Podostemaceae are a clade of aquatic flowering plants that form important components of tropical river ecosystems. Species in the family exhibit highly derived growth forms and high vegetative phenotypic plasticity, both of which contribute to taxonomic confusion. The backbone phylogeny of the family remains poorly resolved, many species remain to be included in a molecular phylogenetic analysis, and the monophyly of many taxa remains to be tested. To address these issues, we assembled sequence data for 73 protein-coding plastid genes from 132 samples representing 68 species (∼23% of described species) that span the breadth of most major taxonomic, morphological, and biogeographic groups of Podostemaceae. With these data, we conducted the first plastid phylogenomic analysis of the family with broad taxon sampling. These analyses resolved most nodes with high support, including relationships not recovered in previous analyses. No evidence of widespread, well-supported conflict among individual plastid genes and the concatenated phylogeny was observed. We present new evidence that four genera (Apinagia, Marathrum, Oserya, and Podostemum), as well as four species, are not monophyletic. In particular, we show that Podostemum flagelliforme should not be included in Podostemum and is better recognized as Devillea flagelliformis, and that Marathrum capillaceum is embedded within Lophogyne s.l. and should be recognized as Lophogyne capillacea. We also place a previously unsampled and undescribed species that likely represents a new genus. In contrast to previous studies, the neotropical genera Diamantina, Ceratolacis, Cipoia, and Podostemum are resolved as successive sister groups to a clade of all paleotropical Podostemoideae taxa sampled, suggesting a single dispersal event from the neotropics to the paleotropics in the history of the subfamily. These results provide a strong basis for improving the classification of Podostemaceae and a framework for future phylogenomic studies of the clade employing data from the nuclear genome.
Podostemaceae, the riverweed family, are a clade of flowering plants in the order Malpighiales (Koi et al. 2012; Xi et al. 2012; Ruhfel et al. 2016; Katayama et al. 2022). They are the largest strictly aquatic flowering plant family, with 51–54 genera and ∼300 species (Ruhfel et al. 2011; Koi et al. 2012; Katayama et al. 2022; Bove et al. 2023; POWO 2023) and are circumscribed into three subfamilies: Podostemoideae, Tristichoideae, and Weddellinoideae. Each subfamily is well-supported by molecular and morphological evidence (Engler 1930; Cook and Rutishauser 2007; Ruhfel et al. 2011; Koi et al. 2012, 2022). Tristichoideae are characterized by tricarpellate ovaries and pantoporate pollen, in contrast to Podostemoideae and Weddellinoideae, which have bicarpellate ovaries and mostly tricolporate or tricolpate pollen. Podostemoideae are characterized by the presence of a spathella that encloses the flower bud prior to anthesis. Weddellinoideae differs from Podostemoideae by the absence of a spathella and the presence of a distinct perianth, which are likely plesiomorphic character states shared with Tristichoideae (Kita and Kato 2001; Cook and Rutishauser 2007; Ruhfel et al. 2011). Many genera in the family are monotypic (42%; Katayama et al. 2022).
The distribution of the family is primarily pantropical with a few species occurring in temperate areas of North America and Japan (Ruhfel et al. 2016). Species of Podostemaceae attach to solid substrates in swift flowing currents of river-rapids and waterfalls. These are unusual habitats for angiosperms and form crucial elements of river biodiversity and ecology. Dense beds of riverweeds are dominant primary producers and important contributors to making river-rapids the most productive regions of rivers (Hynes 1970; Horn and Goldman 1994) while providing food and habitat for fish (Jégu et al. 2002; Chernoff et al. 2003) and invertebrates (Giller and Malmqvist 2002; Giller et al. 2004; Hutchens et al. 2004). Indeed, Podostemaceae are the dominant macrophyte in large stretches of tropical rivers and are often considered to be foundational species for river-rapid ecosystems (sensu Dayton 1972; Ellison et al. 2005; Wood and Freeman 2017). Currently, the construction of hydroelectric dams and the associated massive reservoirs is destroying river-rapid habitat along hundreds of kilometers of rivers at a time when many species of Podostemaceae are of conservation concern (Philbrick et al. 2010; Cheek et al. 2017, 2022; Bidault et al. 2023).
The vegetative plant body of Podostemaceae species is atypical compared to other angiosperms and interpretations of structural homology vary considerably (Jäger-Zürn 1997, 2005; Rutishauser 1997, 2015; Stevens 2001; Cook and Rutishauser 2007; Koi and Kato 2007, 2010; Sehgal et al. 2007). It has been suggested that “conventional morphology cannot cope with the family” (Stevens 2001) and that Podostemaceae be recognized in their own class, equal in rank to monocots and dicots (Cusset and Cusset 1988). Difficulty interpreting the various components of the plant body may be in part due to the presence of organs of mixed shoot-leaf identity in species of Podostemoideae (Katayama et al. 2010; Rutishauser 2020). Another example of this difficulty is evident in an investigation by Jäger-Zürn et al. (2016) where they reinterpreted pinnately compound leaves as foliate roots. These highly derived growth forms along with vegetative phenotypic plasticity exhibited by species in the family (Rutishauser et al. 1999; Philbrick et al. 2010; Jäger-Zürn et al. 2016) contribute to taxonomic confusion.
Functional taxonomies do not exist for many genera of Podostemaceae, and their absence hinders meaningful assessment of species distributions and endemism important for conservation efforts (Philbrick et al. 2010; Bidault et al. 2023). For example, although van Royen's taxonomic monograph of neotropical species (Van Royen 1951, 1953, 1954) has been important to our understanding of some genera of these plants, his treatments of the largest neotropical genera, including Apinagia, are largely unworkable. Indeed, the most basic of taxonomic questions, ‘What species does this specimen represent?’ often remains difficult to answer, especially for members of Podostemoideae (Berry 2004; Philbrick et al. 2010; Bidault et al. 2023). This problem is also evident elsewhere in the literature. For example, 75% of accessions of two genera (Apinagia s.l. and Rhyncholacis) in an important molecular phylogenetic analysis of the family (Koi et al. 2012) were not identified to species. More than a decade later, taxonomic treatments of these genera remain absent, and researchers would not have greater success identifying these specimens today.
Understanding of the evolutionary relationships in Podostemaceae has improved considerably over the last several years due to molecular phylogenetic studies. Well-sampled phylogenies have relied primarily on plastid data, and particularly the matK region (e.g. Koi et al. 2012, 2019; Werukamkul et al. 2018; Wu et al. 2022). Other studies have utilized multiple genes, combining plastid data with the nuclear marker ITS (Tippery et al. 2011; Da Costa et al. 2018; Koi et al. 2022) or the mitochondrial gene matR (Ruhfel et al. 2011, 2016; Sun et al. 2016). Phylogenies based on plastid genome (plastome) datasets of Podostemaceae are becoming available as more plastomes are sequenced (Bedoya et al. 2019, 2020; Jin et al. 2020; Trad et al. 2021). However, to date, these plastome phylogenies of Podostemaceae have had limited taxonomic sampling. Furthermore, conflict between individual plastid gene trees and trees constructed using concatenated plastome data remains relatively unexplored at the family level (but see Trad et al. 2021 who included three Podostemaceae taxa in their analyses) and can be important to examine when inferring plastid phylogenies (Gonçalves et al. 2019; Walker et al. 2019). A single study using a large nuclear data set has been published (Katayama et al. 2022), but as in previous plastome-level studies, its taxon sampling is limited (< 10 taxa) and thus comparison to studies with more taxon-rich phylogenetic analyses is difficult. Hybridization has been reported in the family (Cheek et al. 2017; Bedoya et al. 2021) but it remains unclear how widespread this may be and if it impacts phylogenetic reconstruction.
Despite recent advances, several key areas of the Podostemaceae phylogeny remain unresolved, particularly in Podostemoideae, and many species remain to be sampled with molecular data. These issues inhibit our ability to assess their classification, elucidate patterns of character evolution, and infer their biogeography. For example, placement of the monotypic Diamantina, a neotropical lineage in the subfamily, remains unclear and is weakly supported in all previous studies. Diamantina has been placed either as sister to all other Podostemoideae (Ruhfel et al. 2011; Koi et al. 2012; Sun et al. 2016), as sister to a clade containing the neotropical genera Ceratolacis, Cipoia, and Podostemum (Da Costa et al. 2018), or as sister to a strictly neotropical subclade (Ruhfel et al. 2016) that did not contain Ceratolacis, Cipoia, and Podostemum. Similarly, support remains weak for the relationships among several strongly-supported subclades, each of which are restricted to one of three main geographic areas (North and South America, Asia/Australia, and Africa/Madagascar; Koi et al. 2012; Ruhfel et al. 2016). Non-monophyletic genera are also common with many in need of further taxonomic adjustments and monographic work (e.g. Apinagia s.l., Ledermanniella s.l., Marathrum, Zeylanidium; Philbrick et al. 2010, 2016, 2018; Koi et al. 2012, 2019; Schenk et al. 2015), and the circumscription of other genera remains to be tested with molecular data. For example, the monophyly of Podostemum sensu Philbrick and Novelo (2004) remains to be tested with molecular data, particularly regarding the inclusion of Podostemum flagelliforme (Tul. & Wedd.) C.T.Philbrick & Novelo, which was previously placed in the monotypic genus Devillea. This species was included in Podostemum based on phylogenetic analysis of morphological data, but has characteristics not seen in other members of the genus (Philbrick and Novelo 2004). Remaining questions such as these can be addressed with increased taxon and character sampling and thus a well-sampled plastid phylogenomic analysis may offer further insight.
In this study, we analyzed 73 coding regions of the plastome to conduct the first taxonomically well-sampled plastid phylogenomic analysis of Podostemaceae. The data analyzed comprised 132 samples, including eight publicly available plastomes and 124 samples newly sequenced for this study, obtained using a genome skimming approach. Our main goals were to 1) resolve backbone relationships within the family, 2) test the monophyly of several genera and species, and 3) examine conflict between individual plastid gene trees and a concatenated dataset of plastid coding regions. The results presented here allow for a better understanding of generic and species boundaries in Podostemaceae, provide evidence to support nomenclatural and taxonomic changes, and highlight areas of the phylogeny that need better sampling. The insights provided regarding the evolutionary history of the plastome in Podostemaceae will complement future phylogenomic studies of the clade using data from the nuclear genome.
Materials and Methods
Taxon Sampling—Our sampling comprised 132 accessions including outgroups. Voucher information and GenBank numbers are provided in Appendix 1. Of these, 130 were Podostemaceae representing the three subfamilies, 31 genera, and 68 species (∼23% of the described species diversity in the family; Ruhfel et al. 2011; Koi et al. 2012; Katayama et al. 2022; POWO 2023). Also included was one specimen that could not be placed in a recognized genus (Philbrick et al. 6055, [MICH]). All new specimens gathered for this study were collected legally with appropriate permits and permissions, where required. Following Tippery et al. (2011), Apinagia nana was retained as a distinct species, although Van Royen (1951) placed it in synonymy with A. pilgeri Mildbr. Apinagia pilgeri has since been transferred to Oserya as O. pilgeri by Philbrick et al. (2016). The name Apinagia riedelii (Bong.) Tul. has been widely used in the literature since 1849, but this species is more correctly referred to as A. fucoides (Bove and Philbrick 2016; Philbrick et al. 2016). A proposal to conserve the more commonly used name (A. riedelii) was put forward (Bove and Philbrick 2016) but rejected (Wilson 2022). Thus, we use the name Apinagia fucoides here. This issue is particularly important to consider when comparing the results of this study to previous works that used the name A. riedelii (e.g. Tippery et al. 2011; Koi et al. 2012; Bove and Philbrick 2016 and references therein; Bedoya et al. 2019). Author citations for each name at the generic rank or below are listed in Appendix 1, if the name is not listed in the appendix, the authority is listed the first time the name is mentioned in the main text. Representatives of all major clades recovered in a densely sampled phylogenetic study using the plastid gene matK (clades A–M in Koi et al. 2012) are included here except Saxicolella p.p. (clade H in Koi et al. 2012; Pohliella Engl. sensu Cheek 2020). Sampling focused on neotropical taxa, and several species had more than one representative included to help test generic and species boundaries. Most genera not included in our sampling are from Africa/Madagascar (Angolaea Wedd., Endocaulos C.Cusset, Lebbiea Cheek, Leiothylax Warm., Letestuella G.Taylor, Paleodicraeia C.Cusset, Pohiella, Saxicolella s.s., Sphaerothylax Bisch. ex C.Krauss, Winklerella Engl., and Zehnderia C.Cusset) or Asia/Australia (Cussetia M. Kato, Farmeria Willis ex Hook.f., Griffithella (Tul.) Warm, Hanseniella C.Cusset, Hydrodiscus Koi & M.Kato, Indodalzellia Koi & M. Kato, Indotristicha P. Royen, Paradalzellia Koi, P.L.Uniyal & M.Kato, Thawatchaia M.Kato, Koi & Y.Kita, and Willisia Warm.). The only genus from the Americas not sampled was Wettsteiniola Suess., which may be closely related to Rhyncholacis (Da Costa et al. 2018). These missing genera were not sampled because specimens or tissues were not available, or extractions were unsuccessful. Many of the samples included here were sequenced for at most four gene regions in previous studies by the authors (Ruhfel et al. 2011, 2016; Tippery et al. 2011). Hypericaceae is the sister group to Podostemaceae (Koi et al. 2012; Xi et al. 2012; Ruhfel et al. 2016; Li et al. 2021; Katayama et al. 2022) and as such, Cratoxylum cochinchinense and Vismia guianensis were included as outgroups. Data from eight taxa were downloaded from GenBank including six species of Podostemaceae and the two outgroups (Xi et al. 2012; Bedoya et al. 2019; Jin et al. 2020).
DNA Extraction and Sequencing—Total genomic DNA (gDNA) was extracted from tissue from herbarium specimens or tissue preserved in silica gel or a saturated CTAB solution (Rogstad 1992) using the DNeasy Plant Mini Kit (Qiagen, Germantown, Maryland) following a modified protocol (Ruhfel et al. 2011). DNA quantification was performed with a Qubit Fluorometer 2.0 using a dsDNA BR Assay kit (Thermo Fisher, Waltham, Massachusetts) to ensure appropriate amounts of gDNA were available for high-throughput sequencing. DNA quantity and quality were also assessed using agarose gel electrophoresis. Samples of gDNA were submitted to Rapid Genomics (Gainesville, Florida) for library preparation and sequencing using a genome skimming approach (low-coverage shotgun-sequencing; Straub et al. 2012). Samples were sequenced either on the Illumina HiSeq 2500 Ultra-High-Throughput platform using single-end 100 bp reads (46 samples) or the Illumina HiSeq X platform (78 samples) using 150 bp paired-end reads (see Table S1).
Assembly of Plastid Exons—Demultiplexed reads were cleaned of adapters and trimmed using Trimmomatic 0.39 (Bolger et al. 2014) and quality checked with FastQC (Andrews 2010). During trimming, leading and trailing bases with a quality score less than 10 were removed and then reads were scanned with a 4-bp sliding window and cut when the average quality fell below 20. Reads less than 30 bp after cleaning were discarded.
Coding regions (exons only) from the plastid genome were assembled with HybPiper 1.3.1 (Johnson et al. 2016) using the Burrows-Wheeler alignment method (Li and Durbin 2009) to map reads to targets using the default coverage cutoff (–cov_cutoff = 8). The targets for gene assembly were plastid coding regions extracted from seven taxa of Podostemaceae available on GenBank: Apinagia fucoides, Lophogyne capillacea (syn. Marathrum capillaceum (Pulle) P.Royen), Lophogyne royenella (syn. Monostylis capillacea Tul.), Marathrum foeniculaceum, Marathrum utile, Podostemum ceratophyllum, and Tristicha trifaria (Xi et al. 2012; Bedoya et al. 2019; Jin et al. 2020). Paralogs were assessed using the method implemented in HybPiper which uses MAFFT (Katoh and Standley 2013) and FastTree (Price et al. 2009). Coding regions with an internal stop codon present were removed from further analysis and loci with fewer than 50% of the taxa present were not used in phylogenetic analyses.
Alignments and Phylogenetic Analyses—We aligned each gene with MUSCLE 3.8.425 (Edgar 2004) as implemented in Geneious 2022.2.1 (Bio-matters, Ltd., Auckland, New Zealand) using the default options. Alignments were then manually inspected and adjusted by hand to retain the correct reading frame based on plastomes of Podostemaceae published in Bedoya et al. (2019). Alignments were then trimmed of codons that were missing data in > 40% of the samples using pxclsq in Phyx (Brown et al. 2017).
