Callianthe (Malvaceae): A New Genus of Neotropical Malveae

Author(s): Aliya A. Donnell, Harvey E. Ballard Jr. and Philip D. Cantino

Source: Systematic Botany , July-September 2012, Vol. 37, No. 3 (July-September 2012),
pp. 712-722
Published by: American Society of Plant Taxonomists

Stable URL: https://www.jstor.org/stable/41515161

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Systematic Botany (2012), 37(3): pp. 712-722
© Copyright 2012 by the American Society of Plant Taxonomists
DOI 10.1600/036364412X648689

Callianthe (Malvaceae): A New Genus of Neotropical Malveae

Aliya A. Donnell,1'2 Harvey E. Ballard Jr.,1 and Philip D. Cantino1

department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701, U. S. A.
2Author for correspondence (AliyaA.Donnell.l@ohio.edu)

Communicating Editor: Mark P. Simmons

Abstract - The boundaries between the genera Bakeridesia and Abutilón have long been debated. Results from recent phylogenetic studies
using the rDNA ITS in tribe Malveae strongly suggest that these two genera as currently delimited are polyphyletic. Some species previously
included in each genus form a well-supported clade that is phylogenetically removed from both Bakeridesia and Abutilón. The congruence of
morphological and karyological distinctions with this molecular evidence provides compelling support for recognition of the clade as a new
genus, here described as Callianthe. In the present study we transfer 40 species to Callianthe based on ITS data (including the synapomorphy of a
25-base pair deletion in ITS2) and /or morphological evidence, including the character states of four or more ovules per carpel, toothed and /or
lobed leaves, and petals with impressed veins (the latter is a synapomorphy). In addition, species of Callianthe share a base chromosome number
of X = 8 (versus 7 in Abutilón and 15 in Bakeridesia). Not only is Callianthe phylogenetically removed from both Bakeridesia and Abutilón, but it is
also morphologically distinct from its closest phylogenetic relative, Gaya, which has a base chromosome number of x = 6. Callianthe has a
Neotropical distribution with a center of diversity in eastern Brazil. It includes all species previously referred to Bakeridesia subg. Dipteron and
several large-flowered species previously referred to Abutilón. The following new combinations are made Callianthe antoena, C. andrade-limae,
C. bedfordiana, C. bezerrae, C. brenesii, C. cycloneruosa, C. darwinii, C. elegans, C. fluviatilis, C. geminiflora, C.glaziovii, C. inaequalis, C. jaliscana,
C. jujuiensis, C. lanata, С. latipetala, С. laxa, С. longifolia, С. macrantha, С. malmeana, С. mexiae, С. monteiroi, С. mouraei, С. muelleri-
friderici, С. pachecoana, С. petiolaris, С. pickellii, С. pieta, С. purpusii, С. regnellii, С. rufinerva, С. rufivela, С. scabrida, С. schenckii,
С. sellowiana, С. senilis, С. striata, С. torrendii, С. tridens, and С. vexillaris.

Keywords - Abutilón , Bakeridesia, Brazil, ITS.

Abutilón Mill. (Malvaceae) is a large genus of approximatelythe pluriovulate species may warrant elevation to generic
160 species. It is extremely variable in morphology and hasrank a (Fryxell 1997b, 2002). The pluriovulate species of
Abutilón share a base chromosome number of x = 8 (Bates
broad geographic distribution, with representatives on all con-
tinents except Antarctica. Abutilón is one of several poorly 1968; Bates and Blanchard 1970), in contrast to the remainder
delimited genera of Malvaceae that have historically beenof the genus, which has a base chromosome number of x-7
"dumping grounds" for difficult-to-place species. In the past,(Bates 1968; Bates and Blanchard 1970). Notable exceptions
there has been a tendency to assign to Abutilón any species are
of the weedy, annual species of Abutilón , such as A.
tribe Malveae that lacked an epicalyx and had multiple ovules theophrasti and A. indicum, which have been reported to have
up to six ovules per mericarp but have a base chromosome
per carpel if it did not easily fit into another, better circum-
scribed genus. Abutilón has been described as "the most diffi-number of x = 7; in all other morphological features they fit in
cult of the genera of Malvaceae" (Kearney 1958, p. 201) and with
is the pauciovulate species.
considered to be heterogeneous with "many taxonomie prob-Though Abutilón has never been treated as a whole, several
lems" (Fryxell 2002, p. 79). The genus as a whole has never genera have been segregated from it over the years, including
been monographed, but there have been several treatments Pseudabutilon
of R. E. Fr., Corynabutilon (K. Schum.) Kearney,
Tetrasida Ulbr., and Bastardia Kunth. Another such segregate
regional groups (St.-Hilaire 1825; St.-Hilaire and Naudin 1842;
Schumann 1891; Standley 1937; Standley and Steyermark 1949; is Bakeridesia Hochr., a genus of tall shrubs native to Latin
Kearney 1955, 1958; Robyns 1965; Krapovickas 1969; P. A. America. Bakeridesia is distinguished from other genera in
tribe Malveae by having a lacerate wing on the dorsal margin
Fryxell 1976, 1988, 1992; J. E. Fryxell 1983), and several genera
have been segregated from Abutilón over the years (Kearney of the mericarp. For this reason, Bakeridesia galeotta Hochr.
1949; Fryxell and Fuertes 1992; Fryxell 1997a). Currently,was removed from Abutilón (Hochreutiner 1913). Subse-
Abutilón is characterized within tribe Malveae by its lack of quently, 27 species were added to Bakeridesia (Hochreutiner
an epicalyx, mericarps lacking an internal constriction 1920; Monteiro Filho 1955, 1973; Bates 1973; Fryxell 2002;
(endoglossum), five or more carpels containing at least two Fryxell and Olivera 2001).
ovules each, capitate stigmas, mericarps lacking a dorsalCurrently, 28 species are assigned to Bakeridesia and two
"wing," and non-inflated fruits. subgenera are recognized: one that ranges from Mexico to
Traditionally, the number of ovules per carpel has been aEcuador (subgenus Bakeridesia) and another that is exclu-
diagnostic character for species groups within Abutilón. In sively Brazilian (subgenus Dipteron Hochr.). Though both
subgenera have winged mericarps, many differences exist
many of the older Neotropical keys, this character was the first
major dichotomy (St.-Hilaire 1825; St.-Hilaire and Naudin between them. Species included in subgenus Bakeridesia dif-
1842; Schumann 1891; Robyns 1965), and species were fer from Abutilón in several characters. In addition to the
grouped into one of two categories: three or fewer ovules per winged mericarps, plants in this group have entire leaves,
carpel ("pauciovulate") or four or more ovules per carpel yellow or orange flowers (petals various colors in Abutilón ),
("pluriovulate"). Though many taxonomists viewed the pollen with three apertures (Jiménez-Reyes 2003), compared
pluriovulate species as a natural group (St.-Hilaire 1825; with 2-6 and 8-15 apertures reported in Abutilón (Fryxell
St.-Hilaire and Naudin 1842; Schumann 1891; Robyns 1965;1997b), and a base chromosome number of x = 15 (Bates and
Bates and Blanchard 1970), it was not formally recognized untilBlanchard 1970; Bates 1973). In contrast, species included in
Fryxell (1988) gave the group sectional status in his treatmentsubgenus Dipteron are more similar to Abutilón in leaf margin
(toothed and /or lobed) and flower color, though the base
of Mexican Abutilón (sect. Pluriovulata). He later suggested that

