Callianthe (Malvaceae): A New Genus of Neotropical Malveae Author(s): Aliya A. Donnell, Harvey E. Ballard Jr., and Philip D. Cantino Source: Systematic Botany, 37(3):712-722. 2012. Published By: The American Society of Plant Taxonomists URL: http://www.bioone.org/doi/full/10.1600/036364412X648689 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. 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 1 Department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701, U. S. A. 2 Author for correspondence (Aliya.A.Donnell.1@ohio.edu) Communicating Editor: Mark P. Simmons Abstract—The boundaries between the genera Bakeridesia and Abutilon 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 Abutilon. 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 Abutilon and 15 in Bakeridesia). Not only is Callianthe phylogenetically removed from both Bakeridesia and Abutilon, 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 Abutilon. The following new combinations are made Callianthe amoena, C. andrade-limae, C. bedfordiana, C. bezerrae, C. brenesii, C. cyclonervosa, C. darwinii, C. elegans, C. fluviatilis, C. geminiflora, C. glaziovii, C. inaequalis, C. jaliscana, C. jujuiensis, C. lanata, C. latipetala, C. laxa, C. longifolia, C. macrantha, C. malmeana, C. mexiae, C. monteiroi, C. mouraei, C. muellerifriderici, C. pachecoana, C. petiolaris, C. pickellii, C. picta, C. purpusii, C. regnellii, C. rufinerva, C. rufivela, C. scabrida, C. schenckii, C. sellowiana, C. senilis, C. striata, C. torrendii, C. tridens, and C. vexillaris. Keywords—Abutilon, Bakeridesia, Brazil, ITS. Abutilon Mill. (Malvaceae) is a large genus of approximately 160 species. It is extremely variable in morphology and has a broad geographic distribution, with representatives on all continents except Antarctica. Abutilon is one of several poorly delimited genera of Malvaceae that have historically been “dumping grounds” for difficult-to-place species. In the past, there has been a tendency to assign to Abutilon any species of tribe Malveae that lacked an epicalyx and had multiple ovules per carpel if it did not easily fit into another, better circumscribed genus. Abutilon has been described as “the most difficult of the genera of Malvaceae” (Kearney 1958, p. 201) and is considered to be heterogeneous with “many taxonomic problems” (Fryxell 2002, p. 79). The genus as a whole has never been monographed, but there have been several treatments of regional groups (St.-Hilaire 1825; St.-Hilaire and Naudin 1842; Schumann 1891; Standley 1937; Standley and Steyermark 1949; Kearney 1955, 1958; Robyns 1965; Krapovickas 1969; P. A. Fryxell 1976, 1988, 1992; J. E. Fryxell 1983), and several genera have been segregated from Abutilon over the years (Kearney 1949; Fryxell and Fuertes 1992; Fryxell 1997a). Currently, Abutilon is characterized within tribe Malveae by its lack of an epicalyx, mericarps lacking an internal constriction (endoglossum), five or more carpels containing at least two ovules each, capitate stigmas, mericarps lacking a dorsal “wing,” and non-inflated fruits. Traditionally, the number of ovules per carpel has been a diagnostic character for species groups within Abutilon. In many of the older Neotropical keys, this character was the first major dichotomy (St.-Hilaire 1825; St.-Hilaire and Naudin 1842; Schumann 1891; Robyns 1965), and species were grouped into one of two categories: three or fewer ovules per carpel (“pauciovulate”) or four or more ovules per carpel (“pluriovulate”). Though many taxonomists viewed the pluriovulate species as a natural group (St.-Hilaire 1825; St.-Hilaire and Naudin 1842; Schumann 1891; Robyns 1965; Bates and Blanchard 1970), it was not formally recognized until Fryxell (1988) gave the group sectional status in his treatment of Mexican Abutilon (sect. Pluriovulata). He later suggested that the pluriovulate species may warrant elevation to generic rank (Fryxell 1997b, 2002). The pluriovulate species of Abutilon share a base chromosome number of x = 8 (Bates 1968; Bates and Blanchard 1970), in contrast to the remainder of the genus, which has a base chromosome number of x = 7 (Bates 1968; Bates and Blanchard 1970). Notable exceptions are the weedy, annual species of Abutilon, such as A. theophrasti and A. indicum, which have been reported to have up to six ovules per mericarp but have a base chromosome number of x = 7; in all other morphological features they fit in with the pauciovulate species. Though Abutilon has never been treated as a whole, several genera have been segregated from it over the years, including Pseudabutilon R. E. Fr., Corynabutilon (K. Schum.) Kearney, Tetrasida Ulbr., and Bastardia Kunth. Another such segregate is Bakeridesia Hochr., a genus of tall shrubs native to Latin America. Bakeridesia is distinguished from other genera in tribe Malveae by having a lacerate wing on the dorsal margin of the mericarp. For this reason, Bakeridesia galeottii Hochr. was removed from Abutilon (Hochreutiner 1913). Subsequently, 27 species were added to Bakeridesia (Hochreutiner 1920; Monteiro Filho 1955, 1973; Bates 1973; Fryxell 2002; Fryxell and Olivera 2001). Currently, 28 species are assigned to Bakeridesia and two subgenera are recognized: one that ranges from Mexico to Ecuador (subgenus Bakeridesia) and another that is exclusively Brazilian (subgenus Dipteron Hochr.). Though both subgenera have winged mericarps, many differences exist between them. Species included in subgenus Bakeridesia differ from Abutilon in several characters. In addition to the winged mericarps, plants in this group have entire leaves, yellow or orange flowers (petals various colors in Abutilon), pollen with three apertures (Jiménez-Reyes 2003), compared with 2–6 and 8–15 apertures reported in Abutilon (Fryxell 1997b), and a base chromosome number of x = 15 (Bates and Blanchard 1970; Bates 1973). In contrast, species included in subgenus Dipteron are more similar to Abutilon in leaf margin (toothed and/or lobed) and flower color, though the base 712 DONNELL ET AL.: CALLIANTHE: A NEW GENUS OF MALVEAE Materials and Methods Plant Material and Taxon Sampling—A total of 104 species and 35 genera were included in this study, representing ten of the 14 generic alliances of tribe Malveae recognized by Kubitzki and Bayer (2003). Outgroup selection was based on results from prior molecular analyses of Malveae using the ITS region (Tate et al. 2005). To assess the monophyly of Bakeridesia and Abutilon, a proportionately greater number of species from the clades most closely related to Bakeridesia subg. Bakeridesia and to Abutilon were sampled. Sixteen species of Bakeridesia subg. Bakeridesia (all 14 previously described species and two new species to be described by the first author) and six of the 13 species of subg. Dipteron (sensu Bates 1973) were included. The 10 Abutilon sequences available in GenBank, including two accessions of the type species, Abutilon theophrasti, were included. Ten Abutilon species with winged mericarps, some of which have never been allied with Bakeridesia, were selected to determine their proper placement. Some Abutilon species with mericarps containing more than four ovules were also included. A total of 49 sequences were newly generated for this study. Voucher specimens, their