All model testing, partition strategy selection, phylogenetic analyses, and branch support measurements were performed using IQ-TREE 1.6.12 (Nguyen et al. 2015). Maximum likelihood phylogenetic analyses of individual genes were conducted with standard model selection (-m TEST, Kalyaanamoorthy et al. 2017) using AICc followed by tree reconstruction with 20 independent runs and 1000 standard non-parametric bootstrap replicates. To analyze the full data set, we first concatenated the individual gene alignments and produced a partition file using pxcat in Phyx (Brown et al. 2017). Next, we determined the optimal partitioning scheme for the concatenated dataset with IQ-TREE and the “-m TESTMERGEONLY” option and reconstructed the phylogeny using the edge-linked proportional partition model (Chernomor et al. 2016) with 20 independent runs and 1000 standard non-parametric bootstrap replicates. To further examine branch support in the concatenated analyses we also calculated ultra-fast bootstrap support (UFBS; 1000 replicates, Hoang et al. 2017) and SH-aLRT support (1000 replicates, Guindon et al. 2010) using the optimized partition scheme and search strategy above. Each of these measures of support (BS, UFBS, and SH-aLRT) has its own strengths and weaknesses under various conditions (e.g. the presence of polytomies, model violations, and weak phylogenetic signal in the data; Hillis and Bull 1993; Guindon et al. 2010; Minh et al. 2013; Hoang et al. 2017). For this reason, we considered a branch to be well supported if the following three conditions were met: 1) BS value ≥ 80%; 2) SH-aLRT ≥ 80; and 3) UFBS value ≥ 95%.
Conflict Analysis—To examine conflict between individual gene trees and the concatenated phylogeny, we mapped the gene trees against the concatenated tree using the bipartition method implemented in PhyParts v. 0.0.1 (Smith et al. 2015) using either no support cutoff or a 70% BS support cutoff, representing moderate support (Hillis and Bull 1993). Prior to the PhyParts analysis, all trees were rooted on the outgroups using pxrr in Phyx (Brown et al. 2017). When no support cutoff is used, the node is evaluated as conflicting or concordant regardless of node support. When using a support cutoff, any node in the gene tree with support less than the value specified is considered uninformative. Using either approach, if a taxon is missing in the gene alignment, and is relevant to a particular node, that gene is considered uninformative for that node. PhyParts results were visualized using PhyPartsPieCharts (Johnson 2021).
Results
Data Set Characteristics—One hundred and twenty-four accessions were newly sequenced for this study. The number of reads obtained and the proportion of the target gene length recovered for each accession are shown in Table S1 and Fig. S1, respectively. The final concatenated alignment contained 73 coding gene regions, 132 samples, and was 53,943 bps in length with 3.71% missing data (gaps and completely undetermined characters). Details on each individual coding region alignment, the concatenated alignment, and their best fit models can be found in Table S2.
Phylogenetic Analyses—Our maximum likelihood analysis of the concatenated dataset (Fig. 1, summary phylogeny; Figs. 2A, B, full phylogeny; Fig. S2, full phylogeny with branch lengths) recovered strong support for the monophyly of the family as well as relationships among major clades, genera, and species of Podostemaceae. Of the 129 internal branches of the phylogeny, only nine received < 70 BS with all but one of these between accessions of the same species. Of the 120 branches that received ≥ 70 BS, only eight were not also well-supported using our other measures of support (≥ 80SH-aLRT; ≥ 95% UFBS). Because most branches in the phylogeny (112/129; ∼87%) were well-supported by all metrics (i.e. ≥ 80 BS, ≥ 80 SH-aLRT, and ≥ 95 UFBS), we limit our discussion of branch support to BS values for well-supported branches and specifically mention SH-aLRT and UFBS values only if one or more of the three measures of support disagree. See Fig. S3 for SH-aLRT and UFBoot values.
Fig. 1.
Maximum likelihood summary phylogeny of Podostemaceae inferred from a concatenated dataset of 73 protein-coding genes of the plastid genome. Standard bootstrap support (BS) values ≥ 50% are indicated above the branches. An asterisk (*) next to the BS value indicates that the branch was also supported by ≥ 95% ultra-fast bootstrap support and ≥ 80 SH-aLRT branch tests. Clades labeled with “A” or “B” represent major subclades of Podostemoideae containing either both neotropical and paleotropical taxa or strictly neotropical taxa, respectively. See Figs. 2A and 2B for the complete tree. Shaded squares next to each terminal indicate their broad biogeographic distribution. Subfamilies of Podostemaceae and outgroups are indicated to the right of the phylogeny. Wed. = Weddellinoideae.
Fig. 2A.
Maximum likelihood phylogeny of Podostemaceae inferred from a concatenated dataset of 73 protein-coding genes of the plastid genome. Voucher collection numbers are given after each species name. Standard bootstrap support (BS) values ≥ 50% are indicated above the branches. An asterisk (*) next to the BS value indicates that the branch was also supported by ≥ 95% ultra-fast bootstrap support and ≥ 80 SH-aLRT branch tests. The clade labeled with an “A” represents a major subclade of Podostemoideae containing both neotropical and paleotropical taxa. Subfamilies of Podostemaceae and outgroups are indicated to the right of the phylogeny. Out. = Outgroups; Tristich. = Tristichoideae; Wed. = Weddellinoideae. Phylogeny continues in Fig. 2B.
We recovered maximum support for the monophyly of the three subfamilies of Podostemaceae and the relationships among them. Tristichoideae was placed sister to Podostemoideae + Weddellinoideae. Within Tristichoideae, Terniopsis brevis and Dalzellia ceylanica were successive sister groups (100 BS) to a clade (100 BS) of three Tristicha trifaria samples. The two samples of Weddellina squamulosa formed a clade with 100 BS. Within Podostemoideae, Devillea flagelliformis (syn. Podostemum flagelliforme) was strongly supported (98 BS) as sister to the remaining taxa, which were separated into two major subclades. The first subclade of Podostemoideae (Fig. 1, clade A) was moderately supported (74 BS) by standard bootstrap methods but did not receive support from the other methods used (72.6 SH-aLRT, 77 UFBS) and is comprised of four neotropical genera (Ceratolacis, Cipoia, Diamantina, and Podostemum) and all sampled paleotropical taxa. The second major subclade of Podostemoideae (Fig. 1, clade B) was strictly comprised of neotropical taxa and received maximum support (100 BS).
In the first subclade, containing both neotropical and paleotropical Podostemoideae taxa (Fig. 1, clade A), neotropical Diamantina was sister to all other taxa which formed a strongly supported clade (100 BS). The neotropical genera Ceratolacis, Cipoia, and Podostemum (each monophyletic with 100 BS support), formed strongly supported (98 BS) successive sister groups to a strongly supported (96 BS) clade of all paleotropical taxa containing two subclades. The first, with Asian and Malagasy taxa (71 BS, not supported by other metrics: 68.6 SH-aLRT, 81 UFBS) and the second with African (100 BS) taxa. In the Asian/Malagasy clade, the Malagasy Thelethylax minutiflora was sister to a strongly supported (100 BS) clade of the Asian species. Here, Zeylanidium subulatum + Polypleurum stylosum (100 BS) was sister (71 BS, not supported by other metrics: 63 SH-aLRT, 83 UFBS) to a clade in which Hydrobryum japonicum was sister to Cladopus doianus + Paracladopus chanthaburiensis (100 BS). In the African clade, Monandriella linearifolia was sister to a poorly supported clade (58 BS) of the remaining African taxa. Among those, Inversodicraea bosii was sister to a strongly supported clade (100 BS) containing Stonesia ghoguei + Macropodiella heteromorpha (100 BS) which was sister to a clade (97 BS) containing Ledermanniella bowlingii, Djinga felicis, Ledermaniella letouzii, and Dicraeanthus zehnderi. Relationships within this subclade of African taxa were all strongly supported (≥ 99 BS).
The strictly neotropical subclade of Podostemoideae (Fig. 1, clade B) is represented here by Apinagia, Autana, Castelnavia, Lophogyne, Marathrum, Mourera, Noveloa, Oserya, Rhyncholacis, and an undescribed species that likely represents a new genus (Philbrick et al. 6055, [MICH]). Within this clade, Mourera, Lophogyne s.l. (including Marathrum capillaceum [syn. Lophogyne capillacea Pulle]), and Rhyncholacis were each strongly supported as monophyletic (100 BS) and successive sister groups to the remaining neotropical taxa. Within Lophogyne s.l., two species, L. fimbriata and L. royenella, were strongly supported (100 BS) as not monophyletic. Lophogyne fimbriata was paraphyletic, with one sample supported (100 BS) as more closely related to a clade containing L. wilsonii and two samples of an unidentified Lophogyne species (L. sp. 2). Similarly, samples of L. royenella form a paraphyletic grade with one sample being strongly placed as sister to L. ceratophylla + an unidentified Lophogyne species (L. sp. 1). Among the remaining taxa, a strongly supported clade (94 BS) containing Autana (100 BS) + Castelnavia (100 BS), was sister to a clade (100 BS) containing three lineages whose relationships were not well supported (70 BS, not supported by other metrics: 71 SH-aLRT, 88 UFBS). The first of these three groups was an undescribed species (‘Genus nov.’) that likely represents a new genus. The second clade received strong support from standard bootstrapping (94 BS) and the SH-aLRT test (97.3) but fell just below the cutoff for the UFBS values representing a well-supported branch (94 UFBS). This clade contained subclades representing Oserya s.s., Noveloa, Marathrum and a clade of Apinagia nana + Oserya pilgeri), thus making Apinagia and Oserya non-monophyletic. Each of these four clades was supported with 100 BS. Oserya perpusilla + Noveloa is strongly supported (92 BS) and was sister to Marathrum + (Apinagia nana + Oserya pilgeri) which received 100 BS. The third clade was strongly supported (100 BS) and represented Apinagia s.s. This clade contained samples from six species of Apinagia s.s., two of which were strongly supported (> 98 BS) as non-monophyletic. Apinagia longifolia was paraphyletic with regard to A. richardiana and A. fucoides was paraphyletic with A. tenuifolia + A. ‘crowii’ embedded within.
Fig. 2B.
Continuation of Fig. 2A. Maximum likelihood phylogeny of Podostemaceae inferred from a concatenated dataset of 73 protein-coding genes of the plastid genome. The clade labeled with a “B” represents a major subclade of Podostemoideae containing only neotropical taxa. Subfamily Podostemoideae, Apinagia s.s., and Oserya s.s. are indicated to the right of the phylogeny.
Conflict Analysis—When no support cutoff was used, we observed high levels of gene tree conflict throughout the phylogeny (Fig. S4). However, with the 70% BS cutoff (Fig. S5), the majority of individual genes were uninformative at most nodes and little conflict was evident. Of the 129 nodes of Podostemaceae in the 70% BS cutoff analysis (Fig. S5), 110 nodes (∼85%) had from one to 68 (mean = 14.6) concordant genes with BS support over 70%; 56 nodes (∼43%) had from one to 15 (mean = 0.9) conflicting genes with support over 70%; and 14 nodes (∼11%) had no conflicting or concordant genes. Of the 56 nodes with conflict present: 19 involved branches with near-zero lengths (Fig. S2, six nodes) and/or were within a clade representing a single species or closely related species (13 nodes). For example, within the clade of Oserya perpusilla, one internal node had five genes in conflict, and in the clade of Tristicha trifaria one internal node had 15 genes in conflict. Genes with the most conflict were rps7 (471 bps, 17 conflicts) and clpp (687 bps, 15 conflicts). Four conflicts were observed involving matK, with each of these present within a clade representing a single species.
Alignments, partition files, and tree files are available in the Dryad Data Repository (Ruhfel et al. 2024).
Discussion
Phylogenetic Relationships and Implications for Taxonomy, Nomenclature, Morphological Evolution, and Biogeography—The results presented here provide new insights on the evolutionary relationships of Podostemaceae using the largest plastome dataset for the family to date. We also provide the first examination of phylogenetic conflict of protein-coding genes within the plastome of the family. Previous plastome phylogenies of the family have included ≤ 5 species (Bedoya et al. 2019; Jin et al. 2020; Trad et al. 2021) while the most taxonomically well-sampled phylogeny of the clade (∼132 species representing ∼43 genera) was reconstructed using one plastid coding region (matK; Koi et al. 2012). In comparison to previous studies based on fewer genes and the current taxonomy of the group, the phylogeny presented here is mostly in agreement with notable exceptions discussed below. Our results strongly support the monophyly of the family as well as the three established subfamilies and agree with the relationships among them as recovered in previous analyses. However, we provide greater support for the backbone and relationships among genera, with the majority of the branches in the phylogeny being well supported, including many relationships that were unsupported in previous studies. In addition, our sampling includes several taxa that previously have not been included in molecular phylogenetic studies, which allows for important novel insights. We also show that four genera (Apinagia s.l., Marathrum, Oserya, and Podostemum) and four species are not monophyletic as currently circumscribed, and place an undescribed species that likely represents a new genus (Philbrick et al. 6055, [MICH]). Our phylogenetic results provide a strong foundation for reassessing the current classification of the family and set the stage for future monographic and molecular phylogenetic work. We discuss the most important implications of our results below.
Placement of Podostemum flagelliforme and the Non-Monophyly of Podostemum—Until the present study, the position of Podostemum flagelliforme (syn. Devillea flagelliformis) was untested with molecular data. Rediscovered in 2013 at a locality in the Balsas River in the Brazilian state of Tocantins approximately 120 km from the type locality, this species was previously known only from the type material collected over 175 yr ago from the Tocantins River also in the Brazilian state of Tocantins (Bove and Philbrick 2014). A phylogenetic analysis using morphological data (Philbrick and Novelo 2004) determined that the species belonged within Podostemum s.l. (including P. ceratophyllum, the type species of the genus [Van Royen 1954; Philbrick and Novelo 2004]), due to three homoplasious synapomorphies recovered in their analysis: an anisolobous ovary, a capsule at a 30–40° angle on the pedicel, and one deciduous capsule valve. However, the authors noted that it lacked morphological features (two stamens with an andropodium and pollen in dyads) shared by all other members of the genus Podostemum; Podostemum flagelliforme has one stamen per flower (thus no andropodium) and pollen in monads. Other treatments of the group (e.g. Moline et al. 2006; Cook and Rutishauser 2007) continued to recognize Devillea as a separate monotypic genus due to these differences in morphology. The strongly-supported placement of Podostemum flagelliforme as sister to all other Podostemoideae (Fig. 1; Fig. 2A as Devillea flagelliformis) indicates that Podostemum (sensu Philbrick and Novelo 2004) is not monophyletic and that Devillea should be recognized as a monotypic genus distantly related to Podostemum and that Podostemum flagelliforme is better recognized as Devillea flagelliformis.
Placement of Diamantina—Placement of Diamantina as sister to all other taxa in the mixed neotropical and paleotropical clade (Fig. 1, clade A) as recovered here agrees with the placement recovered in the molecular analysis of Da Costa et al. (2018) but contrasts with the results of previous phylogenetic analyses based on molecular (Ruhfel et al. 2011; Koi et al. 2012; Ruhfel et al. 2016; Sun et al. 2016) or morphological data (Philbrick and Novelo 2004; Philbrick et al. 2004). These studies variously placed Diamantina as sister to all other Podostemoideae, as sister to the strictly neotropical Podostemoideae clade (Fig. 1, clade B), or as a member of a paraphyletic grade of paleotropical taxa. Maximum likelihood bootstrap support for the placement presented here (74 BS) is higher than support for its placement in previous studies, however, our other two methods of support for this placement did not pass our threshold for being well supported (72.6 SH-aLRT; 77 UFBS). Morphological features of the pollen and silica bodies of Diamantina lend support to the phylogenetic position presented here and in Da Costa et al. (2018). Diamantina has previously been reported as the only genus within the family to produce pollen in tetrads (Philbrick et al. 2004; Rutishauser et al. 2005). However, those studies examined a limited number of specimens. Further examination of newly collected specimens, including one specimen included in this study (Bove et al. 2253) revealed that initial observations were based on underdeveloped anthers and that Diamantina regularly produces both tetrads and dyads (De Barros Marinho 2013; De Barros Marinho et al. 2014; C. T. Philbrick unpubl. data), a condition rare in angiosperms (Furness 2012). Most Podostemaceae in this mixed neotropical and paleotropical clade (Fig. 1, clade A) produce pollen in dyads, including all Asian, Australian, and many African taxa, as well as the neotropical genera Ceratolacis, Cipoia, and Podostemum (Philbrick et al. 2004; Cook and Rutishauser 2007). This suggests that dyad pollen may be a synapomorphy for this clade and has implications for reconstructing the evolutionary shift from monad to dyad pollen, or vice versa (De Barros Marinho et al. 2014) as some African taxa nested within this clade possess monad pollen (Thiv et al. 2009; Koi et al. 2012). The only known genus with dyad pollen that occurs outside of this clade is Lophogyne (Bove et al. 2011; De Barros Marinho 2013; De Barros Marinho et al. 2014) which suggests that dyad pollen has arisen independently at least twice in the family. Morphology of silica bodies found in the vegetative tissues of some Podostemaceae also supports the placement of Diamantina as a member of this mixed neotropical and paleotropical clade. Although only neotropical Podostemoideae taxa were included in the study by Da Costa et al. (2018), their recovered clade containing Ceratolacis, Cipoia, Diamantina, and Podostemum (here found to be the first several diverging lineages of the mixed neotropical and paleotropical clade [Fig. 1, clade A]) had a combination of silica body features not found elsewhere in neotropical Podostemoideae. These features include silica bodies present and located in the epidermis, undulate or perforated silica body morphology, and the presence of a lumen. An expanded investigation of silica bodies in the family, particularly including all paleotropical taxa (but see Ameka et al. 2002) and Devillea flagelliformis, is warranted.