712

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2012] DONNELL ET AL.: CALLIANTHE: A NEW GENUS OF MALVEAE 713

chromosome number is inferred to be x = 8amplification

(see discussion
primers was conducted on an Applied Biosystems Genetic
Analyzer 3130x at the Genomics Facility at Ohio University, Athens, Ohio.
section). Because of these differences, some authors have
The following primers were used for sequencing: ITS5 (White et al. 1990) for
questioned whether the Brazilian species actually belongand
the full sequence inITSl, and ITS3B for ITS2. Due to the high quality of
Bakeridesia (Bates 1973; Fryxell 1997b), but due to insufficient
sequences obtained from forward sequencing alone, either of the full ITS or
chromosome, pollen, and molecular data, it hasITSl been difficult
and ITS2 separately, reverse sequencing was not necessary.
Sequence Alignment
to test this. Other authors accept the genus Bakeridesia with-and Phylogenetic Analysis - Sequences were
edited using Sequence Scanner (v. 1.0, Applied Biosystems™) and
out including the species of subgenus Dipteron (Mabberley
aligned using the CLUSTAL W interface in BioEdit (Hall 1999), followed
1997; Kubitzki and Bayer 2003). by minimal manual corrections. The majority of the minor adjustments
The boundary between Bakeridesia subg. Dipteron
made attempted toand
minimize gaps, and where feasible to align gaps to be
Abutilón has been unclear. Several pluriovulate species
simultaneous of
across groups of taxa (nested gaps). The remaining correc-
Abutilón have a lacerate wing on the dorsal tions were toof
margin ensure
thethat the computer-generated alignment reflected
actual biological processes (i.e. transitions are more likely to occur than
mericarps but have not been transferred to Bakeridesia. On the
transversions). The highly conserved 5.8S region was not available for all
other hand, several species of Bakeridesia subg. Dipteron have
sequences in GenBank, and visual inspection of available 5.8S sequences
leaf morphology and flower characteristics that arelow
revealed similar
variationto
relative to the two spacers. Of the 104 sequences
the pluriovulate species of Abutilón. In addition, all 14
included, species of from GenBank lacked the 5.8S region. An addi-
downloaded
tional
subgenus Dipteron have more than four ovules per34 sequences
mericarp had only partial 5.8S regions due to lack of primer
region overlap
and are therefore pluriovulate. Here we present in taxa that were amplified in two parts (ITSl and ITS2).
evidence,
Because nearly half of the sequences lacked a complete 5.8S sequence,
based on the ITS region of the 18S-26S nuclear ribosomal
and the percentages of variable (10.2%) and parsimony-informative
repeat, morphology, and chromosome numbers,
(4.2%) that neither
characters were relatively low (compared to 58.3% of characters
Bakeridesia nor Abutilón is monophyletic as currently in the ITSl and ITS2), the region was excluded
parsimony-informative
delimited. We demonstrate that most of the from
pluriovulate spe-analysis. The percentage of characters in the data
the phylogenetic
matrix
cies of Abutilón (i.e. excluding the weedy (with species
annual the 5.8S removed) scored as missing was 8.7% including
gaps, which were treated as missing data. Parsimony analyses were
mentioned above) and the species of Bakeridesia subgenus
performed with NONA ver. 2.0 (Goloboff 1999) and implemented
Dipteron together form a previously unrecognized clade, to be
throughWinclada 1.00.08 (Nixon 2002) using the Ratchet (Island Hopper)
described here as a new genus, Callianthe Donnell.
(Nixon 1999) option with 200 iterations /rep, 5 trees held per iteration,
and 10% of the characters sampled. Bootstrap analysis (Felsenstein 1985)
was performed to assess the amount of support for monophyletic groups
Materials and Methods
(1,000 replicates each with 10 search reps and the "do not do max*"
Plant Material and Taxon Sampling - A total of option). Maximum
104 species andlikelihood (ML) analyses (Felsenstein 1973) were
performed
35 genera were included in this study, representing ten of using GARLI 0.96 (Zwickl 2006) under the TVM + 1 + G model
the 14 generic
alliances of tribe Malveae recognized by Kubitzki and of substitution,
Bayer (2003).which was selected as the best model to fit the data by
jModelTestanalyses
Outgroup selection was based on results from prior molecular (Guindon and Gascuel 2003; Posada 2008) based on Akaike's
of Malveae using the ITS region (Tate et al. 2005). To information
assess the criterion
mono- (Akaike 1974). The ML analyses were conducted
with thenumber
phyly of Bakeridesia and Abutilón, a proportionately greater followingof parameters: state frequencies (statefreq) were set to
equal,
species from the clades most closely related to Bakeridesia subg.the number of relative substitution rate parameters (ratematrix)
Bakeridesia
and to Abutilón were sampled. Sixteen species of Bakeridesia
was set to (0 subg.
1 2 3 1 4), the rate heterogeneity model (ratehetmodel) was
Bakeridesia (all 14 previously described species and twosetnewto gamma,
species theto number
be of categories of invariable rates (numratecats)
described by the first author) and six of the 13 specieswasof set
subg.
to 4,Dipteron
and the proportion of invariable sites (invariantsites) was set
(sensu Bates 1973) were included. The 10 Abutilón sequences available
to estimate. in support values were estimated from 1,000 repli-
Bootstrap
GenBank, including two accessions of the type species, cates underAbutilón
the same model used in the tree searches. Bayesian analyses
theophrasti, were included. Ten Abutilón species with winged
(BI) weremericarps,
conducted using MrBayes 3.1.2 (Ronquist and Huelsenbeck
some of which have never been allied with Bakeridesia,2003)wereusing
selected to
parameters from the substitution model GTR + I + G, since
determine their proper placement. Some Abutilón species TVM with
+ I +mericarps
G cannot be implemented in MrBayes. Two simultaneous and
containing more than four ovules were also included. A total MCMC
independent of 49runs were performed, each with four linked chains,
sequences were newly generated for this study. Voucher specimens,
sampling trees their
every 100 generations. Three of the four chains were
locations, and GenBank accessions for the ITS region are listed in Appen-
heated. The analysis was stopped after 9 x 106 generations when the aver-
dix 1. Base chromosome numbers were inferred by Bates (1968) and Bates
age standard deviation of split frequencies between the two runs lingered
and Blanchard (1970). These authors reportedly used all chromosome
close to 0.01 (std. dev. = 0.011382) during the last million generations. The
numbers for Abutilón (including the Callianthe clade) first available at that
10,200 trees were discarded as burn-in after visual examination of the
time, as well as two species of Bakeridesia subg. Bakeridesia, to infer base
chromosome numbers. likelihood x generation plot. The remaining trees were used to produce a
majority-rule consensus tree and to calculate posterior probabilities. The
DNA Extraction and ITS Amplification - Genomic DNA was isolated
aligned data matrix and the resulting trees are deposited in TreeBASE
from silica gel-dried leaf material (15 samples) and herbarium specimens
(study number SI 21 16).
(34 samples) using a modified СТАВ protocol (Doyle and Doyle 1987).
Ancestral State Reconstruction - To investigate the evolution of the
The ITS region was amplified by PCR using a previously described
winged
amplification program (Aguilar et al. 2003). This region includes ITS1, the mericarp, the parsimony algorithm for reconstructing ancestral
5.8S subunit and ITS2. Double-stranded amplification was performed states
using in Mesquite ver. 2.75 (Maddison and Maddison 2011) was used.
Presence /absence of the winged mericarp was determined from herbar-
primers leul (Baum et al. 1998) and ITS4 (White et al. 1990). In instances
where amplification was initially unsuccessful for the full ITS sequence,ium
the specimens of Abutilón and Bakeridesia species sampled for this study.
two segments of ITS were amplified separately, using primers leulBakeridesia
and species with a wing "remnant" were scored as lacking the
ITS2 (Baum et al. 1998) for ITSl, and ITS3B (Baum et al. 1998) andwing
ITS4 (absent). Absence of the character in all other taxa was inferred from
the literature. The tree used for this reconstruction was the strict consen-
(White et al. 1990) for 5.8S and ITS2. Amplification products were separated
sus of equally most parsimonious trees from the parsimony analysis. The
on a 1.3% agarose gel in 0.5% TBE buffer, stained with ethidium bromide,
unordered state assumption was used, as was the MPRs (most parsimo-
and then visualized with UV on a transilluminator. Of the 49 accessions
amplified, only two showed evidence of weak additional "ghost" nious
bands;reconstructions) mode.
these secondary fragments were shorter than the ITS fragments and are
conceivably results of secondary structures (given the low annealing tem-
perature of the thermal cycler program used). Sequences for these samples, Results
however, showed no evidence of polymorphism. The PCR products were
cleaned using the Wizard® SV gel and PCR cleanup system (PromegaThe final data matrix was 564 nucleotides long with 430
variable characters, 329 of them parsimony-informative. The
Corporation, Madison, Wisconsin). Automated sequencing using forward