locations, and GenBank accessions for the ITS region are listed in Appendix 1. Base chromosome numbers were inferred by Bates (1968) and Bates and Blanchard (1970). These authors reportedly used all chromosome numbers for Abutilon (including the Callianthe clade) available at that time, as well as two species of Bakeridesia subg. Bakeridesia, to infer base chromosome numbers. DNA Extraction and ITS Amplification—Genomic DNA was isolated from silica gel-dried leaf material (15 samples) and herbarium specimens (34 samples) using a modified CTAB protocol (Doyle and Doyle 1987). The ITS region was amplified by PCR using a previously described amplification program (Aguilar et al. 2003). This region includes ITS1, the 5.8S subunit and ITS2. Double-stranded amplification was performed using primers leu1 (Baum et al. 1998) and ITS4 (White et al. 1990). In instances where amplification was initially unsuccessful for the full ITS sequence, the two segments of ITS were amplified separately, using primers leu1 and ITS2 (Baum et al. 1998) for ITS1, and ITS3B (Baum et al. 1998) and ITS4 (White et al. 1990) for 5.8S and ITS2. Amplification products were separated on a 1.3% agarose gel in 0.5% TBE buffer, stained with ethidium bromide, and then visualized with UV on a transilluminator. Of the 49 accessions amplified, only two showed evidence of weak additional ”ghost” bands; these secondary fragments were shorter than the ITS fragments and are conceivably results of secondary structures (given the low annealing temperature of the thermal cycler program used). Sequences for these samples, however, showed no evidence of polymorphism. The PCR products were cleaned using the Wizard® SV gel and PCR cleanup system (Promega Corporation, Madison, Wisconsin). Automated sequencing using forward amplification primers was conducted on an Applied Biosystems Genetic Analyzer 3130x at the Genomics Facility at Ohio University, Athens, Ohio. The following primers were used for sequencing: ITS5 (White et al. 1990) for the full sequence and ITS1, and ITS3B for ITS2. Due to the high quality of sequences obtained from forward sequencing alone, either of the full ITS or ITS1 and ITS2 separately, reverse sequencing was not necessary. Sequence Alignment and Phylogenetic Analysis—Sequences were edited using Sequence Scanner (v. 1.0, Applied BiosystemsÔ) and aligned using the CLUSTAL W interface in BioEdit (Hall 1999), followed by minimal manual corrections. The majority of the minor adjustments made attempted to minimize gaps, and where feasible to align gaps to be simultaneous across groups of taxa (nested gaps). The remaining corrections were to ensure that the computer-generated alignment reflected actual biological processes (i.e. transitions are more likely to occur than transversions). The highly conserved 5.8S region was not available for all sequences in GenBank, and visual inspection of available 5.8S sequences revealed low variation relative to the two spacers. Of the 104 sequences included, 14 downloaded from GenBank lacked the 5.8S region. An additional 34 sequences had only partial 5.8S regions due to lack of primer region overlap in taxa that were amplified in two parts (ITS1 and ITS2). Because nearly half of the sequences lacked a complete 5.8S sequence, and the percentages of variable (10.2%) and parsimony-informative (4.2%) characters were relatively low (compared to 58.3% of characters parsimony-informative in the ITS1 and ITS2), the region was excluded from the phylogenetic analysis. The percentage of characters in the data matrix (with the 5.8S removed) scored as missing was 8.7% including gaps, which were treated as missing data. Parsimony analyses were performed with NONA ver. 2.0 (Goloboff 1999) and implemented throughWinclada 1.00.08 (Nixon 2002) using the Ratchet (Island Hopper) (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 (1,000 replicates each with 10 search reps and the “do not do max*” option). Maximum likelihood (ML) analyses (Felsenstein 1973) were performed using GARLI 0.96 (Zwickl 2006) under the TVM + I + G model of substitution, which was selected as the best model to fit the data by jModelTest (Guindon and Gascuel 2003; Posada 2008) based on Akaike’s information criterion (Akaike 1974). The ML analyses were conducted with the following parameters: state frequencies (statefreq) were set to equal, the number of relative substitution rate parameters (ratematrix) was set to (0 1 2 3 1 4), the rate heterogeneity model (ratehetmodel) was set to gamma, the number of categories of invariable rates (numratecats) was set to 4, and the proportion of invariable sites (invariantsites) was set to estimate. Bootstrap support values were estimated from 1,000 replicates under the same model used in the tree searches. Bayesian analyses (BI) were conducted using MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003) using parameters from the substitution model GTR + I + G, since TVM + I + G cannot be implemented in MrBayes. Two simultaneous and independent MCMC runs were performed, each with four linked chains, sampling trees every 100 generations. Three of the four chains were heated. The analysis was stopped after 9 106 generations when the average standard deviation of split frequencies between the two runs lingered close to 0.01 (std. dev. = 0.011382) during the last million generations. The first 10,200 trees were discarded as burn-in after visual examination of the likelihood generation plot. The remaining trees were used to produce a majority-rule consensus tree and to calculate posterior probabilities. The aligned data matrix and the resulting trees are deposited in TreeBASE (study number S12116). Ancestral State Reconstruction—To investigate the evolution of the winged mericarp, the parsimony algorithm for reconstructing ancestral states in Mesquite ver. 2.75 (Maddison and Maddison 2011) was used. Presence/absence of the winged mericarp was determined from herbarium specimens of Abutilon and Bakeridesia species sampled for this study. Bakeridesia species with a wing “remnant” were scored as lacking the wing (absent). Absence of the character in all other taxa was inferred from the literature. The tree used for this reconstruction was the strict consensus of equally most parsimonious trees from the parsimony analysis. The unordered state assumption was used, as was the MPRs (most parsimonious reconstructions) mode. + chromosome number is inferred to be x = 8 (see discussion section). Because of these differences, some authors have questioned whether the Brazilian species actually belong in Bakeridesia (Bates 1973; Fryxell 1997b), but due to insufficient chromosome, pollen, and molecular data, it has been difficult to test this. Other authors accept the genus Bakeridesia without including the species of subgenus Dipteron (Mabberley 1997; Kubitzki and Bayer 2003). The boundary between Bakeridesia subg. Dipteron and Abutilon has been unclear. Several pluriovulate species of Abutilon have a lacerate wing on the dorsal margin of the mericarps but have not been transferred to Bakeridesia. On the other hand, several species of Bakeridesia subg. Dipteron have leaf morphology and flower characteristics that are similar to the pluriovulate species of Abutilon. In addition, all species of subgenus Dipteron have more than four ovules per mericarp and are therefore pluriovulate. Here we present evidence, based on the ITS region of the 18S-26S nuclear ribosomal repeat, morphology, and chromosome numbers, that neither Bakeridesia nor Abutilon is monophyletic as currently delimited. We demonstrate that most of the pluriovulate species of Abutilon (i.e. excluding the weedy annual species mentioned above) and the species of Bakeridesia subgenus Dipteron together form a previously unrecognized clade, to be described here as a new genus, Callianthe Donnell. 