In summary, the placement of Diamantina has not yet received strong support from phylogenetic analyses of molecular data. However, the shared morphological features of pollen and silica bodies with other taxa in the mixed neotropical and paleotropical clade (Fig. 1, clade A) provide additional evidence supporting the placement of Diamantina presented here. Nevertheless, a more comprehensive examination, including phylogenetic analysis of nuclear data and other morphological features, is warranted.
Relationships in the Mixed Neotropical and Paleotropical Clade—Relationships among the neotropical and paleotropical lineages within Podostemoideae have been unclear in previous studies. For example, in Koi et al. (2012) and Ruhfel et al. (2016), there were four paleotropical clades (2 African, 1 Malagasy, and 1 Asian/Australian), though relationships among those clades and the clades representing neotropical genera Ceratolacis, Cipoia (not included in Koi et al. 2012), and Podostemum were either weakly supported or in conflict with those presented here. In Ruhfel et al. (2016), Ceratolacis, Cipoia, and Podostemum were strongly supported (98 posterior probability) as members of a clade containing African and Malagasy taxa but excluding the Asian/Australian taxa. This conflict may be the result of Bayesian analyses providing inflated levels of support (Suzuki et al. 2002; Douady et al. 2003; Simmons et al. 2004) and/or issues with missing data (e.g. Endocaulos, Pohliella, and Thelethylax were each represented by only one of four genes included in that analysis). Here, support for the relationships among these clades is much improved. We found a strongly supported paleotropical clade embedded in a paraphyletic grade of neotropical taxa (Fig. 1, clade A; Fig. 2A, clade A). In addition to the neotropical Diamantina being placed as sister to all other taxa in this clade as discussed above, clades representing Ceratolacis, Cipoia, and Podostemum are strongly supported as successive sister groups to a strongly-supported clade containing all sampled paleotropical Podostemoideae taxa. Within that paleotropical clade, African and Asian/Australian taxa form strongly supported clades, and the Malagasy Thelethylax is moderately supported (71 BS) as sister to the Asian/ Australian taxa. This set of relationships would likely alter biogeographic reconstructions of ancestral areas in the early history of the family as compared to those reconstructed in Koi et al. (2015) and Ruhfel et al. (2016). Here, in addition to the placement of the neotropical Devillea as sister to all other Podostemoideae, the first several lineages of the mixed neotropical and paleotropical clade are neotropical, suggesting a single dispersal event from the neotropics to the paleotropics within Podostemoideae.
Before a formal reexamination of the biogeographic history of the family is conducted, genera missing in our sampling (see Materials and Methods) should be included in a plastid phylogenomic analysis. Particularly important to include are taxa that may help clarify the relationships between major subclades in the mixed neotropical and paleotropical clade. These taxa include two genera endemic to Africa (i.e. Lebbiea [Cheek and Lebbie 2018]; Pohliella [Cheek 2020]), one endemic to Madagascar (Endocaulos), and two that occur in both Africa and Madagascar (Paleodicraeia and Sphaerothylax, [Koi et al. 2012, R. Rutishauser unpubl. data]). Lebbiea is also important to include in a phylogenetic analysis as several aspects of its morphology are unique within the family (Cheek and Lebbie 2018). Though the Malagasy taxa Endocaulos and Thelethylax have been included in previous studies, Paleodicraeia and Sphaerothylax have never been included in a molecular analysis and thus it is not known if the four Podostemoideae genera that occur in Madagascar form a monophyletic group. In previous studies, a clade containing Endocaulos + Thelethylax has been variously placed as 1) sister to a clade of Asian and Australian taxa (Koi et al. 2012), 2) sister to a clade comprised of Cipoia, Ceratolacis, and Podostemum (Ruhfel et al. 2016), or 3) sister to other African species (Moline et al. 2006; Schenk et al. 2015). Finally, it is important to sample Pohliella in future plastid phylogenomic analyses, as Cheek (2020) stated that Cipoia and Pohliella are identical morphologically and recommended that they be included in a future molecular phylogenetic analysis to determine if taxa in these genera should be recognized as one amphi-Atlantic genus. However, initial investigations of the morphology of Cipoia and Pohliella by R. Rutishauser (unpubl. data) have revealed that the similarities between these two genera may be the result of shared plesiomorphic or convergent character states common in other Podostemoideae taxa such as thread-like roots, a single stamen per flower, presence of a gynophore, and bilocular ovaries (Ameka et al. 2002; Philbrick and Novelo 2004; Philbrick et al. 2004; Cook and Rutishauser 2007). Furthermore, Ruhfel et al. (2016) included samples representing both Cipoia and Pohliella (i.e. Pohliella amicorum [J.B.Hall] Cheek, syn. Saxicolella amicorum J.B. Hall) in their molecular phylogenetic analysis, the results of which suggest that Cipoia and Pohliella are not closely related. Given that the placement of Cipoia presented here conflicts with its position in Ruhfel et al. (2016), and that data for only one gene were available for Pohliella in that study, the relationship of Cipoia and Pohliella should be reexamined when additional plastome-level data are available. Other taxa missing in our sampling (e.g. Saxicolella s.s., Koi et al. 2012; Cheek et al. 2022) are likely embedded in more nested subclades of Asian or African taxa and while important to include in future studies, are less so for determining major relationships and biogeographic patterns in the family.
Relationships in the Strictly Neotropical Clade—Relationships within the strictly neotropical clade of Podostemoideae (Fig. 1, clade B; Fig. 2B, clade B) are much improved compared to previous studies, with most sampled, currently recognized genera strongly supported as monophyletic. Particularly with the nomenclatural changes suggested below, some of which include changes to be made in future publications, taxonomy at the genus level in the neotropical clade is moving closer to reflecting evolutionary history. In contrast, work below the genus level is needed to define species boundaries and relationships, particularly in the genera Apinagia s.s., the recently expanded Lophogyne s.l., and Rhyncholacis. For example, in Apinagia s.s. and Rhyncholacis initial investigations for monographic work (C. T. Philbrick unpubl. data) suggest that there are many fewer species than currently accepted names in these genera. This is likely because extreme variation in vegetative form has led to the over-description of species (Philbrick et al. 2010). This reduction in species number has occurred in previously published monographs of (e.g. Ceratolacis, Philbrick et al. 2009; Podostemum, Philbrick and Novelo 2004) and other nomenclatural adjustments (Philbrick et al. 2010, 2016).
Lophogyne s.l. and the Non-Monophyly of Marathrum—Our results support broadening the circumscription of Lophogyne as proposed by Philbrick and Bove (2019) and recover strongly-supported relationships within the clade. Along with the previously recognized Lophogyne species (Bove et al. 2011), Lophogyne s.l. now includes taxa that were formerly placed in the genera Apinagia, Jenmaniella Engl., Marathrum, and Monostylis Tul. Philbrick and Bove (2019) list four morphological characters that define Lophogyne s.l.: 1) the ovary is oriented vertically or obliquely at anthesis, 2) the flower projects from the ruptured spathella, 3) stamens and tepals occur on only one side of the ovary, and 4) silica bodies are absent from the leaves. For the first time, we present evidence that the South American species currently recognized as Marathrum capillaceum (Pulle) P.Royen is embedded within Lophogyne s.l. and should be recognized as Lophogyne capillacea Pulle as originally described (see Bove et al. 2019 regarding the illegitimate later homonym Lophogyne capillacea (Tul.) C.T.Philbrick & C.P.Bove). This species possesses the three floral features of Lophogyne s.l. given by Philbrick and Bove (2019). However, the presence of silica bodies needs further study. Schnell (1967) reports the presence of silica bodies in the petiole but does not mention their presence elsewhere in the leaf. Philbrick and Bove (2019) cite Da Costa et al. (2018) regarding the absence of silica bodies in the leaves of Lophogyne s.l. species, but L. capillacea was not included in that study. For those species they did examine, their methods state that they sampled “apical, medial and basal portions of fully developed leaves,” thus it is unclear if they sampled the same portion of the leaf that Schnell (1967) refers to as the petiole. Initial investigations (C. T. Philbrick unpubl. data) suggest that Marathrum s.s., including the type species of the genus, M. foeniculaceum (Van Royen 1951; Tippery et al. 2011), is found in Mexico, Central America, Columbia, and Cuba; other species once placed in Marathrum that are found outside of those areas have been transferred to other genera (Philbrick et al. 2018). Our results also strongly support that two Lophogyne species, L. fimbriata and L. royenella, are not monophyletic. Lophogyne ceratophylla and an unidentified Lophogyne species (L. sp1) are embedded within a paraphyletic L. royenella while L. wilsonii (paratype sampled here, Bove and Philbrick 2474; Bove et al. 2020) and another unidentified Lophogyne species (L. sp2) are embedded within a paraphyletic L. fimbriata. Paraphyletic species are not uncommon in molecular phylogenetic studies of Podostemaceae and may be attributed to allopatric speciation (e.g. Koi et al. 2012; Kato et al. 2019). Paraphyletic or non-monophyletic species may also be the result of misidentifications due the challenge of placing names on poorly known taxa, such as some species of Lophogyne s.l., that lack information regarding their morphological variability combined with a limited understanding of species distributions (Bidault et al. 2023). Monographic work in Lophogyne s.l. is clearly needed.
Possible New Genus—We place an undescribed species collected in Venezuela in 2007 (Philbrick et al. 6055, [MICH]). As far as we are aware, this species has been collected at only one locality. Initial attempts by C. T. Philbrick to identify this specimen to genus based on morphology were inconclusive and given its placement in the phylogeny, it likely represents a new genus. This species is strongly placed within a clade containing two other subclades, the first representing Apinagia s.s. (100 BS) and the second (94 BS) containing several taxa, including Oserya s.s., Noveloa, Marathrum s.s., and Apinagia nana and Oserya pilgeri. Relationships among these three lineages are not well supported (70 BS). Initial investigations by C. T. Philbrick (unpubl. data) suggest this undescribed taxon has prostrate stems divided into two regions including disklike hold-fasts from which leaves do not arise, and a region where the leaves arise. Thus the stems of this taxon are not similar to those in Apinagia s.s. which has upright stems (Tippery et al. 2011). Species of the second subclade have either upright or prostrate stems (Tippery et al. 2011). Morphological investigations, including comparisons to other closely related taxa, are ongoing and a formal description of this taxon will be presented in a future publication.
Oserya is Not Monophyletic—Our results place the genera Oserya s.s. and Noveloa as strongly supported sister taxa. Tippery et al. (2011) separated two species from Oserya s.l. and placed them in the new genus Noveloa based on their phylogenetic results and differences in morphology and biogeographic distribution (Noveloa coulteriana; N. longifollia). Oserya s.s. has pinnate leaf divisions, unistaminate flowers, five nonsuture ribs per capsule valve, and is distributed in South America, while Noveloa has dichotomous or subdichotomous leaf divisions, stamens numbering 1–3, three nonsuture ribs per capsule valve, and is distributed in Central America and Mexico (Tippery et al. 2011). While our results suggest that Oserya s.s. and Noveloa could be circumscribed into a broad Oserya s.l., we prefer to recognize these two genera separately due to their morphological and biogeographic differences. Our results also provide evidence that the recent transfer of Apinagia pilgeri Mildbr. into Oserya s.s. as Oserya pilgeri by Philbrick et al. (2016) is not warranted and would make Oserya s.s. nonmonophyletic. Oserya pilgeri is strongly supported as sister to Apinagia nana, and that clade is strongly supported as sister to Marathrum. As Apinagia nana and Oserya pilgeri are clearly not at home in either Oserya s.s. or Apinagia s.s., they should either be transferred to a new genus or into Marathrum. We defer these taxonomic changes pending further sampling from Marathrum. These two species are morphologically quite different from Marathrum species, so if this clade remains sister to Marathrum in future studies we suggest creating a new genus. An additional nomenclatural issue remains unaddressed as Philbrick et al. (2016) considered Oserya flabellifera Tul. & Wedd., the type species of Oserya (Van Royen 1954), and Castelnavia monandra synonyms. They did not propose combinations in Castelnavia for other names in Oserya s.s. (e.g. O. perpusilla, sampled here). Based on our results, O. perpusilla (and perhaps other species still included in Oserya s.s. [Tippery et al. 2011]) is not closely related to Castelnavia. We will address the necessary nomenclatural adjustments in a future publication.
Apinagia s.s.—The taxonomic impediment in Podostemaceae is especially acute in Apinagia s.l., the largest (34–37 spp.) and most taxonomically troublesome genus of neotropical Podostemaceae (Van Royen 1951; Berry 2004; Philbrick et al. 2010, 2016; POWO 2023). The genus is also of special conservation concern as over 50% of its species are documented from only one or two rivers, yet taxonomic uncertainty severely clouds the understanding of species distributions. Two factors contribute to this taxonomic uncertainty. First is the application of a largely typological species concept (cf. Cracraft 2000) by many previous workers, where idealized types represent a species and population level variation is disregarded. Second is the heavy reliance by previous workers on vegetative characters (e.g. leaf form) to distinguish species. Highly plastic vegetative forms, common in aquatic macrophytes and in Podostemaceae in particular (e.g. Sculthorpe 1967; Philbrick and Novelo 2004), combined with species delineations that emphasize vegetative features, have resulted in a taxonomy of dubious utility. The solution, we believe, rests on the use of characters that do not exhibit phenotypic plasticity (e.g. reproductive characters) for species recognition. Initial investigations by C. T. Philbrick suggest that characters important for species recognition in Apinagia include stamen arrangement around the ovary, tepal form, pedicel apex shape, and features of the mature capsules including the number and form of ribs that occur on capsule valves.
Results presented here agree with Tippery et al. (2011) who revealed that Apinagia s.l. is not monophyletic. Only Apinagia s.l. species with upright stems (one attachment point), as opposed to prostrate stems (multiple attachment points) should be included in Apinagia s.s. as the type for Apinagia, A. fucoides (Van Royen 1951; Tippery et al. 2011; Bove and Philbrick 2016), is a member of this upright-stem clade. Upright stems are not unique to Apinagia s.s., and are also present in species of Castelnavia, Oserya, and Lophogyne (Tippery et al. 2011). Many Apinagia species with prostrate stems have been transferred to other genera (Philbrick and Bove 2019) or will be in the future (e.g. Apinagia nana; Ruhfel et al. in prep), and other names have been placed into synonymy (Philbrick et al. 2016). Furthermore, preliminary investigations based on more than 20 yr of field work and hundreds of specimens collected by C. T. Philbrick and colleagues along with inspection of type material suggest that many fewer species exist than the number of accepted names (Philbrick et al. 2016). Preliminary work by C. T. Philbrick (unpubl. data) suggests that Apinagia s.s. contains ∼15 species, at least two of which are undescribed, though we defer making any taxonomic revisions until more species can be sampled and nuclear data can be examined. Two species of Apinagia s.s. are strongly supported as not monophyletic in our results. Apinagia richardiana is embedded within Apinagia longifolia and Apinagia tenuifolia and the undescribed species A. ‘crowii’ are embedded within A. fucoides. While no morphological differences were found between the two clades of A. fucoides, there is a difference in their geographic distribution. The clade containing A. fucoides specimens C.P. Bove et al. 2236, 2276, and 2284, and C.T. Philbrick et al. 5992 is distributed in the northeast of Brazil (Piauí, Tocantins) while the other A. fucoides clade is distributed in central Brazil (Tocantins, Goias, Mato Grosso, Mato Grosso do Sul). Overall, further work is needed on species boundaries within Apinagia. Results presented here provide critical information for developing a functional taxonomy for this genus to be included in an upcoming monograph, necessary for implementing conservation strategies to save these threatened species. Clarified species boundaries will allow for critically needed evaluation of local species endemism and the impact of expanding hydroelectric generation capacity (Philbrick et al. 2010).