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 714 SYSTEMATIC BOTANY [Volume 37

maximum sequence and Bakeridesia s. s. is nested well within

divergence was the clade. Within
29.9%, an
С content was 54.4%. Within the Callianthe
Bakeridesia s. s., species relationships are poorly resolved. In
below), 73 characters were
likelihood and Bayesianvariable
analyses, B. exalata (Oaxacaand accession)22 w
informative. The maximum
is moderately supported as sistersequence
to the remainder of the diverg
clade
Callianthe clade was (LB 7.3%,
= 58%, PP = 0.94), and but its the
position is mean
unresolved in the G +
parsimony analysis. A clade including
53.4%. In the final alignment, a 25six bp exemplars from
deletio
coastal Veracruz, Mexico clade,
all species in the Callianthe (B. integerrima, Вand . notolophium,
could В.
molecular synapomorphy ferruginea (2) and В. sp. nov. 1 the
for (2)) was strongly
clade. supported in Par
yielded 499 equally all three analyses (LB
most = 89%, PB = 90%, PP = 1.0), but very few
parsimonious tree
tency index (Kluge otherandspecies-levelFarris
relationships were consistently
1969) recovered.
of 0.
uninformative characters included),
Ancestral reconstructions a retent
of the "winged" mericarp yielded
1989) of 0.73, and a two most parsimonious
length of reconstructions
1,921 (MPRs). Both MPRs
steps. Li
showed a wingless
yielded a best tree with a InL common ancestor
score with one ofgain and two
-10,11
mental online Fig. 1). losses ofBootstrap
the mericarp wing in Bakeridesia s. s. For the Callianthe
values from
parsimony analyses clade,(> two 50%) MPRs wereand recovered, Bayesian
both with a winged com-pos
mon ancestor.
ties of > 0.5 (calculated from One MPR showed one loss and three gains of
a majority-rule
the winged mericarp,
are presented on a strict consensus while the other showed
of two losses and
equally
ous trees, shown in two Fig.gains of the
1. character.
The Species with winged mericarps are re
topologies
indicated on Fig. 1 by
three analyses were mostly a large asterisk.
congruent, with f
Both Bakeridesia and Abutilón as currently ci
polyphyletic. Species included Discussionin these gener
clades consistently recovered in all three an
the " Bakeridesia s. s. All three
clade/' topologies (MP, ML and BI) recovered
composed sole in th
study are similar
subg. Bakeridesia species; the to the"results obtained by Tate et
Abutilón s. al. s.
(20
the type of Abutilónfor andtaxa represented
species in both. of As determined
Bastardia in the earli a
(K. Schum.) Hassl.; and
study, many of what we currently
the generic alliances are recognized
nam
Malveae (Kubitzkiof
Callianthe clade, composed and Bayer
species2003) are not monophyle
curren
(Fig. 2). The Anoda alliance,
either Abutilón or Bakeridesia subg. composed of Anoda Cav. a
Dipteron
clades have Bayesian Períptera
posterior DC., is the soleprobability
monophyletic alliance recovered sco
least 95% bootstrap this analysis (though the
support for monophylyboth of some alliances w
likeli
mony analyses (Fig. not 1). tested). The Abutilón alliance, containing Abutilón, Sida
The "Bakeridesia s. and s. their
clade," " Abutilón
respective segregates, is non-monophyletic, s. as s. a
" Callianthe clade," arethe Batesimalva,
part Gaya,of Sidalcea, and Sphaeralcea clade
a larger alliances. S
is polyphyletic as
three analyses (likelihood commonly circumscribed
bootstrap [LB] (Fig. 1). = 8
bootstrap [PB] = 58%, Our Bayesian
results confirm that neither posteriorBakeridesia nor Abutilónpro
0.83, Node 1 in Fig. monophyletic
1) consisting as currently delimited. of Athe previously follunnam
clade, hereby referred
Bakeridesia s. s. + Períptera , Anoda, to as the Callianthe clade, is composed
Horsfordia
Wissadula , Tetrasida species
, and currently assigned to Bakeridesia subg. (here
Pseudabutilon Dipteron a
as the "B" clade); Abutilón
"pluriovulate" species s.ofs.; Abutilónand (Fig. 1).the Cal
The Callianthe c
Gaya , Briquetia, Hochr is well supported
eutiner by Bayesiana, (PP =Dirhamp
1.0), likelihood (L
Billieturnera (hereafter 95%), and parsimony
referred (PB = 98%) analyses.
to as In addition,
the spec "
parsimony analysis (Fig. included in 1),
the Callianthe clade share a synapomorphy
Billieturnera, th o
a clade comprising Abutilón 25-bp deletion in ITS2.s.The s.clade
+is the
united by"В"
the follow
c
chotomy. In contrast, suite of the
morphological characters: carpels
Bayesian andcontaining
likefour
place Billieturnera as more
the ovules,basal
a shrubby habit,
member a glabrous staminal
of columth
with weak support (LB petals with
= impressed
< 50%, veins, a PP
pubescent
= inner mericarp w
0.72).
Within the "C" clade, and leavesa that are toothed group
sister and /or lobed. The 25-bp del
relatio
Gaya and the Callianthe clade is well suppor tion, the shrubby habit and the petals with impressed veins
PB = 92%, PP = 1.0). synapomorphies Species for the clade, and further examination
relationships with m
clade are poorly resolved. reveal that theInpubescent
the inner mericarp wall is also a syn
parsimony a
amoenum is recovered as sister to the remainder of the new morphy. The remaining diagnostic features are symples
genus, with moderate support (PB = 67%), but in Bayesian morphies, however no other clade included in this st
and likelihood analyses, A. amoenum is placed in a derived possesses this particular combination of characters. We in
position, with a clade comprising Abutilón regnellii + the base chromosome number for the clade to be x = 8, ba
Bakeridesia esculenta (LB = 70%, PB = 74%, PP = 0.88) being on sporic chromosome counts of 2n = 16 for eight members
sister to the remaining species, though this topology is not the clade (Krapovickas 1957; Bates 1976; Fernández 19
strongly supported (LB = < 50%, PP = 0.56). Most other rela- Fernández et al. 2003). The Callianthe clade is sister to Gay
tionships within Callianthe have little to no support. Though the two clades are phylogenetically closely related
In all three analyses, Abutilón s. s. and the "В" clade show a prefer not to assign the Callianthe clade to Gaya because t
moderately supported sister group relationship (LB = 64%, share few morphological similarities (see Table 1), and Gaya
PB = 78%, PP = 1.0, Node 2 in Fig. 1). The topologies of the a base chromosome number of x = 6 (Bates 1968).
"B" clade outside of Bakeridesia s. s. are identical in all three The Abutilón s. s. clade is composed of Abutilón species t
analyses: Pseudabutilon is sister to the rest of the "B" clade have an herbaceous habit, fewer than four ovules per car

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2012] DONNELL ET AL.: CALLIANTHE: A NEW GENUS OF MALVEAE 715

Fig. 1. Strict consensus tree of 499 equally most parsimonious trees (CI = 0.40; RI = 0.73; L = 1921). Bootstrap suppo
shown above the branches (maximum likelihood / maximum parsimony), and Bayesian posterior probabilities are sh
of greater than 50% and posterior probabilities of greater than 0.5 are shown; a small asterisk indicates a topolog
posterior probability) support. Topologies not supported by greater than 50% in any analyses are left blank. Large as
dorsal mericarp wing, a feature until now thought to be diagnostic of Bakeridesia.

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 716 SYSTEMATIC BOTANY [Volume 37

Fig. 2. Simplified majority-rule Bayesian consensus tr

are listed to the right of the genera.

or both. Also included in the Abutilón s. s. clade are Bastardia Abutilón (Bates 1968; Bates and Blanchard 1970; Fryxell
1997b; Kubitzki and Bayer 2003), but they have never been
bivalvis and Bastardiopsis densiflora, two Neotropical species
with uniovulate carpels and pseudocapsular fruits. Both included within Abutilón due to their uniovulate carpels.
The placement of these species within a clade comprised of
Bastardia and Bastardiopsis have been previously allied with

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2012] DONNELL ET AL.: CALLIANTHE: A NEW GENUS OF MALVEAE 717

Table 1. Distinguishing characteristics and distribution of Callianthe, Bakeridesia s. s., Abutilón s.