713 + 2012] Results The final data matrix was 564 nucleotides long with 430 variable characters, 329 of them parsimony-informative. The 714 SYSTEMATIC BOTANY maximum sequence divergence was 29.9%, and the mean G + C content was 54.4%. Within the Callianthe clade (described below), 73 characters were variable and 22 were parsimonyinformative. The maximum sequence divergence within the Callianthe clade was 7.3%, and the mean G + C content was 53.4%. In the final alignment, a 25 bp deletion was revealed in all species in the Callianthe clade, and could be considered a molecular synapomorphy for the clade. Parsimony analysis yielded 499 equally most parsimonious trees, with a consistency index (Kluge and Farris 1969) of 0.40 (parsimonyuninformative characters included), a retention index (Farris 1989) of 0.73, and a length of 1,921 steps. Likelihood analysis yielded a best tree with a lnL score of –10,112.808927 (supplemental online Fig. 1). Bootstrap values from likelihood and parsimony analyses (> 50%) and Bayesian posterior probabilities of > 0.5 (calculated from a majority-rule consensus tree) are presented on a strict consensus of equally most parsimonious trees, shown in Fig. 1. The topologies recovered from all three analyses were mostly congruent, with few differences. Both Bakeridesia and Abutilon as currently circumscribed are polyphyletic. Species included in these genera fall into three clades consistently recovered in all three analyses (Fig. 1): the “Bakeridesia s. s. clade,” composed solely of Bakeridesia subg. Bakeridesia species; the “Abutilon s. s. clade” including the type of Abutilon and species of Bastardia and Bastardiopsis (K. Schum.) Hassl.; and what we are naming here the Callianthe clade, composed of species currently assigned to either Abutilon or Bakeridesia subg. Dipteron (Fig. 1). All three clades have Bayesian posterior probability scores of 1.0, and at least 95% bootstrap support for both likelihood and parsimony analyses (Fig. 1). The “Bakeridesia s. s. clade,” “Abutilon s. s. clade,” and the “Callianthe clade,” are part of a larger clade recovered in all three analyses (likelihood bootstrap [LB] = 80%, parsimony bootstrap [PB] = 58%, Bayesian posterior probabilities [PP] = 0.83, Node 1 in Fig. 1) consisting of the following subclades: Bakeridesia s. s. + Periptera, Anoda, Horsfordia, Bastardiastrum, Wissadula, Tetrasida, and Pseudabutilon (hereafter referred to as the “B” clade); Abutilon s. s.; and the Callianthe clade + Gaya, Briquetia, Hochreutinera, Dirhamphis, and perhaps Billieturnera (hereafter referred to as the “C” clade). In the parsimony analysis (Fig. 1), Billieturnera, the “C” clade, and a clade comprising Abutilon s. s. + the “B” clade form a trichotomy. In contrast, the Bayesian and likelihood analyses place Billieturnera as the basal member of the “C” clade, albeit with weak support (LB = < 50%, PP = 0.72). Within the “C” clade, a sister group relationship between Gaya and the Callianthe clade is well supported (LB = 92%, PB = 92%, PP = 1.0). Species relationships within the Callianthe clade are poorly resolved. In the parsimony analysis, Abutilon amoenum is recovered as sister to the remainder of the new genus, with moderate support (PB = 67%), but in Bayesian and likelihood analyses, A. amoenum is placed in a derived position, with a clade comprising Abutilon regnellii + Bakeridesia esculenta (LB = 70%, PB = 74%, PP = 0.88) being sister to the remaining species, though this topology is not strongly supported (LB = < 50%, PP = 0.56). Most other relationships within Callianthe have little to no support. In all three analyses, Abutilon s. s. and the “B” clade show a moderately supported sister group relationship (LB = 64%, PB = 78%, PP = 1.0, Node 2 in Fig. 1). The topologies of the “B” clade outside of Bakeridesia s. s. are identical in all three analyses: Pseudabutilon is sister to the rest of the “B” clade [Volume 37 and Bakeridesia s. s. is nested well within the clade. Within Bakeridesia s. s., species relationships are poorly resolved. In likelihood and Bayesian analyses, B. exalata (Oaxaca accession) is moderately supported as sister to the remainder of the clade (LB = 58%, PP = 0.94), but its position is unresolved in the parsimony analysis. A clade including six exemplars from coastal Veracruz, Mexico (B. integerrima, B. notolophium, B. ferruginea (2) and B. sp. nov. 1 (2)) was strongly supported in all three analyses (LB = 89%, PB = 90%, PP = 1.0), but very few other species-level relationships were consistently recovered. Ancestral reconstructions of the “winged” mericarp yielded two most parsimonious reconstructions (MPRs). Both MPRs showed a wingless common ancestor with one gain and two losses of the mericarp wing in Bakeridesia s. s. For the Callianthe clade, two MPRs were recovered, both with a winged common ancestor. One MPR showed one loss and three gains of the winged mericarp, while the other showed two losses and two gains of the character. Species with winged mericarps are indicated on Fig. 1 by a large asterisk. Discussion All three topologies (MP, ML and BI) recovered in this study are similar to the results obtained by Tate et al. (2005) for taxa represented in both. As determined in the earlier study, many of the generic alliances currently recognized in Malveae (Kubitzki and Bayer 2003) are not monophyletic (Fig. 2). The Anoda alliance, composed of Anoda Cav. and Periptera DC., is the sole monophyletic alliance recovered in this analysis (though the monophyly of some alliances was not tested). The Abutilon alliance, containing Abutilon, Sida L., and their respective segregates, is non-monophyletic, as are the Batesimalva, Gaya, Sidalcea, and Sphaeralcea alliances. Sida is polyphyletic as commonly circumscribed (Fig. 1). Our results confirm that neither Bakeridesia nor Abutilon is monophyletic as currently delimited. A previously unnamed clade, hereby referred to as the Callianthe clade, is composed of species currently assigned to Bakeridesia subg. Dipteron and “pluriovulate” species of Abutilon (Fig. 1). The Callianthe clade is well supported by Bayesian (PP = 1.0), likelihood (LB = 95%), and parsimony (PB = 98%) analyses. In addition, species included in the Callianthe clade share a synapomorphy of a 25-bp deletion in ITS2. The clade is united by the following suite of morphological characters: carpels containing four or more ovules, a shrubby habit, a glabrous staminal column, petals with impressed veins, a pubescent inner mericarp wall, and leaves that are toothed and/or lobed. The 25-bp deletion, the shrubby habit and the petals with impressed veins are synapomorphies for the clade, and further examination may reveal that the pubescent