Phylogenetic Conflict and the Use of Concatenated Plastid Genes for Phylogenetic Reconstruction in Podostemaceae—We observed very little supported conflict between individual plastid gene trees and the concatenated phylogeny. Furthermore, most individual gene trees lacked strong support for the majority of branches (Fig. S5), indicating that concatenation of the plastid coding regions was appropriate for inferring the species phylogeny. Most supported conflict was within clades corresponding to single species or closely related groups of species, suggesting that there was no evidence of widespread conflict among plastid gene trees in Podostemaceae due to biological causes (e.g. heteroplasmic recombination) or systematic and stochastic error (see Walker et al. 2019 and references therein). There were four nodes where the gene tree for matK, the gene region most often used to infer the phylogeny of this family in previous publications, conflicted with the concatenated tree. However, each of these conflicts was within a clade representing a single species and thus not relevant to relationships among genera and species in the family. This suggests that individual plastid gene regions such as matK may continue to be useful for placing taxa within the Podostemaceae phylogeny, provided they contain sufficient phylogenetic information. Hybridization has been suggested in Inversodicraea based on morphology (Cheek et al. 2017) and demonstrated in Marathrum with molecular data (Bedoya et al. 2021) and thus it is possible that introgression as well as incomplete lineage sorting could cause phylogenetic conflict among nuclear genes and the plastome. In such a scenario, plastome data may or may not reflect the overall species divergence history. The extent to which this is an issue for phylogenetic inference in Podostemaceae remains to be tested with broadly sampled, nuclear phylogenomic data.
Conclusions and Future Directions—The phylogeny presented here is the first well-sampled plastid phylogenomic analysis of the family and provides novel insights on Podostemaceae evolutionary history and plastome evolution. Our analyses resolve several recalcitrant nodes compared to previous studies, provide evidence for taxonomic and nomenclatural adjustments, place a previously unsampled and undescribed taxon that likely represents a new genus and, in contrast to previously published results, suggest that there has been only one dispersal event from the neotropics to the paleotropics within Podostemoideae. Additionally, we show that there is no evidence of widespread conflict among plastid genes and the concatenated phylogeny. While taxon sampling here was adequate to address several important questions, expanded sampling is needed to complete an assessment of phylogenetic relationships of all described taxa of Podostemaceae. Particularly important to include in future studies are representatives of genera not sampled here, and an expanded sampling of large and taxonomically confusing genera such as Apinagia s.s., Inversodicraea, Ledermanniella, Lophogyne s.l., and Rhyncholacis, all of which are greatly in need of monographic work. Particularly needed are more comparative morphological studies on several populations per species in order to evaluate the range of phenotypic plasticity (e.g. Bidault et al. 2023). Now that the phylogenetic history of the Podostemaceae plastome is becoming clear, a more complete understanding of the evolutionary history of the family will require analyses using nuclear genomic data.
Acknowledgments
This manuscript is dedicated to the memories of Bruno Bove da Costa, C. Thomas Philbrick, and Zhenxiang Xi. We thank Deise J. P. Goncalves, Lucas Majure, Kurt Neubig, James Pease, Stephen A. Smith, and Gregory W. Stull for discussions regarding analytical approaches; Christiane Anderson, Fred Barrie, Kanchi Gandhi, and Anton A. Reznicek for advice on nomenclatural issues; and Allison Harrington and Kyle Lough for help with the figures. We also thank the reviewers for their helpful comments on the manuscript. Finally, we thank the herbaria that allowed us access to their collections (see Appendix 1). Funding was provided by the National Science Foundation to BRR (DEB-1754329), CTP (DEB-0444589 & DEB-1754199), and DAL (IOS-2109716). The University Research Committee at Eastern Kentucky University provided funding to BRR. This research was not preregistered with an independent, institutional registry.
© Copyright 2024 by the American Society of Plant Taxonomists
Author Contributions
BRR and CTP conceived and designed the study. BRR was the primary author of the manuscript. BRR, CPB, CTP, and RR provided important samples and vouchers. BRR and NK extracted the gDNA and prepared it for sequencing. BRR and DAL developed and utilized the pipeline to trim, filter, assemble, and organize the raw data. BRR analyzed the final datasets and produced the figures. All authors contributed to the final manuscript.
Literature Cited
Appendices
Appendix 1.
Voucher information and GenBank accessions for sequences used in this study. Data produced for this study have GenBank numbers beginning with “OR” or “PP”. Data for species not sequenced in this study list the GenBank numbers for each coding region or the full plastome accession number. An em dash (—) indicates that the sequence was not available. Herbarium acronyms follow Index Herbariorum (Thiers 2023). Order of data in the appendix below is as follows, except for those with only a full plastome accession number: Family. Genus. Species: voucher (herbarium), GenBank accessions: atpA, atpB, atpE, atpF, atpH, atpI, ccsA, cema, clpP, matK, ndhA, ndhB, ndhC, ndhD, ndhE, ndhF, ndhG, ndhH, ndhI, ndhJ, ndhK, petA, petB, petD, petG, petL, petN, psaA, psaB, psaC, psaI, psaJ, psbA, psbB, psbC, psbD, psbE, psbF, psbH, psbI, psbJ, psbK, psbL, psbM, psbN, psbT, psbZ, rbcL, rpl2, rpl14, rpl16, rpl20, rpl22, rpl32, rpl33, rpl36, rpoA, rpoB, rpoC1, rpoC2, rps11, rps12, rps14, rps15, rps18, rps19, rps2, rps3, rps4, rps7, rps8, ycf3, and ycf4.
Hypericaceae (outgroup). Cratoxylum Blume. C. cochinchinense (Lour.) Blume: unknown CNShis0047329 (CNS), MK995180.
Vismia Vand. V. guianensis (Aubl.) Pers.: A. M. Amorim 7659 (CEPEC), JX662421, JX663828, JX663747, JX664444, JX662935, JX665032, JX662304, JX664834, —, JX661967, JX663657, JX664325, JX661839, JX664366, JX664647, JX662779, JX663435, JX662826, JX662090, JX662735, JX664561, JX664114, JX663943, JX663015, JX662454, JX662486, —, JX664235, JX662981, JX663129, JX664189, JX664679, JX663305, JX662222, JX662603, JX662343, JX664515, JX663861, JX663222, JX663895, JX664030, JX661926, JX662262, —, JX662648, JX664606, JX664149, JX664076, JX664280, JX663393, JX663990, JX662567, JX662863, —, —, JX663059, JX662048, JX663504, JX664967, JX662692, JX664799, JX663610, JX661885, JX663534, —, JX662384, JX663177, JX663787, JX664877, —, JX663349, JX664482, JX663262.
Podostemaceae (ingroup). Apinagia Tul. A. ‘crowii’: C. T. Philbrick et al. 6320 (MICH, VEN), PP242918, PP245408, PP246237, PP244218, PP242314, PP242798, PP242195, PP243494, PP241351, PP245884, PP244816, PP241110, PP244098, PP243041, PP242078, PP245170, PP245291, PP246603, PP244697, PP241720, PP243611, PP244458, PP241597, PP242558, PP245770, PP243278, PP245053, PP248280, PP247448, PP247690, PP248403, PP247213, PP246849, PP243732, PP241473, PP242435, PP242676, PP245649, PP244935, PP246001, PP243855, PP241233, PP241840, PP243382, PP245529, PP247931, PP247330, OR885176, PP246971, PP240989, PP246358, PP249106, PP247095, PP248634, PP248519, PP248167, PP244338, PP246480, PP248046, PP249338, PP246115, PP243975, PP241957, PP243161, PP244578, PP246726, PP247570, PP247810, PP249461, PP249226, PP248864, PP248984, PP248743; C. T. Philbrick et al. 6321 (MICH, VEN), PP242919, PP245409, PP246238, PP244219, PP242315, PP242799, PP242196, PP243495, PP241352, PP245885, PP244817, PP241111, PP244099, PP243042, PP242079, PP245171, PP245292, PP246604, PP244698, PP241721, PP243612, PP244459, PP241598, PP242559, PP245771, PP243279, PP245054, PP248281, PP247449, PP247691, PP248404, PP247214, PP246850, PP243733, PP241474, PP242436, PP242677, PP245650, PP244936, PP246002, PP243856, PP241234, PP241841, PP243383, PP245530, PP247932, PP247331, OR885177, PP246972, PP240990, PP246359, PP249107, PP247096, PP248635, PP248520, PP248168, PP244339, PP246481, PP248047, PP249339, PP246116, PP243976, PP241958, PP243162, PP244579, PP246727, PP247571, PP247811, PP249462, PP249227, PP248865, PP248985, PP248744. A. fucoides (Mart.) Tul.: C. P. Bove et al. 2200 (R), PP242881, PP245370, PP246199, PP244180, PP242276, PP242760, PP242157, PP243456, PP241313, PP245846, PP244778, PP241072, PP244060, PP243003, PP242040, PP245132, PP245253, PP246565, PP244659, PP241682, PP243573, PP244420, PP241559, PP242520, PP245732, PP243240, PP245015, PP248243, PP247411, PP247652, PP248365, PP247176, PP246811, PP243694, PP241435, PP242397, PP242638, PP245611, PP244897, PP245963, PP243817, PP241195, PP241802, PP243345, PP245491, PP247893, PP247292, OR885138, PP246933, PP240951, PP246320, PP249068, PP247057, PP248596, PP248481, PP248129, PP244300, PP246442, PP248009, PP249301, PP246077, PP243937, PP241919, PP243123, PP244540, PP246688, PP247532, PP247772, PP249423, PP249188, PP248826, PP248946, PP248705; C. P. Bove et al. 2210 (MICH, R), PP242875, PP245364, PP246193, PP244174, PP242270, PP242754, PP242151, PP243450, PP241307, PP245840, PP244772, PP241066, PP244054, PP242997, PP242034, PP245126, PP245247, PP246559, PP244653, PP241676, PP243567, PP244414, PP241553, PP242514, PP245726, PP243234, PP245009, PP248237, PP247405, PP247646, PP248359, PP247170, PP246805, PP243688, PP241429, PP242391, PP242632, PP245605, PP244891, PP245957, PP243811, PP241189, PP241796, PP243339, PP245485, PP247887, PP247286, OR885132, PP246927, PP240945, PP246314, PP249062, PP247051, PP248590, PP248475, PP248123, PP244294, PP246436, PP248003, PP249295, PP246071, PP243931, PP241913, PP243117, PP244534, PP246682, PP247526, PP247766, PP249417, PP249182, PP248820, PP248940, PP248699; C. P. Bove et al. 2227 (MICH, R), PP242910, PP245399, PP246228, PP244209, PP242305, PP242789, PP242186, PP243485, PP241342, PP245875, PP244807, PP241101, PP244089, PP243032, PP242069, PP245161, PP245282, PP246594, PP244688, PP241711, PP243602, PP244449, PP241588, PP242549, PP245761, PP243269, PP245044, PP248271, PP247439, PP247681, PP248394, PP247204, PP246840, PP243723, PP241464, PP242426, PP242667, PP245640, PP244926, PP245992, PP243846, PP241224, PP241831, PP243373, PP245520, PP247922, PP247321, OR885167, PP246962, PP240980, PP246349, PP249097, PP247086, PP248625, PP248510, PP248158, PP244329, PP246471, PP248037, PP249329, PP246106, PP243966, PP241948, PP243152, PP244569, PP246717, PP247561, PP247801, PP249452, PP249217, PP248855, PP248975, PP248734; C. P. Bove et al. 2236 (MICH, R), PP242907, PP245396, PP246225, PP244206, PP242302, PP242786, PP242183, PP243482, PP241339, PP245872, PP244804, PP241098, PP244086, PP243029, PP242066, PP245158, PP245279, PP246591, PP244685, PP241708, PP243599, PP244446, PP241585, PP242546, PP245758, PP243266, PP245041, PP248268, PP247436, PP247678, PP248391, PP247201, PP246837, PP243720, PP241461, PP242423, PP242664, PP245637, PP244923, PP245989, PP243843, PP241221, PP241828, PP243370, PP245517, PP247919, PP247318, OR885164, PP246959, PP240977, PP246346, PP249094, PP247083, PP248622, PP248507, PP248155, PP244326, PP246468, PP248034, PP249326, PP246103, PP243963, PP241945, PP243149, PP244566, PP246714, PP247558, PP247798, PP249449, PP249214, PP248852, PP248972, PP248731; C. P. Bove et al. 2237 (MICH, R), PP242911, PP245400, PP246229, PP244210, PP242306, PP242790, PP242187, PP243486, PP241343, PP245876, PP244808, PP241102, PP244090, PP243033, PP242070, PP245162, PP245283, PP246595, PP244689, PP241712, PP243603, PP244450, PP241589, PP242550, PP245762, PP243270, PP245045, PP248272, PP247440, PP247682, PP248395, PP247205, PP246841, PP243724, PP241465, PP242427, PP242668, PP245641, PP244927, PP245993, PP243847, PP241225, PP241832, PP243374, PP245521, PP247923, PP247322, OR885168, PP246963, PP240981, PP246350, PP249098, PP247087, PP248626, PP248511, PP248159, PP244330, PP246472, PP248038, PP249330, PP246107, PP243967, PP241949, PP243153, PP244570, PP246718, PP247562, PP247802, PP249453, PP249218, PP248856, PP248976, PP248735; C. P. Bove et al. 2276 (MICH, R), PP242876, PP245365, PP246194, PP244175, PP242271, PP242755, PP242152, PP243451, PP241308, PP245841, PP244773, PP241067, PP244055, PP242998, PP242035, PP245127, PP245248, PP246560, PP244654, PP241677, PP243568, PP244415, PP241554, PP242515, PP245727, PP243235, PP245010, PP248238, PP247406, PP247647, PP248360, PP247171, PP246806, PP243689, PP241430, PP242392, PP242633, PP245606, PP244892, PP245958, PP243812, PP241190, PP241797, PP243340, PP245486, PP247888, PP247287, OR885133, PP246928, PP240946, PP246315, PP249063, PP247052, PP248591, PP248476, PP248124, PP244295, PP246437, PP248004, PP249296, PP246072, PP243932, PP241914, PP243118, PP244535, PP246683, PP247527, PP247767, PP249418, PP249183, PP248821, PP248941, PP248700; C. P. Bove et al. 2284 (MICH, R), PP242916, PP245406, PP246235, PP244216, PP242312, PP242796, PP242193, PP243492, PP241349, PP245882, PP244814, PP241108, PP244096, PP243039, PP242076, PP245168, PP245289, PP246601, PP244695, PP241718, PP243609, PP244456, PP241595, PP242556, PP245768, PP243276, PP245051, PP248278, PP247446, PP247688, PP248401, PP247211, PP246847, PP243730, PP241471, PP242433, PP242674, PP245647, PP244933, PP245999, PP243853, PP241231, PP241838, PP243380, PP245527, PP247929, PP247328, OR885174, PP246969, PP240987, PP246356, PP249104, PP247093, PP248632, PP248517, PP248165, PP244336, PP246478, PP248044, PP249336, PP246113, PP243973, PP241955, PP243159, PP244576, PP246724, PP247568, PP247808, PP249459, PP249224, PP248862, PP248982, PP248741; C. P. Bove