Callianthe Abutilón s. s. Bakeridesia s. s. Gaya

Distribution Primarily South American Pantropical; a few Primarily Mesoamerican, Neotropical, from
with a center of diversity temperate species with center of diversity Mexico and West
in eastern Brazil in southern Mexico Indes to Bolivia
Number of ovules /carpel 4-13 3 (- 6) 2-7 1
Growth habit Shrubs to small trees Annuals, woody perennials, Tall shrubs to small trees Erect herbs or subshrubs
sub-shrubs & shrubs
Base chromosome no. x = 8 x = 7 x = 15 x = 6
Petal color Various Various Yellow to orange, Usually yellow
often red at base
Petal venation Veins impressed, often Veins not impressed Veins not impressed Veins not impressed
a different color
from petals
Mericarp winged? Some species Never Most species Never
Leaf margin Toothed and /or lobed; Toothed; rarely entire Entire Toothed to subentire
rarely entire
Staminal column Glabrous Glabrous to densely pubescent Glabrous to densely pubescent Glabrous
Inner wall of mericarp Pubescent Glabrous Glabrous Unknown

Abutilón species was previously shown by but

Abutilón varies, Tate et al.
the inner (2005),
mericarp wall is glabrous and
though only three Abutilón species shinywere sampled.
in the specimens Thirteen
examined by the first author (rep-
species of Abutilón were sampled for the
resenting present
~ 60% study
of the species). and in Callianthe
The mericarps
our results further confirm that Bastardia may or may and not have a lacerate dorsal margin
Bastardiopsis are or remnant,
nested among Abutilón species in the but nowell-supported
species in the Abutilón s. Abutilón
s. clade possess this charac-
s. s. clade. The base chromosome number for the Abutilón s. s. teristic. Finally, Callianthe differs from both Bakeridesia and
clade is x = 7, as reported for several Abutilón species as well Abutilón by its petals with impressed veins (this feature
as for Bastardia and Bastardiopsis (Fryxell 1988,1997b). absent in the other two genera). From Gaya, its closest rela-
The Bakeridesia s. s. clade has 100% bootstrap support fortive, Callianthe differs in habit ( Gaya is largely herbaceous),
both ML and MP, and a 1.0 PP score for Bayesian analysis. number of ovules per mericarp (mericarps in Gaya are
The clade is distinctive morphologically, as it is composed of uniovulate), corolla venation (veins not impressed in Gaya)
large shrubs (or small trees) with yellow to orange flowers, and base chromosome number (x = 6 in Gaya).
entire leaves, and winged mericarp margins. The base chromo- Our results are consistent with recent molecular studies
some number is x = 15 (Bates 1973; Bates and Blanchard 1970). revealing that many of the genera in Malveae traditionally
Based on the totality of evidence from ITS sequences, mor- characterized by a single morphological trait are not mono-
phology, chromosome counts, and geography, we propose phyletic. The lacerate mericarp wing, previously thought to
that the Callianthe clade be recognized at the generic level. It be a distinguishing feature of the genus Bakeridesia s. 1., has
is substantially different from its closest relative, Gaya, in apparently evolved more than once in the Malveae (Fig. 1).
morphology and chromosome number. Furthermore, it is Similar results have been reported for genera of the Malva
not closely related to the genera with which its members alliance (Alcea L., Althaea L., Lavatera L., and Malva L.), which
have previously been affiliated, Bakeridesia and Abutilón . To were traditionally separated from each other based on the
maintain Bakeridesia and Abutilón as monophyletic genera, degree of fusion of the epicalyx. Phylogenetic analyses using
the Callianthe clade must be segregated. It is described heremolecular markers showed that delimiting these genera
as the new genus Callianthe. solely based on epicalyx fusion resulted in artificial genera
Table 1 summarizes some geographical, morphological, that were not monophyletic (Ray 1995; Garcia et al. 2009). In
and karyological characteristics of Callianthe and other gen- Palaua, heavy reliance on superimposed carpels as a defining
era associated with it. Callianthe differs from Bakeridesia s. s. character of the genus nearly resulted in the exclusion of
by its toothed and/or lobed leaves (entire in Bakeridesia s. s.), P. sandemanii, which has uniseriate carpels. Phylogenetic
pubescent inner mericarp wall and its chromosome number. studies using nuclear and cpDNA showed that P. sandemanii
In addition, Callianthe has a wider variety of flower colors, is actually nested within a monophyletic Palaua and repre-
including purple, white, yellow, pink, and cream. In contrast, sents the sole loss of the superimposed carpels in the genus
Bakeridesia s. s. has uniformly yellow to orange flowers with (Huertas et al. 2007).
or without red marks at the base of the petals. Callianthe is a In our case, the winged mericarp is a synapomorphy for
bit more difficult to distinguish morphologically from Abutilón Bakeridesia s. s., as all of the species included in that clade
s. s. Callianthe has a woody habit (herbaceous or woody in possess the wing (or a remnant) but none of its closest phylo-
Abutilón), and the mericarps contain four or more ovules genetic relatives do. However, it is also present in several
(versus usually three in Abutilón, but a few weedy herba- species of the Callianthe clade. Based on results from the
ceous species of Abutilón can have up to six). In general, ancestral state reconstruction, the winged mericarp evolved
Callianthe has larger flowers than Abutilón, with petals sel- once in Bakeridesia s. s., but was lost three times. In Callianthe,
dom shorter than 1.5 cm (petals of Abutilón can be less than the mericarps of the common ancestor were winged, and two
four mm in length), and there is a predominance of small, evolutionary scenarios are possible for the development of the
stellate pubescence on all parts, especially the mericarps trait within the clade. The first is that the mericarp wing was lost
(including the inner mericarp wall). The pubescence in once and regained three times. The second option is that it was

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 718 SYSTEMATIC BOTANY [Volume 37

lost twice and gainednarrowly

twice oblong or ovate, 0.3-1.5
(see Fig. cm long,
1).0.1-0.6 cm wide.
Better
Callianthe may helpPetioles
clarify the
2-20 cm long, with evolutionary
the longer petioles on older parts
of the plant.
winged mericarp within the Lamina elliptic to broadly
clade. In ovate (broadly
addition,
using more variable obovate
gene in species with lobed leaves),
regions areunlobed, shallowly
needed
species relationships 3-lobed or 5-7-lobed,
within basally cordate to lobate,
Bakeridesia s.rounded,
s. and
Included within Callianthe are
cuneate, near-truncate or the
oblique, margins "floweri
serrate-crenate to
"parlor maples/' long prized
serrulate-crenulate as dentate
or subentire (rarely horticultu
or entire), apex
majority of flowering maples
acute to on
acuminate, leaf surface the market
stellate-pubescent, often more t
of unknown parentage denselyandon abaxial surface;sold
are leaves sometimes
as " discolorous,
Abutil
a name that has not venation
yet been Flowers
brochidodromous. validly publishe
solitary, paired or in clus-
have likely contributed
ters of 3 to
flowers,the genetic
axillary, on makeup
pedicels to 10 cm long. Involucel
maples including, but not
absent. Calyces be limited
gamosepalous, to:
5-lobed, the lobes 1^4.5 cm, Ab
(Gillies often heavily veined
ex Hook. & Arn.) Walp.,or keeled, the midvein prominent, st
Abutilón
Lindl., minutely toHook,
Abutilón
darwinii densely stellatef.,
pubescent
and (sometimes den-
Abutiló
(Hook.) A.
St.-Hil. & droid,
Naud.,rarely simple),all of which
the indumentum often brown, tanareor h
to Callianthe. With theferrugineous
exception (rarely silvery-tan
of or white), the lobes ovate- pic
Abutilón
species were first described
lanceolate, free from from
the tube 2/8-7/8cultivated
of the way to the acute m
apex. Corolla rotate or campanulate, exceeding calyx in
length, the petals obovate and often clawed at the base, 1.5-
Taxonomic Treatment
5.5 cm long, 0.5-5 cm wide; red, white, cream, yellow, purple,
lavender, pink, or burgundy, usually prominently veined, the
Callianthe Donnell gen. nov.-TYPE SPECIES: Callianthe
rufinerva (A. St.-Hil.) Donnell. veins sometimes of another color. Staminal column usually
exserted but sometimes included, 0.7-5 cm long, glabrous.
Frútices vel arbores parvi, flores sine epicalyce. Columna
Carpels 8-14(-16), 4-13-ovulate. Styles equal in number
staminalis glabra, pétala nervosa. Carpella unilocularia,
to carpels, surpassing the anther mass by up to 1 cm or
pariete interior stellato-pubescenti. Differì a Abutilón Mill,
included. Stigmas capitate. Fruits schizocarpic, to 5 cm in
petalis comparate longioribus, habitu nunquam herbáceo,
diameter. Mericarps unilocular, sclerotic and brown to
carpellis plus quam 3-ovulatis. Differt a Bakeridesia Hochr.
black at maturity, 1-4.8 cm long, sparsely to densely stellate-
foliis dentatis vel lobatis vel uterque. Chromosomatum
numerus: x = 8. pubescent on outer wall, minutely stellate-pubecent on inner
wall, dehiscing dorsally, the dorsal margin entire, subentire,
Shrubs or treelets 1-8 m. tall, stems densely pubescentapically cuspidate, or with a lacerate, wing-like projection.
when young and minutely pubescent at maturity, the tri- Seeds to 4 mm long, stellate pubescent or pilose. Base chromo-
some
chomes stellate or simple, rarely dendroid. Leaves simple, to number: x = 8.
22 cm long and 19 cm wide, generally decreasing in size
Etymology - Callianthe is derived from the Greek for
"beautiful flower."
towards the stem apex. Stipules often early deciduous, linear,

