1084
(RNG), GU817819; Phaneroglossa bolusii (Oliv.) B.Nord., AF384765; Pippenalia
delphinifolia (Rydb.) MacVaugh, Spellenberg & Bacon 11048 (TEX), GU817894;
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Pittocaulon praecox (Cav.) H.Rob. & Brettell, EF537986; Pladaroxylon leucadendron
(G.Forst.) Hook.f., Cairns-Wicks s.n., GU817895; Pojarkovia pojarkovae (Schischk.)
Greuter, Pelser cult. 191 (L), GU817896; Polymnia canadensis L., Tepe 1665 (MU),
GU817897; AF384769; Psacaliopsis purpusii (Greenm. ex Brandegee) H.Rob. &
Brettell, Panero 2607, Davila & Tenorio (TEX), GU817898; Psacalium cirsiifolium
(Zucc.) H.Rob. & Brettell, EF537979; Pseudogynoxys haenkei (DC.) Cabr., Molina R.
18431 (U), GU817899; Rainiera stricta (Greene) Greene, Dennis 2317 (U), GU817900;
Robinsonecio gerberifolius (Sch.Bip. ex Hemsl.) T.M.Barkley & Janovec, Garcia P.
171 (MO), GU817901; Robinsonia berteroi (DC.) R.W.Sanders, Stuessy & Martic.,
EF537987; R. evenia Phil., Stuessy et al. 11308 (CONC), GU817902; R. gracilis Decne.,
EF537988; Stuessy et al. 11282 (CONC), GU817903; R. masafuerae Skottsb., Stuessy et
al. s.n. (CONC), GU817818; Roldana suffulta (Greenm.) H.Rob. & Brettell,
Rzedowski 36569 (UC), GU817800; Rugelia nudicaulis Shuttlew. ex Chapm., Feist,
Phillippe, Molano-Flores, Busemeyer & Carroll 714 (TENN), GU817904; Scrobicaria
ilicifolia (L.f.) B.Nord., Cleef, Garcia-B. & Jaramillo-M. 3528 (U), GU817814; Senecio
acanthifolius Hombr. & Jacq., ex Decne., EF537970; S. adamantinus Bong.,
EF537989; S. algens Wedd., Beck 2879 (S), GU817905; S. arnicoides Hook. & Arn.,
Zöllner 3474 (L), GU817801; S. boyacensis (Cuatrec.) Cuatrec., Gonzalez 180 (S),
GU817906; S. brasiliensis (Spreng.) Less., UEC 50.171 (UEC), GU817820; S. cadiscus
B.Nord. & Pelser, Goldblatt & Manning 10690 (MO), GU817794; S. chilensis Less.,
EF537992; S. ctenophyllus Phil., Zöllner 3959 (L), GU817804; S. culcitioides Wedd.,
Øllgaard & Balslev 8822 (U), GU817907; S. deltoideus Less., EF537993; S. elegans L.,
Cron & Goodman 687 (J), GU817908; S. fistulosus Poepp. ex Less., Beck & Liberman
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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9672 (S), GU817909; S. flaccidus Less. var. flaccidus, Jardin Thuret cult. s.n. (MJG),
GU817910; S. flavus (Decne.) Sch.Bip., EF537994; S. gayanus (Colla) Cabr., Rosas
2157 (INIA), GU817911; S. gramineus Harv., Hoener 2104 (WAG), GU817802; S.
gregorii F.Muell., Albrecht 7091 (NT), GU817912; S. hemmendorffii Malme,
EF537996; S. hieracium Remy, Baeza & Finot 3695 (CONC), GU817913; S.
hispidissimus I.Thomps., Thompson 927 (MEL), GU817914; S. hollandii Compton,
Germishuizen 6586 (WAG), GU817821; S. hypsobates Wedd., Øllgaard & Balslev 9863
(U), GU817822; S. ilicifolius Thunb., Cron & Goodman 686 (J), GU817915; S.
integerrimus Nutt. var. exaltatus (Nutt.) Cronquist, Crockett 437 (MU), GU817916; S.
involucratus (Kunth) DC., EF537952; S. jarae Phil., EF537997; S. lastarrianus Remy,
Ricardi 3230 (B), GU817823; S. latifolius DC., EF537998; S. lineatus DC., EF537999;
S. mairetianus DC., EF538001; S. medley-woodii Hutch. (Caputia medley-woodii
ined.), EF538002; S. meuselii Rauh, EF538003; S. nemorensis L., EF538004;
AY723209; S. nevadensis Boiss. & Reut., Vogt 4172 (B), GU817917; S. niveo-aureus
Cuatrec., Cleef 6665 (S), GU817918; S. oerstedianus Benth. ex Oerst., Nordenstam
9160 (S), GU817919; S. oreophyton Remy, Beck 21589 (S), GU817920; S. otites Kunze
ex DC., EF538005; S. patens (Kunth) DC., Zak & Jaramillo 3427 (L), GU817795; S.
pflanzii (Perkins) Cuatrec., EF537960; S. pinifolius (L.) Lam., EF538006; S.
pinnatifolius A.Rich. var. lanceolatus (Benth.) I.Thomps., Thompson 904 (MEL),
GU817922; S. polygaloides Phil., Zöllner 5442 (L), GU817923; S. prenanthoides
A.Rich., Thompson 915 (MEL), GU817924; S. psilocarpus R.O. Belcher & D.E.
Albrecht, Thompson 937 (MEL), GU817925; S. retrorsus DC., EF538007; S. roseus
Sch.Bip., Garcia P. 250 (L), GU817926; S. saxatilis Wall. ex DC., EF538009; S.
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scandens Buch.-Ham. ex D.Don, EF538010; AY723214; S. scaposus DC. (Caputia
scaposa ined.), EF538011; S. squarrosus A.Rich., Thompson 907 (MEL), GU817927; S.
stigophlebius Baker, EF538012; S. suaveolens (L.) Ell., EF537968; S. superandinus
Cuatrec., EF537973; S. szyszylowiczii Hiern., Sagastegui A. et al. 15797 (S),
GU817928; S. tauricola V.A.Matthews, Budak 1735, Aksoy & Hamazaoǧlu (Yozgat
Türkiye Florası Herbaryumu), GU817929; S. thapsoides DC., EF538014; S. triodon
Phil. var. triodon, Luebert & Teillier 2266 (CONC), GU817930; S. triqueter Less.,
EF538015; S. vestitus P.J.Bergius, Greuter 21766 (B), GU817824; S. viscosus L.,
EF538016; Shafera platyphylla Greenm., Nordenstam s.n. (S), GU817921; Sinacalia
tangutica (Maxim.) B.Nord., AY723243; Sinosenecio euosmus (Hand.-Mazz.)
B.Nord., Bouford, Donoghue & Ree 27625 (MO), GU817788; Solanecio biafrae (Oliv.
& Hiern) C.Jeffrey, Brunel 6220 (B), GU817931; Steirodiscus capillaceus (Thunb.)
Less., Cron & Goodman 653 (J), GU817932; Stilpnogyne bellidioides DC., Goldblatt &
Porter 11729 (MO), GU817933; Symphyotrichum novae-angliae (L.) G.L.Nesom, Tepe
1675 (MU), GU817797; Syneilesis palmata (Thunb.) Maxim., L39432; Synotis
nagensium (C.B.Clarke) C.Jeffrey & Y.L.Chen, EF538018; Tagetes sp., Vincent
13224 (MU), GU817934; Talamancalia boquetensis (Standl.) H.Rob. & Cuatrec.,
Wilbur 15397 (NY), GU817816; Telanthophora grandifolia (Less.) H.Rob. & Brettell,
EF538019; Tephroseris integrifolia (L.) Holub ssp. aucheri (DC.) B.Nord., Budak
1688 & Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu), GU817936; Tetradymia
filifolia Greene, EF538020; Tussilago farfara L., AY723239; Urostemon kirkii
(Hook.f. ex Kirk) B.Nord., Cooper & Nickerson s.n. (US), GU817791; Vernonia
altissima Nutt., Tepe 1668 (MU), GU817937; Villasenoria orcuttii (Greenm.)
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B.L.Clark, Robles 389 (XAL), GU817815; Werneria caespitosa Wedd., Funk 11324
(US), GU817825; Xenophyllum poposum (Phil.) V.A.Funk, Funk 11351 (US),
GU817938; Yermo xanthocephalus Dorn, Anderson 13691 (MO), GU817939; Zemisia
discolor (Sw.) B.Nord., Webster et al. 8420 (S), GU817803.
psbA-trnH. Abrotanella emarginata (Gaudich.) Cass., EF538021; Acrisione cymosa
(J.Rémy) B.Nord., Lammers, Rodriques & Baeza 6447 (MU), GU818329; Adenostyles
alpina (L.) Bluff & Fingerh., Schwerdtfeger 12251 (B), GU818330; Aequatorium
asterotrichum B.Nord., Asplund 18263 (S), GU818331; Anacyclus valentinus L., Tepe
1186 & Moreno-Paez (MU), GU818332; Antillanthus almironcillo (M.Gómez)
B.Nord., El-Ghazaby et al. 23.608 (S), GU818333; Arbelaezaster ellsworthii (Cuatrec.)
Cuatrec., Kilip & Smith 19413 (US), GU818334; Arnoglossum atriplicifolium (L.)
H.Rob., EF538056; Arrhenechthites mixta (A.Rich.) Belcher, EF538062; Austrosynotis
rectirama (Baker) C.Jeffrey, LaCroix 4001 (MO), GU818336; Bahia dissecta (Gray)
Britton, Tuhy 3868 (MU), GU818337; Barkleyanthus salicifolius (Kunth) H.Rob. &
Brettell, Genelle & Fleming 861 (B), GU818338; Barnadesia sp., Hunziker & Gamerro
12561 (MU), GU818339; Bedfordia arborescens Hochr., Greuter 21319 (B),
GU818340; DQ131861; Bethencourtia palmensis (Nees) Choisy, Nordenstam 9326 (S),
GU818341; Blennosperma nanum S.F.Blake, EF538023; DQ131853;
Blepharispermum zanguebaricum Oliv. & Hiern., Medley 570 (MU), GU818342;
Bolandia pedunculosa (DC.) Cron, Cron & Goodman 681 (J), GU818343;
Brachyglottis repanda J.R.Forst. & G.Forst., Schwerdtfeger 17407 (B), GU818344;
Cabreriella oppositicordia (Cuatrec.) Cuatrec., Romero-Castañeda 7428 (US),
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GU818345; Cacaliopsis nardosmia (A.Gray) A.Gray, DQ131859; Calendula arvensis
L., Watson 95-13C (MU), GU818348; Capelio caledonica B.Nord., Nordenstam 9644
(S), GU818349; Caucasalia parviflora (M.Bieb.) B.Nord., EF538024; Caxamarca
sanchezii M.O.Dillon & Sagást., Sagástegui et al. 15548 (F), GU818350; Centropappus
brunonis Hook.f., Wapstra MW2 (MEL), GU818351; Chaenactis douglasii (Hook.)
Hook. & Arn., Vincent 8582 (MU), GU818352; Charadranaetes durandii (Klatt)
Janovec & H.Rob., Gomez P. 2242 (S), GU818353; Chersodoma jodopappa (Sch.Bip.)
Cabr., EF538025; Cichorium intybus L., Tepe 1673 (MU), GU818354; Cineraria
abyssinica Sch.Bip. ex A.Rich., EF538026; Cirsium discolor (Muhl.) Spreng., Tepe
1670 (MU), GU818355; Cissampelopsis volubilis (Blume) Miq., Carvalho 3175 (MU),
GU818356; Coreopsis sp., Vincent 13226 (MU), GU818357; Corymbium enerve
Markötter, Trinder-Smith 124 (US), GU818358; Cotula coronopifolia L., Watson &
Panero 94-26 (MU), GU818359; Crassocephalum crepidioides (Benth.) S.Moore,
AY155640; Cremanthodium humile Maxim., EF538061; Crocidium multicaule Hook.,
Bartholomew 5749 (MO), GU818360; Curio rowleyanus (H.Jacobsen) P.V.Heath,
EF538047; Dauresia alliariifolia (O.Hoffm.) B.Nord & Pelser, Müller & Tilson 907
(M), GU818361; Delairea odorata Lem., Tepe 2180, Pelser & Marticorena (MU),
GU818362; Dendrocacalia crepidifolia (Nakai) Nakai, Kobayashi 190493 (MO),
GU818363; Dendrophorbium bomanii (R.E.Fr.) C.Jeffrey, EF538028; Dendrosenecio
kilimanjari (Mildbr.) E.B.Knox ssp. cottonii (Hutch. & G.Taylor) E.B.Knox, Knox 50
(MICH), GU818364; Digitacalia jatrophoides (Kunth) Pippen, Cruden 1086 (UC),
GU818365; Dolichoglottis lyallii (Hook.f.) B.Nord., Strid 22172 (MO), GU818366;
Dolichorrhiza caucasica (M.Bieb.) Galushko, Schneeweiss 44 (WU), GU818367;
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Dorobaea pimpinellifolia (Kunth) B.Nord., Tepe 1467 (MU), GU818368; Doronicum
pardalianches L., EF538088; Ekmaniopappus mikanioides (Urb. & Ekman) Borhidi,
Zanoni et al. 33908 (JBSD), GU818369; Elekmania picardae (Krug & Urb.) B.Nord.,
EF538070; Emilia coccinea (Sims) G.Don., EF538029; Endocellion sibiricum
(J.F.Gmel.) J.Toman, Schönswetter & Tribsch T480 (WU), GU818370; Erato
polymnoides DC., Tepe 1456 (MU), GU818371; Erechtites valerianifolius (Link ex
Spreng.) Less. ex DC., EF538030; Eriothrix lycopodioides DC., Nordenstam 9208 (S),
GU818372; Eupatorium serotinum Michx., Tepe 1667 (MU), GU818373; Euryops
pectinatus (L.) Cass., EF538031; Farfugium japonicum (L.) Kitam., Wu 1439 (MO),
GU818374; Faujasia squamosa (Bory) C.Jeffrey, Nordenstam 9210 (S), GU818375;
Faujasiopsis flexuosa (Lam.) C.Jeffrey ssp. bourbonensis C.Jeffrey, Nordenstam 9202
(S), GU818376; Galinsoga quadriradiata Cav., Vincent 13227 (MU), GU818377;
Garcibarrigoa telembina (Cuatrec.) Cuatrec., Holm-Nielsen et al. 6211 (S), GU818378;
Gerbera sp., Vincent 13223 (MU), GU818379; Graphistylis dichroa (Bong.)