et al. 2311 (R), PP242908, PP245397, PP246226, PP244207, PP242303, PP242787, PP242184, PP243483, PP241340, PP245873, PP244805, PP241099, PP244087, PP243030, PP242067, PP245159, PP245280, PP246592, PP244686, PP241709, PP243600, PP244447, PP241586, PP242547, PP245759, PP243267, PP245042, PP248269, PP247437, PP247679, PP248392, PP247202, PP246838, PP243721, PP241462, PP242424, PP242665, PP245638, PP244924, PP245990, PP243844, PP241222, PP241829, PP243371, PP245518, PP247920, PP247319, OR885165, PP246960, PP240978, PP246347, PP249095, PP247084, PP248623, PP248508, PP248156, PP244327, PP246469, PP248035, PP249327, PP246104, PP243964, PP241946, PP243150, PP244567, PP246715, PP247559, PP247799, PP249450, PP249215, PP248853, PP248973, PP248732; C. P. Bove et al. 2513 (MICH, R), MN165812; C. P. Bove et al. 2517 (MICH, R), PP242912, PP245401, PP246230, PP244211, PP242307, PP242791, PP242188, PP243487, PP241344, PP245877, PP244809, PP241103, PP244091, PP243034, PP242071, PP245163, PP245284, PP246596, PP244690, PP241713, PP243604, PP244451, PP241590, PP242551, PP245763, PP243271, PP245046, PP248273, PP247441, PP247683, PP248396, PP247206, PP246842, PP243725, PP241466, PP242428, PP242669, PP245642, PP244928, PP245994, PP243848, PP241226, PP241833, PP243375, PP245522, PP247924, PP247323, OR885169, PP246964, PP240982, PP246351, PP249099, PP247088, PP248627, PP248512, PP248160, PP244331, PP246473, PP248039, PP249331, PP246108, PP243968, PP241950, PP243154, PP244571, PP246719, PP247563, PP247803, PP249454, PP249219, PP248857, PP248977, PP248736; C. P. Bove et al. 2523 (MICH, R), PP242913, PP245402, PP246231, PP244212, PP242308, PP242792, PP242189, PP243488, PP241345, PP245878, PP244810, PP241104, PP244092, PP243035, PP242072, PP245164, PP245285, PP246597, PP244691, PP241714, PP243605, PP244452, PP241591, PP242552, PP245764, PP243272, PP245047, PP248274, PP247442, PP247684, PP248397, PP247207, PP246843, PP243726, PP241467, PP242429, PP242670, PP245643, PP244929, PP245995, PP243849, PP241227, PP241834, PP243376, PP245523, PP247925, PP247324, OR885170, PP246965, PP240983, PP246352, PP249100, PP247089, PP248628, PP248513, PP248161, PP244332, PP246474, PP248040, PP249332, PP246109, PP243969, PP241951, PP243155, PP244572, PP246720, PP247564, PP247804, PP249455, PP249220, PP248858, PP248978, PP248737; C. P. Bove et al. 2527 (MICH, R), PP242914, PP245403, PP246232, PP244213, PP242309, PP242793, PP242190, PP243489, PP241346, PP245879, PP244811, PP241105, PP244093, PP243036, PP242073, PP245165, PP245286, PP246598, PP244692, PP241715, PP243606, PP244453, PP241592, PP242553, PP245765, PP243273, PP245048, PP248275, PP247443, PP247685, PP248398, PP247208, PP246844, PP243727, PP241468, PP242430, PP242671, PP245644, PP244930, PP245996, PP243850, PP241228, PP241835, PP243377, PP245524, PP247926, PP247325, OR885171, PP246966, PP240984, PP246353, PP249101, PP247090, PP248629, PP248514, PP248162, PP244333, PP246475, PP248041, PP249333, PP246110, PP243970, PP241952, PP243156, PP244573, PP246721, PP247565, PP247805, PP249456, PP249221, PP248859, PP248979, PP248738; C. T. Philbrick et al. 5960 (MICH, R), PP242877, PP245366, PP246195, PP244176, PP242272, PP242756, PP242153, PP243452, PP241309, PP245842, PP244774, PP241068, PP244056, PP242999, PP242036, PP245128, PP245249, PP246561, PP244655, PP241678, PP243569, PP244416, PP241555, PP242516, PP245728, PP243236, PP245011, PP248239, PP247407, PP247648, PP248361, PP247172, PP246807, PP243690, PP241431, PP242393, PP242634, PP245607, PP244893, PP245959, PP243813, PP241191, PP241798, PP243341, PP245487, PP247889, PP247288, OR885134, PP246929, PP240947, PP246316, PP249064, PP247053, PP248592, PP248477, PP248125, PP244296, PP246438, PP248005, PP249297, PP246073, PP243933, PP241915, PP243119, PP244536, PP246684, PP247528, PP247768, PP249419, PP249184, PP248822, PP248942, PP248701; C. T. Philbrick et al. 5969 (MICH, R), —, PP245404, PP246233, PP244214, PP242310, PP242794, PP242191, PP243490, PP241347, PP245880, PP244812, PP241106, PP244094, PP243037, PP242074, PP245166, PP245287, PP246599, PP244693, PP241716, PP243607, PP244454, PP241593, PP242554, PP245766, PP243274, PP245049, PP248276, PP247444, PP247686, PP248399, PP247209, PP246845, PP243728, PP241469, PP242431, PP242672, PP245645, PP244931, PP245997, PP243851, PP241229, PP241836, PP243378, PP245525, PP247927, PP247326, OR885172, PP246967, PP240985, PP246354, PP249102, PP247091, PP248630, PP248515, PP248163, PP244334, PP246476, PP248042, PP249334, PP246111, PP243971, PP241953, PP243157, PP244574, PP246722, PP247566, PP247806, PP249457, PP249222, PP248860, PP248980, PP248739; C. T. Philbrick et al. 5970 (MICH, R), PP242909, PP245398, PP246227, PP244208, PP242304, PP242788, PP242185, PP243484, PP241341, PP245874, PP244806, PP241100, PP244088, PP243031, PP242068, PP245160, PP245281, PP246593, PP244687, PP241710, PP243601, PP244448, PP241587, PP242548, PP245760, PP243268, PP245043, PP248270, PP247438, PP247680, PP248393, PP247203, PP246839, PP243722, PP241463, PP242425, PP242666, PP245639, PP244925, PP245991, PP243845, PP241223, PP241830, PP243372, PP245519, PP247921, PP247320, OR885166, PP246961, PP240979, PP246348, PP249096, PP247085, PP248624, PP248509, PP248157, PP244328, PP246470, PP248036, PP249328, PP246105, PP243965, PP241947, PP243151, PP244568, PP246716, PP247560, PP247800, PP249451, PP249216, PP248854, PP248974, PP248733; C. T. Philbrick et al. 5992 (MICH, R), PP242915, PP245405, PP246234, PP244215, PP242311, PP242795, PP242192, PP243491, PP241348, PP245881, PP244813, PP241107, PP244095, PP243038, PP242075, PP245167, PP245288, PP246600, PP244694, PP241717, PP243608, PP244455, PP241594, PP242555, PP245767, PP243275, PP245050, PP248277, PP247445, PP247687, PP248400, PP247210, PP246846, PP243729, PP241470, PP242432, PP242673, PP245646, PP244932, PP245998, PP243852, PP241230, PP241837, PP243379, PP245526, PP247928, PP247327, OR885173, PP246968, PP240986, PP246355, PP249103, PP247092, PP248631, PP248516, PP248164, PP244335, PP246477, PP248043, PP249335, PP246112, PP243972, PP241954, PP243158, PP244575, PP246723, PP247567, PP247807, PP249458, PP249223, PP248861, PP248981, PP248740. A. guyanensis (Pulle) P. Royen: C. P. Bove and C. T. Philbrick 2402 (MICH, R), PP242886, PP245375, PP246204, PP244185, PP242281, PP242765, PP242162, PP243461, PP241318, PP245851, PP244783, PP241077, PP244065, PP243008, PP242045, PP245137, PP245258, PP246570, PP244664, PP241687, PP243578, PP244425, PP241564, PP242525, PP245737, PP243245, PP245020, PP248248, PP247416, PP247657, PP248370, PP247181, PP246816, PP243699, PP241440, PP242402, PP242643, PP245616, PP244902, PP245968, PP243822, PP241200, PP241807, PP243350, PP245496, PP247898, PP247297, OR885143, PP246938, PP240956, PP246325, PP249073, PP247062, PP248601, PP248486, PP248134, PP244305, PP246447, PP248014, PP249306, PP246082, PP243942, PP241924, PP243128, PP244545, PP246693, PP247537, PP247777, PP249428, PP249193, PP248831, PP248951, PP248710; C. P. Bove and C. T. Philbrick 2407 (MICH, R), PP242887, PP245376, PP246205, PP244186, PP242282, PP242766, PP242163, PP243462, PP241319, PP245852, PP244784, PP241078, PP244066, PP243009, PP242046, PP245138, PP245259, PP246571, PP244665, PP241688, PP243579, PP244426, PP241565, PP242526, PP245738, PP243246, PP245021, PP248249, PP247417, PP247658, PP248371, PP247182, PP246817, PP243700, PP241441, PP242403, PP242644, PP245617, PP244903, PP245969, PP243823, PP241201, PP241808, PP243351, PP245497, PP247899, PP247298, OR885144, PP246939, PP240957, PP246326, PP249074, PP247063, PP248602, PP248487, PP248135, PP244306, PP246448, PP248015, PP249307, PP246083, PP243943, PP241925, PP243129, PP244546, PP246694, PP247538, PP247778, PP249429, PP249194, PP248832, PP248952, PP248711; C. P. Bove and C. T. Philbrick 2410 (MICH, R), PP242899, PP245388, PP246217, PP244198, PP242294, PP242778, PP242175, PP243474, PP241331, PP245864, PP244796, PP241090, PP244078, PP243021, PP242058, PP245150, PP245271, PP246583, PP244677, PP241700, PP243591, PP244438, PP241577, PP242538, PP245750, PP243258, PP245033, PP248260, PP247428, PP247670, PP248383, PP247194, PP246829, PP243712, PP241453, PP242415, PP242656, PP245629, PP244915, PP245981, PP243835, PP241213, PP241820, PP243363, PP245509, PP247911, PP247310, OR885156, PP246951, PP240969, PP246338, PP249086, PP247075, PP248614, PP248499, PP248147, PP244318, PP246460, PP248027, PP249318, PP246095, PP243955, PP241937, PP243141, PP244558, PP246706, PP247550, PP247790, PP249441, PP249206, PP248844, PP248964, PP248723; C. P. Bove and C. T. Philbrick 2411 (MICH, R), PP242888, PP245377, PP246206, PP244187, PP242283, PP242767, PP242164, PP243463, PP241320, PP245853, PP244785, PP241079, PP244067, PP243010, PP242047, PP245139, PP245260, PP246572, PP244666, PP241689, PP243580, PP244427, PP241566, PP242527, PP245739, PP243247, PP245022, PP248250, PP247418, PP247659, PP248372, PP247183, PP246818, PP243701, PP241442, PP242404, PP242645, PP245618, PP244904, PP245970, PP243824, PP241202, PP241809, PP243352, PP245498, PP247900, PP247299, OR885145, PP246940, PP240958, PP246327, PP249075, PP247064, PP248603, PP248488, PP248136, PP244307, PP246449, PP248016, —, PP246084, PP243944, PP241926, PP243130, PP244547, PP246695, PP247539, PP247779, PP249430, PP249195, PP248833, PP248953, PP248712; C. P. Bove and C. T. Philbrick 2412 (MICH, R), PP242890, PP245379, PP246208, PP244189, PP242285, PP242769, PP242166, PP243465, PP241322, PP245855, PP244787, PP241081, PP244069, PP243012, PP242049, PP245141, PP245262, PP246574, PP244668, PP241691, PP243582, PP244429, PP241568, PP242529, PP245741, PP243249, PP245024, PP248252, PP247419, PP247661, PP248374, PP247185, PP246820, PP243703, PP241444, PP242406, PP242647, PP245620, PP244906, PP245972, PP243826, PP241204, PP241811, PP243354, PP245500, PP247902, PP247301, OR885147, PP246942, PP240960, PP246329, PP249077, PP247066, PP248605, PP248490, PP248138, PP244309, PP246451, PP248018, PP249309, PP246086, PP243946, PP241928, PP243132, PP244549, PP246697, PP247541, PP247781, PP249432, PP249197, PP248835, PP248955, PP248714; C. P. Bove and C. T. Philbrick 2420 (MICH, R), PP242898, PP245387, PP246216, PP244197, PP242293, PP242777, PP242174, PP243473, PP241330, PP245863, PP244795, PP241089, PP244077, PP243020, PP242057, PP245149, PP245270, PP246582, PP244676, PP241699, PP243590, PP244437, PP241576, PP242537, PP245749, PP243257, PP245032, —, PP247427, PP247669, PP248382, PP247193, PP246828, PP243711, PP241452, PP242414, PP242655, PP245628, PP244914, PP245980, PP243834, PP241212, PP241819, PP243362, PP245508, PP247910, PP247309, OR885155, PP246950, PP240968, PP246337, PP249085, PP247074, PP248613, PP248498, PP248146, PP244317, PP246459, PP248026, PP249317, PP246094, PP243954, PP241936, PP243140, PP244557, PP246705, PP247549, PP247789, PP249440, PP249205, PP248843, PP248963, PP248722; C. P. Bove and C. T. Philbrick 2434 (MICH, R), PP242917, PP245407, PP246236, PP244217, PP242313, PP242797, PP242194, PP243493, PP241350, PP245883, PP244815, PP241109, PP244097, PP243040, PP242077, PP245169, PP245290, PP246602, PP244696, PP241719, PP243610, PP244457, PP241596, PP242557, PP245769, PP243277, PP245052, PP248279, PP247447, PP247689, PP248402, PP247212, PP246848, PP243731, PP241472, PP242434, PP242675, PP245648, PP244934, PP246000, PP243854, PP241232, PP241839, PP243381, PP245528, PP247930, PP247329, OR885175, PP246970, PP240988, PP246357, PP249105, PP247094, PP248633, PP248518, PP248166, PP244337, PP246479, PP248045, PP249337, PP246114, PP243974, PP241956, PP243160, PP244577, PP246725, PP247569, PP247809, PP249460, PP249225, PP248863, PP248983, PP248742; C. P. Bove and C. T. Philbrick 2437 (R), PP242897, PP245386, PP246215, PP244196, PP242292, PP242776, PP242173, PP243472, PP241329, PP245862, PP244794, PP241088, PP244076, PP243019, PP242056, PP245148, PP245269, PP246581, PP244675, PP241698, PP243589, PP244436, PP241575, PP242536, PP245748, PP243256, PP245031, PP248259, PP247426, PP247668, PP248381, PP247192, PP246827, PP243710, PP241451, PP242413, PP242654, PP245627, PP244913, PP245979, PP243833, PP241211, PP241818, PP243361, PP245507, PP247909, PP247308, OR885154, PP246949, PP240967, PP246336, PP249084, PP247073, PP248612, PP248497, PP248145, PP244316, PP246458, PP248025, PP249316, PP246093, PP243953, PP241935, PP243139, PP244556, PP246704, PP247548, PP247788, PP249439, PP249204, PP248842, PP248962, PP248721; C. P. Bove and C. T. Philbrick 2487 (MICH, R), PP242896, PP245385, PP246214, PP244195, PP242291, PP242775, PP242172, PP243471, PP241328, PP245861, PP244793, PP241087, PP244075, PP243018, PP242055, PP245147, PP245268, PP246580, PP244674, PP241697, PP243588, PP244435, PP241574, PP242535, PP245747, PP243255, PP245030, PP248258, PP247425, PP247667, PP248380, PP247191, PP246826, PP243709, PP241450, PP242412, PP242653, PP245626, PP244912, PP245978, PP243832, PP241210, PP241817, PP243360, PP245506, PP247908, PP247307, OR885153, PP246948, PP240966, PP246335, PP249083, PP247072, PP248611, PP248496, PP248144, PP244315, PP246457, PP248024, PP249315, PP246092, PP243952, PP241934, PP243138, PP244555, PP246703, PP247547, PP247787, PP249438, PP249203, PP248841, PP248961, PP248720; C. P. Bove and C. T. Philbrick 2489 (MICH, R), PP242889, PP245378, PP246207, PP244188, PP242284, PP242768, PP242165, PP243464, PP241321, PP245854, PP244786, PP241080, PP244068, PP243011, PP242048, PP245140, PP245261, PP246573, PP244667, PP241690, PP243581, PP244428, PP241567, PP242528, PP245740, PP243248, PP245023, PP248251, —, PP247660, PP248373, PP247184, PP246819, PP243702, PP241443, PP242405, PP242646, PP245619, PP244905, PP245971, PP243825, PP241203, PP241810, PP243353, PP245499, PP247901, PP247300, OR885146, PP246941, PP240959, PP246328, PP249076, PP247065, PP248604, PP248489, PP248137, PP244308, PP246450, PP248017, PP249308, PP246085, PP243945, PP241927, PP243131, PP244548, PP246696, PP247540, PP247780, PP249431, PP249196, PP248834, PP248954, PP248713; C. P. Bove