Key to Genera

1. Leaves entire; tall shrubs or small trees; petals yellow to orange, with or without a red basal spot; pubescence markedly
stellate-ferruginous; mericarps almost always with a conspicuous lacerate dorsal "wing"; southern Mexico to Central America
and northwestern South America

1. Leaves toothed and/

without a red basal sp
with some temperate species
2. Plants herbaceous

2. Plants woody

3. Ovules 4-
staminal column glabrous; primarily South American (~5 Mesoamerican species

3. Ovules 3 per mericarp; petals lacking deeply impressed ve

staminal column glabrous or pubescent; pantropical, few temperate species

Species Requiring
NewnotCombinations - New comb
all of the diagnostic m
listed below discerned
following meet from 1)
criteria: the original
The the
first
seen the type specimenspecimens.
(holotype Nevertheless,
and /or isotype),
thes
person or via high resolution
becausephoto,
they and
havecan confirm
reported s
type specimen fits the first
of 2 author's concept
n = 16 and of the t
they possess n
Callianthe; 2) The original species
ical description
features. has been
An Abutilón nom
the type specimen viewed has been confirmed
an invaluable tocom
resource for b
cited in the protologue; and 3) the original species d
Callianthe amoena (K. Schum
and /or the type specimen conforms to the generic dia
amoenum К. Schum., Fl. Br
Neotropical with four or more ovules per mericarp, a
TYPE: BRAZIL. "Brasilia mer
habit, glabrous staminal column, and heavily veined p
some species ( Callianthe
Callianthe andrade-limae (Monteiro)C.
fluviatilis, Donnell, comb. nov.
jujuiense, C.
frideňci, С. pieta Bakeridesia С.
, С. purpusii, andrade-limae Monteiro, Anais.С.
regnellii, Soc. Bot. Brasil
striata, an

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2012] DONNELL ET AL.: CALLIANTHE: A NEW GENUS OF MALVEAE 719

23: 121 (1973). -TYPE: BRAZIL. Buique, (1913).-TYPE:

Andrade-Lima ARGENTINA.
55- Jujuy, Celilegua, Cerro
2111 (holotype: IPA!; isotype: RBR!). San Fransisco, 20 June 1911, M. Lillo 10800 (lectotype,
Callianthe bedfordiana (Hook.) Donnell, designated
comb.Fryxell
nov.2002:
SidaLIL!; isolectotype: G, LIL!).
bedfordiana Hook., Bot. Mag. 68: plate 3892
Callianthe (1841).
lanata (Miq.) Donnell, comb. nov. Abutilón
Abutilón bedfordianum (Hook.) A. St.-Hil.
lanatum& Miq.,
Naud., Ann.
Linnaea 22: 553 (1849). Bakeridesia lanata
Soc. Nat. Bot. sér 2, 18: 48 (1842).-TYPE: BRAZIL.
(Miq.) Organ
Leite & Monteiro, Bol. Soc. Portug. Ci. Nat. 5(2):
123 (19 55). designated
Mountains, 1837, G. Gardner 320 (lectotype, -TYPE: BRAZIL. Minas Gerais, Ciadas, 1844,
Fryxell 2002: G-DEL; isolectotype: NY!).Л. F. Regnell 1-13 (holotype: S; isotypes: С, F!, K!, U!,US!).
Callianthe bezerrae (Monteiro) Donnell, comb. nov. Callianthe latipetala (G. L. Esteves & Krapov.) Donnell, comb,
Bakeridesia bezerrae Monteiro, Anais. Soc. Bot. Brasil nov. Abutilón latipetalum G. L. Esteves & Krapov., Kew.
23:118 (1973). -TYPE: BRAZIL. Ceará, Serra de Baturité, Buli. 57(2): 479 (2002).-TYPE: BRAZIL. São Paulo,
Guaramiranga, 29 June 1941, P. Bezzera 288 (holotype: Pindamonhagaba, towards Campos do Jordão, 9 May
RBR!; isotype: NY!). 2000, Esteves & Pastore 2758 (holotype: SP; isotypes: SPSF,
Callianthe brenesii (Standi.) Donnell, comb. nov. Abutilón SPF, CTES!).
brenesii Standi., Pubi. Field Mus. Nat. Hist., Bot. Ser. 18: Callianthe laxa (Rusby) Donnell, comb. nov. Abutilón laxum
664 (1937). -TYPE: COSTA RICA. Río Jesús de San Rusby, Mem. New York Bot. Gard. 7: 296 (1927).-TYPE:
Ramón, A. Brenes 3587 (holotype: F!). BOLIVIA. Río Bopi Valley, September 1927, H. H. Rusby
Callianthe cyclonervosa (Hochr.) Donnell, comb. nov. 658 (Holotype: NY!; Isotypes: K!, BKL!).
Abutilón cyclonervosum Hochr., Annuaire Conserv. Jard. Callianthe longifolia (K. Schum.) Donnell, comb. nov. Abutilón
Bot. Geneve 6: 27 (1902). Abutilón bakeri Rusby, Bull. longifolium К. Schum., Fl. Bras. (Martius) 12(3): 402 (1891).
New York Bot. Gard. 4: 329 (1907), nom. superfl.-TYPE: Pro syn.: Weldena longifolia (first published as synonym of
BOLIVIA. Yungas, Coroico, 22 July 1894, M. Bang 2537 A. longifolium) - TYPE: BRAZIL. Minas Gerais, Serra do
(holotype: NY!; isotypes: С, CAS (fragment)!, CM, E!, F!, Chumbo, Pohl 3001 (Lectotype, designated by Fryxell
GH, K!, LIL!, MO!, NY!, PH!, US!). 2002: W; isolectotype: F!)
Callianthe darwinii (Hook, f.) Donnell, comb. nov. Abutilón Callianthe macrantha (A. St.-Hil.) Donnell, comb. nov. Abutilón
darwinii Hook, f., Bot. Mag. 97: t. 5917 (1871).-TYPE: macranthum A. St.-Hil., Fl. Bras. Merid. 1: 208 (1827).
BRAZIL (holotype: plate 5917, loc. cit.). Bakeridesia macrantha (A. St.-Hil.) Leite & Monteiro, Bol.
Callianthe elegans (A. St.-Hil.) Donnell, comb. nov. Abutilón Soc. Portug. Ci. Nat. 5(2): 124 (1955).-TYPE: BRAZIL.
elegans A. St.-Hil., FI. Bras. Merid. 1: 207 (1827).-TYPE: Minas Gerais, prope Onça, A. St.-Hil B1700 (holotype:
BRAZIL. In nemoribus montis dicti Serra Negra ad P; isotypes: B!,US!).
limites provinciarum Rio de Janeiro et Minas Geraes, A. Callianthe malmeana (R. E. Fries) Donnell, comb. nov.
St.-Hil. s. n. (holotype: P!; isotype: P!). Abutilón malmeanum R.E. Fries, Kongl. Svenska Vetensk.
Callianthe fluviatilis (Veil.) Donnell, comb. nov. Sida Acad. Handl. 42(12): 31 (1908).-TYPE: BRAZIL. Matto
fluviatilis Veli., Fl. Flumin. 278 (1825) [1827]. Abutilón Grosso, Santa Anna de Chapada, collected between
fluviatile (Veil.) К. Schum., Fl. Bras. (Martius) 12(3): 399 1902 and 1903, Malme 11:1989 (lectotype, designated by
(1891).-TYPE: Lectotype, designated by Fryxell, 2002: Fryxell 2002: S!).
plate 7, loc. cit.; Epitype, designated by Fryxell, 2002:
Callianthe mexiae (R. E. Fries) Donnell, comb. nov. Abutilón
BRAZIL. Aquae duct bei Rio de Janeiro, А. С. V. Schott mexiae R. E. Fries, Kongl. Svenska Vetensk. Acad. Handl.
s. п. (В!). 24(2): 7 (1947). -TYPE: BRAZIL. Minas Gerais, Viçosa,
Callianthe geminiflora (Kunth) Donnell, comb. nov. Abutilón agricultural College lands, north-west hill, abandoned
geminiflorum Kunth, Nov. Gen. Sp. 5: 274 (1822).-TYPE: road up Chacka Valley, 4 July 1930, Mexia 4838 (holo-
VENEZUELA. Near Caracas, F. W. H. A. Humboldt & type: S!; isotypes: CAS!, PH!, UC,US!, WIS).
A. J. A. Bonpland 1132 (holotype: P; isotype: B!). Callianthe monteiroi (Krapov.) Donnell, comb. nov. Abutilón
Callianthe glaziovii (K. Schum.) Donnell, comb. nov. Abutilón monteiroi Krapov., Hickenia 1(51): 273 (1982).-TYPE:
glaziovii К. Schum., Fl. Bras. (Martius) 12(3): 408 (1891).- BRAZIL. Minas Gerais, Serra do Espinhaço, 11 February
TYPE: BRAZIL. Near Rio de Janeiro, November 1879, 1972, W.A. Andersonetal. 35793 (holotype: UB; isotype: NY!).
Glaziou 10307 (holotype: B!; isotype: С, K!). Callianthe mouraei (K. Schum.) Donnell, comb. nov.
Callianthe inaequalis (Link & Otto) Donneil, comb. nov. Sida Abutilón mouraei К. Schum., Fl. Bras. (Martius) 12(3): 410
inaequalis Link & Otto, Icon. Pl. Select. 75, t. 34 (1825). (1891). -TYPE: BRAZIL. Near Rio de Janeiro, 1882,
Abutilón inaequale (Link & Otto) К. Schum., Fl. Bras. Glaziou 13542 (lectotype, designated by Fryxell 2002: P;
(Martius) 12(3): 408 (1891).-TYPE: BRAZIL, (lectotype, isolée totypes: C, K!, P).
designated by Fryxell 2002: plate 34, loc. cit.). Callianthe muelleri-friderici (Gíirke & K. Schum.) Donnell,
Callianthe jaliscana (Standi.) Donnell, comb. nov. Abutilóncomb. nov. Abutilón muelleri-friderici Gürke & К. Schum.,
jaliscanum Standi., Pubi. Field Mus. Nat. Hist., Bot. Ser. 4: Fl. Bras. (Martius) 12(3): 410 (1891).-TYPE: BRAZIL.
229 (1929). -TYPE: MEXICO. Jalisco, Hacienda de Ototal, Santa Catarina, prope Blumenau, 1881, J. H. Schenck 497
Arroyo de los Palos Blancos, W of San Sebastián, March (holotype: B!).
8 1927, Ynes Mexia 1842a (holotype: F!; isotypes: BM!, Callianthe pachecoana (Standi. & Steyerm.) Donnell, comb,
CAS!, DS!, GH, MICH!, MO!, NY!, US!). nov. Abutilón pachecoanum Standi. & Steyerm., Pubi.
Callianthe jujuiensis (Hassl.) Donneil, comb. nov. AbutilónField Mus. Nat. Hist., Bot. Ser. 23(2): 61 (1944).-TYPE:
jujuiense Hassl., Repert. Spec. Nov. Regni Veg. 12: 499 GUATEMALA. Quezaltenango, region of Las Nubes,