D.J.N.Hind, Silva 140 & Zelma (S), GU818380; Gymnodiscus capillaris (L.f.) Less.,
Coppejans EC1116 (U), GU818381; Nordenstam 9631 (S), GU818382; Cron &
Goodman 650 (J), GU818383; Gynoxys soukupii Cuatrec., EF538051; Gynura
divaricata (L.) DC. ssp. formosana (Kitam.) F.G.Davies, Panero & Hsiao 6457 (TEX),
GU818384; Helenium autumnale L., Tepe 1664 (MU), GU818385; Helianthus
tuberosus L., Tepe 1672 (MU), GU818386; Helichrysum stoechas DC., Tepe 1092
(MU), GU818387; Herodotia haitiensis Urb. & Ekman, EF538033; Herreranthus
rivalis (Greenm.) B.Nord., Nordenstam s.n. (2/12/2007) (S), GU818388; Hertia pallens
(DC.) Kuntze, Koekemoer & Funk 1963 (US), GU818389; Hoehnephytum trixoides
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(Gard.) Cabr., Carmo 131 (BHCB), GU818390; Homogyne alpina (L.) Cass., Uhink
98-56 (MJG), GU818391; Hubertia riparia (DC.) C.Jeffrey, Labat, Phillipson & Lowry
II 2022 (WAG), GU818392; Humbertacalia sp., Phillipson et al. 5641 (P), GU818393;
Inula viscosa (L.) Aiton, Tepe 1142 (MU), GU818394; Io ambondrombeensis
(Humbert) B.Nord., Malcomber et al. 1380 (MO), GU818395; Iranecio cariensis
(Boiss.) C.Jeffrey, Budak 1724, Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası
Herbaryumu), GU818396; Ischnea elachoglossa F.Muell., Beaman 12129 (MO),
GU818397; Jacmaia incana (Sw.) B.Nord., Nordenstam 7914 (S), GU818398;
Jacobaea vulgaris Gaertn., AY155657; Jessea multivenia (Benth.) H.Rob. &
Cuatrec., Nordenstam 9161 (S), GU818399; Kleinia neriifolia Haw., AY155642;
Lachanodes arborea (Roxb.) B.Nord., Cairns-Wicks s.n., GU818400; Lactuca
canadensis L., Vincent 13128 (MU), GU818401; Lamprocephalus montanus B.Nord.,
Nordenstam 9542 (S), GU818402; Leonis trineurus (Griseb.) B.Nord., Smith & al. 3238
(S), GU818403; Lepidospartum burgessii B.L.Turner, EF538053; Ligularia
stenocephala (Maxim.) Matsum. & Koidz., EF538078; Lomanthus fosbergii
(Cuatrec.) B. Nord. & Pelser, Øllgaard et al. 90697 (AAU), GU818480; Lopholaena
coriifolia (Sond.) Phillips & C.A.Sm., Van der Westhuizen & Liamé; s.n. (PREM),
GU818404; Lordhowea insularis (Benth.) B.Nord., Trodd & Thorne 3703 (L),
GU818405; Luina hypoleuca Benth., Greuter 17706 (B), GU818406; DQ131858;
Lundinia plumbea (Griseb.) B.Nord., Zanoni 45816 (JBSD), GU818407; Madia sp.,
Vincent 12912 (MU), GU818408; Mattfeldia triplinervis Urb., Ekman H7492 (S),
GU818409; Mesogramma apiifolium DC., Giess 16074 (M), GU818410; Mikaniopsis
clematoides (Sch.Bip. ex A.Rich.) Milne-Redh., De Wilde & De Wilde-Duyfjes 9006
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(B), GU818411; Misbrookea strigosissima (A.Gray) V.A.Funk, EF538036; Monticalia
abietina (Willd. ex Wedd.) C.Jeffrey, Cleef 8411 (U), GU818412; Monticalia apiculata
(Sch.Bip. ex Wedd.) C.Jeffrey, Alston 6935 (S), GU818413; Monticalia arbutifolia
(Kunth.) C.Jeffrey, EF538037; Nelsonianthus tapianus (B.L.Turner) C.Jeffrey, Pérez
& Kendizabal 457 (XAL), GU818414; Nemosenecio nikoensis (Miq.) B.Nord., Koyama
4079 (L), GU818415; Nesampelos lucens (Poir. in Lam.) B.Nord., EF538054;
Neurolaena lobata (L.) R.Br., Acevedo 9316 (US), GU818416; Nordenstamia kingii
(H.Rob. & Cuatrec.) B.Nord., Ståhl 5572A (S), GU818417; Oldfeltia polyphlebia
(Griseb.) B.Nord. & Lundin, EF538058; Oresbia heterocarpa Cron & B.Nord.,
Nordenstam 9628 (S), GU818418; Othonna capensis Bailey, EF538039; Packera
eurycephala (Torr. & A.Gray) W.A.Weber & Á.Löve, EF538040; Papuacalia
dindondl (P.Royen) Veldkamp, EF538041; Paracalia jungioides (Hook. & Arn.)
Cuatrec., King & Collins 9017 (UC), GU818419; Parafaujasia fontinalis (Cordem.)
C.Jeffrey, Lorence 2602 (MO), GU818420; Parasenecio adenostyloides (Franch. et
Sav. ex Maxim.) H.Koyama, EF538055; Pentacalia arborea (Kunth) H.Rob. &
Cuatrec., Øllgaard & Balslev 8298 (U), GU818421; Pericallis murrayi (Bornm.)
B.Nord., EF538060; Perityle emoryi Torr., Atwood 26483 (MU), GU818422; Petasites
albus (L.) Gaertn., OPTIMA Iter VIII 1084 (RNG), GU818423; Phaneroglossa bolusii
(Oliv.) B.Nord., EF538069; Pippenalia delphinifolia (Rydb.) MacVaugh, Spellenberg
& Bacon 11048 (TEX), GU818424; Pittocaulon praecox (Cav.) H.Rob. & Brettell,
EF538042; DQ131854; Pladaroxylon leucadendron (G.Forst.) Hook.f., Cairns-Wicks
s.n., GU818425; Pojarkovia pojarkovae (Schischk.) Greuter, Pelser cult. 191 (L),
GU818426; Polymnia canadensis L., Tepe 1665 (MU), GU818427; Psacaliopsis
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purpusii (Greenm. ex Brandegee) H.Rob. & Brettell, Panero 2607, Davila & Tenorio
(TEX), GU818428; Psacalium cirsiifolium (Zucc.) H.Rob. & Brettell, EF538038;
Pseudogynoxys haenkei (DC.) Cabr., Molina R. 18431 (U), GU818429; Rainiera stricta
(Greene) Greene, Dennis 2317 (U), GU818430; Robinsonecio gerberifolius (Sch.Bip.
ex Hemsl.) T.M.Barkley & Janovec, Garcia P. 171 (MO), GU818431; Robinsonia
berteroi (DC.) R.W.Sanders, Stuessy & Martic., EF538082; R. evenia Phil., Stuessy et
al. 11308 (CONC), GU818432; R. gracilis Decne., EF538068; Stuessy et al. 11282
(CONC), GU818433; Stuessy et al. 11312 (CONC), GU818434; R. masafuerae Skottsb.,
Stuessy et al. s.n. (CONC), GU818435; Roldana suffulta (Greenm.) H.Rob. & Brettell,
Rzedowski 36569 (UC), GU818436; Rugelia nudicaulis Shuttlew. ex Chapm., Feist,
Phillippe, Molano-Flores, Busemeyer & Carroll 714 (TENN), GU818437; Scrobicaria
ilicifolia (L.f.) B.Nord., Cleef, Garcia-B. & Jaramillo-M. 3528 (U), GU818470; Senecio
acanthifolius Hombr. & Jacq., ex Decne., EF538034; S. adamantinus Bong.,
EF538059; S. algens Wedd., Beck 2879 (S), GU818438; S. arnicoides Hook. & Arn.,
Zöllner 3474 (L), GU818439; S. boyacensis (Cuatrec.) Cuatrec., Gonzalez 180 (S),
GU818440; S. brasiliensis (Spreng.) Less., UEC 50.171 (UEC), GU818441; S. cadiscus
B.Nord. & Pelser, Goldblatt & Manning 10690 (MO), GU818346; Rourke 1118 (S),
GU818347; S. chilensis Less., EF538043; S. ctenophyllus Phil., Zöllner 3959 (L),
GU818443; S. culcitioides Wedd., Øllgaard & Balslev 8822 (U), GU818442; S.
deltoideus Less., EF538076; S. elegans L., Cron & Goodman 687 (J), GU818444; S.
fistulosus Poepp. ex Less., Beck & Liberman 9672 (S), GU818445; S. flaccidus Less.,
DQ131873; S. flavus (Decne.) Sch.Bip., EF538087; S. gayanus (Colla) Cabr., Rosas
2157 (INIA), GU818446; S. gramineus Harv., Hoener 2104 (WAG), GU818447; S.
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gregorii F.Muell., Albrecht 7091 (NT), GU818448; S. hemmendorffii Malme,
EF538063; S. hieracium Remy, Baeza & Finot 3695 (CONC), GU818449; S.
hispidissimus I.Thomps., Thompson 927 (MEL), GU818450; S. hollandii Compton,
Germishuizen 6586 (WAG), GU818451; S. hypsobates Wedd., Øllgaard & Balslev 9863
(U), GU818452; S. ilicifolius Thunb., Cron & Goodman 686 (J), GU818453; S.
integerrimus Nutt., DQ131871; S. involucratus (Kunth) DC., EF538022; S. jarae Phil.,
EF538044; S. lastarrianus Remy, Ricardi 3230 (B), GU818454; S. latifolius DC.,
EF538084; S. lineatus DC., EF538075; S. mairetianus DC., EF538045; S. medleywoodii Hutch. (Caputia medley-woodii ined.), EF538080; S. meuselii Rauh, EF538073;
S. nemorensis L., EF538046; S. nevadensis Boiss. & Reut., Vogt 4172 (B), GU818455;
S. niveo-aureus Cuatrec., Cleef 6665 (S), GU818456; S. oerstedianus Benth. ex Oerst.,
Nordenstam 9160 (S), GU818457; S. oreophyton Remy, Beck 21589 (S), GU818458; S.
otites Kunze ex DC., EF538065; S. patens (Kunth) DC., Zak & Jaramillo 3427 (L),
GU818459; S. pflanzii (Perkins) Cuatrec., EF538027; S. pinifolius (L.) Lam.,
EF538079; S. pinnatifolius A.Rich. var. lanceolatus (Benth.) I.Thomps., Thompson
904 (MEL), GU818460; S. polygaloides Phil., Zöllner 5442 (L), GU818461; S.
prenanthoides A.Rich., Thompson 915 (MEL), GU818462; S. psilocarpus R.O. Belcher
& D.E. Albrecht, Thompson 937 (MEL), GU818463; S. retrorsus DC., EF538077; S.
roseus Sch.Bip., Garcia P. 250 (L), GU818464; S. saxatilis Wall. ex DC., EF538086; S.
scandens Buch.-Ham. ex D.Don, EF538048; S. scaposus DC. (Caputia scaposa ined.),
EF538049; S. squarrosus A.Rich., Thompson 907 (MEL), GU818465; S. stigophlebius
Baker, EF538072; S. suaveolens (L.) Ell., EF538032; S. superandinus Cuatrec.,
EF538035; S. szyszylowiczii Hiern., Sagastegui A. et al. 15797 (S), GU818466; S.
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tauricola V.A.Matthews, Budak 1735, Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası
Herbaryumu), GU818467; S. thapsoides DC., EF538085; S. triodon Phil. var. triodon,
Luebert & Teillier 2266 (CONC), GU818468; S. triqueter Less., EF538074; S. vestitus
P.J.Bergius, Greuter 21766 (B), GU818469; S. viscosus L., AY155666; Shafera
platyphylla Greenm., Nordenstam s.n. (S), GU818471; Sinacalia tangutica (Maxim.)
B.Nord., Lian et al 93-54 (MO), GU818472; Sinosenecio euosmus (Hand.-Mazz.)
B.Nord., Bouford, Donoghue & Ree 27625 (MO), GU818473; Solanecio biafrae (Oliv.