et al. 2206 (MICH, R), PP242883, PP245372, PP246201, PP244182, PP242278, PP242762, PP242159, PP243458, PP241315, PP245848, PP244780, PP241074, PP244062, PP243005, PP242042, PP245134, PP245255, PP246567, PP244661, PP241684, PP243575, PP244422, PP241561, PP242522, PP245734, PP243242, PP245017, PP248245, PP247413, PP247654, PP248367, PP247178, PP246813, PP243696, PP241437, PP242399, PP242640, PP245613, PP244899, PP245965, PP243819, PP241197, PP241804, PP243347, PP245493, PP247895, PP247294, OR885140, PP246935, PP240953, PP246322, PP249070, PP247059, PP248598, PP248483, PP248131, PP244302, PP246444, PP248011, PP249303, PP246079, PP243939, PP241921, PP243125, PP244542, PP246690, PP247534, PP247774, PP249425, PP249190, PP248828, PP248948, PP248707. A. longifolia (Tul.) P.Royen.: C. P. Bove and C. T. Philbrick 2431 (MICH, R), PP242878, PP245367, PP246196, PP244177, PP242273, PP242757, PP242154, PP243453, PP241310, PP245843, PP244775, PP241069, PP244057, PP243000, PP242037, PP245129, PP245250, PP246562, PP244656, PP241679, PP243570, PP244417, PP241556, PP242517, PP245729, PP243237, PP245012, PP248240, PP247408, PP247649, PP248362, PP247173, PP246808, PP243691, PP241432, PP242394, PP242635, PP245608, PP244894, PP245960, PP243814, PP241192, PP241799, PP243342, PP245488, PP247890, PP247289, OR885135, PP246930, PP240948, PP246317, PP249065, PP247054, PP248593, PP248478, PP248126, PP244297, PP246439, PP248006, PP249298, PP246074, PP243934, PP241916, PP243120, PP244537, PP246685, PP247529, PP247769, PP249420, PP249185, PP248823, PP248943, PP248702; C. P. Bove and C. T. Philbrick 2450 (MICH, R), PP242879, PP245368, PP246197, PP244178, PP242274, PP242758, PP242155, PP243454, PP241311, PP245844, PP244776, PP241070, PP244058, PP243001, PP242038, PP245130, PP245251, PP246563, PP244657, PP241680, PP243571, PP244418, PP241557, PP242518, PP245730, PP243238, PP245013, PP248241, PP247409, PP247650, PP248363, PP247174, PP246809, PP243692, PP241433, PP242395, PP242636, PP245609, PP244895, PP245961, PP243815, PP241193, PP241800, PP243343, PP245489, PP247891, PP247290, OR885136, PP246931, PP240949, PP246318, PP249066, PP247055, PP248594, PP248479, PP248127, PP244298, PP246440, PP248007, PP249299, PP246075, PP243935, PP241917, PP243121, PP244538, PP246686, PP247530, PP247770, PP249421, PP249186, PP248824, PP248944, PP248703; C. P. Bove and C. T. Philbrick 2452 (MICH, R), PP242880, PP245369, PP246198, PP244179, PP242275, PP242759, PP242156, PP243455, PP241312, PP245845, PP244777, PP241071, PP244059, PP243002, PP242039, PP245131, PP245252, PP246564, PP244658, PP241681, PP243572, PP244419, PP241558, PP242519, PP245731, PP243239, PP245014, PP248242, PP247410, PP247651, PP248364, PP247175, PP246810, PP243693, PP241434, PP242396, PP242637, PP245610, PP244896, PP245962, PP243816, PP241194, PP241801, PP243344, PP245490, PP247892, PP247291, OR885137, PP246932, PP240950, PP246319, PP249067, PP247056, PP248595, PP248480, PP248128, PP244299, PP246441, PP248008, PP249300, PP246076, PP243936, PP241918, PP243122, PP244539, PP246687, PP247531, PP247771, PP249422, PP249187, PP248825, PP248945, PP248704; C. P. Bove and C. T. Philbrick 2455 (MICH, R), PP242874, PP245363, PP246192, PP244173, PP242269, PP242753, PP242150, PP243449, PP241306, PP245839, PP244771, PP241065, PP244053, PP242996, PP242033, PP245125, PP245246, PP246558, PP244652, PP241675, PP243566, PP244413, PP241552, PP242513, PP245725, PP243233, PP245008, PP248236, PP247404, PP247645, PP248358, PP247169, PP246804, PP243687, PP241428, PP242390, PP242631, PP245604, PP244890, PP245956, PP243810, PP241188, PP241795, PP243338, PP245484, PP247886, PP247285, OR885131, PP246926, PP240944, PP246313, PP249061, PP247050, PP248589, PP248474, PP248122, PP244293, PP246435, PP248002, PP249294, PP246070, PP243930, PP241912, PP243116, PP244533, PP246681, PP247525, PP247765, PP249416, PP249181, PP248819, PP248939, PP248698; C. T. Philbrick 6259 (BBS, MICH), PP242873, PP245362, PP246191, PP244172, PP242268, PP242752, —, —, PP241305, —, —, PP241064, —, PP242995, —, PP245124, —, PP246557, PP244651, PP241674, PP243565, PP244412, PP241551, —, —, —, —, PP248235, PP247403, PP247644, —, —, PP246803, PP243686, PP241427, PP242389, PP242630, PP245603, —, —, —, —, —, —, —, —, —, OR885130, PP246925, PP240943, PP246312, PP249060, PP247049, —, PP248473, —, —, PP246434, PP248001, PP249293, —, PP243929, PP241911, —, —, PP246680, —, —, PP249415, PP249180, —, PP248938, —. A. nana Went: C. T. Philbrick et al. 6180 (BBS, MICH), PP242901, PP245390, PP246219, PP244200, PP242296, PP242780, PP242177, PP243476, PP241333, PP245866, PP244798, PP241092, PP244080, PP243023, PP242060, PP245152, PP245273, PP246585, PP244679, PP241702, PP243593, PP244440, PP241579, PP242540, PP245752, PP243260, PP245035, PP248262, PP247430, PP247672, PP248385, —, PP246831, PP243714, PP241455, PP242417, PP242658, PP245631, PP244917, PP245983, PP243837, PP241215, PP241822, —, PP245511, PP247913, PP247312, OR885158, PP246953, PP240971, PP246340, PP249088, PP247077, PP248616, PP248501, PP248149, PP244320, PP246462, PP248028, PP249320, PP246097, PP243957, PP241939, PP243143, PP244560, PP246708, PP247552, PP247792, PP249443, PP249208, PP248846, PP248966, PP248725; C. T. Philbrick et al. 6322 (MICH, VEN), PP242900, PP245389, PP246218, PP244199, PP242295, PP242779, PP242176, PP243475, PP241332, PP245865, PP244797, PP241091, PP244079, PP243022, PP242059, PP245151, PP245272, PP246584, PP244678, PP241701, PP243592, PP244439, PP241578, PP242539, PP245751, PP243259, PP245034, PP248261, PP247429, PP247671, PP248384, PP247195, PP246830, PP243713, PP241454, PP242416, PP242657, PP245630, PP244916, PP245982, PP243836, PP241214, PP241821, PP243364, PP245510, PP247912, PP247311, OR885157, PP246952, PP240970, PP246339, PP249087, PP247076, PP248615, PP248500, PP248148, PP244319, PP246461, —, PP249319, PP246096, PP243956, PP241938, PP243142, PP244559, PP246707, PP247551, PP247791, PP249442, PP249207, PP248845, PP248965, PP248724. A. richardiana (Tul.) P. Royen: C. P. Bove et al. 1945 (MICH, R), PP242903, PP245392, PP246221, PP244202, PP242298, PP242782, PP242179, PP243478, PP241335, PP245868, PP244800, PP241094, PP244082, PP243025, PP242062, PP245154, PP245275, PP246587, PP244681, PP241704, PP243595, PP244442, PP241581, PP242542, PP245754, PP243262, PP245037, PP248264, PP247432, PP247674, PP248387, PP247197, PP246833, PP243716, PP241457, PP242419, PP242660, PP245633, PP244919, PP245985, PP243839, PP241217, PP241824, PP243366, PP245513, PP247915, PP247314, OR885160, PP246955, PP240973, PP246342, PP249090, PP247079, PP248618, PP248503, PP248151, PP244322, PP246464, PP248030, PP249322, PP246099, PP243959, PP241941, PP243145, PP244562, PP246710, PP247554, PP247794, PP249445, PP249210, PP248848, PP248968, PP248727; C. P. Bove et al. 1956 (MICH, R), PP242904, PP245393, PP246222, PP244203, PP242299, PP242783, PP242180, PP243479, PP241336, PP245869, PP244801, PP241095, PP244083, PP243026, PP242063, PP245155, PP245276, PP246588, PP244682, PP241705, PP243596, PP244443, PP241582, PP242543, PP245755, PP243263, PP245038, PP248265, PP247433, PP247675, PP248388, PP247198, PP246834, PP243717, PP241458, PP242420, PP242661, PP245634, PP244920, PP245986, PP243840, PP241218, PP241825, PP243367, PP245514, PP247916, PP247315, OR885161, PP246956, PP240974, PP246343, PP249091, PP247080, PP248619, PP248504, PP248152, PP244323, PP246465, PP248031, PP249323, PP246100, PP243960, PP241942, PP243146, PP244563, PP246711, PP247555, PP247795, PP249446, PP249211, PP248849, PP248969, PP248728; C. P. Bove et al. 1971 (MICH, R), PP242905, PP245394, PP246223, PP244204, PP242300, PP242784, PP242181, PP243480, PP241337, PP245870, PP244802, PP241096, PP244084, PP243027, PP242064, PP245156, PP245277, PP246589, PP244683, PP241706, PP243597, PP244444, PP241583, PP242544, PP245756, PP243264, PP245039, PP248266, PP247434, PP247676, PP248389, PP247199, PP246835, PP243718, PP241459, PP242421, PP242662, PP245635, PP244921, PP245987, PP243841, PP241219, PP241826, PP243368, PP245515, PP247917, PP247316, OR885162, PP246957, PP240975, PP246344, PP249092, PP247081, PP248620, PP248505, PP248153, PP244324, PP246466, PP248032, PP249324, PP246101, PP243961, PP241943, PP243147, PP244564, PP246712, PP247556, PP247796, PP249447, PP249212, PP248850, PP248970, PP248729; C. P. Bove et al. 1972 (MICH, R), PP242906, PP245395, PP246224, PP244205, PP242301, PP242785, PP242182, PP243481, PP241338, PP245871, PP244803, PP241097, PP244085, PP243028, PP242065, PP245157, PP245278, PP246590, PP244684, PP241707, PP243598, PP244445, PP241584, PP242545, PP245757, PP243265, PP245040, PP248267, PP247435, PP247677, PP248390, PP247200, PP246836, PP243719, PP241460, PP242422, PP242663, PP245636, PP244922, PP245988, PP243842, PP241220, PP241827, PP243369, PP245516, PP247918, PP247317, OR885163, PP246958, PP240976, PP246345, PP249093, PP247082, PP248621, PP248506, PP248154, PP244325, PP246467, PP248033, PP249325, PP246102, PP243962, PP241944, PP243148, PP244565, PP246713, PP247557, PP247797, PP249448, PP249213, PP248851, PP248971, PP248730; C. T. Philbrick et al. 6153 (MICH), PP242882, PP245371, PP246200, PP244181, PP242277, PP242761, PP242158, PP243457, PP241314, PP245847, PP244779, PP241073, PP244061, PP243004, PP242041, PP245133, PP245254, PP246566, PP244660, PP241683, PP243574, PP244421, PP241560, PP242521, PP245733, PP243241, PP245016, PP248244, PP247412, PP247653, PP248366, PP247177, PP246812, PP243695, PP241436, PP242398, PP242639, PP245612, PP244898, PP245964, PP243818, PP241196, PP241803, PP243346, PP245492, PP247894, PP247293, OR885139, PP246934, PP240952, PP246321, PP249069, PP247058, PP248597, PP248482, PP248130, PP244301, PP246443, PP248010, PP249302, PP246078, PP243938, PP241920, PP243124, PP244541, PP246689, PP247533, PP247773, PP249424, PP249189, PP248827, PP248947, PP248706. A. tenuifolia P. Royen: C. P. Bove et al. 1943 (MICH, R), PP242891, PP245380, PP246209, PP244190, PP242286, PP242770, PP242167, PP243466, PP241323, PP245856, PP244788, PP241082, PP244070, PP243013, PP242050, PP245142, PP245263, PP246575, PP244669, PP241692, PP243583, PP244430, PP241569, PP242530, PP245742, PP243250, PP245025, PP248253, PP247420, PP247662, PP248375, PP247186, PP246821, PP243704, PP241445, PP242407, PP242648, PP245621, PP244907, PP245973, PP243827, PP241205, PP241812, PP243355, PP245501, PP247903, PP247302, OR885148, PP246943, PP240961, PP246330, PP249078, PP247067, PP248606, PP248491, PP248139, PP244310, PP246452, PP248019, PP249310, PP246087, PP243947, PP241929, PP243133, PP244550, PP246698, PP247542, PP247782, PP249433, PP249198, PP248836, PP248956, PP248715; C. P. Bove et al. 1951 (MICH, R), PP242892, PP245381, PP246210, PP244191, PP242287, PP242771, PP242168, PP243467, PP241324, PP245857, PP244789, PP241083, PP244071, PP243014, PP242051, PP245143, PP245264, PP246576, PP244670, PP241693, PP243584, PP244431, PP241570, PP242531, PP245743, PP243251, PP245026, PP248254, PP247421, PP247663, PP248376, PP247187, PP246822, PP243705, PP241446, PP242408, PP242649, PP245622, PP244908, PP245974, PP243828, PP241206, PP241813, PP243356, PP245502, PP247904, PP247303, OR885149, PP246944, PP240962, PP246331, PP249079, PP247068, PP248607, PP248492, PP248140, PP244311, PP246453, PP248020, PP249311, PP246088, PP243948, PP241930, PP243134, PP244551, PP246699, PP247543, PP247783, PP249434, PP249199, PP248837, PP248957, PP248716; C. P. Bove et al. 1952 (MICH, R), PP242893, PP245382, PP246211, PP244192, PP242288, PP242772, PP242169, PP243468, PP241325, PP245858, PP244790, PP241084, PP244072, PP243015, PP242052, PP245144, PP245265, PP246577, PP244671, PP241694, PP243585, PP244432, PP241571, PP242532, PP245744, PP243252, PP245027, PP248255, PP247422, PP247664, PP248377, PP247188, PP246823, PP243706, PP241447, PP242409, PP242650, PP245623, PP244909, PP245975, PP243829, PP241207, PP241814, PP243357, PP245503, PP247905, PP247304, OR885150, PP246945, PP240963, PP246332, PP249080, PP247069, PP248608, PP248493, PP248141, PP244312, PP246454, PP248021, PP249312, PP246089, PP243949, PP241931, PP243135, PP244552, PP246700, PP247544, PP247784, PP249435, PP249200, PP248838, PP248958, PP248717; C. P. Bove et al. 1954 (MICH, R), PP242894, PP245383, PP246212, PP244193, PP242289, PP242773, PP242170, PP243469, PP241326, PP245859, PP244791, PP241085, PP244073, PP243016, PP242053, PP245145, PP245266, PP246578, PP244672, PP241695, PP243586, PP244433, PP241572, PP242533, PP245745, PP243253, PP245028, PP248256, PP247423, PP247665, PP248378, PP247189, PP246824, PP243707, PP241448, PP242410, PP242651, PP245624, PP244910, PP245976, PP243830, PP241208, PP241815, PP243358, PP245504, PP247906, PP247305, OR885151, PP246946, PP240964, PP246333, PP249081, PP247070, PP248609, PP248494, PP248142, PP244313, PP246455, PP248022, PP249313, PP246090, PP243950, PP241932, PP243136, PP244553, PP246701, PP247545, PP247785, PP249436, PP249201, PP248839, PP248959, PP248718; C. P. Bove et al. 1975 (MICH, R), PP242895, PP245384, PP246213, PP244194, PP242290, PP242774, PP242171, PP243470, PP241327, PP245860, PP244792, PP241086, PP244074, PP243017, PP242054, PP245146, PP245267, PP246579, PP244673, PP241696, PP243587, PP244434, PP241573, PP242534, PP245746, PP243254, PP245029, PP248257, PP247424, PP247666, PP248379, PP247190, PP246825, PP243708, PP241449, PP242411, PP242652, PP245625, PP244911, PP245977, PP243831, PP241209, PP241816, PP243359, PP245505, PP247907, PP247306, OR885152, PP246947, PP240965, PP246334, PP249082, PP247071, PP248610, PP248495, PP248143, PP244314, PP246456, PP248023, PP249314, PP246091, PP243951, PP241933, PP243137, PP244554, PP246702, PP247546, PP247786, PP249437, PP249202, PP248840, PP248960, PP248719.