inner mericarp wall is also a synapomorphy. The remaining diagnostic features are symplesiomorphies, however no other clade included in this study possesses this particular combination of characters. We infer the base chromosome number for the clade to be x = 8, based on sporic chromosome counts of 2n = 16 for eight members of the clade (Krapovickas 1957; Bates 1976; Fernández 1981; Fernández et al. 2003). The Callianthe clade is sister to Gaya. Though the two clades are phylogenetically closely related, we prefer not to assign the Callianthe clade to Gaya because they share few morphological similarities (see Table 1), and Gaya has a base chromosome number of x = 6 (Bates 1968). The Abutilon s. s. clade is composed of Abutilon species that have an herbaceous habit, fewer than four ovules per carpel, 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 support values from 1,000 replicates are shown above the branches (maximum likelihood / maximum parsimony), and Bayesian posterior probabilities are shown below. Only bootstrap values of greater than 50% and posterior probabilities of greater than 0.5 are shown; a small asterisk indicates a topology with less than 50% bootstrap (0.5 posterior probability) support. Topologies not supported by greater than 50% in any analyses are left blank. Large asterices indicate species posessing the dorsal mericarp wing, a feature until now thought to be diagnostic of Bakeridesia. 716 SYSTEMATIC BOTANY [Volume 37 Fig. 2. Simplified majority-rule Bayesian consensus tree with nodes collapsed to genus. The generic alliances (following Kubitzki and Bayer 2003) are listed to the right of the genera. or both. Also included in the Abutilon s. s. clade are Bastardia bivalvis and Bastardiopsis densiflora, two Neotropical species with uniovulate carpels and pseudocapsular fruits. Both Bastardia and Bastardiopsis have been previously allied with Abutilon (Bates 1968; Bates and Blanchard 1970; Fryxell 1997b; Kubitzki and Bayer 2003), but they have never been included within Abutilon due to their uniovulate carpels. The placement of these species within a clade comprised of 2012] Table 1. DONNELL ET AL.: CALLIANTHE: A NEW GENUS OF MALVEAE 717 Distinguishing characteristics and distribution of Callianthe, Bakeridesia s. s., Abutilon s. s., and Gaya. Callianthe Distribution Number of ovules/carpel Growth habit Primarily South American with a center of diversity in eastern Brazil 4–13 Shrubs to small trees Base chromosome no. Petal color x=8 Various Petal venation Veins impressed, often a different color from petals Some species Toothed and/or lobed; rarely entire Glabrous Pubescent Mericarp winged? Leaf margin Staminal column Inner wall of mericarp Abutilon s. s. Bakeridesia s. s. Gaya Primarily Mesoamerican, with center of diversity in southern Mexico 2–7 Tall shrubs to small trees Neotropical, from Mexico and West Indes to Bolivia 1 Erect herbs or subshrubs x=6 Usually yellow Veins not impressed x = 15 Yellow to orange, often red at base Veins not impressed Never Toothed; rarely entire Most species Entire Never Toothed to subentire Glabrous to densely pubescent Glabrous Glabrous to densely pubescent Glabrous Glabrous Unknown Pantropical; a few temperate species 3 (– 6) Annuals, woody perennials, sub-shrubs & shrubs x=7 Various Abutilon species was previously shown by Tate et al. (2005), though only three Abutilon species were sampled. Thirteen species of Abutilon were sampled for the present study and our results further confirm that Bastardia and Bastardiopsis are nested among Abutilon species in the well-supported Abutilon s. s. clade. The base chromosome number for the Abutilon s. s. clade is x = 7, as reported for several Abutilon species as well as for Bastardia and Bastardiopsis (Fryxell 1988,1997b). The Bakeridesia s. s. clade has 100% bootstrap support for both ML and MP, and a 1.0 PP score for Bayesian analysis. The clade is distinctive morphologically, as it is composed of large shrubs (or small trees) with yellow to orange flowers, entire leaves, and winged mericarp margins. The base chromosome number is x = 15 (Bates 1973; Bates and Blanchard 1970). Based on the totality of evidence from ITS sequences, morphology, chromosome counts, and geography, we propose that the Callianthe clade be recognized at the generic level. It is substantially different from its closest relative, Gaya, in morphology and chromosome number. Furthermore, it is not closely related to the genera with which its members have previously been affiliated, Bakeridesia and Abutilon. To maintain Bakeridesia and Abutilon as monophyletic genera, the Callianthe clade must be segregated. It is described here as the new genus Callianthe. Table 1 summarizes some geographical, morphological, and karyological characteristics of Callianthe and other genera associated with it. Callianthe differs from Bakeridesia s. s. by its toothed and/or lobed leaves (entire in Bakeridesia s. s.), pubescent inner mericarp wall and its chromosome number. In addition, Callianthe has a wider variety of flower colors, including purple, white, yellow, pink, and cream. In contrast, Bakeridesia s. s. has uniformly yellow to orange flowers with or without red marks at the base of the petals. Callianthe is a bit more difficult to distinguish morphologically from Abutilon s. s. Callianthe has a woody habit (herbaceous or woody in Abutilon), and the mericarps contain four or more ovules (versus usually three in Abutilon, but a few weedy herbaceous species of Abutilon can have up to six). In general, Callianthe has larger flowers than Abutilon, with petals seldom shorter than 1.5 cm (petals of Abutilon can be less than four mm in length), and there is a predominance of small, stellate pubescence on all parts, especially the mericarps (including the inner mericarp wall). The pubescence in Veins not impressed Abutilon varies, but the inner mericarp wall is glabrous and shiny in the specimens examined by the first author (representing 60% of the species). The mericarps in Callianthe may or may not have a lacerate dorsal margin or remnant, but no species in the Abutilon s. s. clade possess this characteristic. Finally, Callianthe differs from both Bakeridesia and Abutilon by its petals with impressed veins (this feature absent in the other two genera). From Gaya, its closest relative, Callianthe differs in habit (Gaya is largely herbaceous), number of ovules per mericarp (mericarps in Gaya are uniovulate), corolla venation (veins not impressed in Gaya) and base chromosome number (x = 6 in Gaya). Our results are consistent with recent molecular studies revealing that many of the genera in Malveae traditionally characterized by a single morphological trait are not monophyletic. The lacerate mericarp wing, previously thought to be a distinguishing feature of the genus Bakeridesia s. l., has apparently evolved more than once in the Malveae (Fig. 1). Similar results have been reported for genera of the Malva alliance (Alcea L., Althaea L., Lavatera L., and Malva L.), which were traditionally separated from each other based on the degree of fusion of the epicalyx. Phylogenetic analyses using molecular markers showed that delimiting these genera solely based on epicalyx fusion resulted in artificial genera that were not monophyletic (Ray 1995; Garcı́a et al. 2009). In Palaua, heavy reliance on superimposed carpels as a defining character of the genus nearly resulted in the exclusion of P. sandemanii, which has uniseriate carpels. Phylogenetic studies using nuclear and cpDNA showed that P. sandemanii is actually nested within a monophyletic Palaua and represents the sole loss of the superimposed carpels in the genus (Huertas et al. 2007). In our case, the winged mericarp is a synapomorphy for Bakeridesia s. s., as all of the species included in that clade possess the wing (or a remnant) but none of its closest phylogenetic relatives do. However, it is also present in several species of the Callianthe clade. Based on results from the ancestral state reconstruction, the winged mericarp evolved once in Bakeridesia s. s., but was lost three times. In Callianthe, the mericarps of the common ancestor were winged, and two evolutionary scenarios are possible for the development of the trait within the clade. The first is that the mericarp wing was lost once and regained three times. The second option is that it was 718 SYSTEMATIC BOTANY lost twice and gained twice (see Fig. 1). Better sampling within Callianthe may help clarify the evolutionary history of the winged mericarp within the clade. In addition, further studies using more variable gene regions are needed to better resolve species relationships within Bakeridesia s. s. and Callianthe. Included within Callianthe are the “flowering maples” or “parlor maples,” long prized as horticultural plants. The majority of flowering maples on the market today are hybrids of unknown parentage and are sold as “Abutilon x hybridum”, a name that has not yet been validly published. Several species have likely contributed to the genetic makeup of the flowering maples including, but not be limited to: Abutilon pictum (Gillies ex Hook. & Arn.) Walp., Abutilon striatum Dicks. Ex Lindl., Abutilon darwinii Hook. f., and Abutilon bedfordianum (Hook.) A. St.-Hil. & Naud., all of which are here transferred to Callianthe. With the exception of Abutilon pictum, all of these species were first described from cultivated material. Taxonomic Treatment Callianthe Donnell gen. nov.–TYPE SPECIES: Callianthe rufinerva (A. St.-Hil.) Donnell. Frutices vel arbores parvi, flores sine epicalyce. Columna staminalis glabra, petala nervosa. Carpella unilocularia, pariete interior stellato-pubescenti. Differt a Abutilon Mill. petalis comparate longioribus, habitu nunquam herbaceo, carpellis plus quam 3-ovulatis. Differt a Bakeridesia Hochr. foliis dentatis vel lobatis vel uterque. Chromosomatum numerus: x = 8. Shrubs or treelets 1–8 m. tall, stems densely pubescent when young and minutely pubescent at maturity, the trichomes stellate or simple, rarely dendroid. Leaves simple, to 22 cm long and 19 cm wide, generally decreasing in size towards the stem apex. Stipules often early deciduous, linear, [Volume 37 narrowly oblong or ovate, 0.3–1.5 cm long, 0.1–0.6 cm wide. Petioles 2–20 cm long, with the longer petioles on older parts of the plant. Lamina elliptic to broadly ovate (broadly obovate in species with lobed leaves), unlobed, shallowly 3-lobed or 5–7-lobed, basally cordate to lobate, rounded, cuneate, near-truncate or oblique, margins serrate-crenate to serrulate-crenulate or subentire (rarely dentate or entire), apex acute to acuminate, leaf surface stellate-pubescent, often more densely on abaxial surface; leaves sometimes discolorous, venation brochidodromous. Flowers solitary, paired or in clusters of 3 flowers, axillary, on pedicels to 10 cm long. Involucel absent. Calyces gamosepalous, 5-lobed, the lobes 1–4.5 cm, often heavily veined or keeled, the midvein prominent, minutely to densely stellate pubescent (sometimes dendroid, rarely simple), the indumentum often brown, tan or ferrugineous (rarely silvery-tan or white), the lobes ovatelanceolate, free from the tube 2/8–7/8 of the way to the acute apex. Corolla rotate or campanulate, exceeding calyx in length, the petals obovate and often clawed at the base, 1.5– 5.5 cm long, 0.5–5 cm wide; red, white, cream, yellow, purple, lavender, pink, or burgundy, usually prominently veined, the veins sometimes of another color. Staminal column usually exserted but sometimes included, 0.7–5 cm long, glabrous. Carpels 8–14(–16), 4–13-ovulate. Styles equal in number to carpels, surpassing the anther mass by up to 1 cm or included. Stigmas capitate. Fruits schizocarpic, to 5 cm in diameter. Mericarps unilocular, sclerotic and brown to black at maturity, 1–4.8 cm long, sparsely to densely stellatepubescent on outer wall, minutely stellate-pubecent on inner wall, dehiscing dorsally, the dorsal margin entire, subentire, apically cuspidate, or with a lacerate, wing-like projection. Seeds to 4 mm long, stellate pubescent or pilose. Base chromosome number: x = 8. Etymology—Callianthe is derived from the Greek for “beautiful flower.” Key to Genera 1. 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bakeridesia Leaves toothed and/or lobed (rarely entire); plants herbaceous or woody with toothed and/or lobed leaves; petals of various colors, without a red basal spot; pubescence various; mericarps with or without a conspicuous lacerate dorsal “wing”; pantropical, with some temperate species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Plants herbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Abutilon 2. Plants woody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Ovules 4–13 per mericarp; petals with deeply impressed veins; inner mericarp wall sparsely stellate-pubescent; staminal column glabrous; primarily South American (5 Mesoamerican species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callianthe 3. Ovules 3 per mericarp; petals lacking deeply impressed veins; inner mericarp wall glabrous; staminal column glabrous or pubescent; pantropical, few temperate species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Abutilon Species Requiring New Combinations—New combinations listed below meet the following criteria: 1) The first author has seen the type specimen (holotype and/or isotype), either in person or via high resolution photo, and can confirm that the type specimen fits the first author’s concept of the new genus Callianthe; 2) The original species description has been seen and the type specimen viewed has been confirmed to be the one cited in the protologue; and 3) the original species description and/or the type specimen conforms to the generic diagnosis, i.e. Neotropical with four or more ovules per mericarp, a shrubby habit, glabrous staminal column, and heavily veined petals. For some species (Callianthe fluviatilis, C. jujuiense, C. muellerifriderici, C. picta, C. purpusii, C. regnellii, C. striata, and C. laxa), not all of the diagnostic morphological characters could be discerned from the original species description and/or type specimens. Nevertheless, these species are included in Callianthe because they have reported sporophytic chromosome numbers of 2n = 16 and they possess two or more diagnostic morphological features. An Abutilon nomenclator (Fryxell 2002) proved to be an invaluable resource for compiling the new combinations. Callianthe amoena (K. Schum.) Donnell, comb. nov. Abutilon amoenum K. Schum., Fl. Bras. (Martius) 12(3): 411 (1891).