& Hiern) C.Jeffrey, Brunel 6220 (B), GU818474; Steirodiscus capillaceus (Thunb.)
Less., Cron & Goodman 653 (J), GU818475; Stilpnogyne bellidioides DC., Goldblatt &
Porter 11729 (MO), GU818476; Symphyotrichum novae-angliae (L.) G.L.Nesom, Tepe
1675 (MU), GU818335; Syneilesis palmata (Thunb.) Maxim., Hata s.n. (4-Aug-1979)
(CHR), GU818477; Synotis nagensium (C.B.Clarke) C.Jeffrey & Y.L.Chen,
EF538052; Tagetes sp., Vincent 13224 (MU), GU818478; Talamancalia boquetensis
(Standl.) H.Rob. & Cuatrec., Wilbur 15397 (NY), GU818479; Telanthophora
grandifolia (Less.) H.Rob. & Brettell, EF538050; Tephroseris integrifolia (L.) Holub
ssp. aucheri (DC.) B.Nord., Budak 1688 & Hamazaoǧlu (Yozgat Türkiye Florası
Herbaryumu), GU818481; Tetradymia filifolia Greene, EF538057; Tussilago farfara
L., DQ131857; Urostemon kirkii (Hook.f. ex Kirk) B.Nord., Cooper & Nickerson s.n.
(US), GU818482; Vernonia altissima Nutt., Tepe 1668 (MU), GU818483; Villasenoria
orcuttii (Greenm.) B.L.Clark, Robles 389 (XAL), GU818484; Werneria caespitosa
Wedd., Funk 11324 (US), GU818485; Xenophyllum poposum (Phil.) V.A.Funk, Funk
11351 (US), GU818486; Yermo xanthocephalus Dorn, Anderson 13691 (MO),
GU818487; Zemisia discolor (Sw.) B.Nord., Webster et al. 8420 (S), GU818488.
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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rbcL. Anacyclus valentinus L., Tepe 1186 & Moreno-Paez (MU), GU817737;
Arnoglossum atriplicifolium (L.) H.Rob., Vincent 3925 (MU), GU817738;
Arrhenechthites mixta (A.Rich.) Belcher, Lawrence 1308 (S), GU817739; Bahia
dissecta (Gray) Britton, Tuhy 3868 (MU), GU817741; Barnadesia caryophylla (Vell.)
S.F.Blake, L13859; Calendula arvensis L., Watson 95-13C (MU), GU817742; Capelio
caledonica B.Nord., Nordenstam 9644 (S), GU817743; Caucasalia parviflora (M.Bieb.)
B.Nord., Schneeweiss, Tribsch, Staudinger & Schönswetter 8643 (WU), GU817744;
Caxamarca sanchezii M.O.Dillon & Sagást., Sagástegui et al. 15548 (F), GU817745;
Centropappus brunonis Hook.f., Wapstra MW2 (MEL), GU817746; Chaenactis
douglasii (Hook.) Hook. & Arn., Vincent 8582 (MU), GU817747; Chersodoma
jodopappa (Sch.Bip.) Cabr., Hensen 2617 (S), GU817748; Cirsium discolor (Muhl.)
Spreng., Tepe 1670 (MU), GU817749; Coreopsis sp., Vincent 13226 (MU), GU817750;
Corymbium enerve Markötter, Trinder-Smith 124 (US), GU817751; Cotula
coronopifolia L., Watson & Panero 94-26 (MU), GU817752; Crassocephalum
crepidioides (Benth.) S.Moore, Pelser cult. 354 (L), GU817753; Dauresia alliariifolia
(O.Hoffm.) B.Nord & Pelser, Müller & Tilson 907 (M), GU817754; Dendrophorbium
bomanii (R.E.Fr.) C.Jeffrey, Dematteis & Seijo 722 (U), GU817755; Dendrosenecio
kilimanjari (Mildbr.) E.B.Knox ssp. cottonii (Hutch. & G.Taylor) E.B.Knox, Knox 50
(MICH), GU817756; Doronicum pardalianches L., Cubr 40487 (B), GU817757; Emilia
coccinea (Sims) G.Don., Pelser cult. 126 (L), GU817758; Erato polymnoides DC., Tepe
1456 (MU), GU817759; Eupatorium serotinum Michx., Tepe 1667 (MU), GU817760;
Faujasia squamosa (Bory) C.Jeffrey, Nordenstam 9210 (S), GU817761; Galinsoga
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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quadriradiata Cav., Vincent 13227 (MU), GU817762; Gerbera sp., Vincent 13223
(MU), GU817763; Gymnodiscus capillaris (L.f.) Less., AM234876; Helenium
autumnale L., Tepe 1664 (MU), GU817764; Helianthus tuberosus L., Tepe 1672 (MU),
GU817765; Helichrysum stoechas DC., Tepe 1092 (MU), GU817766; Homogyne
alpina (L.) Cass., Uhink 98-56 (MJG), GU817767; Inula viscosa (L.) Aiton, Tepe 1142
(MU), GU817768; Jacobaea vulgaris Gaertn., Pelser cult. 6 (L), GU817769; Kleinia
neriifolia Haw., Pelser cult. 216 (L), GU817770; Lactuca canadensis L., Vincent 13128
(MU), GU817771; Lamprocephalus montanus B.Nord., Nordenstam 9542 (S),
GU817772; Lopholaena coriifolia (Sond.) Phillips & C.A.Sm., Van der Westhuizen &
Liamé; s.n. (PREM), GU817773; Luina hypoleuca Benth., Greuter 17706 (B),
GU817774; Madia sp., Vincent 12912 (MU), GU817775; Mikaniopsis clematoides
(Sch.Bip. ex A.Rich.) Milne-Redh., Friis, Hounde & Jacobsen 499 (WAG), GU817776;
Othonna capensis Bailey, Pelser cult. 106 (L), GU817777; Packera eurycephala (Torr.
& A.Gray) W.A.Weber & Á.Löve, Vincent 8581 (MU), GU817778; Perityle emoryi
Torr., Atwood 26483 (MU), GU817779; Phaneroglossa bolusii (Oliv.) B.Nord., Watson
& Panero 94-62 (TEX), GU817780; Polymnia canadensis L., Tepe 1665 (MU),
GU817781; Senecio deltoideus Less., Sloet S.A.105 (U), GU817782; S. medley-woodii
Hutch. (Caputia medley-woodii ined.), Jeffrey Kew cult. 14 (K), GU817783;
Steirodiscus capillaceus (Thunb.) Less., Cron & Goodman 653 (J), GU817784;
Symphyotrichum novae-angliae (L.) G.L.Nesom, Tepe 1675 (MU), GU817740; Tagetes
sp., Vincent 13224 (MU), GU817785; Tephroseris integrifolia (L.) Holub ssp. aucheri
(DC.) B.Nord., Budak 1688 & Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu),
GU817786; Vernonia altissima Nutt., Tepe 1668 (MU), GU817787.
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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trnL and trnL-F. Abrotanella emarginata (Gaudich.) Cass., EF538089; Acrisione
cymosa (J.Rémy) B.Nord., Lammers, Rodriques & Baeza 6447 (MU), GU817965;
Adenostyles alpina (L.) Bluff & Fingerh., Schwerdtfeger 12251 (B), GU817966;
Aequatorium asterotrichum B.Nord., Asplund 18263 (S), GU817967; Anacyclus
valentinus L., Tepe 1186 & Moreno-Paez (MU), GU817959; Antillanthus almironcillo
(M.Gómez) B.Nord., El-Ghazaby et al. 23.608 (S), GU817968; Arbelaezaster
ellsworthii (Cuatrec.) Cuatrec., Kilip & Smith 19413 (US), GU817969; Arnoglossum
atriplicifolium (L.) H.Rob., EF538091; Arrhenechthites mixta (A.Rich.) Belcher,
EF538092; Austrosynotis rectirama (Baker) C.Jeffrey, LaCroix 4001 (MO),
GU817971; Bahia dissecta (Gray) Britton, Tuhy 3868 (MU), GU817954, GU817944;
Barkleyanthus salicifolius (Kunth) H.Rob. & Brettell, Genelle & Fleming 861 (B),
GU817972; Barnadesia sp., Hunziker & Gamerro 12561 (MU), GU817973; Bedfordia
arborescens Hochr., Greuter 21319 (B), GU817974; AF452498; Bethencourtia
palmensis (Nees) Choisy, Nordenstam 9326 (S), GU817975; Blennosperma nanum
S.F.Blake, AF460174; EF028720; Blepharispermum zanguebaricum Oliv. & Hiern.,
Medley 570 (MU), GU817976; AY216147; Bolandia pedunculosa (DC.) Cron,
AY952915; Brachyglottis repanda J.R.Forst. & G.Forst., Schwerdtfeger 17407 (B),
GU817977; Cabreriella oppositicordia (Cuatrec.) Cuatrec., Romero-Castañeda 7428
(US), GU817978; Cacaliopsis nardosmia (A.Gray) A.Gray, Ertter 6502 (UC),
GU817979; Calendula arvensis L., Watson 95-13C (MU), GU817981; Capelio
caledonica B.Nord., Nordenstam 9644 (S), GU817982; Caucasalia parviflora (M.Bieb.)
B.Nord., EF538093; Caxamarca sanchezii M.O.Dillon & Sagást., Sagástegui et al.
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15548 (F), GU817983; Centropappus brunonis Hook.f., Wapstra MW2 (MEL),
GU817984; Chaenactis douglasii (Hook.) Hook. & Arn., Vincent 8582 (MU),
GU817985; Charadranaetes durandii (Klatt) Janovec & H.Rob., Gomez P. 2242 (S),
GU817986; Chersodoma jodopappa (Sch.Bip.) Cabr., EF538094; Cichorium intybus
L., Tepe 1673 (MU), GU817987; Cineraria abyssinica Sch.Bip. ex A.Rich., AF460173;
EF028721; Cirsium discolor (Muhl.) Spreng., Tepe 1670 (MU), GU817988;
Cissampelopsis volubilis (Blume) Miq., Carvalho 3175 (MU), GU817989; Coreopsis
sp., Vincent 13226 (MU), GU817943, GU817953; Cotula coronopifolia L., Watson &
Panero 94-26 (MU), GU817960; Crassocephalum crepidioides (Benth.) S.Moore,
AF460138; EF028722; Cremanthodium humile Maxim., EF538095; AY723195;
Crocidium multicaule Hook., Bartholomew 5749 (MO), GU817990; Curio rowleyanus
(H.Jacobsen) P.V.Heath, AF460148; EF028731; Dauresia alliariifolia (O.Hoffm.)
B.Nord & Pelser, Müller & Tilson 907 (M), GU817991; Delairea odorata Lem., Ertter
5766 (UC), GU817992; Dendrophorbium bomanii (R.E.Fr.) C.Jeffrey, EF538097;
Dendrosenecio kilimanjari (Mildbr.) E.B.Knox ssp. cottonii (Hutch. & G.Taylor)
E.B.Knox, AF460172; AY952923; Digitacalia jatrophoides (Kunth) Pippen, Cruden
1086 (UC), GU817993; Dolichoglottis lyallii (Hook.f.) B.Nord., Strid 22172 (MO),
GU817994; Dolichorrhiza caucasica (M.Bieb.) Galushko, Schneeweiss 44 (WU),
GU817995; Dorobaea pimpinellifolia (Kunth) B.Nord., Tepe 1467 (MU), GU817996;
Doronicum pardalianches L., EF538098; AJ400125; Ekmaniopappus mikanioides
(Urb. & Ekman) Borhidi, Zanoni et al. 33908 (JBSD), GU817997; Elekmania picardae
(Krug & Urb.) B.Nord., EF538099; Emilia coccinea (Sims) G.Don., AF460171;
EF028723; Endocellion sibiricum (J.F.Gmel.) J.Toman, Schönswetter & Tribsch T480
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(WU), GU817998; Erato polymnoides DC., Tepe 1456 (MU), GU817999; Erechtites
valerianifolius (Link ex Spreng.) Less. ex DC., EF538100; Eriothrix lycopodioides
DC., Nordenstam 9208 (S), GU818000; Eupatorium serotinum Michx., Tepe 1667
(MU), GU818001; Euryops pectinatus (L.) Cass., AF460170; EF028724; Farfugium
japonicum (L.) Kitam., AF468163; AF468177; Faujasia squamosa (Bory) C.Jeffrey,
Nordenstam 9210 (S), GU818002; Faujasiopsis flexuosa (Lam.) C.Jeffrey ssp.
bourbonensis C.Jeffrey, Nordenstam 9202 (S), GU818003; Galinsoga quadriradiata
Cav., Vincent 13227 (MU), GU817942, GU817956; Galinsoga quadriradiata Cav.,
AY216179; Garcibarrigoa telembina (Cuatrec.) Cuatrec., Holm-Nielsen et al. 6211 (S),
GU817940, GU817951; Gerbera sp., Vincent 13223 (MU), GU817941; GU817955;
Graphistylis dichroa (Bong.) D.J.N.Hind, Silva 140 & Zelma (S), GU818004;
Gymnodiscus capillaris (L.f.) Less., Coppejans EC1116 (U), GU818005; Gynoxys
soukupii Cuatrec., EF538101; Gynura divaricata (L.) DC. ssp. formosana (Kitam.)