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 720 SYSTEMATIC BOTANY [Volume 37

south of San Martin Veg.Chile 13: 501 (1915), nom. illegit (superfluous,
Verde, January Art. 19
52.1).-TYPE: BRAZIL.
Standley 83528 (lectotype, Rio de Janeiro, February
designated by 1882,
Fryxe
isolectotypes: EAP!, F!-2, NY!,
A. F. M. Glaziou US!).
12438 (holotype: B!; isotypes: С, G-DC,
K!, P.).
Callianthe petiolaris (Kunth) Donnell, comb. no
Callianthe striataGen.
petiolare Kunth, Nov. (Dicks, ex Lindl.) Donnell,
Sp. comb.
5: nov. 272 (182
COLOMBIA. Humboldt & Dicks,
Abutilón striatum Bonpland
ex Lindl., Edward's Bot. Reg. s. n. (ho
isotype: P!). 25: mise 39 (1839). -TYPE: BRAZIL (holotype: Dickson in
Callianthe pickellii (Monteiro) Donnell, comb. nov. Maund, The Botanist 3: plate 144, 1839). Callianthe
Bakeridesia pickellii Monteiro, Anais. Soc. Bot. Brasil 23: torrendii (Monteiro) Donnell, comb. nov. Bakeridesia
116 (1973).-TYPE: BRAZIL. Pernambuco, Tapera, São torrendii Monteiro, Anais. Soc. Bot. Brasil 23: 125 (1973).-
Bento, 18 August 1923, B. }. Pickel 150 (holotype: IPA!). TYPE: BRAZIL. Bahía, Vitoria da Conquista, С. Torrend 5
(holotype: RBR!).
Callianthe picta (Gilles ex Hook. & Arn.) Donnell, comb,
nov. Sida picta Gilles ex Hook. & Arn., Bot. Misc. 3: 154 Callianthe tridens (Standi. & Steyerm.) Donnell, comb. nov.
(1833). Abutilón pictum (Gilles ex Hook. & Arn.) Walp., Abutilón tridens Standi. & Steyerm., Pub. Field. Mus. Nat.
Repert. Bot. Syst. 1: 324 (1824).-TYPE: ARGENTINA. Hist., Bot. Ser. 23: 173 (1944).-TYPE: GUATEMALA. El
Buenos Aires, J. Gilles , s. n. (lectotype, designated by Progreso, hills between Finca Piamonte and the slopes
Fryxell 2002: K!; isolectotype: OXF). southeast, Sierra de las Minas, 4 February 1942, J. A.
Steyermark 43439 (lectotype, designated by Fryxell 1988:
Callianthe purpusii (Standi.) Donnell, comb. nov. Abutilón
F!; isolectotypes- F!, US!).
purpusii Standi. Contr. U.S. Natl. Herb. 23: 750 (1923).-
TYPE: MEXICO. Veracruz, Zacuapan, Barranca de Callianthe vexillaris (E. Morren) Donnell, comb. nov. Abutilón
Tenampa, January 1910, C. A. Purpus 4332 (holotype: vexillarum E. Morren, Belgique Hort. 14: 289 (1864).-
US!; isotypes: BM!, F!, GH, MO!). TYPE: SOUTH AMERICA (holotype: loc. cit., plate 16).