– TYPE: BRAZIL. “Brasilia meridionalis,” 1836, Sellow s. n. (B!). Callianthe andrade-limae (Monteiro) Donnell, comb. nov. Bakeridesia andrade-limae Monteiro, Anais. Soc. Bot. Brasil 2012] DONNELL ET AL.: CALLIANTHE: A NEW GENUS OF MALVEAE 23: 121 (1973).–TYPE: BRAZIL. Buique, Andrade-Lima 55– 2111 (holotype: IPA!; isotype: RBR!). Callianthe bedfordiana (Hook.) Donnell, comb. nov. Sida bedfordiana Hook., Bot. Mag. 68: plate 3892 (1841). Abutilon bedfordianum (Hook.) A. St.-Hil. & Naud., Ann. Soc. Nat. Bot. sér 2, 18: 48 (1842).–TYPE: BRAZIL. Organ Mountains, 1837, G. Gardner 320 (lectotype, designated Fryxell 2002: G-DEL; isolectotype: NY!). Callianthe bezerrae (Monteiro) Donnell, comb. nov. Bakeridesia bezerrae Monteiro, Anais. Soc. Bot. Brasil 23:118 (1973).–TYPE: BRAZIL. Ceará, Serra de Baturité, Guaramiranga, 29 June 1941, P. Bezzera 288 (holotype: RBR!; isotype: NY!). Callianthe brenesii (Standl.) Donnell, comb. nov. Abutilon brenesii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 664 (1937).–TYPE: COSTA RICA. Rı́o Jesús de San Ramón, A. Brenes 3587 (holotype: F!). Callianthe cyclonervosa (Hochr.) Donnell, comb. nov. Abutilon cyclonervosum Hochr., Annuaire Conserv. Jard. Bot. Geneve 6: 27 (1902). Abutilon bakeri Rusby, Bull. New York Bot. Gard. 4: 329 (1907), nom. superfl.–TYPE: BOLIVIA. Yungas, Coroico, 22 July 1894, M. Bang 2537 (holotype: NY!; isotypes: C, CAS (fragment)!, CM, E!, F!, GH, K!, LIL!, MO!, NY!, PH!, US!). Callianthe darwinii (Hook. f.) Donnell, comb. nov. Abutilon darwinii Hook. f., Bot. Mag. 97: t. 5917 (1871).–TYPE: BRAZIL (holotype: plate 5917, loc. cit.). Callianthe elegans (A. St.-Hil.) Donnell, comb. nov. Abutilon elegans A. St.-Hil., Fl. Bras. Merid. 1: 207 (1827).–TYPE: BRAZIL. In nemoribus montis dicti Serra Negra ad limites provinciarum Rio de Janeiro et Minas Geraes, A. St.-Hil. s. n. (holotype: P!; isotype: P!). Callianthe fluviatilis (Vell.) Donnell, comb. nov. Sida fluviatilis Vell., Fl. Flumin. 278 (1825) [1827]. Abutilon fluviatile (Vell.) K. Schum., Fl. Bras. (Martius) 12(3): 399 (1891).–TYPE: Lectotype, designated by Fryxell, 2002: plate 7, loc. cit.; Epitype, designated by Fryxell, 2002: BRAZIL. Aquae duct bei Rio de Janeiro, A. C. V. Schott s. n. (B!). Callianthe geminiflora (Kunth) Donnell, comb. nov. Abutilon geminiflorum Kunth, Nov. Gen. Sp. 5: 274 (1822).–TYPE: VENEZUELA. Near Caracas, F. W. H. A. Humboldt & A. J. A. Bonpland 1132 (holotype: P; isotype: B!). Callianthe glaziovii (K. Schum.) Donnell, comb. nov. Abutilon glaziovii K. Schum., Fl. Bras. (Martius) 12(3): 408 (1891).– TYPE: BRAZIL. Near Rio de Janeiro, November 1879, Glaziou 10307 (holotype: B!; isotype: C, K!). Callianthe inaequalis (Link & Otto) Donnell, comb. nov. Sida inaequalis Link & Otto, Icon. Pl. Select. 75, t. 34 (1825). Abutilon inaequale (Link & Otto) K. Schum., Fl. Bras. (Martius) 12(3): 408 (1891).–TYPE: BRAZIL. (lectotype, designated by Fryxell 2002: plate 34, loc. cit.). Callianthe jaliscana (Standl.) Donnell, comb. nov. Abutilon jaliscanum Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 229 (1929).–TYPE: MEXICO. Jalisco, Hacienda de Ototal, Arroyo de los Palos Blancos, W of San Sebastián, March 8 1927, Ynes Mexia 1842a (holotype: F!; isotypes: BM!, CAS!, DS!, GH, MICH!, MO!, NY!, US!). Callianthe jujuiensis (Hassl.) Donnell, comb. nov. Abutilon jujuiense Hassl., Repert. Spec. Nov. Regni Veg. 12: 499 719 (1913).–TYPE: ARGENTINA. Jujuy, Celilegua, Cerro San Fransisco, 20 June 1911, M. Lillo 10800 (lectotype, designated Fryxell 2002: LIL!; isolectotype: G, LIL!). Callianthe lanata (Miq.) Donnell, comb. nov. Abutilon lanatum Miq., Linnaea 22: 553 (1849). Bakeridesia lanata (Miq.) Leite & Monteiro, Bol. Soc. Portug. Ci. Nat. 5(2): 123 (1955). –TYPE: BRAZIL. Minas Gerais, Cladas, 1844, A. F. Regnell I-13 (holotype: S; isotypes: C, F!, K!, U!,US!). Callianthe latipetala (G. L. Esteves & Krapov.) Donnell, comb. nov. Abutilon latipetalum G. L. Esteves & Krapov., Kew. Bull. 57(2): 479 (2002).–TYPE: BRAZIL. São Paulo, Pindamonhagaba, towards Campos do Jordão, 9 May 2000, Esteves & Pastore 2758 (holotype: SP; isotypes: SPSF, SPF, CTES!). Callianthe laxa (Rusby) Donnell, comb. nov. Abutilon laxum Rusby, Mem. New York Bot. Gard. 7: 296 (1927).–TYPE: BOLIVIA. Rı́o Bopi Valley, September 1927, H. H. Rusby 658 (Holotype: NY!; Isotypes: K!, BKL!). Callianthe longifolia (K. Schum.) Donnell, comb. nov. Abutilon longifolium K. Schum., Fl. Bras. (Martius) 12(3): 402 (1891). Pro syn.: Weldena longifolia (first published as synonym of A. longifolium).–TYPE: BRAZIL. Minas Gerais, Serra do Chumbo, Pohl 3001 (Lectotype, designated by Fryxell 2002: W; isolectotype: F!) Callianthe macrantha (A. St.-Hil.) Donnell, comb. nov. Abutilon macranthum A. St.-Hil., Fl. Bras. Merid. 1: 208 (1827). Bakeridesia macrantha (A. St.-Hil.) Leite & Monteiro, Bol. Soc. Portug. Ci. Nat. 5(2): 124 (1955).–TYPE: BRAZIL. Minas Gerais, prope Onça, A. St.-Hil. B1700 (holotype: P; isotypes: B!,US!). Callianthe malmeana (R. E. Fries) Donnell, comb. nov. Abutilon malmeanum R.E. Fries, Kongl. Svenska Vetensk. Acad. Handl. 42(12): 31 (1908).–TYPE: BRAZIL. Matto Grosso, Santa Anna de Chapada, collected between 1902 and 1903, Malme II:1989 (lectotype, designated by Fryxell 2002: S!). Callianthe mexiae (R. E. Fries) Donnell, comb. nov. Abutilon mexiae R. E. Fries, Kongl. Svenska Vetensk. Acad. Handl. 24(2): 7 (1947).–TYPE: BRAZIL. Minas Gerais, Viçosa, agricultural College lands, north-west hill, abandoned road up Chacka Valley, 4 July 1930, Mexia 4838 (holotype: S!; isotypes: CAS!, PH!, UC,US!, WIS). Callianthe monteiroi (Krapov.) Donnell, comb. nov. Abutilon monteiroi Krapov., Hickenia 1(51): 273 (1982).–TYPE: BRAZIL. Minas Gerais, Serra do Espinhaço, 11 February 1972,W.A.Andersonetal.35793(holotype:UB;isotype:NY!). Callianthe mouraei (K. Schum.) Donnell, comb. nov. Abutilon mouraei K. Schum., Fl. Bras. (Martius) 12(3): 410 (1891). –TYPE: BRAZIL. Near Rio de Janeiro, 1882, Glaziou 13542 (lectotype, designated by Fryxell 2002: P; isolectotypes: C, K!, P). Callianthe muelleri-friderici (Gürke & K. Schum.) Donnell, comb. nov. Abutilon muelleri-friderici Gürke & K. Schum., Fl. Bras. (Martius) 12(3): 410 (1891).–TYPE: BRAZIL. Santa Catarina, prope Blumenau, 1881, J. H. Schenck 497 (holotype: B!). Callianthe pachecoana (Standl. & Steyerm.) Donnell, comb. nov. Abutilon pachecoanum Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23(2): 61 (1944).–TYPE: GUATEMALA. Quezaltenango, region of Las Nubes, 720 SYSTEMATIC BOTANY south of San Martı́n Chile Verde, January 1941, P. C. Standley 83528 (lectotype, designated by Fryxell 2002: F!; isolectotypes: EAP!, F!-2, NY!, US!). Callianthe petiolaris (Kunth) Donnell, comb. nov. Abutilon petiolare Kunth, Nov. Gen. Sp. 5: 272 (1822).–TYPE: COLOMBIA. Humboldt & Bonpland s. n. (holotype: P; isotype: P!). Callianthe pickellii (Monteiro) Donnell, comb. nov. Bakeridesia pickellii Monteiro, Anais. Soc. Bot. Brasil 23: 116 (1973).–TYPE: BRAZIL. Pernambuco, Tapera, São Bento, 18 August 1923, B. J. Pickel 150 (holotype: IPA!). Callianthe picta (Gilles ex Hook. & Arn.) Donnell, comb. nov. Sida picta Gilles ex Hook. & Arn., Bot. Misc. 3: 154 (1833). Abutilon pictum (Gilles ex Hook. & Arn.) Walp., Repert. Bot. Syst. 1: 324 (1824).