Autana C.T.Philbrick. A. andersonii C.T.Philbrick: C. T. Philbrick et al. 5862 (MEXU, MICH, VEN), PP242920, PP245410, PP246239, PP244220, PP242316, PP242800, PP242197, PP243496, PP241353, PP245886, PP244818, PP241112, PP244100, PP243043, PP242080, PP245172, PP245293, PP246605, PP244699, PP241722, PP243613, PP244460, PP241599, PP242560, PP245772, PP243280, PP245055, PP248282, PP247450, PP247692, PP248405, PP247215, PP246851, PP243734, PP241475, PP242437, PP242678, PP245651, PP244937, PP246003, PP243857, PP241235, PP241842, PP243384, PP245531, PP247933, PP247332, OR885178, PP246973, PP240991, PP246360, PP249108, PP247097, PP248636, PP248521, PP248169, PP244340, PP246482, PP248048, PP249340, PP246117, PP243977, PP241959, PP243163, PP244580, PP246728, PP247572, PP247812, PP249463, PP249228, PP248866, PP248986, PP248745; C. T. Philbrick et al. 5867 (MEXU, MICH, VEN), PP242921, PP245411, PP246240, PP244221, PP242317, PP242801, PP242198, PP243497, PP241354, PP245887, PP244819, PP241113, PP244101, PP243044, PP242081, PP245173, PP245294, PP246606, PP244700, PP241723, PP243614, PP244461, PP241600, PP242561, PP245773, PP243281, PP245056, PP248283, PP247451, PP247693, PP248406, PP247216, PP246852, PP243735, PP241476, PP242438, PP242679, PP245652, PP244938, PP246004, PP243858, PP241236, PP241843, PP243385, PP245532, PP247934, PP247333, OR885179, PP246974, PP240992, PP246361, PP249109, PP247098, PP248637, PP248522, PP248170, PP244341, PP246483, PP248049, PP249341, PP246118, PP243978, PP241960, PP243164, PP244581, PP246729, PP247573, PP247813, PP249464, PP249229, PP248867, PP248987, PP248746.
Castelnavia Tul & Wedd. C. fluitans Tul. & Wedd.: C. T. Philbrick et al. 5840 (MICH, R), PP242922, PP245412, PP246241, PP244222, PP242318, PP242802, PP242199, PP243498, PP241355, PP245888, PP244820, PP241114, PP244102, PP243045, PP242082, PP245174, PP245295, PP246607, PP244701, PP241724, PP243615, PP244462, PP241601, PP242562, PP245774, PP243282, PP245057, PP248284, PP247452, PP247694, PP248407, PP247217, PP246853, PP243736, PP241477, PP242439, PP242680, PP245653, PP244939, PP246005, PP243859, PP241237, PP241844, PP243386, PP245533, PP247935, PP247334, OR885180, PP246975, PP240993, PP246362, PP249110, PP247099, PP248638, PP248523, PP248171, PP244342, PP246484, PP248050, PP249342, PP246119, PP243979, PP241961, PP243165, PP244582, PP246730, PP247574, PP247814, PP249465, PP249230, PP248868, PP248988, PP248747. C. monandra Tul. & Wedd.: C. T. Philbrick et al. 5982 (R), PP242923, PP245413, PP246242, PP244223, PP242319, PP242803, PP242200, PP243499, PP241356, PP245889, PP244821, PP241115, PP244103, PP243046, PP242083, PP245175, PP245296, PP246608, PP244702, PP241725, PP243616, PP244463, PP241602, PP242563, PP245775, PP243283, PP245058, PP248285, PP247453, PP247695, PP248408, PP247218, PP246854, PP243737, PP241478, PP242440, PP242681, PP245654, PP244940, PP246006, PP243860, PP241238, PP241845, PP243387, PP245534, PP247936, PP247335, OR885181, PP246976, PP240994, PP246363, PP249111, PP247100, PP248639, PP248524, PP248172, PP244343, PP246485, PP248051, PP249343, PP246120, PP243980, PP241962, PP243166, PP244583, PP246731, PP247575, PP247815, PP249466, PP249231, PP248869, PP248989, PP248748. C. princeps Tul. & Wedd.: C. P. Bove et al. 2211 (MICH, R), PP242924, PP245414, PP246243, PP244224, PP242320, PP242804, PP242201, PP243500, PP241357, PP245890, PP244822, PP241116, PP244104, PP243047, PP242084, PP245176, PP245297, PP246609, PP244703, PP241726, PP243617, PP244464, PP241603, PP242564, PP245776, —, PP245059, PP248286, PP247454, PP247696, PP248409, PP247219, PP246855, PP243738, PP241479, PP242441, PP242682, PP245655, PP244941, PP246007, PP243861, PP241239, PP241846, PP243388, PP245535, PP247937, PP247336, OR885182, PP246977, PP240995, PP246364, PP249112, PP247101, —, PP248525, —, PP244344, PP246486, PP248052, PP249344, PP246121, PP243981, PP241963, PP243167, PP244584, PP246732, PP247576, PP247816, PP249467, PP249232, PP248870, PP248990, PP248749.
Ceratolacis (Tul.) Wedd. C. pedunculatum C.T.Philbrick, Novelo & Irgang: C. P. Bove and C. T. Philbrick 2336 (MICH, R), PP242925, PP245415, PP246244, PP244225, PP242321, PP242805, PP242202, PP243501, PP241358, PP245891, PP244823, PP241117, PP244105, PP243048, PP242085, PP245177, PP245298, PP246610, PP244704, PP241727, PP243618, PP244465, PP241604, PP242565, PP245777, PP243284, PP245060, PP248287, PP247455, PP247697, PP248410, PP247220, PP246856, PP243739, PP241480, PP242442, PP242683, PP245656, PP244942, PP246008, PP243862, PP241240, PP241847, PP243389, PP245536, PP247938, PP247337, OR885183, PP246978, PP240996, PP246365, PP249113, PP247102, PP248640, PP248526, PP248173, PP244345, PP246487, PP248053, PP249345, PP246122, PP243982, PP241964, PP243168, PP244585, PP246733, PP247577, PP247817, PP249468, PP249233, PP248871, PP248991, PP248750; C. P. Bove et al. 2199 (MICH, R), PP242926, PP245416, PP246245, PP244226, PP242322, PP242806, PP242203, PP243502, PP241359, PP245892, PP244824, PP241118, PP244106, PP243049, PP242086, PP245178, PP245299, PP246611, PP244705, PP241728, PP243619, PP244466, PP241605, PP242566, PP245778, PP243285, PP245061, PP248288, PP247456, PP247698, PP248411, PP247221, PP246857, PP243740, PP241481, PP242443, PP242684, PP245657, PP244943, PP246009, PP243863, PP241241, PP241848, PP243390, PP245537, PP247939, PP247338, OR885184, PP246979, PP240997, PP246366, PP249114, PP247103, PP248641, PP248527, PP248174, PP244346, PP246488, PP248054, PP249346, PP246123, PP243983, PP241965, PP243169, PP244586, PP246734, PP247578, PP247818, PP249469, PP249234, PP248872, PP248992, PP248751.
Cipoia C.T.Philbrick, Novelo, and Irgang. C. inserta C.T.Philbrick, Novelo & Irgang: C. P. Bove et al. 2205 (MICH, R), PP242927, PP245417, PP246246, PP244227, PP242323, PP242807, PP242204, PP243503, PP241360, PP245893, PP244825, PP241119, PP244107, PP243050, PP242087, PP245179, PP245300, PP246612, PP244706, PP241729, PP243620, PP244467, PP241606, PP242567, PP245779, PP243286, PP245062, PP248289, PP247457, PP247699, PP248412, PP247222, PP246858, PP243741, PP241482, PP242444, PP242685, PP245658, PP244944, PP246010, PP243864, PP241242, PP241849, PP243391, PP245538, PP247940, PP247339, OR885185, PP246980, PP240998, PP246367, PP249115, PP247104, PP248642, PP248528, PP248175, PP244347, PP246489, PP248055, PP249347, PP246124, PP243984, PP241966, PP243170, PP244587, PP246735, PP247579, PP247819, PP249470, PP249235, PP248873, PP248993, PP248752. C. ramosa C.P.Bove: C.T.Philbrick & Novelo: C. P. Bove et al. 2251 (MICH, R), PP242928, PP245418, PP246247, PP244228, PP242324, PP242808, PP242205, PP243504, PP241361, PP245894, PP244826, PP241120, PP244108, PP243051, PP242088, PP245180, PP245301, PP246613, PP244707, PP241730, PP243621, PP244468, PP241607, PP242568, PP245780, PP243287, PP245063, PP248290, PP247458, PP247700, PP248413, PP247223, PP246859, PP243742, PP241483, PP242445, PP242686, PP245659, PP244945, PP246011, PP243865, PP241243, PP241850, PP243392, PP245539, PP247941, PP247340, OR885186, PP246981, PP240999, PP246368, PP249116, PP247105, —, PP248529, PP248176, PP244348, PP246490, PP248056, PP249348, PP246125, PP243985, PP241967, PP243171, PP244588, PP246736, PP247580, PP247820, PP249471, PP249236, PP248874, PP248994, PP248753.
Cladopus H.A.Möller. C. doianus (Koidz.) Koriba: S. Koi & N. Katayama jp404 (TNS), PP242929, PP245419, PP246248, PP244229, PP242325, PP242809, PP242206, PP243505, PP241362, PP245895, —, PP241121, PP244109, PP243052, PP242089, PP245181, —, PP246614, PP244708, PP241731, PP243622, PP244469, PP241608, PP242569, —, —, PP245064, PP248291, PP247459, PP247701, —, PP247224, PP246860, PP243743, PP241484, PP242446, PP242687, PP245660, PP244946, PP246012, PP243866, PP241244, PP241851, —, PP245540, —, PP247341, OR885187, PP246982, PP241000, PP246369, PP249117, PP247106, —, PP248530, —, PP244349, PP246491, PP248057, PP249349, PP246126, PP243986, PP241968, —, PP244589, PP246737, PP247581, PP247821, PP249472, PP249237, PP248875, PP248995, PP248754.
Dalzellia Wight. D. ceylanica (Gardner) Wight: M. Kato & N. Katayama sl101 (TNS), PP242930, PP245420, PP246249, PP244230, PP242326, PP242810, PP242207, —, PP241363, —, PP244827, PP241122, PP244110, —, PP242090, —, —, PP246615, PP244709, PP241732, PP243623, PP244470, PP241609, —, —, —, —, PP248292, PP247460, PP247702, —, —, PP246861, PP243744, PP241485, PP242447, PP242688, PP245661, PP244947, —, —, —, —, —, —, —, PP247342, OR885188, PP246983, PP241001, PP246370, —, —, —, —, —, —, PP246492, PP248058, PP249350, PP246127, PP243987, PP241969, —, —, PP246738, —, PP247822, PP249473, PP249238, —, PP248996, PP248755.
Devillea Tul. & Wedd. D. flagelliformis Tul. & Wedd.: C. P. Bove et al. 2368 (MICH, R), PP242973, PP245462, PP246291, PP244273, PP242368, PP242853, PP242248, PP243547, PP241405, PP245938, PP244870, PP241166, PP244152, PP243095, PP242132, PP245225, PP245343, PP246659, PP244751, PP241774, PP243664, PP244512, PP241653, PP242611, PP245821, PP243323, PP245106, PP248336, PP247503, PP247744, PP248456, PP247266, PP246903, PP243788, PP241529, PP242491, PP242731, PP245704, PP244988, PP246053, PP243909, PP241286, PP241893, PP243431, PP245582, PP247983, PP247385, OR885232, PP247027, PP241043, PP246413, PP249160, PP247149, PP248682, PP248570, PP248217, PP244392, PP246535, PP248101, PP249393, PP246170, PP244031, PP242011, PP243214, PP244632, PP246782, PP247623, PP247865, PP249516, PP249273, PP248918, PP249039, PP248798.
Diamantina Novelo, C.T.Philbrick & Irgang. D. lombardii Novelo, C.T.Philbrick & Irgang: C. P. Bove et al. 2131 (R), PP242931, PP245421, PP246250, PP244231, PP242327, PP242811, PP242208, PP243506, PP241364, PP245896, PP244828, PP241123, PP244111, PP243053, PP242091, PP245183, PP245302, PP246616, PP244710, PP241733, PP243624, PP244471, PP241610, PP242570, PP245781, PP243288, PP245065, PP248293, PP247461, PP247703, PP248414, PP247225, PP246862, PP243745, PP241486, PP242448, PP242689, PP245662, PP244948, PP246013, PP243867, PP241245, PP241852, PP243393, PP245541, PP247942, PP247343, OR885189, PP246984, PP241002, PP246371, PP249118, PP247107, PP248643, PP248531, PP248177, PP244350, PP246493, PP248059, PP249351, PP246128, PP243988, PP241970, PP243172, PP244590, PP246739, PP247582, PP247823, PP249474, —, PP248876, PP248997, PP248756; C. P. Bove et al. 2253 (MICH, R), PP242932, PP245422, PP246251, PP244232, PP242328, PP242812, PP242209, PP243507, PP241365, PP245897, PP244829, PP241124, PP244112, PP243054, PP242092, PP245184, PP245303, PP246617, PP244711, PP241734, PP243625, PP244472, PP241611, PP242571, PP245782, PP243289, PP245066, PP248294, PP247462, PP247704, PP248415, PP247226, PP246863, PP243746, PP241487, PP242449, PP242690, PP245663, PP244949, PP246014, PP243868, PP241246, PP241853, PP243394, PP245542, PP247943, PP247344, OR885190, PP246985, PP241003, PP246372, PP249119, PP247108, PP248644, PP248532, PP248178, PP244351, PP246494, PP248060, PP249352, PP246129, PP243989, PP241971, PP243173, PP244591, PP246740, PP247583, PP247824, PP249475, —, PP248877, PP248998, PP248757.
Dicraeanthus Engl. D. zehnderi H.Hess: J. Ghogue gho1650 (YA, Z/ ZT), PP242933, PP245423, PP246252, PP244233, PP242329, PP242813, PP242210, PP243508, PP241366, PP245898, PP244830, PP241125, PP244113, PP243055, PP242093, PP245185, PP245304, PP246618, PP244712, PP241735, PP243626, PP244473, PP241612, PP242572, PP245783, PP243290, PP245067, PP248295, PP247463, PP247705, PP248416, PP247227, PP246864, PP243747, PP241488, PP242450, PP242691, PP245664, PP244950, PP246015, PP243869, PP241247, PP241854, PP243395, PP245543, PP247944, PP247345, OR885191, PP246986, PP241004, PP246373, PP249120, PP247109, PP248645, PP248533, PP248179, PP244352, PP246495, PP248061, PP249353, PP246130, PP243990, PP241972, PP243174, PP244592, PP246741, PP247584, PP247825, PP249476, PP249239, PP248878, PP248999, PP248758.
Djinga C.Cusset. D. felicis C.Cusset: J. Ghogue et al. gar09 (YA, Z/ ZT), PP242934, PP245424, PP246253, PP244234, PP242330, PP242814, PP242211, PP243509, PP241367, PP245899, PP244831, PP241126, PP244114, PP243056, PP242094, PP245186, PP245305, PP246619, PP244713, PP241736, PP243627, PP244474, PP241613, PP242573, PP245784, —, PP245068, PP248296, PP247464, PP247706, PP248417, PP247228, PP246865, PP243748, PP241489, PP242451, PP242692, PP245665, PP244951, PP246016, PP243870, PP241248, PP241855, PP243396, PP245544, PP247945, PP247346, OR885192, PP246987, PP241005, PP246374, PP249121, PP247110, PP248646, PP248534, PP248180, PP244353, PP246496, PP248062, PP249354, PP246131, PP243991, PP241973, PP243175, PP244593, PP246742, PP247585, PP247826, PP249477, PP249240, PP248879, PP249000, PP248759.
Genus nov.: C. T. Philbrick et al. 6055 (MICH), PP242970, PP245458, PP246287, PP244269, PP242365, PP242849, PP242244, PP243544, PP241402, PP245934, PP244866, PP241162, PP244149, PP243091, PP242129, PP245221, PP245340, PP246655, PP244748, PP241771, PP243660, PP244509, PP241649, PP242608, PP245819, PP243321, PP245103, PP248332, PP247499, PP247741, PP248452, PP247263, PP246900, PP243784, PP241525, PP242487, PP242727, PP245701, PP244985, PP246051, PP243905, PP241283, PP241890, PP243427, PP245579, PP247980, PP247381, OR885228, PP247023, PP241040, PP246409, PP249156, PP247145, —, PP248567, PP248213, PP244388, PP246531, PP248097, PP249389, PP246166, PP244027, PP242008, PP243210, PP244628, PP246778, PP247620, PP247861, PP249512, PP249270, PP248914, PP249035, PP248794.
Hydrobryum Endl. H. japonicum Imamura: S. Koi & N. Katayama jp401 (TNS), PP242935, PP245425, PP246254, PP244235, PP242331, PP242815, PP242212, PP243510, PP241368, PP245900, PP244832, PP241127, PP244115, PP243057, PP242095, PP245187, PP245306, PP246620, PP244714, PP241737, PP243628, PP244475, PP241614, PP242574, PP245785, PP243291, PP245069, PP248297, PP247465, PP247707, PP248418, PP247229, PP246866, PP243749, PP241490, PP242452, PP242693, PP245666, PP244952, PP246017, PP243871, PP241249, PP241856, PP243397, PP245545, PP247946, PP247347, OR885193, PP246988, PP241006, PP246375, PP249122, PP247111, PP248647, PP248535, PP248181, PP244354, PP246497, PP248063, PP249355, PP246132, PP243992, PP241974, PP243176, PP244594, PP246743, PP247586, PP247827, PP249478, —, PP248880, PP249001, PP248760.