F.G.Davies, AF460136; Panero & Hsiao 6457 (TEX), GU818006; Helenium
autumnale L., Tepe 1664 (MU), GU818007; Helianthus tuberosus L., Tepe 1672 (MU),
GU818008; Helichrysum stoechas DC., Tepe 1092 (MU), GU818009; Herodotia
haitiensis Urb. & Ekman, EF538103; Herreranthus rivalis (Greenm.) B.Nord.,
Nordenstam s.n. (2/12/2007) (S), GU818010; Hertia pallens (DC.) Kuntze, Koekemoer
& Funk 1963 (US), GU818011; Hoehnephytum trixoides (Gard.) Cabr., Carmo 131
(BHCB), GU818012; Homogyne alpina (L.) Cass., Uhink 98-56 (MJG), GU818013;
Hubertia riparia (DC.) C.Jeffrey, Labat, Phillipson & Lowry II 2022 (WAG),
GU818014; Humbertacalia sp., Phillipson et al. 5641 (P), GU818015; Ignurbia
constanzae (Urb.) B.Nord., Nordenstam & Lundin 560 (S), GU818016; Inula viscosa
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(L.) Aiton, Tepe 1142 (MU), GU818017; Io ambondrombeensis (Humbert) B.Nord.,
Malcomber et al. s.n. (P), GU818019; Malcomber et al. 1380 (MO), GU818018;
Iranecio cariensis (Boiss.) C.Jeffrey, Budak 1724, Aksoy & Hamazaoǧlu (Yozgat
Türkiye Florası Herbaryumu), GU818020; Ischnea elachoglossa F.Muell., Beaman
12129 (MO), GU818021; Jacmaia incana (Sw.) B.Nord., Nordenstam 7914 (S),
GU818022; Jacobaea vulgaris Gaertn., AF460153; EF028725; AY155621; Jessea
multivenia (Benth.) H.Rob. & Cuatrec., Nordenstam 9161 (S), GU818023; Kleinia
neriifolia Haw., AF460169; EF028726; Lachanodes arborea (Roxb.) B.Nord., CairnsWicks s.n., GU818024; Lactuca canadensis L., Vincent 13128 (MU), GU818025;
Lamprocephalus montanus B.Nord., Nordenstam 9542 (S), GU818026; Leonis
trineurus (Griseb.) B.Nord., Smith & al. 3238 (S), GU818027; Lepidospartum
burgessii B.L.Turner, EF538106; Ligularia stenocephala (Maxim.) Matsum. &
Koidz., AF460168; EF538107; Lomanthus fosbergii (Cuatrec.) B. Nord. & Pelser,
Øllgaard et al. 90697 (AAU), GU818099; Lopholaena coriifolia (Sond.) Phillips &
C.A.Sm., Van der Westhuizen & Liamé; s.n. (PREM), GU818028; Lordhowea insularis
(Benth.) B.Nord., AF460135; Trodd & Thorne 3703 (L), GU818029; Luina hypoleuca
Benth., Greuter 17706 (B), GU818030; Lundinia plumbea (Griseb.) B.Nord., Zanoni
45816 (JBSD), GU818031; Madia sp., Vincent 12912 (MU), GU818032; Mesogramma
apiifolium DC., Giess 16074 (M), GU817948, GU817957; Mikaniopsis clematoides
(Sch.Bip. ex A.Rich.) Milne-Redh., De Wilde & De Wilde-Duyfjes 9006 (B),
GU818033; Miricacalia makineana (Yatabe) Kitam., Kobayashi 33484 (HYO),
GU818034; Misbrookea strigosissima (A.Gray) V.A.Funk, EF538108; Monticalia
abietina (Willd. ex Wedd.) C.Jeffrey, Cleef 8411 (U), GU818035; Monticalia apiculata
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(Sch.Bip. ex Wedd.) C.Jeffrey, Alston 6935 (S), GU818036; Monticalia arbutifolia
(Kunth.) C.Jeffrey, EF538109; Nelsonianthus tapianus (B.L.Turner) C.Jeffrey, Pérez
& Kendizabal 457 (XAL), GU818037; Nemosenecio nikoensis (Miq.) B.Nord., Koyama
4079 (L), GU818038; Nesampelos lucens (Poir. in Lam.) B.Nord., EF538110;
Neurolaena lobata (L.) R.Br., Acevedo 9316 (US), GU818039; AY216209;
Nordenstamia kingii (H.Rob. & Cuatrec.) B.Nord., Ståhl 5572A (S), GU818040;
Oldfeltia polyphlebia (Griseb.) B.Nord. & Lundin, EF538112; Oresbia heterocarpa
Cron & B.Nord., Nordenstam 9628 (S), GU818041; AY952925; Othonna capensis
Bailey, AF460167; EF028727; Packera eurycephala (Torr. & A.Gray) W.A.Weber &
Á.Löve, EF538113; Papuacalia dindondl (P.Royen) Veldkamp, EF538114; Paracalia
jungioides (Hook. & Arn.) Cuatrec., King & Collins 9017 (UC), GU818042;
Parafaujasia fontinalis (Cordem.) C.Jeffrey, Lorence 2602 (MO), GU817949,
GU817958; Paragynoxys neodendroides (Cuatrec.) Cuatrec., Cleef 9852a (U),
GU818043; Parasenecio adenostyloides (Franch. et Sav. ex Maxim.) H.Koyama,
AF460139; EF028728; Pentacalia arborea (Kunth) H.Rob. & Cuatrec., Øllgaard &
Balslev 8298 (U), GU818044; Pericallis murrayi (Bornm.) B.Nord., EF538115;
AY952922; Perityle emoryi Torr., Atwood 26483 (MU), GU818045; Petasites albus (L.)
Gaertn., Cubr 40398 (B), GU818046; OPTIMA Iter VIII 1084 (RNG), GU818047;
Phaneroglossa bolusii (Oliv.) B.Nord., AF460131; EF538116; AY216221; Pippenalia
delphinifolia (Rydb.) MacVaugh, Spellenberg & Bacon 11048 (TEX), GU818048;
Pittocaulon praecox (Cav.) H.Rob. & Brettell, EF538117; Pladaroxylon leucadendron
(G.Forst.) Hook.f., Cairns-Wicks s.n., GU818049; Pojarkovia pojarkovae (Schischk.)
Greuter, Pelser cult. 191 (L), GU818050; Polymnia canadensis L., Tepe 1665 (MU),
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GU818051; AY216224; Psacaliopsis purpusii (Greenm. ex Brandegee) H.Rob. &
Brettell, Panero 2607, Davila & Tenorio (TEX), GU818052; Psacalium cirsiifolium
(Zucc.) H.Rob. & Brettell, EF538111; Pseudogynoxys haenkei (DC.) Cabr., Molina R.
18431 (U), GU818053; Rainiera stricta (Greene) Greene, Dennis 2317 (U), GU817952;
Robinsonecio gerberifolius (Sch.Bip. ex Hemsl.) T.M.Barkley & Janovec, Garcia P.
171 (MO), GU818054; Robinsonia berteroi (DC.) R.W.Sanders, Stuessy & Martic.,
AF460129; EF538142; R. evenia Phil., Stuessy et al. 11308 (CONC), GU818055; R.
gracilis Decne., EF538118; Stuessy et al. 11282 (CONC), GU817947, GU817962;
Stuessy et al. 11312 (CONC), GU817946, GU817963; R. masafuerae Skottsb., Stuessy
et al. s.n. (CONC), GU817945; GU817961; Roldana suffulta (Greenm.) H.Rob. &
Brettell, Rzedowski 36569 (UC), GU818056; Rugelia nudicaulis Shuttlew. ex Chapm.,
Feist, Phillippe, Molano-Flores, Busemeyer & Carroll 714 (TENN), GU818057;
Scrobicaria ilicifolia (L.f.) B.Nord., Cleef, Garcia-B. & Jaramillo-M. 3528 (U),
GU818090; Senecio acanthifolius Hombr. & Jacq., ex Decne., EF538104; S.
adamantinus Bong., EF538119; S. algens Wedd., Beck 2879 (S), GU818058; S.
arnicoides Hook. & Arn., Zöllner 3474 (L), GU818059; S. boyacensis (Cuatrec.)
Cuatrec., Gonzalez 180 (S), GU818060; S. brasiliensis (Spreng.) Less., UEC 50.171
(UEC), GU818061; S. cadiscus B.Nord. & Pelser, Rourke 1118 (S), GU817980; S.
chilensis Less., EF538122; S. ctenophyllus Phil., Zöllner 3959 (L), GU818063; S.
culcitioides Wedd., Øllgaard & Balslev 8822 (U), GU818062; S. deltoideus Less.,
AY952917; S. elegans L., Cron & Goodman 687 (J), GU818064; S. fistulosus Poepp. ex
Less., Beck & Liberman 9672 (S), GU818065; S. flaccidus Less. var. flaccidus, Jardin
Thuret cult. s.n. (MJG), GU818066; S. flavus (Decne.) Sch.Bip., DQ208176; EF028729;
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S. gayanus (Colla) Cabr., Rosas 2157 (INIA), GU818067; S. gramineus Harv., Hoener
2104 (WAG), GU818068; S. gregorii F.Muell., Albrecht 7091 (NT), GU818069; S.
hemmendorffii Malme, EF538124; S. hieracium Remy, Baeza & Finot 3695 (CONC),
GU818070; S. hispidissimus I.Thomps., Thompson 927 (MEL), GU818071; S. hollandii
Compton, Germishuizen 6586 (WAG), GU818072; S. hypsobates Wedd., Øllgaard &
Balslev 9863 (U), GU818073; S. ilicifolius Thunb., Cron & Goodman 686 (J),
GU818074; S. integerrimus Nutt. var. exaltatus (Nutt.) Cronquist, Crockett 437 (MU),
GU818075; S. involucratus (Kunth) DC., EF538090; S. jarae Phil., EF538125; S.
lastarrianus Remy, Ricardi 3230 (B), GU818076; S. latifolius DC., EF538126; S.
lineatus DC., AF098852; AF100515; S. mairetianus DC., EF538128; S. medley-woodii
Hutch. (Caputia medley-woodii ined.), AF460151; EF538129; S. meuselii Rauh,
EF538130; S. nemorensis L., AF460150; EF028730; S. nevadensis Boiss. & Reut., Vogt
4172 (B), GU818077; S. niveo-aureus Cuatrec., Cleef 6665 (S), GU818078; S.
oerstedianus Benth. ex Oerst., Nordenstam 9160 (S), GU817950; S. oreophyton Remy,
Beck 21589 (S), GU818079; S. otites Kunze ex DC., EF538131; S. patens (Kunth) DC.,
Zak & Jaramillo 3427 (L), GU818080; S. pflanzii (Perkins) Cuatrec., EF538096; S.
pinifolius (L.) Lam., EF538132; S. pinnatifolius A.Rich. var. lanceolatus (Benth.)
I.Thomps., Thompson 904 (MEL), GU818081; S. polygaloides Phil., Zöllner 5442 (L),
GU818082; S. prenanthoides A.Rich., Thompson 915 (MEL), GU818083; S.
psilocarpus R.O. Belcher & D.E. Albrecht, Thompson 937 (MEL), GU818084; S.
retrorsus DC., EF538133; S. roseus Sch.Bip., Garcia P. 250 (L), GU818085; S.
saxatilis Wall. ex DC., EF538134; S. scandens Buch.-Ham. ex D.Don, AF460147;
EF028732; S. scaposus DC. (Caputia scaposa ined.), AF460146; EF028733; S.
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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squarrosus A.Rich., Thompson 907 (MEL), GU817964; S. stigophlebius Baker,
EF538135; S. suaveolens (L.) Ell., EF538102; S. superandinus Cuatrec., EF538105; S.
szyszylowiczii Hiern., Sagastegui A. et al. 15797 (S), GU818086; S. tauricola
V.A.Matthews, Budak 1735, Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası
Herbaryumu), GU818087; S. thapsoides DC., EF538137; S. triodon Phil. var. triodon,
Luebert & Teillier 2266 (CONC), GU818088; S. triqueter Less., EF538138; S. vestitus
P.J.Bergius, Greuter 21766 (B), GU818089; S. viscosus L., AF460142; EF028734;
Shafera platyphylla Greenm., Nordenstam s.n. (S), GU818091; Sinacalia tangutica
(Maxim.) B.Nord., AY176088; AY723199; Sinosenecio euosmus (Hand.-Mazz.)
B.Nord., Bouford, Donoghue & Ree 27625 (MO), GU818092; Solanecio biafrae (Oliv.
& Hiern) C.Jeffrey, Brunel 6220 (B), GU818093; Steirodiscus capillaceus (Thunb.)
Less., Cron & Goodman 653 (J), GU818094; Stilpnogyne bellidioides DC., Goldblatt &
Porter 11729 (MO), GU818095; Symphyotrichum novae-angliae (L.) G.L.Nesom, Tepe
1675 (MU), GU817970; Syneilesis palmata (Thunb.) Maxim., Hata s.n. (4-Aug-1979)
(CHR), GU818096; Synotis nagensium (C.B.Clarke) C.Jeffrey & Y.L.Chen,
AF460121; EF028735; Tagetes sp., Vincent 13224 (MU), GU818097; Talamancalia
boquetensis (Standl.) H.Rob. & Cuatrec., Wilbur 15397 (NY), GU818098;
Telanthophora grandifolia (Less.) H.Rob. & Brettell, EF538140; Tephroseris
integrifolia (L.) Holub ssp. aucheri (DC.) B.Nord., Budak 1688 & Hamazaoǧlu
(Yozgat Türkiye Florası Herbaryumu), GU818100; Tetradymia filifolia Greene,
EF538141; Tussilago farfara L., AF468166; AJ400103; Urostemon kirkii (Hook.f. ex
Kirk) B.Nord., Cooper & Nickerson s.n. (US), GU818101; Vernonia altissima Nutt.,
Tepe 1668 (MU), GU818102; Villasenoria orcuttii (Greenm.) B.L.Clark, Robles 389
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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(XAL), GU818103; Werneria caespitosa Wedd., Funk 11324 (US), GU818104;
Xenophyllum poposum (Phil.) V.A.Funk, Funk 11351 (US), GU818105; Yermo
xanthocephalus Dorn, Anderson 13691 (MO), GU818106; Zemisia discolor (Sw.)