Callianthe regnellii (Miq.) Donnell, comb. nov. Abutilón The following names represent species that will likely be
transferred to Callianthe once more source information is
regnellii Miq., Linnaea 22: 554 (1849).-TYPE: BRAZIL.
available and /or nomenclatural issues are resolved: Abutilón
Minas Gerais, Caldas, A. F. Regneil 11-17 (holotype: U;
isotypes: F!, NY!). appendiculatum K. Schum., Abutilón arboreum Sweet., Abutilón
carneum A. St.-Hil., Abutilón carinatus Krapov., Abutilón
Callianthe rufinerva (A. St.-Hil.) Donnell, comb. nov. Abutilón
dianthum C. Presi, Abutilón esculentum A. St.-Hil., Abutilón
rufìnerve A. St.-Hil., Fl. Bras. Merid. 1: 205 (1827).
falcatum A. St.-Hil & Naud., Abutilón macrocarpum St. - Hil. &
Bakeridesia rufinerva (A. St.-Hil.) Monteiro, Bol. Soc. Portug.
Naud., Abutilón megapotamicum (A. Spreng.) St.-Hil. & Naud.,
Ci. Nat. 5(2): 124 (1955).-TYPE: BRAZIL. Villa do Principe,
Abutilón montanum A. St.-Hil., Abutilón nigricans G. L. Esteves
propéque praedium Domingos Alfonso, A. St.-Hil s. n.
& Krapov., Abutilón niveum Gris., Abutilón pauciflorum A.
(holotype: P; isotypes: P, MPU!-2).
St.-Hil., Abutilón pedrae-brancae K. Schum., Abutilón peltatum
Callianthe rufivela (Hochr.) Donnell, comb. nov. Bakeridesia K. Schum., Abutilón peruvianum (Lam.) Kearney, Abutilón
rufivela Hochr., Annuaire Conserv. Jard. Bot. Geneve 21: piurense Ulbr., Abutilón sordidum K. Schum., Abutilón sylvaticum
419 (1920). Abutilón rufivelum К. Schum. ex. Baker f., J. (Cav.) K. Schum., Abutilón weberbaueri Ulbr., Bakeridesia
Bot. 31: 271 (1893), nom. nud..-TYPE: BRAZIL. Rio de purpurascens (Link.) Monteiro.
Janeiro, Nova Friburgo, Alto Macahé, 1891, Glaziou
18136 (holotype: G-DC; isotypes: B!, F!, K!).
Acknowledgments. There are many people without whom this
Callianthe scabrida (K. Schum.) Donnell, comb. nov. Abutilón research would not have been possible. The first author thanks Greg
scabridum К. Schum., Fl. Bras. (Martius) 12(3): 413 (1891). Wahlert and Daryl Lam for their guidance in lab work protocols; Tara
Killen and Vij ay Nadella for invaluable assistance with sequencing;
Bakeridesia scabrida Kearney ex Fryxell., Lundellia 5: 106
Melanie Schori for guidance in DNA extraction techniques and help with
(2002).-TYPE: BRAZIL. Prope Estiva ut lego, Sellow 744 botanical Latin; my dissertation committee members, Gar Rothwell and
(syntype: B!). Alycia Stigall for their helpful comments on this manuscript; Ricardo
Madrigal and Hector Gómez for their incredible assistance with specimen
Callianthe schenckii (K. Schum.) Donnell, comb. nov.
collection in Mexico; Sergio Avendaño and Dr. Gonzalo Castillo of the
Abutilón schenckii К. Schum., Fl. Bras. (Martius) 12(3):
Instituto de Ecología in Xalapa, Veracruz for hosting me and allowing me
412 (1891). -TYPE: BRAZIL. Madre de Dios, June 1833,to use their supplies for field collection; Dr. Miguel Angel Perez-Ferrara
for helping me arrange my collecting time in Chiapas; Dr. Jennifer Tate
L. Riedel 1316 (lectotype, designated by Fryxell 2002: P;
isolectotype: K!). and the late Dr. Paul Fryxell for helping me explore options for
Malvaceae research; and the Malvaceae yahoo group members (espe-
Callianthe sellowiana (Klotzsch) Donnell, comb. nov. Sida cially Stewart R. Hinsley). Finally, I must acknowledge Ohio University,
sellowiana Klotzsch, Allg. Gartenzeitung (Otto & Dietrich) which partially funded my research through a Student Enhancement
4: 9 (1836). Synonyms: Abutilón sellowianum (Klotzsch) Grant and two Graduate Student Senate Original Work Grants.
Regel., Ann. Sci. Nat., Bot. sér. 4, 12: 379 (1859); Bakeridesia
sellowiana (Klotzsch) Monteiro, Bol. Soc. Portug. Ci. Nat.
Literature Cited
5(2): 127 (1955) -TYPE: Hort. Bot. Berlin, ex Brazil. Sellow
s. n. (holotype: B!). Akaike, H. 1974. A new look at the statistical model identification. IEEE
Callianthe senilis (K. Schum.) Donnell, comb. nov. Abutilón Transactions on Automatic Control 19: 716-723.

senile К. Schum., Fl. Bras. (Martius) 12(3): 424 (1891). Aguilar, J. F., P. A. Fryxell, and R. K. Jansen. 2003. Phylogenetic relation-
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editor and analysis program for Windows 95/98/NT. Nucleictaxon, Acids country, state or department, collection locality, voucher collection
Symposium Series 41: 95-98. (herbarium): GenBank accession number (s). Taxa with only one GenBank
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accession number correspond to a complete ITS sequence (ITS1 + 5.8S +
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15-16: others were amplified or downloaded as ITS1 and ITS2 separately
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and are indicated as such. Voucher information is given only for
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21: 405-428.

Huertas, M. L., J. V. Schneider, and G. Zizka. 2007. Phylogenetic analysis Abutilón abutiloides (Jacq.) Garcke ex Hochr., Mexico, Nayarit, G. Flores-
of Palaua (Malveae, Malvaceae) based on plastid and nuclear Franco et al. 2966 (MO): JQ753294; Abutilón amoenum К. Schum., Brazil,
sequences. Systematic Botany 32: 157-165. Paraná, União da Vitoria, A. Krapovickas & С. L. Cristobal 39596 (MO):
Jiménez-Reyes, N. 2003. Morfología de los granos de polen de laJQ753306; familia Abutilón andrewsianum W. Fitzg.: AY591807; Abutilón costicalyx
Malvaceae de Jalisco, México. II: Anoda, Bakeridesia, Bastrardia, К. Schum., Brazil, São Paulo, Caraguatatuba, E. L. Esteves et al. 2760 (MO):
Bastar diastrum, Briquetia, Gaya y Gossypium. Scientia-CUCBA 5: 1-30. JQ753295; Abutilón dianthum С. Presi., Bolivia, La Paz, Franz Tamayo,
Kearney, T. H. 1949. Malvaceae: a new subtribe and genus and new Cayóla et al. 890 (MO): JQ753291; Abutilón dianthum С. Presi., Bolivia, La
combinations. Leaflets of Western Botany 5: 189-191. Paz, Franz Tamayo, A. Fuentes & R. Alvares 4810 (MO): JQ753292; Abutilón
Kearney, T. H. 1955. A tentative key to the North American species of eremitopetalum Caum: EF219363; Abutilón geminiflorum Kunth, Venezuela,
Abutilón Miller. Leaflets of Western Botany 7: 241-254. Aragua, Distrito Ricaurte, W. Meier et al. 5892 (US): JQ753293; A.
Kearney, T. H. 1958. A tentative key to the South American species of grandifolium (Willd.) Sweet: EF219369; Abutilón x hybridům hort. ex Voss,
Abutilón Miller. Leaflets of Western Botany 8: 201-216. cultivated origin, A. Donnell 132 (NY): JQ753296; Abutilón incanum (Link)

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 722 SYSTEMATIC BOTANY [Volume 37