–TYPE: ARGENTINA. Buenos Aires, J. Gilles, s. n. (lectotype, designated by Fryxell 2002: K!; isolectotype: OXF). Callianthe purpusii (Standl.) Donnell, comb. nov. Abutilon purpusii Standl. Contr. U.S. Natl. Herb. 23: 750 (1923).– TYPE: MEXICO. Veracruz, Zacuapan, Barranca de Tenampa, January 1910, C. A. Purpus 4332 (holotype: US!; isotypes: BM!, F!, GH, MO!). Callianthe regnellii (Miq.) Donnell, comb. nov. Abutilon regnellii Miq., Linnaea 22: 554 (1849).–TYPE: BRAZIL. Minas Gerais, Caldas, A. F. Regnell II-17 (holotype: U; isotypes: F!, NY!). Callianthe rufinerva (A. St.-Hil.) Donnell, comb. nov. Abutilon rufinerve A. St.-Hil., Fl. Bras. Merid. 1: 205 (1827). Bakeridesia rufinerva (A. St.-Hil.) Monteiro, Bol. Soc. Portug. Ci. Nat. 5(2): 124 (1955).–TYPE: BRAZIL. Villa do Principe, propéque praedium Domingos Alfonso, A. St.-Hil. s. n. (holotype: P; isotypes: P, MPU!-2). Callianthe rufivela (Hochr.) Donnell, comb. nov. Bakeridesia rufivela Hochr., Annuaire Conserv. Jard. Bot. Geneve 21: 419 (1920). Abutilon rufivelum K. Schum. ex. Baker f., J. Bot. 31: 271 (1893), nom. nud..–TYPE: BRAZIL. Rio de Janeiro, Nova Friburgo, Alto Macahé, 1891, Glaziou 18136 (holotype: G-DC; isotypes: B!, F!, K!). Callianthe scabrida (K. Schum.) Donnell, comb. nov. Abutilon scabridum K. Schum., Fl. Bras. (Martius) 12(3): 413 (1891). Bakeridesia scabrida Kearney ex Fryxell., Lundellia 5: 106 (2002).–TYPE: BRAZIL. Prope Estiva ut lego, Sellow 744 (syntype: B!). Callianthe schenckii (K. Schum.) Donnell, comb. nov. Abutilon schenckii K. Schum., Fl. Bras. (Martius) 12(3): 412 (1891).–TYPE: BRAZIL. Madre de Dios, June 1833, L. Riedel 1316 (lectotype, designated by Fryxell 2002: P; isolectotype: K!). Callianthe sellowiana (Klotzsch) Donnell, comb. nov. Sida sellowiana Klotzsch, Allg. Gartenzeitung (Otto & Dietrich) 4: 9 (1836). Synonyms: Abutilon sellowianum (Klotzsch) Regel., Ann. Sci. Nat., Bot. sér. 4, 12: 379 (1859); Bakeridesia sellowiana (Klotzsch) Monteiro, Bol. Soc. Portug. Ci. Nat. 5(2): 127 (1955).–TYPE: Hort. Bot. Berlin, ex Brazil. Sellow s. n. (holotype: B!). Callianthe senilis (K. Schum.) Donnell, comb. nov. Abutilon senile K. Schum., Fl. Bras. (Martius) 12(3): 424 (1891). Synonyms: Bakeridesia senilis (K. Schum.) Hochr., Annuaire Conserv. Jard. Bot. Geneve 21: 421 (1920); Abutilon quinquelobum Ulbr., Repert. Nov. Spec. Regni [Volume 37 Veg. 13: 501 (1915), nom. illegit (superfluous, Art. 52.1).–TYPE: BRAZIL. Rio de Janeiro, February 1882, A. F. M. Glaziou 12438 (holotype: B!; isotypes: C, G-DC, K!, P.). Callianthe striata (Dicks. ex Lindl.) Donnell, comb. nov. Abutilon striatum Dicks. ex Lindl., Edward’s Bot. Reg. 25: misc 39 (1839).–TYPE: BRAZIL (holotype: Dickson in Maund, The Botanist 3: plate 144, 1839). Callianthe torrendii (Monteiro) Donnell, comb. nov. Bakeridesia torrendii Monteiro, Anais. Soc. Bot. Brasil 23: 125 (1973).– TYPE: BRAZIL. Bahı́a, Vitória da Conquista, C. Torrend 5 (holotype: RBR!). Callianthe tridens (Standl. & Steyerm.) Donnell, comb. nov. Abutilon tridens Standl. & Steyerm., Pub. Field. Mus. Nat. Hist., Bot. Ser. 23: 173 (1944).–TYPE: GUATEMALA. El Progreso, hills between Finca Piamonte and the slopes southeast, Sierra de las Minas, 4 February 1942, J. A. Steyermark 43439 (lectotype, designated by Fryxell 1988: F!; isolectotypes– F!, US!). Callianthe vexillaris (E. Morren) Donnell, comb. nov. Abutilon vexillarum E. Morren, Belgique Hort. 14: 289 (1864).– TYPE: SOUTH AMERICA (holotype: loc. cit., plate 16). The following names represent species that will likely be transferred to Callianthe once more source information is available and/or nomenclatural issues are resolved: Abutilon appendiculatum K. Schum., Abutilon arboreum Sweet., Abutilon carneum A. St.-Hil., Abutilon carinatus Krapov., Abutilon dianthum C. Presl, Abutilon esculentum A. St.-Hil., Abutilon falcatum A. St.-Hil & Naud., Abutilon macrocarpum St. - Hil. & Naud., Abutilon megapotamicum (A. Spreng.) St.-Hil. & Naud., Abutilon montanum A. St.-Hil., Abutilon nigricans G. L. Esteves & Krapov., Abutilon niveum Gris., Abutilon pauciflorum A. St.-Hil., Abutilon pedrae-brancae K. Schum., Abutilon peltatum K. Schum., Abutilon peruvianum (Lam.) Kearney, Abutilon piurense Ulbr., Abutilon sordidum K. Schum., Abutilon sylvaticum (Cav.) K. Schum., Abutilon weberbaueri Ulbr., Bakeridesia purpurascens (Link.) Monteiro. Acknowledgments. There are many people without whom this research would not have been possible. The first author thanks Greg Wahlert and Daryl Lam for their guidance in labwork protocols; Tara Killen and Vijay Nadella for invaluable assistance with sequencing; Melanie Schori for guidance in DNA extraction techniques and help with botanical Latin; my dissertation committee members, Gar Rothwell and Alycia Stigall for their helpful comments on this manuscript; Ricardo Madrigal and Hector Gómez for their incredible assistance with specimen collection in Mexico; Sergio Avendaño and Dr. Gonzalo Castillo of the Instituto de Ecologı́a in Xalapa, Veracruz for hosting me and allowing me to use their supplies for field collection; Dr. Miguel Angel Perez-Ferrara for helping me arrange my collecting time in Chiapas; Dr. Jennifer Tate and the late Dr. Paul Fryxell for helping me explore options for Malvaceae research; and the Malvaceae yahoo group members (especially Stewart R. Hinsley). 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Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. Pp. 315– 322 in PCR protocols: A guide to methods and applications. eds. M. Innis, D. Gelfand, J. Sninsky, and T. White. San Diego: Academic Press. Zwickl, D. J. 2006. Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph. D. dissertation. Austin: The University of Texas. Appendix 1. GenBank accession numbers and voucher information for taxa included in the present study, arranged in the following order: taxon, country, state or department, collection locality, voucher collection (herbarium): GenBank accession number(s). Taxa with only one GenBank accession number correspond to a complete ITS sequence (ITS1 + 5.8S + ITS2); others were amplified or downloaded as ITS1 and ITS2 separately and are indicated as such. Voucher information is given only for sequences newly generated in this study; these novel sequences are indicated by bold GenBank accession numbers. Abutilon abutiloides ( Jacq.) Garcke ex Hochr., Mexico, Nayarit, G. FloresFranco et al. 2966 (MO): JQ753294; Abutilon amoenum K. Schum., Brazil, Paraná, União da Vitoria, A. Krapovickas & C. L. Cristóbal 39596 (MO): JQ753306; Abutilon andrewsianum W. Fitzg.: AY591807; Abutilon costicalyx K. Schum., Brazil, São Paulo, Caraguatatuba, E. L. Esteves et al. 2760 (MO): JQ753295; Abutilon dianthum C. Presl., Bolivia, La Paz, Franz Tamayo, Cayola et al. 890 (MO): JQ753291; Abutilon dianthum C. Presl., Bolivia, La Paz, Franz Tamayo, A. Fuentes & R. Alvares 4810 (MO): JQ753292; Abutilon eremitopetalum Caum: EF219363; Abutilon geminiflorum Kunth, Venezuela, Aragua, Distrito Ricaurte, W. Meier et al. 5892 (US): JQ753293; A. grandifolium (Willd.) Sweet: EF219369; Abutilon x hybridum hort. ex Voss, cultivated origin, A. Donnell 132 (NY): JQ753296; Abutilon incanum (Link) 722 SYSTEMATIC BOTANY Sweet: EF219368; Abutilon indicum (L.) Sweet: AY863059; Abutilon malacum S. Watson, Mexico, Sonora, Sonoyta, R. S. Felger 92-81 (MO): JQ753298; Abutilon menziesii Seem.: EF219365; Abutilon monteiroi Krapov. Brazil, Minas Gerais, Grão Mogol, G. M. Hatschbach 54157 (MO): JQ753299; Abutilon muelleri-friderici Gürke & K. Schum., Brazil, Rio Grande do Sul, Caxias do Sul, L. Scur 828 (MO): JQ753300; Abutilon pauciflorum A. St.