B.Nord., Webster et al. 8420 (S), GU818107.
ITS. Abrotanella emarginata (Gaudich.) Cass., EF538143; AY929883; Acrisione
cymosa (J.Rémy) B.Nord., EF538144; Adenostyles alpina (L.) Bluff & Fingerh.,
EF538146; Aequatorium asterotrichum B.Nord., Asplund 18263 (S), GU818489;
EF538147; Anacyclus valentinus L., Tepe 1186 & Moreno-Paez (MU), GU818490;
Antillanthus almironcillo (M.Gómez) B.Nord., EF538153; Arbelaezaster ellsworthii
(Cuatrec.) Cuatrec., Kilip & Smith 19413 (US), GU818491; Arnoglossum
atriplicifolium (L.) H.Rob., EF538154; Arrhenechthites mixta (A.Rich.) Belcher,
EF538156; Austrosynotis rectirama (Baker) C.Jeffrey, EF538157; Bahia dissecta
(Gray) Britton, Tuhy 3868 (MU), GU818493; Barkleyanthus salicifolius (Kunth)
H.Rob. & Brettell, Genelle & Fleming 861 (B), GU817578; AF161598; AF161648;
Barnadesia caryophylla (Vell.) S.F.Blake, AY504686; Bedfordia arborescens Hochr.,
Greuter 21319 (B), GU818494; Bethencourtia palmensis (Nees) Choisy, EF538160;
Blennosperma nanum S.F.Blake, AF459971; EF538158; AF161597; AF161647;
Blepharispermum zanguebaricum Oliv. & Hiern., AF229259; AF229260; Bolandia
pedunculosa (DC.) Cron, Cron & Goodman 681 (J), GU818495; AY953925;
Brachyglottis repanda J.R.Forst. & G.Forst., AY554103; Cabreriella oppositicordia
(Cuatrec.) Cuatrec., Romero-Castañeda 7428 (US), GU818496; Cacaliopsis nardosmia
(A.Gray) A.Gray, Ertter 6502 (UC), GU818497; AF161599; AF161649; Calendula
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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arvensis L., Watson 95-13C (MU), GU818507; Capelio caledonica B.Nord.,
Nordenstam 9644 (S), GU818508; Caucasalia parviflora (M.Bieb.) B.Nord., EF538163;
EF538162; Caxamarca sanchezii M.O.Dillon & Sagást., Sagástegui et al. 15548 (F),
GU818509; Centropappus brunonis Hook.f., Wapstra MW2 (MEL), GU818510;
AY554095; Chaenactis douglasii (Hook.) Hook. & Arn., Vincent 8582 (MU),
GU818511; Charadranaetes durandii (Klatt) Janovec & H.Rob., EF538164;
Chersodoma jodopappa (Sch.Bip.) Cabr., EF538167; Cineraria abyssinica Sch.Bip. ex
A.Rich., AF459969; Pelser cult. 208 (L), GU818512- GU818521; Cirsium discolor
(Muhl.) Spreng., AF443692; Cissampelopsis volubilis (Blume) Miq., EF538172;
Coreopsis sp., Vincent 13226 (MU), GU818522-GU818531; Corymbium enerve
Markötter, Trinder-Smith 124 (US), GU818532-GU818540; EF581385; Cotula
coronopifolia L., AF422118; Crassocephalum crepidioides (Benth.) S.Moore,
AF459968; EF538173; Cremanthodium humile Maxim., EF538175; AY723270;
Crocidium multicaule Hook., Bartholomew 5749 (MO), GU818541; AY929902; Curio
rowleyanus (H.Jacobsen) P.V.Heath, AF459933; DQ915888; Dauresia alliariifolia
(O.Hoffm.) B.Nord & Pelser, AF457413; Delairea odorata Lem., Ertter 5766 (UC),
GU817579; Tepe 2180, Pelser & Marticorena (MU), GU818542; Dendrocacalia
crepidifolia (Nakai) Nakai, Kobayashi 190493 (MO), GU818543; AY723280;
Dendrophorbium bomanii (R.E.Fr.) C.Jeffrey, EF538181; Dendrosenecio kilimanjari
(Mildbr.) E.B.Knox, AF155963; AF155996; D. kilimanjari (Mildbr.) E.B.Knox ssp.
cottonii (Hutch. & G.Taylor) E.B.Knox, AF459967; AY953933; Digitacalia
jatrophoides (Kunth) Pippen, Panero 2330 & Salinas (TEX), GU818544; Cruden 1086
(UC), GU818545; Dolichoglottis lyallii (Hook.f.) B.Nord., Strid 22172 (MO),
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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GU818546; Dolichorrhiza caucasica (M.Bieb.) Galushko, Schneeweiss 44 (WU),
GU818547; Dorobaea pimpinellifolia (Kunth) B.Nord., Tepe 1467 (MU), GU818548;
AF155964; AF155997; Doronicum pardalianches L., EF538188; AJ400032; AJ400033;
AJ400078; AJ400079; Ekmaniopappus mikanioides (Urb. & Ekman) Borhidi,
EF538189; Zanoni et al. 33908 (JBSD), GU817577; Elekmania picardae (Krug &
Urb.) B.Nord., EF538193; Emilia coccinea (Sims) G.Don., AF459966; EF538194;
Endocellion sibiricum (J.F.Gmel.) J.Toman, EF538197; EF538198; Erechtites
valerianifolius (Link ex Spreng.) Less. ex DC., EF538199; Eriothrix lycopodioides
DC., EF538200; Eupatorium serotinum Michx., AB032050; DQ236176; Euryops
pectinatus (L.) Cass., AF459964; AF155965; AF155998; Farfugium japonicum (L.)
Kitam., Wu 1439 (MO), GU817572; AY176139; DQ272323; Faujasia squamosa
(Bory) C.Jeffrey, EF538210; Faujasiopsis flexuosa (Lam.) C.Jeffrey ssp.
bourbonensis C.Jeffrey, Nordenstam 9202 (S), GU818549; Galinsoga quadriradiata
Cav., Vincent 13227 (MU), GU818550; Garcibarrigoa telembina (Cuatrec.) Cuatrec.,
EF538211; Gerbera sp., Vincent 13223 (MU), GU818551; Graphistylis dichroa (Bong.)
D.J.N.Hind, EF538212; Gymnodiscus capillaris (L.f.) Less., EF538217; Esterhuysen
34744 (S), GU817575; Cron & Goodman 650 (J), GU818552; Gynoxys soukupii
Cuatrec., AF459963; Gynura divaricata (L.) DC. ssp. formosana (Kitam.) F.G.Davies,
AF155966; AF155999; Haastia pulvinaris Hook.f., AF422122; Helenium autumnale
L., Tepe 1664 (MU), GU818553; Helianthus tuberosus L., AF047953; Helichrysum
stoechas DC. ssp. stoechas, AY445193; Herodotia haitiensis Urb. & Ekman,
EF538224; Herreranthus rivalis (Greenm.) B.Nord., Nordenstam s.n. (2/12/2007) (S),
GU818554; Hertia pallens (DC.) Kuntze, EF538229; Hoehnephytum trixoides (Gard.)
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Cabr., Carmo 131 (BHCB), GU818555; Homogyne alpina (L.) Cass., Uhink 98-56
(MJG), GU817576; EF538230; EF538231; Hubertia riparia (DC.) C.Jeffrey, Labat,
Phillipson & Lowry II 2022 (WAG), GU818556; Humbertacalia sp., Phillipson et al.
5641 (P), GU818557; Ignurbia constanzae (Urb.) B.Nord., Nordenstam & Lundin 560
(S), GU817574; Inula viscosa (L.) Aiton, Tepe 1142 (MU), GU818558; Io
ambondrombeensis (Humbert) B.Nord., Malcomber et al. s.n. (P), GU818560;
Malcomber et al. 1380 (MO), GU818559; Iranecio cariensis (Boiss.) C.Jeffrey, Budak
1724, Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu), GU818561; Ischnea
elachoglossa F.Muell., Beaman 12129 (MO), GU818562; Jacobaea vulgaris Gaertn.,
AF459941; Pelser cult. 6 (L), GU818563-GU818567; AY155610; AF161642;
AF161692; AY554112; Jessea multivenia (Benth.) H.Rob. & Cuatrec., EF538246;
Kleinia neriifolia Haw., AF459962; Pelser cult. 216 (L), GU818568-GU818573;
DQ915868; Lachanodes arborea (Roxb.) B.Nord., Cairns-Wicks s.n., GU818574;
Lactuca canadensis L., Vincent 13128 (MU), GU818575; Lamprocephalus montanus
B.Nord., Nordenstam 9542 (S), GU818576; Leonis trineurus (Griseb.) B.Nord.,
EF538249; Smith & al. 3238 (S), GU818577-GU818583; Lepidospartum burgessii
B.L.Turner, EF538250; Ligularia stenocephala (Maxim.) Matsum. & Koidz.,
AF459961; Lomanthus bangii (Rusby) B. Nord. & Pelser, Beck 3527 (S), GU818633GU818640; L. fosbergii (Cuatrec.) B. Nord. & Pelser, Øllgaard et al. 90697 (AAU),
GU818723; Lopholaena coriifolia (Sond.) Phillips & C.A.Sm., EF538251; Lordhowea
insularis (Benth.) B.Nord., Nordenstam 8615 (S), GU818584-GU818592; Luina
hypoleuca Benth., Greuter 17706 (B), GU818593; AF161601; AF161651; Lundinia
plumbea (Griseb.) B.Nord., EF538252; Madia sp., Vincent 12912 (MU), GU818594;
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Mattfeldia triplinervis Urb., EF538253; Mesogramma apiifolium DC., AF457412;
Mikaniopsis clematoides (Sch.Bip. ex A.Rich.) Milne-Redh., Friis, Hounde & Jacobsen
499 (WAG), GU817581; De Wilde & De Wilde-Duyfjes 9006 (B), GU818595;
Miricacalia makineana (Yatabe) Kitam., Kobayashi 33484 (HYO), GU818596;
AY723281; Misbrookea strigosissima (A.Gray) V.A.Funk, EF538254; Monticalia
abietina (Willd. ex Wedd.) C.Jeffrey, EF538255; Monticalia apiculata (Sch.Bip. ex
Wedd.) C.Jeffrey, EF538256; Monticalia arbutifolia (Kunth.) C.Jeffrey, EF538257;
Nelsonianthus tapianus (B.L.Turner) C.Jeffrey, Pérez & Kendizabal 457 (XAL),
GU817580; Nemosenecio nikoensis (Miq.) B.Nord., EF538264; EF538265; AY723279;
Nesampelos lucens (Poir. in Lam.) B.Nord., EF538266; Neurolaena lobata (L.) R.Br.,
Acevedo 9316 (US), GU818597; Nordenstamia kingii (H.Rob. & Cuatrec.) B.Nord.,
EF538267; Oldfeltia polyphlebia (Griseb.) B.Nord. & Lundin, EF538271; Oresbia
heterocarpa Cron & B.Nord., Nordenstam 9628 (S), GU818598-GU818607;
AY953935; Othonna capensis Bailey, AF459960; DQ915865; Packera eurycephala
(Torr. & A.Gray) W.A.Weber & Á.Löve, EF538276; Vincent 8581 (MU), GU818608GU818617; Papuacalia dindondl (P.Royen) Veldkamp, EF538279; Paracalia
jungioides (Hook. & Arn.) Cuatrec., King & Collins 9017 (UC), GU818618;
Parafaujasia fontinalis (Cordem.) C.Jeffrey, Lorence 2602 (MO), GU818619;
Paragynoxys neodendroides (Cuatrec.) Cuatrec., EF538280; Parasenecio
adenostyloides (Franch. et Sav. ex Maxim.) H.Koyama, AF459970; Pentacalia
arborea (Kunth) H.Rob. & Cuatrec., EF538283; Pericallis aurita (L'Hér.) B.Nord.,
AF155974; AF155975; AF156007; AF156008; P. hansenii (G.Kunkel) Sunding,
AF155979; AF156012; P. murrayi (Bornm.) B.Nord., EF538285; AF155983;
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AF156016; AY953932; Perityle emoryi Torr., AF374868; Petasites albus (L.) Gaertn.,
Cubr 40398 (B), GU818620; OPTIMA Iter VIII 1084 (RNG), GU818621; Phaneroglossa
bolusii (Oliv.) B.Nord., AF155991; AF156024; Watson & Panero 94-62 (TEX),
GU818622-GU818626; Pippenalia delphinifolia (Rydb.) MacVaugh, Breedlove 59032
& Almeda (MO), GU817571; Spellenberg & Bacon 11048 (TEX), GU818627;
Pittocaulon praecox (Cav.) H.Rob. & Brettell, AF161603; AF161653; Pladaroxylon
leucadendron (G.Forst.) Hook.f., Cairns-Wicks s.n., GU818628; Pojarkovia pojarkovae
(Schischk.) Greuter, EF538286; Polymnia canadensis L., AF465876; Psacaliopsis
purpusii (Greenm. ex Brandegee) H.Rob. & Brettell, Panero 2607, Davila & Tenorio
(TEX), GU818629; Psacalium cirsiifolium (Zucc.) H.Rob. & Brettell, EF538270;
Pseudogynoxys haenkei (DC.) Cabr., EF538288; Rainiera stricta (Greene) Greene,
EF538289; AF161605; AF161655; Robinsonecio gerberifolius (Sch.Bip. ex Hemsl.)