Sweet: EF219368; Abutilón indicum (L.) Sweet: AY863059; Abutilón (MO): JQ753266; Bakeridesia notolophium (A. Gray) Hochr., Mexico,
malacum S. Watson, Mexico, Sonora, Sonoyta, R. S. Feiger 92-81 (MO): Veracruz, Tancoco, M. Nee & K. Taylor 28735 (NY): JQ753286; Bakeridesia
JQ753298; Abutilón menziesii Seem.: EF219365; Abutilón monteiroi Krapov. pickellii Monteiro, Brazil, Paraíba, Maturéia, M. F. Agra et al. 4913 (MO):
Brazil, Minas Gerais, Grão Mogol, G. M. Hatschbach 54157 (MO): JQ753303; Bakeridesia pittieri (Donn. Sm.) D. M. Bates, Mexico, Chiapas,
JQ753299; Abutilón muelleri-friderici Giirke & К. Schum., Brazil, Rio Ocozocoautla, A. Donnell 94 w/ H. Gomez-Dominguez (NY): JQ753276;
Grande do Sul, Caxias do Sul, L. Scur 828 (MO): JQ753300; Abutilón Bakeridesia pittieri (Donn. Sm.) D. M. Bates, Mexico, Chiapas, Union Juárez,
pauciflorum A. St.-Hil., Bolivia, Cochabamba, Carrasco, E. Fernández et al. D. E. Breedlove & Smith 31560 (NY): JQ753303; Bakeridesia rufinerva (A. St.-
35226 (MO): JQ753259; Abutilón peruvianum (Lam.) Kearney, Peru, Cusco, Hil.) Monteiro, Brazil, Paraná, Guaraqueçaba, G. Gatti 450 (NY): JQ753267;
La Convención, L. Valenzuela et al. 69 (MO): JQ753261; Abutilón Bakeridesia rufivela Hochr., Brazil, Espirito Santo, Feixe de Pedras, M. M.
peruvianum (Lam.) Kearney, Peru, Contumazá, Cajamarca, A. Sagástegui Arbo et al. 5587 (TEX): JQ753304; Bakeridesia scabrida (К. Schum.) Kearney ex
A. 14878 (MO): JQ753262; Abutilón peruvianum (Lam.) Kearney, Bolivia, Fryxell, Brazil, Rio de Janeiro, Valença, A. M. Amorim 3075 (MO): JQ753305;
Santa Cruz, Manuel M. Caballero, M. Muñoz 165 (MO): JQ753263; Bakeridesia sp. nov. la, Mexico, Veracruz, Vega de Alatorre, A. Donnell
Abutilón pubistamineum Ulbr.: AJ251049 (ITS1), AJ274993 (ITS2); Abutilón 78 w/ R. Madrigal-Chavero (NY): JQ753290; Bakeridesia sp. nov. lb, Mexico,
purpusii Standi., Costa Rica, San José, Canton de Acosta, }. F. MoralesVeracruz, Alto Lucero, A. Donnell 105 w/ R. Madrigal-Chavero (NY):
2343 (MO): JQ753301; Abutilón regnellii Miq., Brazil, São Paulo, Serra JQ753268;
da Bakeridesia sp. nov. 2a, Mexico, Oaxaca, Santo Domingo
Tehuantepec, T. MacDougall s. n. (NY): JQ753281; Bakeridesia sp. nov. 2b,
Mantiqueira, J. R. Piraní 2510 (SPF): JQ753302; Abutilón sandwicense (О.
Mexico, Oaxaca, Santo Domingo Tehuantepec, A. Donnell 112 w/ R. Madrigal-
Deg.) Christoph.: EF219367; Abutilón sp., Argentina, Misiones, San Pedro,
Chavero (NY): JQ753280; Bakeridesia subcordata (Hochr.) D. M. Bates, Mexico,
N. B. Deginani et al. 1662 (MO): JQ753260; Abutilón sp. 1: AY591808;
Abutilón sp. 2, Paraguay, Central, San Antonio, A. Krapovickas & C. L. Oaxaca, San Juan Bautista Cuicatlan, A. Donnell 119 w/ R. Madrigal-
Cristóbal 44548 (MO): JQ753307 (ITS1), JQ753308 (ITS2); Abutilón Chavero (NY): JQ753289; Bakeridesia vulcanicola (Standi.) D. M. Bates, El
theophrasti Medik. (1): DQ287984; Abutilón theophrasti Medik. (2): Salvador, Santa Ana, Parque Nacional Montecristo, J. Monterrosa 436 (US):
DQ006017; Alcea rosea L.: EF679714; Andeimalva mandonii (Baker f.) J. JQ753284;
A. Bakeridesia yucatana (Standi.) D. M. Bates, Mexico, Quintana Roo,
Tate: AY172220; Anoda crenatiflora Ortega: AJ251043 (ITS1), AJ274987 Akumal, E. Cabrera 4016 (MO): JQ753269; Bastardia bivalvis (Cav.) Kunth ex
Griseb.: AY591813; Bastardiastrum cinctum (Brandegee) D. M. Bates:
(ITS2); Bakeridesia amoena Fryxell, Mexico, Chiapas, Cintalapa, A. Donnell
113 w/ R. Madrigal-Chavero (NY): JQ753283; Bakeridesia bakeriana (Rose)AY591814; Bastardiopsis densiflora (Hook. & Arn.) Hassl.: AY591815;
D. M. Bates, Mexico, Jalisco, La Huerta, E. J. Lott 4029 (MICH): JQ753275;
Billieturnera helleri (Rose ex A. Heller) Fryxell: AY591817; Briquetia sonorae
Bakeridesia cf. gloriosa, Mexico, Oaxaca, Santa María Chimalapa, A.Fryxell: AY591818; Callirhoe involucrata (Torr. & A. Gray) A. Gray:
AY591819; Corynabutilon virifolium (Cav.) Kearney: AJ274970 (ITS1),
Donnell 111 w / R. Madrigal-Chavero (NY): JQ753279; Bakeridesia cf.
AJ275001 (ITS2); Cristaria andícola Gay: AY591821; Dendrosida breedlovii
integerrima, Mexico, Oaxaca, Santo Domingo Tehuantepec, M. L. Torres
855 (NY): JQ753271; Bakeridesia cf. pittieri, Mexico, Chiapas, Amatenango Fryxell: AJ251032 (ITS1), AJ274976 (ITS2); Dirhamphis mexicana Fryxell:
de la Frontera, A. Donnell 88 w/ H. Gomez-Dominguez (NY): JQ753282; AY591822; Eremalche parryi (Greene) Greene: AJ304938; Gaya atiquipana
Bakeridesia esculenta (A. St.-Hil.), Monteiro, Brazil, Rio de Janeiro, São Krapov.: AY591825; Gaya calyptrata (Cav.) Kunth ex K. Schum.: AJ251048
Pedro de Aldeia, J. A. Kallunki & J. R. Pirani 633 (NY): JQ753264;(ITS1), AJ274992 (ITS2); Hochreutinera amplexifolia (DC.) Fryxell: AY591827;
Bakeridesia exalata D. M. Bates: AY591812; Bakeridesia exalata D. M. Bates,Horsfordia exalata Fryxell: AY591831; Malvastrum americanum (L.) Torr.:
Honduras, El Paraíso, A. Molina R. 31253 (MO): JQ753285; Bakeridesia AY591842; Malvella sagittifolia (A. Gray) Fryxell: AJ25104 5 (1TS1), AJ274989
ferruginea (Martyn) Krapov., Mexico, Veracruz, Tlaltetela, F. Ventura A. (ITS2; Neobrittonia acerifolia (G. Don) Hochr.: AY591844; Palaua rhombifolia
15735 (NY): JQ753287; Bakeridesia ferruginea (Martyn) Krapov., Mexico, Graham: AY591846, Períptera punicea (Lag.) DC.: AY591847; Phymosia
Oaxaca, Temascal, L. Cortes 725 (MO): JQ753265; Bakeridesia gaumeri umbellata (Cav.) Kearney: AY591848; Plagianthus divaricatus J. R. Forst. &
(Standi.) D. M. Bates: AY591859; Bakeridesia gloriosa D. M. Bates, Mexico, G. Forst.: AY591849; Pseudabutilon umbellatum (L.) Fryxell: AJ274964 (ITS1),
Chiapas, Tuxtla Gutierrez, A. Donnell 83 zv/ H. Gomez-Dominguez (NY): AJ274995 (ITS2); Robinsonella lindeniana (Turcz.) Rose & Baker f.: AY591851;
JQ753288; Bakeridesia integerrima (Hook, f.) D. M. Bates, Mexico, Oaxaca, Sida abutifolia Mill.: AJ274961 (ITS1), AJ251617 (ITS2); Sida aggregata С.
Asunción Ixaltepec, E. A. Pérez-García 2046 (MO): JQ753270; Bakeridesia Presi.: AJ274943 (ITSl), AJ251599 (ITS2); Sida cordifolia L.: AJ274945 (ITS1),
AJ251601 (ITS2); Sida glutinosa Comm. ex Cav.: AJ251037 (ITSl), AJ274981
integerrima (Hook, f.) D. M. Bates, Venezuela, Falcon, Distrito Bolivar, van
der Werjf & R. Wingfield 7433 (MO): JQ753273; Bakeridesia integerrima (ITS2); Sida hookeriana Miq.: AJ274967 (ITSl), AJ274998 (ITS2); Sida
(Hook, f.) D. M. Bates, Mexico, Veracruz, Ozuluama de Mascareñas, platycalyx
A. F. Muell. ex Benth.: AJ251041 (ITSl), AJ274985 (ITS2); Sida
Donnell 72 w/ R. Madrigal-Chavero (NY): JQ753272; Bakeridesia macrantharhombifolia L.: AJ274953 (ITSl), AJ251609 (ITS2); Sidalcea hartwegii A. Gray
(A. St.-Hil.) Leite & Monteiro, Brazil, Rio Grande do Sul, São Francisco ex
de Benth.: AJ304890; Sidalcea stipularis J. T. Howell & G. H. True: AJ304932;
Paula-Linha, R. Wasum 178 (US): JQ753297; Bakeridesia molinae D. M. Sidastrum paniculatum (L.) Fryxell: AJ251040 (ITSl), AJ274984 (ITS2) ; Tarasa
Bates, Honduras, Yoro, Olanchito, Davidse et al. 35493 (TEX): JQ753277; albertii Reiche: AY172200; Tarasa trisecta (Griseb.) Krapov.: AY172236;
Bakeridesia nelsonii (Rose) D. M. Bates, Mexico, Chiapas, La Concordia, A. Tetrasida chachapoyensis (Baker f.) Fryxell & Fuertes: AY591854; Tetrasida
weberbaueri (Ulbr.) Fryxell & Fuertes: AY591855; Wissadula boliviana R. E.
Donnell 100 w / H. Gomez-Dominguez (NY): JQ753278; Bakeridesia nelsonii (Rose)
Fr:
D. M. Bates, Guatemala, Huehuetenango, "El Tapon", J. Castillo 1743 AY591856.

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