-Hil., Bolivia, Cochabamba, Carrasco, E. Fernández et al. 35226 (MO): JQ753259; Abutilon peruvianum (Lam.) Kearney, Peru, Cusco, La Convención, L. Valenzuela et al. 69 (MO): JQ753261; Abutilon peruvianum (Lam.) Kearney, Peru, Contumazá, Cajamarca, A. Sagástegui A. 14878 (MO): JQ753262; Abutilon peruvianum (Lam.) Kearney, Bolivia, Santa Cruz, Manuel M. Caballero, M. Muñoz 165 (MO): JQ753263; Abutilon pubistamineum Ulbr.: AJ251049 (ITS1), AJ274993 (ITS2); Abutilon purpusii Standl., Costa Rica, San José, Cantón de Acosta, J. F. Morales 2343 (MO): JQ753301; Abutilon regnellii Miq., Brazil, São Paulo, Serra da Mantiqueira, J. R. Piranı́ 2510 (SPF): JQ753302; Abutilon sandwicense (O. Deg.) Christoph.: EF219367; Abutilon sp., Argentina, Misiones, San Pedro, N. B. Deginani et al. 1662 (MO): JQ753260; Abutilon sp. 1: AY591808; Abutilon sp. 2, Paraguay, Central, San Antonio, A. Krapovickas & C. L. Cristóbal 44548 (MO): JQ753307 (ITS1), JQ753308 (ITS2); Abutilon theophrasti Medik. (1): DQ287984; Abutilon theophrasti Medik. (2): DQ006017; Alcea rosea L.: EF679714; Andeimalva mandonii (Baker f.) J. A. Tate: AY172220; Anoda crenatiflora Ortega: AJ251043 (ITS1), AJ274987 (ITS2); Bakeridesia amoena Fryxell, Mexico, Chiapas, Cintalapa, A. Donnell 113 w/ R. Madrigal-Chavero (NY): JQ753283; Bakeridesia bakeriana (Rose) D. M. Bates, Mexico, Jalisco, La Huerta, E. J. Lott 4029 (MICH): JQ753275; Bakeridesia cf. gloriosa, Mexico, Oaxaca, Santa Maria Chimalapa, A. Donnell 111 w/ R. Madrigal-Chavero (NY): JQ753279; Bakeridesia cf. integerrima, Mexico, Oaxaca, Santo Domingo Tehuantepec, M. L. Torres 855 (NY): JQ753271; Bakeridesia cf. pittieri, Mexico, Chiapas, Amatenango de la Frontera, A. Donnell 88 w/ H. Gomez-Dominguez (NY): JQ753282; Bakeridesia esculenta (A. St.-Hil.), Monteiro, Brazil, Rio de Janeiro, São Pedro de Aldeia, J. A. Kallunki & J. R. Pirani 633 (NY): JQ753264; Bakeridesia exalata D. M. Bates: AY591812; Bakeridesia exalata D. M. Bates, Honduras, El Paraı́so, A. Molina R. 31253 (MO): JQ753285; Bakeridesia ferruginea (Martyn) Krapov., Mexico, Veracruz, Tlaltetela, F. Ventura A. 15735 (NY): JQ753287; Bakeridesia ferruginea (Martyn) Krapov., Mexico, Oaxaca, Temascal, L. Cortes 725 (MO): JQ753265; Bakeridesia gaumeri (Standl.) D. M. Bates: AY591859; Bakeridesia gloriosa D. M. Bates, Mexico, Chiapas, Tuxtla Gutierrez, A. Donnell 83 w/ H. Gomez-Dominguez (NY): JQ753288; Bakeridesia integerrima (Hook. f.) D. M. Bates, Mexico, Oaxaca, Asunción Ixaltepec, E. A. Pérez-Garcı́a 2046 (MO): JQ753270; Bakeridesia integerrima (Hook. f.) D. M. Bates, Venezuela, Falcon, Distrito Bolivar, van der Werff & R. Wingfield 7433 (MO): JQ753273; Bakeridesia integerrima (Hook. f.) D. M. Bates, Mexico, Veracruz, Ozuluama de Mascareñas, A. Donnell 72 w/ R. Madrigal-Chavero (NY): JQ753272; Bakeridesia macrantha (A. St.-Hil.) Leite & Monteiro, Brazil, Rio Grande do Sul, São Francisco de Paula-Linha, R. Wasum 178 (US): JQ753297; Bakeridesia molinae D. M. Bates, Honduras, Yoro, Olanchito, Davidse et al. 35493 (TEX): JQ753277; Bakeridesia nelsonii (Rose) D. M. Bates, Mexico, Chiapas, La Concordia, A. Donnell 100 w/ H. Gomez-Dominguez (NY): JQ753278; Bakeridesia nelsonii (Rose) D. M. Bates, Guatemala, Huehuetenango, “El Tapon”, J. Castillo 1743 [Volume 37 (MO): JQ753266; Bakeridesia notolophium (A. Gray) Hochr., Mexico, Veracruz, Tancoco, M. Nee & K. Taylor 28735 (NY): JQ753286; Bakeridesia pickellii Monteiro, Brazil, Paraı́ba, Maturéia, M. F. Agra et al. 4913 (MO): JQ753303; Bakeridesia pittieri (Donn. Sm.) D. M. Bates, Mexico, Chiapas, Ocozocoautla, A. Donnell 94 w/ H. Gomez-Dominguez (NY): JQ753276; Bakeridesia pittieri (Donn. Sm.) D. M. Bates, Mexico, Chiapas, Union Juárez, D. E. Breedlove & Smith 31560 (NY): JQ753303; Bakeridesia rufinerva (A. St.Hil.) Monteiro, Brazil, Paraná, Guaraqueçaba, G. Gatti 450 (NY): JQ753267; Bakeridesia rufivela Hochr., Brazil, Espirito Santo, Feixe de Pedras, M. M. Arbo et al. 5587 (TEX): JQ753304; Bakeridesia scabrida (K. Schum.) Kearney ex Fryxell, Brazil, Rio de Janeiro, Valença, A. M. Amorim 3075 (MO): JQ753305; Bakeridesia sp. nov. 1a, Mexico, Veracruz, Vega de Alatorre, A. Donnell 78 w/ R. Madrigal-Chavero (NY): JQ753290; Bakeridesia sp. nov. 1b, Mexico, Veracruz, Alto Lucero, A. Donnell 105 w/ R. Madrigal-Chavero (NY): JQ753268; Bakeridesia sp. nov. 2a, Mexico, Oaxaca, Santo Domingo Tehuantepec, T. MacDougall s. n. (NY): JQ753281; Bakeridesia sp. nov. 2b, Mexico, Oaxaca, Santo Domingo Tehuantepec, A. Donnell 112 w/ R. MadrigalChavero (NY): JQ753280; Bakeridesia subcordata (Hochr.) D. M. Bates, Mexico, Oaxaca, San Juan Bautista Cuicatlan, A. Donnell 119 w/ R. MadrigalChavero (NY): JQ753289; Bakeridesia vulcanicola (Standl.) D. M. Bates, El Salvador, Santa Ana, Parque Nacional Montecristo, J. Monterrosa 436 (US): JQ753284; Bakeridesia yucatana (Standl.) D. M. Bates, Mexico, Quintana Roo, Akumal, E. Cabrera 4016 (MO): JQ753269; Bastardia bivalvis (Cav.) Kunth ex Griseb.: AY591813; Bastardiastrum cinctum (Brandegee) D. M. Bates: AY591814; Bastardiopsis densiflora (Hook. & Arn.) Hassl.: AY591815; Billieturnera helleri (Rose ex A. Heller) Fryxell: AY591817; Briquetia sonorae Fryxell: AY591818; Callirhoe involucrata (Torr. & A. Gray) A. Gray: AY591819; Corynabutilon virifolium (Cav.) Kearney: AJ274970 (ITS1), AJ275001 (ITS2); Cristaria andicola Gay: AY591821; Dendrosida breedlovii Fryxell: AJ251032 (ITS1), AJ274976 (ITS2); Dirhamphis mexicana Fryxell: AY591822; Eremalche parryi (Greene) Greene: AJ304938; Gaya atiquipana Krapov.: AY591825; Gaya calyptrata (Cav.) Kunth ex K. Schum.: AJ251048 (ITS1), AJ274992 (ITS2); Hochreutinera amplexifolia (DC.) Fryxell: AY591827; Horsfordia exalata Fryxell: AY591831; Malvastrum americanum (L.) Torr.: AY591842; Malvella sagittifolia (A. Gray) Fryxell: AJ251045 (ITS1), AJ274989 (ITS2; Neobrittonia acerifolia (G. Don) Hochr.: AY591844; Palaua rhombifolia Graham: AY591846, Periptera punicea (Lag.) DC.: AY591847; Phymosia umbellata (Cav.) Kearney: AY591848; Plagianthus divaricatus J. R. Forst. & G. Forst.: AY591849; Pseudabutilon umbellatum (L.) Fryxell: AJ274964 (ITS1), AJ274995 (ITS2); Robinsonella lindeniana (Turcz.) Rose & Baker f.: AY591851; Sida abutifolia Mill.: AJ274961 (ITS1), AJ251617 (ITS2); Sida aggregata C. Presl.: AJ274943 (ITS1), AJ251599 (ITS2); Sida cordifolia L.: AJ274945 (ITS1), AJ251601 (ITS2); Sida glutinosa Comm. ex Cav.: AJ251037 (ITS1), AJ274981 (ITS2); Sida hookeriana Miq.: AJ274967 (ITS1), AJ274998 (ITS2); Sida platycalyx F. Muell. ex Benth.: AJ251041 (ITS1), AJ274985 (ITS2); Sida rhombifolia L.: AJ274953 (ITS1), AJ251609 (ITS2); Sidalcea hartwegii A. Gray ex Benth.: AJ304890; Sidalcea stipularis J. T. Howell & G. H. True: AJ304932; Sidastrum paniculatum (L.) Fryxell: AJ251040 (ITS1), AJ274984 (ITS2) ; Tarasa albertii Reiche: AY172200; Tarasa trisecta (Griseb.) Krapov.: AY172236; Tetrasida chachapoyensis (Baker f.) Fryxell & Fuertes: AY591854; Tetrasida weberbaueri (Ulbr.) Fryxell & Fuertes: AY591855; Wissadula boliviana R. E. Fr: AY591856.