T.M.Barkley & Janovec, Garcia P. 171 (MO), GU818630; AF161606; AF161656;
Robinsonia berteroi (DC.) R.W.Sanders, Stuessy & Martic., AF459957; EF028712;
EF028719; R. evenia Phil., EF028706; EF028707; EF028713; EF028714; R. gracilis
Decne., EF538290; EF028709; EF028716; R. masafuerae Skottsb., EF028710;
EF028717; Roldana suffulta (Greenm.) H.Rob. & Brettell, Rzedowski 36569 (UC),
GU818631; Rugelia nudicaulis Shuttlew. ex Chapm., Feist, Phillippe, Molano-Flores,
Busemeyer & Carroll 714 (TENN), GU818632; Scrobicaria ilicifolia (L.f.) B.Nord.,
EF538399; Senecio acanthifolius Hombr. & Jacq., ex Decne., EF538238; S.
adamantinus Bong., EF538294; S. algens Wedd., EF538296; S. arnicoides Hook. &
Arn., EF538298; S. boyacensis (Cuatrec.) Cuatrec., EF538176; S. brasiliensis
(Spreng.) Less., UEC 50.171 (UEC), GU818641; AF457434; S. cadiscus B.Nord. &
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Pelser, Goldblatt & Manning 10690 (MO), GU818498-GU818505; Rourke 1118 (S),
GU818506; S. chilensis Less., EF538313; S. ctenophyllus Phil., EF538322; S.
culcitioides Wedd., EF538312; S. deltoideus Less., EF538326; AY953927; S. elegans
L., Cron & Goodman 687 (J), GU818642; S. fistulosus Poepp. ex Less., EF538335; S.
flaccidus Less., AF161640; AF161690; S. flaccidus Less. var. flaccidus, EF538336; S.
flavus (Decne.) Sch.Bip., Merxmüller & Giess 28206 (M), GU818643-GU818648;
AF457414-AF457416; S. flavus (Decne.) Sch.Bip. ssp. flavus, AJ400782; S. gayanus
(Colla) Cabr., Rosas 2157 (INIA), GU818649; S. gramineus Harv., Hoener 2104
(WAG), GU818650; S. gregorii F.Muell., Albrecht 7091 (NT), GU818651; S.
hemmendorffii Malme, EF538346; S. hieracium Remy, Baeza & Finot 3695 (CONC),
GU818652; S. hispidissimus I.Thomps., Thompson 927 (MEL), GU818653-GU818660;
S. hollandii Compton, Germishuizen 6586 (WAG), GU818661; S. hypsobates Wedd.,
EF538348; S. ilicifolius Thunb., Cron & Goodman 686 (J), GU818662; S. integerrimus
Nutt. var. exaltatus (Nutt.) Cronquist, EF538349; S. involucratus (Kunth) DC.,
EF538150; S. jarae Phil., EF538350; S. lastarrianus Remy, Ricardi 3230 (B),
GU818663; S. latifolius DC., EF538354; S. lineatus DC., AF459939; S. mairetianus
DC., EF538359; S. medley-woodii Hutch. (Caputia medley-woodii ined.), DQ915861;
S. meuselii Rauh, DQ915899; S. nemorensis L., AF459937; S. nevadensis Boiss. &
Reut., Vogt 4172 (B), GU818664; S. nevadensis Boiss. & Reut. ssp. malacitanus
(Huter) Greuter, AJ400813; S. niveo-aureus Cuatrec., EF538178; S. oerstedianus
Benth. ex Oerst., EF538362; S. oreophyton Remy, EF538393; S. otites Kunze ex DC.,
EF538363; Plowman 2627 (S), GU818665-GU818670; S. patens (Kunth) DC.,
EF538151; S. pflanzii (Perkins) Cuatrec., EF538179; S. pinifolius (L.) Lam.,
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EF538366; S. pinnatifolius A.Rich. var. lanceolatus (Benth.) I.Thomps., Thompson
904 (MEL), GU818671-GU818680; S. pinnatifolius A.Rich. var. maritimus (Ali)
I.Thomps.,, U93203; S. polygaloides Phil., EF538367; S. prenanthoides A.Rich.,
Thompson 915 (MEL), GU818681; S. psilocarpus R.O. Belcher & D.E. Albrecht,
Thompson 937 (MEL), GU818682-GU818692; S. retrorsus DC., EF538372; S. roseus
Sch.Bip., EF538373; S. saxatilis Wall. ex DC., EF538376; S. scandens Buch.-Ham. ex
D.Don, AF459932; S. scaposus DC. (Caputia scaposa ined.), AF459931; Pelser cult.
332 (L), GU818693-GU818697; S. squarrosus A.Rich., Thompson 907 (MEL),
GU818698-GU818704; S. stigophlebius Baker, EF538384; S. suaveolens (L.) Ell.,
EF538222; EF538223; S. superandinus Cuatrec., EF538248; S. szyszylowiczii Hiern.,
Sagastegui A. et al. 15797 (S), GU818705; S. tauricola V.A.Matthews, Budak 1735,
Aksoy & Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu), GU817570; S. thapsoides
DC., EF538388; Akeroyd 570 (RNG), GU818706; S. triodon Phil. var. triodon, Luebert
& Teillier 2266 (CONC), GU818707; S. triqueter Less., EF538392; S. vestitus
P.J.Bergius, Greuter 21766 (B), GU818708; S. viscosus L., AF459925; AF097539;
AJ400808; Shafera platyphylla Greenm., Nordenstam s.n. (S), GU818709; Sinacalia
tangutica (Maxim.) B.Nord., Lian et al 93-54 (MO), GU817573; AY176157;
Sinosenecio euosmus (Hand.-Mazz.) B.Nord., Bouford, Donoghue & Ree 27625 (MO),
GU818710; Solanecio biafrae (Oliv. & Hiern) C.Jeffrey, Brunel 6220 (B), GU818711;
Steirodiscus capillaceus (Thunb.) Less., Cron & Goodman 653 (J), GU818712GU818720; Stilpnogyne bellidioides DC., Goldblatt & Porter 11729 (MO), GU818721;
Symphyotrichum novae-angliae (L.) G.L.Nesom, Tepe 1675 (MU), GU818492;
Syneilesis palmata (Thunb.) Maxim., EF538401; Synotis nagensium (C.B.Clarke)
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C.Jeffrey & Y.L.Chen, AF459922; Tagetes sp., Vincent 13224 (MU), GU818722;
Talamancalia boquetensis (Standl.) H.Rob. & Cuatrec., EF538403; Telanthophora
grandifolia (Less.) H.Rob. & Brettell, EF538405; EF538406; Tephroseris integrifolia
(L.) Holub ssp. aucheri (DC.) B.Nord., Budak 1688 & Hamazaoǧlu (Yozgat Türkiye
Florası Herbaryumu), GU818724; Tetradymia filifolia Greene, EF538411; Traversia
baccharoides Hook.f., AF422139; Tussilago farfara L., AY176167; Urostemon kirkii
(Hook.f. ex Kirk) B.Nord., Cooper & Nickerson s.n. (US), GU818725; AY554101;
Vernonia altissima Nutt., AY142949; Villasenoria orcuttii (Greenm.) B.L.Clark,
Robles 389 (XAL), GU818726; Werneria caespitosa Wedd., EF538412; Xenophyllum
poposum (Phil.) V.A.Funk, EF538415; Yermo xanthocephalus Dorn, Anderson 13691
(MO), GU818727; Zemisia discolor (Sw.) B.Nord., EF538416.
ETS. Abrotanella emarginata (Gaudich.) Cass., Goodall & Wood 3352 (MU),
GU818108; Acrisione cymosa (J.Rémy) B.Nord., Lammers, Rodriques & Baeza 6447
(MU), GU818109; Adenostyles alpina (L.) Bluff & Fingerh., Schwerdtfeger 12251 (B),
GU818110; Aequatorium asterotrichum B.Nord., Asplund 18263 (S), GU818111;
Anacyclus valentinus L., Tepe 1186 & Moreno-Paez (MU), GU818112; Antillanthus
almironcillo (M.Gómez) B.Nord., El-Ghazaby et al. 23.608 (S), GU818113;
Arbelaezaster ellsworthii (Cuatrec.) Cuatrec., Kilip & Smith 19413 (US), GU818114;
Arnoglossum atriplicifolium (L.) H.Rob., Vincent 3925 (MU), GU818115;
Arrhenechthites mixta (A.Rich.) Belcher, Lawrence 1308 (S), GU818116;
Austrosynotis rectirama (Baker) C.Jeffrey, LaCroix 4001 (WAG), GU818118; Bahia
dissecta (Gray) Britton, Tuhy 3868 (MU), GU818119; Barkleyanthus salicifolius
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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(Kunth) H.Rob. & Brettell, Genelle & Fleming 861 (B), GU818120; Bedfordia
arborescens Hochr., Greuter 21319 (B), GU818121; Bethencourtia palmensis (Nees)
Choisy, Nordenstam 9326 (S), GU818122; Blepharispermum zanguebaricum Oliv. &
Hiern., Medley 570 (MU), GU818123; Bolandia pedunculosa (DC.) Cron, Cron &
Goodman 681 (J), GU818124; Brachyglottis repanda J.R.Forst. & G.Forst.,
Schwerdtfeger 17407 (B), GU818125; Cabreriella oppositicordia (Cuatrec.) Cuatrec.,
Romero-Castañeda 7428 (US), GU818126; Cacaliopsis nardosmia (A.Gray) A.Gray,
Ertter 6502 (UC), GU818127; Calendula arvensis L., Watson 95-13C (MU),
GU818129; Capelio caledonica B.Nord., Nordenstam 9644 (S), GU818130; Caucasalia
parviflora (M.Bieb.) B.Nord., Schneeweiss, Tribsch, Staudinger & Schönswetter 8643
(WU), GU818131; Caxamarca sanchezii M.O.Dillon & Sagást., Sagástegui et al. 15548
(F), GU818132; Centropappus brunonis Hook.f., Wapstra MW2 (MEL), GU818133;
Chaenactis douglasii (Hook.) Hook. & Arn., Vincent 8582 (MU), GU818134;
Charadranaetes durandii (Klatt) Janovec & H.Rob., Gomez P. 2242 (S), GU818135;
Chersodoma jodopappa (Sch.Bip.) Cabr., Hensen 2617 (S), GU818136; Cichorium
intybus L., Tepe 1673 (MU), GU818137; Cineraria abyssinica Sch.Bip. ex A.Rich.,
Pelser cult. 208 (L), GU818138; Cirsium discolor (Muhl.) Spreng., Tepe 1670 (MU),
GU818139; Cissampelopsis volubilis (Blume) Miq., Carvalho 3175 (MU), GU818140;
Coreopsis sp., Vincent 13226 (MU), GU818141; Corymbium enerve Markötter,
Trinder-Smith 124 (US), GU818142; Cotula coronopifolia L., Watson & Panero 94-26
(MU), GU818143; Crassocephalum crepidioides (Benth.) S.Moore, Pelser cult. 354
(L), GU818144; Cremanthodium humile Maxim., Kürschner & Sonnentag 01-436 (B),
GU818145; Crocidium multicaule Hook., Bartholomew 5749 (MO), GU818146; Curio
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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rowleyanus (H.Jacobsen) P.V.Heath, Pelser cult. 112 (L), GU818147; Dauresia
alliariifolia (O.Hoffm.) B.Nord & Pelser, Müller & Tilson 907 (M), GU818148;
Delairea odorata Lem., Tepe 2180, Pelser & Marticorena (MU), GU818149;
Dendrophorbium bomanii (R.E.Fr.) C.Jeffrey, Dematteis & Seijo 722 (U), GU818150;
Dendrosenecio kilimanjari (Mildbr.) E.B.Knox ssp. cottonii (Hutch. & G.Taylor)
E.B.Knox, Knox 50 (MICH), GU818151; Digitacalia jatrophoides (Kunth) Pippen,
Panero 2330 & Salinas (TEX), GU818152; Dolichoglottis lyallii (Hook.f.) B.Nord.,
Strid 22172 (MO), GU818153; Dolichorrhiza caucasica (M.Bieb.) Galushko,
Schneeweiss 44 (WU), GU818154; Dorobaea pimpinellifolia (Kunth) B.Nord., Tepe
1467 (MU), GU818155; Doronicum pardalianches L., Cubr 40487 (B), GU818156;
Ekmaniopappus mikanioides (Urb. & Ekman) Borhidi, Nordenstam & Lundin 500 (S),
GU818157; Elekmania picardae (Krug & Urb.) B.Nord., Zanoni & García 46967 (S),
GU818158; Emilia coccinea (Sims) G.Don., Pelser cult. 126 (L), GU818159;
Endocellion sibiricum (J.F.Gmel.) J.Toman, Schönswetter & Tribsch T480 (WU),
GU818160; Erato polymnoides DC., Tepe 1456 (MU), GU818161; Erechtites
valerianifolius (Link ex Spreng.) Less. ex DC., De Lange 115 (CHR), GU818162;
Eriothrix lycopodioides DC., Nordenstam 9208 (S), GU818163; Eupatorium serotinum
Michx., Tepe 1667 (MU), GU818164; Euryops pectinatus (L.) Cass., Pelser cult. 222
(L), GU818165; Farfugium japonicum (L.) Kitam., Wu 1439 (MO), GU818166;
Faujasia squamosa (Bory) C.Jeffrey, Nordenstam 9210 (S), GU818167; Faujasiopsis
flexuosa (Lam.) C.Jeffrey ssp. bourbonensis C.Jeffrey, Nordenstam 9202 (S),
GU818168; Galinsoga quadriradiata Cav., Vincent 13227 (MU), GU818169;
Garcibarrigoa telembina (Cuatrec.) Cuatrec., Holm-Nielsen et al. 6211 (S), GU818170;
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Gerbera sp., Vincent 13223 (MU), GU818171; Graphistylis dichroa (Bong.)
D.J.N.Hind, Silva 140 & Zelma (S), GU818172; Gymnodiscus capillaris (L.f.) Less.,
Coppejans EC1116 (U), GU818173; Gynoxys soukupii Cuatrec., Hutchison & Wright
5352 (L), GU818174; Gynura divaricata (L.) DC. ssp. formosana (Kitam.) F.G.Davies,
Panero & Hsiao 6457 (TEX), GU818175; Helenium autumnale L., Tepe 1664 (MU),
GU818176; Helianthus tuberosus L., Tepe 1672 (MU), GU818177; Helichrysum
stoechas DC., Tepe 1092 (MU), GU818178; Herodotia haitiensis Urb. & Ekman,
Ekman H1651 (S), GU818179; Herreranthus rivalis (Greenm.) B.Nord., Nordenstam
s.n. (2/12/2007) (S), GU818180; Hertia pallens (DC.) Kuntze, Koekemoer & Funk 1963
(US), GU818181; Hoehnephytum trixoides (Gard.) Cabr., Carmo 131 (BHCB),
GU818182; Homogyne alpina (L.) Cass., Uhink 98-56 (MJG), GU818183; Hubertia
riparia (DC.) C.Jeffrey, Labat, Phillipson & Lowry II 2022 (WAG), GU818184;
Humbertacalia sp., Phillipson et al. 5641 (P), GU818185; Ignurbia constanzae (Urb.)
B.Nord., Nordenstam & Lundin 560 (S), GU818186; Inula viscosa (L.) Aiton, Tepe
1142 (MU), GU818187; Io ambondrombeensis (Humbert) B.Nord., Malcomber et al.
1380 (MO), GU818188; Iranecio cariensis (Boiss.) C.Jeffrey, Budak 1724, Aksoy &
Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu), GU818189; Jacobaea vulgaris
Gaertn., Pelser cult. 6 (L), GU818190; Jessea multivenia (Benth.) H.Rob. & Cuatrec.,
Liesner & Judziewicz 14750 (MO), GU818191; Kleinia neriifolia Haw., Pelser cult. 216
(L), GU818192; Lachanodes arborea (Roxb.) B.Nord., Cairns-Wicks s.n., GU818193;
Lactuca canadensis L., Vincent 13128 (MU), GU818194; Lamprocephalus montanus
B.Nord., Nordenstam 9542 (S), GU818195; Leonis trineurus (Griseb.) B.Nord., Smith
& al. 3238 (S), GU818196; Lepidospartum burgessii B.L.Turner, Worthington 12382
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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(L), GU818197; Ligularia stenocephala (Maxim.) Matsum. & Koidz., Kowal 3092
(WIS), GU818198; Lomanthus fosbergii (Cuatrec.) B. Nord. & Pelser, Øllgaard et al.
90697 (AAU), GU818317; Lopholaena coriifolia (Sond.) Phillips & C.A.Sm., Van der
Westhuizen & Liamé; s.n. (PREM), GU818199; Lordhowea insularis (Benth.) B.Nord.,
Trodd & Thorne 3703 (L), GU818200; Luina hypoleuca Benth., Greuter 17706 (B),
GU818201; Lundinia plumbea (Griseb.) B.Nord., Zanoni 45816 (JBSD), GU818202;
Madia sp., Vincent 12912 (MU), GU818203; Mesogramma apiifolium DC., Giess
16074 (M), GU818204; Mikaniopsis clematoides (Sch.Bip. ex A.Rich.) Milne-Redh.,
De Wilde & De Wilde-Duyfjes 9006 (B), GU818205; Miricacalia makineana (Yatabe)
Kitam., Kobayashi 33484 (HYO), GU818206; Misbrookea strigosissima (A.Gray)
V.A.Funk, Funk 11388 (US), GU818207; Monticalia abietina (Willd. ex Wedd.)
C.Jeffrey, Cleef 8411 (U), GU818208; Monticalia apiculata (Sch.Bip. ex Wedd.)
C.Jeffrey, Alston 6935 (S), GU818209; Monticalia arbutifolia (Kunth.) C.Jeffrey,
Nordenstam 9436 (S), GU818210; Nelsonianthus tapianus (B.L.Turner) C.Jeffrey,
Pérez & Kendizabal 457 (XAL), GU818211; Nemosenecio nikoensis (Miq.) B.Nord.,
Koyama 4079 (L), GU818212; Nesampelos lucens (Poir. in Lam.) B.Nord., Zanoni
45570 (JBSD), GU818213; Neurolaena lobata (L.) R.Br., Acevedo 9316 (US),
GU818214; Nordenstamia kingii (H.Rob. & Cuatrec.) B.Nord., Ståhl 5572A (S),
GU818215; Oldfeltia polyphlebia (Griseb.) B.Nord. & Lundin, Nordenstam & Lundin
340 (S), GU818216; Oresbia heterocarpa Cron & B.Nord., Nordenstam 9628 (S),
GU818217; Othonna capensis Bailey, Pelser cult. 106 (L), GU818218; Packera
eurycephala (Torr. & A.Gray) W.A.Weber & Á.Löve, Vincent 8581 (MU),
GU818219; Papuacalia dindondl (P.Royen) Veldkamp, Johns 9249 (K), GU818220;
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Paracalia jungioides (Hook. & Arn.) Cuatrec., King & Collins 9017 (UC), GU818221;
Parafaujasia fontinalis (Cordem.) C.Jeffrey, Lorence 2602 (MO), GU818222;
Paragynoxys neodendroides (Cuatrec.) Cuatrec., Cleef 9852a (U), GU818223;
Parasenecio adenostyloides (Franch. et Sav. ex Maxim.) H.Koyama, Koyama 408 (L),
GU818224; Pentacalia arborea (Kunth) H.Rob. & Cuatrec., Øllgaard & Balslev 8298
(U), GU818225; Pericallis murrayi (Bornm.) B.Nord., Royl 446 (B), GU818226;
Perityle emoryi Torr., Atwood 26483 (MU), GU818227; Petasites albus (L.) Gaertn.,
Cubr 40398 (B), GU818228; Phaneroglossa bolusii (Oliv.) B.Nord., Watson & Panero
94-62 (TEX), GU818229; Pippenalia delphinifolia (Rydb.) MacVaugh, Breedlove
59032 & Almeda (MO), GU818230; Pittocaulon praecox (Cav.) H.Rob. & Brettell,
GU818231; Pladaroxylon leucadendron (G.Forst.) Hook.f., Cairns-Wicks s.n.,
GU818232; Pojarkovia pojarkovae (Schischk.) Greuter, Pelser cult. 191 (L),
GU818233; Polymnia canadensis L., Tepe 1665 (MU), GU818234; Psacaliopsis
purpusii (Greenm. ex Brandegee) H.Rob. & Brettell, Panero 2607, Davila & Tenorio
(TEX), GU818235; Psacalium cirsiifolium (Zucc.) H.Rob. & Brettell, Kowal 3053
(WIS), GU818236; Pseudogynoxys haenkei (DC.) Cabr., Molina R. 18431 (U),
GU818237; Rainiera stricta (Greene) Greene, Dennis 2317 (U), GU818238;
Robinsonecio gerberifolius (Sch.Bip. ex Hemsl.) T.M.Barkley & Janovec, Garcia P.
171 (MO), GU818239; Robinsonia berteroi (DC.) R.W.Sanders, Stuessy & Martic.,
Stuessy et al. 11238 (CONC), GU818240; R. evenia Phil., Stuessy et al. 11308 (CONC),
GU818241; R. gracilis Decne., Stuessy, Crawford, Valdebenito & Landeros 6560 (B),
GU818242; Stuessy et al. 11282 (CONC), GU818243; Stuessy et al. 11312 (CONC),
GU818244; R. masafuerae Skottsb., Stuessy et al. s.n. (CONC), GU818245; Roldana
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
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suffulta (Greenm.) H.Rob. & Brettell, Rzedowski 36569 (UC), GU818246; Rugelia
nudicaulis Shuttlew. ex Chapm., Feist, Phillippe, Molano-Flores, Busemeyer & Carroll
714 (TENN), GU818247; Scrobicaria ilicifolia (L.f.) B.Nord., Cleef, Garcia-B. &
Jaramillo-M. 3528 (U), GU818307; Senecio acanthifolius Hombr. & Jacq., ex Decne.,
Björnsäter s.n. (S), GU818248; S. algens Wedd., Beck 2879 (S), GU818249; S.
arnicoides Hook. & Arn., Zöllner 3474 (L), GU818250; S. boyacensis (Cuatrec.)
Cuatrec., Gonzalez 180 (S), GU818251; S. brasiliensis (Spreng.) Less., UEC 50.171
(UEC), GU818252; S. cadiscus B.Nord. & Pelser, Goldblatt & Manning 10690 (MO),
GU818128; S. chilensis Less., Zöllner 2958 (L), GU818254; S. ctenophyllus Phil.,
Zöllner 3959 (L), GU818255; S. culcitioides Wedd., Øllgaard & Balslev 8822 (U),
GU818253; S. deltoideus Less., Sloet S.A.105 (U), GU818256; S. elegans L., Cron &
Goodman 687 (J), GU818257; S. fistulosus Poepp. ex Less., Beck & Liberman 9672 (S),
GU818258; S. flaccidus Less. var. flaccidus, Jardin Thuret cult. s.n. (MJG), GU818259;
S. flavus (Decne.) Sch.Bip., Merxmüller & Giess 28206 (M), GU818260; S. gayanus
(Colla) Cabr., Rosas 2157 (INIA), GU818261; S. gramineus Harv., Hoener 2104
(WAG), GU818262; S. gregorii F.Muell., Albrecht 7091 (NT), GU818263; S.
hemmendorffii Malme, Kummrow 3052 et al. (S), GU818264; S. hieracium Remy,
Baeza & Finot 3695 (CONC), GU818265; S. hispidissimus I.Thomps., Thompson 927
(MEL), GU818266; S. hollandii Compton, Germishuizen 6586 (WAG), GU818267; S.
hypsobates Wedd., Øllgaard & Balslev 9863 (U), GU818268; S. ilicifolius Thunb.,
Cron & Goodman 686 (J), GU818269; S. integerrimus Nutt. var. exaltatus (Nutt.)
Cronquist, Crockett 437 (MU), GU818270; S. jarae Phil., Liberman L54 (S),
GU818271; S. lastarrianus Remy, Ricardi 3230 (B), GU818272; S. latifolius DC.,
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Bayer SAF-01074, Chandler & Koekemoer (PRE), GU818273; S. lineatus DC., Bayer &
Puttock SAF-96246 (CANB), GU818274; S. mairetianus DC., Garcia P. 151 (L),
GU818275; S. medley-woodii Hutch. (Caputia medley-woodii ined.), Jeffrey Kew cult.
14 (K), GU818276; S. meuselii Rauh, Greissl s.n. (18-FEB-1988) (MJG), GU818277; S.
nemorensis L., Pelser cult. 102 (L), GU818278; S. nevadensis Boiss. & Reut., Vogt
4172 (B), GU818279; S. niveo-aureus Cuatrec., Cleef 6665 (S), GU818280; S.
oerstedianus Benth. ex Oerst., Nordenstam 9160 (S), GU818281; S. oreophyton Remy,
Beck 21589 (S), GU818282; S. otites Kunze ex DC., Plowman 2627 (S), GU818283; S.
patens (Kunth) DC., Zak & Jaramillo 3427 (L), GU818284; S. pflanzii (Perkins)
Cuatrec., Beck 9094 (S), GU818285; S. pinifolius (L.) Lam., Coppejans EC810 (U),
GU818286; S. pinnatifolius A.Rich. var. lanceolatus (Benth.) I.Thomps., Thompson
904 (MEL), GU818287; S. polygaloides Phil., Zöllner 5442 (L), GU818288; S.
prenanthoides A.Rich., Thompson 915 (MEL), GU818289; S. psilocarpus R.O. Belcher
& D.E. Albrecht, Thompson 937 (MEL), GU818290; S. retrorsus DC., Van Steenis
23961 (L), GU818291; S. roseus Sch.Bip., Garcia P. 250 (L), GU818292; S. saxatilis
Wall. ex DC., Van Beusekom et al. 4253 (L), GU818293; S. scandens Buch.-Ham. ex
D.Don, Pelser cult. 211 (L), GU818294; S. scaposus DC. (Caputia scaposa ined.),
Pelser cult. 332 (L), GU818295; S. squarrosus A.Rich., Thompson 907 (MEL),
GU818296; S. stigophlebius Baker, UEC 2.974 (UEC), GU818297; S. suaveolens (L.)
Ell., Dister s.n. (27 Jul 2002) (MU), GU818298; S. superandinus Cuatrec., Hekker &
Hekking 10.159a (U), GU818299; S. szyszylowiczii Hiern., Sagastegui A. et al. 15797
(S), GU818300; S. tauricola V.A.Matthews, Budak 1735, Aksoy & Hamazaoǧlu (Yozgat
Türkiye Florası Herbaryumu), GU818301; S. thapsoides DC., Touw 21868 (L),
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GU818302; S. triodon Phil. var. triodon, Luebert & Teillier 2266 (CONC), GU818303;
S. triqueter Less., Robertson 293 & Burman (WAG), GU818304; S. vestitus
P.J.Bergius, Greuter 21766 (B), GU818305; S. viscosus L., Pelser 300 (L), GU818306;
Shafera platyphylla Greenm., Nordenstam s.n. (S), GU818308; Sinacalia tangutica
(Maxim.) B.Nord., Lian et al 93-54 (MO), GU818309; Sinosenecio euosmus (Hand.Mazz.) B.Nord., Bouford, Donoghue & Ree 27625 (MO), GU818310; Solanecio biafrae
(Oliv. & Hiern) C.Jeffrey, Brunel 6220 (B), GU818311; Steirodiscus capillaceus
(Thunb.) Less., Cron & Goodman 653 (J), GU818312; Stilpnogyne bellidioides DC.,
Goldblatt & Porter 11729 (MO), GU818313; Symphyotrichum novae-angliae (L.)
G.L.Nesom, Tepe 1675 (MU), GU818117; Syneilesis palmata (Thunb.) Maxim., Hata
s.n. (4-Aug-1979) (CHR), GU818314; Synotis nagensium (C.B.Clarke) C.Jeffrey &
Y.L.Chen, Sino-American Guizhou Botanical Expedition no. 1991 (L), GU818315;
Talamancalia boquetensis (Standl.) H.Rob. & Cuatrec., Wilbur 15397 (NY),
GU818316; Telanthophora grandifolia (Less.) H.Rob. & Brettell, Greissl cult. s.n. (25Jun-1988) (MJG), GU818318; Tephroseris integrifolia (L.) Holub ssp. aucheri (DC.)
B.Nord., Budak 1688 & Hamazaoǧlu (Yozgat Türkiye Florası Herbaryumu), GU818319;
Tetradymia filifolia Greene, Worthington 12360 (L), GU818320; Tussilago farfara L.,
Kennedy 225 (MU), GU818321; Urostemon kirkii (Hook.f. ex Kirk) B.Nord., Cooper
& Nickerson s.n. (US), GU818322; Vernonia altissima Nutt., Tepe 1668 (MU),
GU818323; Villasenoria orcuttii (Greenm.) B.L.Clark, Robles 389 (XAL), GU818324;
Werneria caespitosa Wedd., Funk 11324 (US), GU818325; Xenophyllum poposum
(Phil.) V.A.Funk, Funk 11351 (US), GU818326; Yermo xanthocephalus Dorn,
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Anderson 13691 (MO), GU818327; Zemisia discolor (Sw.) B.Nord., Webster et al. 8420
(S), GU818328.
PELSER ET AL. – American Journal of Botany 97(5): 856-873. 2010. – Data Supplement S2 – Page 1
PELSER, PIETER B., AARON H. KENNEDY, ERIC J. TEPE, JACOB B. SHIDLER, BERTIL NORDENSTAM, JOACHIM W. KADEREIT, AND LINDA E. WATSON. 2010.
Patterns and causes of incongruence between plastid and nuclear Senecioneae (Asteraceae) phylogenies. American Journal of Botany 97(5): 856-873.
Appendix S2. Bayesian inference phylograms. A, plastid data; B, ITS/ETS data.
A
Crocidium
Blennosperma
Ischnea
Robinsonecio
Barkleyanthus
Arnoglossum
Yermo
Emilia
Jacobaea
Packera
B
Crocidium
Blennosperma
Ischnea
Robinsonecio
Barkleyanthus
Arnoglossum
Yermo
Jacobaea
Emilia
Packera
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
S3 – Page 1
PELSER, PIETER B., AARON H. KENNEDY, ERIC J. TEPE, JACOB B. SHIDLER, BERTIL
NORDENSTAM, JOACHIM W. KADEREIT, AND LINDA E. WATSON. 2010. Patterns and causes
of incongruence between plastid and nuclear Senecioneae (Asteraceae) phylogenies.
American Journal of Botany 97(5): 856-873.
Appendix S3. Age estimates of clades present in plastid and ITS/ETS trees.
R8s ITS/ETS
BEAST ITS/ETS age
R8s plastid
BEAST plastid age (my;
age (my)
(my; 95% HPD interval)
age (my)
95% HPD interval)
mrca Abrotanella-Senecio
24.14
(19.15-)21.44(-23.66)
28.94
(23.77-)26.37(-29.25)
mrca Adenostyles-Caucasalia
5.62
(2.32-)3.97(-5.75)
7.32
(1.63-)4.07(-6.87)
mrca Anacyclus-Cotula
20.48
(6.85-)15.65(-23.4)
20.26
(8.22-)15.69(-21.26)
mrca Antillanthus-Shafera
6.88
(1.5-)3.01(-4.68)
7.39
(0.25-)1.46(-3.03)
mrca Arnoglossum-Yermo
1.17
(0.52-)1.43(-2.4)
1.21
(0.33-)1.12(-2.17)
mrca Asteroideae
26.94
(23.71-)26.58(-29.33)
29.56
(26.21-)28.51(-30.75)
mrca Barkleyanthus-Arnoglossum
4.76
(2.4-)3.77(-5.11)
5.74
(1.7-)3.21(-4.9)
mrca Bedfordia-Centropappus
5.08
(0.75-)1.86(-3.16)
4.29
(0.19-)0.92(-1.86)
mrca Blennosperma-Ischnea
6.07
(2.52-)4.76(-7.1)
9.58
(3.2-)5.97(-8.99)
mrca Blepharispermum-Madia
24.43
(16.59-)19.74(-22.74)
24.52
(17.3-)21.04(-24.98)
mrca Capelio-Senecio
21.71
(17.77-)19.51(-21.55)
25.42
(20.06-)22.91(-25.64)
mrca Caputia
2.28
(0.4-)1.7(-3.42)
2.3
(0.23-)1.86(-4.23)
mrca Caucasalia-Pojarkovia
4.94
(1.73-)3.17(-4.84)
5.97
(0.9-)2.79(-5.16)
mrca Caxamarca-Pseudogynoxys
3.38
(0.91-)2.22(-3.45)
2.07
(0.24-)0.99(-1.86)
mrca Chersodoma-Dolichoglottis
19.41
(16.36-)17.75(-19.22)
23.14
(17.91-)20.32(-22.83)
mrca Cichorium-Vernonia
27.96
(18.83-)25.21(-30.35)
27.44
(15.06-)22.54(-30.2)
7.8
(1.26-)4.12(-7.69)
3.61
(0.54-)2(-3.81)
mrca Corybium-Cichorium
30.41
(28.4-)31.38(-34.08)
32.7
(30.29-)32.47(-34.29)
mrca Crocidium-Blennosperma
12.49
(6.48-)9.25(-11.85)
14.52
(6.93-)9.83(-12.82)
mrca Dolichoglottis-Haastia
9.57
(3.14-)5.32(-7.93)
14.78
(1.78-)4.95(-9.76)
mrca Erato-Vernonia
21.18
(9.15-)16.01(-22.53)
24.46
(8.63-)16.85(-24.78)
mrca Eriothrix-Parafaujasia
8.87
(0.66-)2.36(-4.39)
6.54
(0.47-)1.79(-3.38)
mrca Eupatorium-Perityle
13.85
(5.79-)9.58(-13.29)
14.47
(4.6-)8.92(-12.92)
mrca Graphistylis-Dendrophorbium
11.26
(3.44-)5.73(-7.59)
4.91
(1.31-)2.57(-3.95)
mrca Gymnodiscus-Othonna
10.45
(2.97-)6.19(-9.54)
14.53
(6.02-)9.62(-13.04)
mrca Gynura-Kleinia
9.13
(2.52-)4.61(-6.7)
4.23
(1.48-)3.22(-5)
mrca Herreranthus-Lundinia
8.29
(2.84-)4.35(-5.81)
7.39
(0.41-)1.62(-3.13)
Node:
mrca Cissampelopsis-Mikaniopsis
PELSER ET AL. –American Journal of Botany 97(5): 856-873. 2010. – Data Supplement
S3 – Page 2
mrca Hertia-Lopholaena
5.81
(1.09-)4.09(-7.6)
8.02
(2.2-)5.49(-8.63)
mrca Hertia-Othonna
15.16
(7.84-)11.39(-14.43)
17.26
(11.2-)13.51(-15.79)
mrca Kleinia-Delairea
10.77
(4.1-)6.45(-8.79)
9.36
(3.79-)5.81(-7.76)
mrca Lactuca-Chichorium
19.21
(6.28-)14.46(-22.58)
16.66
(5.22-)11.97(-19.59)
mrca Lamprocephalus-Oresbia
8.85
(0.78-)2.65(-5.05)
11.73
(2.18-)6.68(-10.85)
mrca Lepidospartum-Rainiera
6.33
(3.47-)5.83(-8.82)
14.29
(3.41-)7.51(-11.93)
mrca Lepidospartum-Tetradymia
2.06
(0.48-)1.94(-3.87)
4.05
(0.3-)2.15(-4.61)
mrca Luina-Rugelia
5.57
(2.47-)4.7(-7.04)
11.01
(1.75-)4.36(-7.47)
mrca Lundinia-Oldfeltia
3.93
(0.77-)1.99(-3.33)
4.83
(0.09-)0.82(-1.83)
mrca Madia-Tagetes
17.35
(13.49-)15.99(-18.67)
18.16
(12.99-)15.21(-17.98)
mrca Mesogramma-Stilpnogyne
7.73
(1.65-)3.79(-6.19)
7.86
(2.07-)5.38(-9.05)
mrca Nemosenecio-Sinosenecio
4.12
(0.76-)2.16(-3.71)
1.88
(0.3-)1.45(-2.83)
mrca Papuacalia-Acrisione
7.81
(2.05-)3.49(-5.14)
11.12
(1.41-)3.05(-5.05)
mrca Paragynoxys-Nordenstamia
9.98
(3.56-)6.29(-9.55)
10.12
(1.84-)4.17(-6.62)
mrca Paragynoxys-Psacaliopsis
14.94
(9.2-)11.48(-13.58)
14.71
(6.47-)9.15(-11.9)
mrca Pentacalia arborea-Monticalia abietina
10.51
(2.89-)5.38(-7.77)
7.35
(0.26-)1.76(-3.69)
mrca Pippenalia-Psacalium
3.49
(0.89-)2.84(-5.07)
5.61
(0.49-)2.73(-5.42)
mrca Pittocaulon-Villasenoria
6.72
(1.91-)4.08(-6.41)
8.02
(0.86-)2.87(-4.84)
mrca Rainiera-Rugelia
4.24
(1.09-)2.88(-4.78)
6.24
(0.4-)2.06(-4.06)
mrca Robinsonecio-Pippenalia
11.79
(7.59-)9.5(-11.56)
12.26
(4.98-)7.2(-9.65)
mrca Senecio acanthifolius-S. hieracium
3.61
(1.51-)2.52(-3.56)
2.77
(0.32-)1.57(-2.83)
mrca Senecio brasiliensis-S. niveo-aureus
2.1
(1.03-)1.75(-2.54)
2.21
(1.12-)2.07(-3.1)
mrca Senecio ctenophyllus-S. boyacensis
4.03
(2.77-)3.71(-4.55)
3.08
(0.62-)2.02(-3.54)
mrca Senecio deltoideus-S. scandens
2.36
(0.25-)1.48(-3.07)
8.39
(0.68-)2.39(-4.37)
mrca Senecio flaccidus-S. mairetianus
1.88
(0.66-)1.33(-2.09)
0.45
(0.04-)0.56(-1.25)
mrca Senecio stigophlebius-S. adamantinus
10.02
(3.41-)5.32(-7.11)
4.03
(0.43-)1.56(-2.83)
mrca Talamancalia-Jessea
3.71
(1.08-)2.47(-3.8)
2.18
(0.29-)1.17(-2.12)
mrca Tephroseris-Nemosenecio
6.28
(1.7-)3.61(-5.63)
4.66
(1.4-)3.26(-5.49)
mrca Tussilagininae s. str.
17.97
(12.97-)15(-17)
18.18
(11.85-)14.42(-17.06)
mrca Tussilago-Homogyne
12.26
(3.27-)7.16(-11.22)
11.09
(2.02-)5.94(-10.34)
mrca Xenophyllum-Jessea
10.75
(4.97-)6.26(-7.68)
9.17
(2.81-)4.2(-5.65)
mrca Xenophyllum-Werneria
5.27
(1.96-)3.36(-4.84)
1.57
(0.52-)1.62(-2.8)