Inversodicraea Engl. ex R.E.Fr. I. bosii (C.Cusset) Rutish. & Thiv: J. Ghogue et al. gar01 (YA, Z/ZT), PP242936, PP245426, PP246255, PP244236, PP242332, PP242816, PP242213, PP243511, PP241369, PP245901, PP244833, PP241128, PP244116, PP243058, PP242096, PP245188, PP245307, PP246621, PP244715, PP241738, PP243629, PP244476, PP241615, PP242575, PP245786, PP243292, PP245070, PP248298, PP247466, PP247708, PP248419, PP247230, PP246867, PP243750, PP241491, PP242453, PP242694, PP245667, PP244953, PP246018, PP243872, PP241250, PP241857, PP243398, PP245546, PP247947, PP247348, OR885194, PP246989, PP241007, PP246376, PP249123, PP247112, PP248648, PP248536, PP248182, PP244355, PP246498, PP248064, PP249356, PP246133, PP243993, PP241975, PP243177, PP244595, PP246744, PP247587, PP247828, PP249479, PP249241, PP248881, PP249002, PP248761.
Ledermanniella Engl. L. bowlingii (J.B.Hall) C.Cusset: G. Ameka and R. Rutishauser ar021010 (GC, Z/ZT), PP242945, PP245434, PP246264, PP244245, PP242341, PP242825, PP242221, PP243520, PP241378, PP245910, PP244842, PP241137, PP244125, PP243067, PP242105, PP245197, PP245316, PP246630, PP244724, PP241747, PP243638, PP244485, PP241624, PP242584, PP245795, —, PP245079, PP248307, PP247475, PP247717, PP248428, PP247239, PP246876, PP243759, PP241500, PP242462, PP242703, PP245676, PP244961, PP246027, PP243881, PP241259, PP241866, —, PP245555, PP247956, PP247357, OR885203, PP246998, PP241016, PP246385, PP249132, PP247121, PP248657, PP248544, PP248190, PP244364, PP246507, PP248073, PP249365, PP246142, PP244002, PP241984, PP243186, PP244604, PP246753, PP247596, PP247837, PP249488, PP249250, PP248890, PP249011, PP248770. L. letouzeyi C.Cusset: J. Ghogue et al. gar12 (YA, Z/ZT), PP242946, PP245435, PP246265, PP244246, PP242342, PP242826, PP242222, PP243521, PP241379, PP245911, PP244843, PP241138, PP244126, PP243068, PP242106, PP245198, PP245317, PP246631, PP244725, PP241748, PP243639, PP244486, PP241625, PP242585, PP245796, —, PP245080, PP248308, PP247476, PP247718, PP248429, PP247240, PP246877, PP243760, PP241501, PP242463, PP242704, PP245677, PP244962, PP246028, PP243882, PP241260, PP241867, —, PP245556, PP247957, PP247358, OR885204, PP246999, PP241017, PP246386, PP249133, PP247122, —, PP248545, —, PP244365, PP246508, PP248074, PP249366, PP246143, PP244003, PP241985, PP243187, PP244605, PP246754, PP247597, PP247838, PP249489, PP249251, PP248891, PP249012, PP248771.
Lophogyne Tul. L. aeruginosa (P.Royen) C.T.Philbrick & C.P.Bove: C. T. Phlbrick and S. Jairam-Doerga 6214 (BBS, MICH), PP242949, PP245438, PP246268, PP244249, PP242345, PP242829, PP242225, PP243524, PP241382, PP245914, PP244846, PP241141, PP244129, PP243071, PP242109, PP245201, PP245320, PP246634, PP244728, PP241751, PP243642, PP244489, PP241628, PP242588, PP245799, PP243302, PP245083, PP248311, PP247479, PP247721, PP248432, PP247243, PP246880, PP243763, PP241504, PP242466, PP242707, PP245680, PP244965, PP246031, PP243885, PP241263, PP241870, —, PP245559, PP247960, PP247361, OR885207, PP247002, PP241020, PP246389, PP249136, PP247125, PP248660, PP248548, PP248193, PP244368, PP246511, PP248077, PP249369, PP246146, PP244006, PP241988, PP243190, PP244608, PP246757, PP247600, PP247841, PP249492, PP249253, PP248894, PP249015, PP248774. L. capillacea Pulle: C. P. Bove and C. T. Philbrick 2424 (MICH, R), PP242950, PP245439, PP246269, PP244250, PP242346, PP242830, PP242226, PP243525, PP241383, PP245915, PP244847, PP241142, PP244130, PP243072, PP242110, PP245202, PP245321, PP246635, PP244729, PP241752, PP243643, PP244490, PP241629, PP242589, PP245800, PP243303, PP245084, PP248312, PP247480, PP247722, PP248433, PP247244, PP246881, PP243764, PP241505, PP242467, PP242708, PP245681, PP244966, PP246032, PP243886, PP241264, PP241871, PP243408, PP245560, PP247961, PP247362, OR885208, PP247003, PP241021, PP246390, PP249137, PP247126, PP248661, PP248549, PP248194, PP244369, PP246512, PP248078, PP249370, PP246147, PP244007, PP241989, PP243191, PP244609, PP246758, PP247601, PP247842, PP249493, PP249254, PP248895, PP249016, PP248775; C. P. Bove and C. T. Philbrick 2441 (MICH, R), PP242951, PP245440, PP246270, PP244251, PP242347, PP242831, PP242227, PP243526, PP241384, PP245916, PP244848, PP241143, PP244131, PP243073, PP242111, PP245203, PP245322, PP246636, PP244730, PP241753, PP243644, PP244491, PP241630, PP242590, PP245801, PP243304, PP245085, PP248313, PP247481, PP247723, PP248434, PP247245, PP246882, PP243765, PP241506, PP242468, PP242709, PP245682, PP244967, PP246033, PP243887, PP241265, PP241872, PP243409, PP245561, PP247962, PP247363, OR885209, PP247004, PP241022, PP246391, PP249138, PP247127, PP248662, PP248550, PP248195, PP244370, PP246513, PP248079, PP249371, PP246148, PP244008, PP241990, PP243192, PP244610, PP246759, PP247602, PP247843, PP249494, PP249255, PP248896, PP249017, PP248776; C. P. Bove and C. T. Philbrick 2459 (MICH, R), PP242952, PP245441, PP246271, PP244252, PP242348, PP242832, PP242228, PP243527, PP241385, PP245917, PP244849, PP241144, PP244132, PP243074, PP242112, PP245204, PP245323, PP246637, PP244731, PP241754, PP243645, PP244492, PP241631, PP242591, PP245802, PP243305, PP245086, PP248314, PP247482, PP247724, PP248435, PP247246, PP246883, PP243766, PP241507, PP242469, PP242710, PP245683, PP244968, PP246034, PP243888, PP241266, PP241873, PP243410, PP245562, PP247963, PP247364, OR885210, PP247005, PP241023, PP246392, PP249139, PP247128, PP248663, PP248551, PP248196, PP244371, PP246514, PP248080, PP249372, PP246149, PP244009, PP241991, PP243193, PP244611, PP246760, PP247603, PP247844, PP249495, PP249256, PP248897, PP249018, PP248777; C. P. Bove et al. 2493 (R), MN165813. L. ceratophylla (Engl.) C.T.Philbrick & C.P.Bove: C. P. Bove and C. T. Philbrick 1864 (MICH, R), PP242937, PP245427, PP246256, PP244237, PP242333, PP242817, PP242214, PP243512, PP241370, PP245902, PP244834, PP241129, PP244117, PP243059, PP242097, PP245189, PP245308, PP246622, PP244716, PP241739, PP243630, PP244477, PP241616, PP242576, PP245787, PP243293, PP245071, PP248299, PP247467, PP247709, PP248420, PP247231, PP246868, PP243751, PP241492, PP242454, PP242695, PP245668, PP244954, PP246019, PP243873, PP241251, PP241858, PP243399, PP245547, PP247948, PP247349, OR885195, PP246990, PP241008, PP246377, PP249124, PP247113, PP248649, PP248537, PP248183, PP244356, PP246499, PP248065, PP249357, PP246134, PP243994, PP241976, PP243178, PP244596, PP246745, PP247588, PP247829, PP249480, PP249242, PP248882, PP249003, PP248762. L. fimbriata (P.Royen) C.T.Philbrick & C.P.Bove: C. T. Philbrick et al. 6204 (BBS, MICH), PP242938, PP245428, PP246257, PP244238, PP242334, PP242818, PP242215, PP243513, PP241371, PP245903, PP244835, PP241130, PP244118, PP243060, PP242098, PP245190, PP245309, PP246623, PP244717, PP241740, PP243631, PP244478, PP241617, PP242577, PP245788, PP243294, PP245072, PP248300, PP247468, PP247710, PP248421, PP247232, PP246869, PP243752, PP241493, PP242455, PP242696, PP245669, PP244955, PP246020, PP243874, PP241252, PP241859, PP243400, PP245548, PP247949, PP247350, OR885196, PP246991, PP241009, PP246378, PP249125, PP247114, PP248650, PP248538, PP248184, PP244357, PP246500, PP248066, PP249358, PP246135, PP243995, PP241977, PP243179, PP244597, PP246746, PP247589, PP247830, PP249481, PP249243, PP248883, PP249004, PP248763; C. T. Philbrick et al. 6313 (MICH, VEN), PP242939, PP245429, PP246258, PP244239, PP242335, PP242819, PP242216, PP243514, PP241372, PP245904, PP244836, PP241131, PP244119, PP243061, PP242099, PP245191, PP245310, PP246624, PP244718, PP241741, PP243632, PP244479, PP241618, PP242578, PP245789, PP243295, PP245073, PP248301, PP247469, PP247711, PP248422, PP247233, PP246870, PP243753, PP241494, PP242456, PP242697, PP245670, PP244956, PP246021, PP243875, PP241253, PP241860, PP243401, PP245549, PP247950, PP247351, OR885197, PP246992, PP241010, PP246379, PP249126, PP247115, PP248651, PP248539, PP248185, PP244358, PP246501, PP248067, PP249359, PP246136, PP243996, PP241978, PP243180, PP244598, PP246747, PP247590, PP247831, PP249482, PP249244, PP248884, PP249005, PP248764. L. fimbrifolia (P.Royen) C.T.Philbrick & C.P.Bove: C. P. Bove et al. 2234 (MICH, R), PP242884, PP245373, PP246202, PP244183, PP242279, PP242763, PP242160, PP243459, PP241316, PP245849, PP244781, PP241075, PP244063, PP243006, PP242043, PP245135, PP245256, PP246568, PP244662, PP241685, PP243576, PP244423, PP241562, PP242523, PP245735, PP243243, PP245018, PP248246, PP247414, PP247655, PP248368, PP247179, PP246814, PP243697, PP241438, PP242400, PP242641, PP245614, PP244900, PP245966, PP243820, PP241198, PP241805, PP243348, PP245494, PP247896, PP247295, OR885141, PP246936, PP240954, PP246323, PP249071, PP247060, PP248599, PP248484, PP248132, PP244303, PP246445, PP248012, PP249304, PP246080, PP243940, PP241922, PP243126, PP244543, PP246691, PP247535, PP247775, PP249426, PP249191, PP248829, PP248949, PP248708; C. T. Philbrick et al. 6004 (MICH, R), PP242885, PP245374, PP246203, PP244184, PP242280, PP242764, PP242161, PP243460, PP241317, PP245850, PP244782, PP241076, PP244064, PP243007, PP242044, PP245136, PP245257, PP246569, PP244663, PP241686, PP243577, PP244424, PP241563, PP242524, PP245736, PP243244, PP245019, PP248247, PP247415, PP247656, PP248369, PP247180, PP246815, PP243698, PP241439, PP242401, PP242642, PP245615, PP244901, PP245967, PP243821, PP241199, PP241806, PP243349, PP245495, PP247897, PP247296, OR885142, PP246937, PP240955, PP246324, PP249072, PP247061, PP248600, PP248485, PP248133, PP244304, PP246446, PP248013, PP249305, PP246081, PP243941, PP241923, PP243127, PP244544, PP246692, PP247536, PP247776, PP249427, PP249192, PP248830, PP248950, PP248709. L. lacunosa (Gardner) C.P.Bove & C.T.Philbrick: C. P. Bove and C. T. Philbrick 2258 (R), PP242947, PP245436, PP246266, PP244247, PP242343, PP242827, PP242223, PP243522, PP241380, PP245912, PP244844, PP241139, PP244127, PP243069, PP242107, PP245199, PP245318, PP246632, PP244726, PP241749, PP243640, PP244487, PP241626, PP242586, PP245797, PP243301, PP245081, PP248309, PP247477, PP247719, PP248430, PP247241, PP246878, PP243761, PP241502, PP242464, PP242705, PP245678, PP244963, PP246029, PP243883, PP241261, PP241868, PP243407, PP245557, PP247958, PP247359, OR885205, PP247000, PP241018, PP246387, PP249134, PP247123, PP248658, PP248546, PP248191, PP244366, PP246509, PP248075, PP249367, PP246144, PP244004, PP241986, PP243188, PP244606, PP246755, PP247598, PP247839, PP249490, PP249252, PP248892, PP249013, PP248772. L. royenella C.P.Bove & C.T.Philbrick: C. P. Bove et al. 2220 (MICH, R), PP242957, PP245445, PP246275, PP244256, PP242352, PP242836, PP242232, PP243531, PP241389, PP245921, PP244853, PP241149, PP244136, PP243078, PP242116, PP245208, PP245327, PP246642, PP244735, PP241758, PP243649, PP244496, PP241636, PP242595, PP245806, PP243309, PP245090, PP248319, PP247486, PP247728, PP248439, PP247250, PP246887, PP243771, PP241512, PP242474, PP242714, PP245688, PP244972, PP246038, PP243892, PP241270, PP241877, PP243414, PP245566, PP247967, PP247368, OR885215, PP247010, PP241027, PP246396, PP249143, PP247132, PP248667, PP248555, PP248200, PP244375, PP246519, PP248084, PP249376, PP246153, PP244014, PP241995, PP243197, PP244615, PP246765, PP247607, PP247848, PP249499, PP249261, PP248901, PP249022, PP248781; C. P. Bove et al. 2524 (MICH, R), MN165815; C. T. Philbrick et al. 5825 (MICH, R), PP242958, PP245446, PP246276, PP244257, PP242353, PP242837, PP242233, PP243532, PP241390, PP245922, PP244854, PP241150, PP244137, PP243079, PP242117, PP245209, PP245328, PP246643, PP244736, PP241759, PP243650, PP244497, PP241637, PP242596, PP245807, PP243310, PP245091, PP248320, PP247487, PP247729, PP248440, PP247251, PP246888, PP243772, PP241513, PP242475, PP242715, PP245689, PP244973, PP246039, PP243893, PP241271, PP241878, PP243415, PP245567, PP247968, PP247369, OR885216, PP247011, PP241028, PP246397, PP249144, PP247133, PP248668, PP248556, PP248201, PP244376, PP246520, PP248085, PP249377, PP246154, PP244015, PP241996, PP243198, PP244616, PP246766, PP247608, PP247849, PP249500, PP249262, PP248902, PP249023, PP248782. L. sp. 1: C. P. Bove et al. 2233 (R), PP242942, PP245431, PP246261, PP244242, PP242338, PP242822, PP242219, PP243517, PP241375, PP245907, PP244839, PP241134, PP244122, PP243064, PP242102, PP245194, PP245313, PP246627, PP244721, PP241744, PP243635, PP244482, PP241621, PP242581, PP245792, PP243298, PP245076, PP248304, PP247472, PP247714, PP248425, PP247236, PP246873, PP243756, PP241497, PP242459, PP242700, PP245673, PP244959, PP246024, PP243878, PP241256, PP241863, PP243404, PP245552, PP247953, PP247354, OR885200, PP246995, PP241013, PP246382, PP249129, PP247118, PP248654, PP248542, PP248188, PP244361, PP246504, PP248070, PP249362, PP246139, PP243999, PP241981, PP243183, PP244601, PP246750, PP247593, PP247834, PP249485, PP249247, PP248887, PP249008, PP248767. L. sp. 2: C. P. Bove et al. 2291 (MICH, R), PP242943, PP245432, PP246262, PP244243, PP242339, PP242823, —, PP243518, PP241376, PP245908, PP244840, PP241135, PP244123, PP243065, PP242103, PP245195, PP245314, PP246628, PP244722, PP241745, PP243636, PP244483, PP241622, PP242582, PP245793, PP243299, PP245077, PP248305, PP247473, PP247715, PP248426, PP247237, PP246874, PP243757, PP241498, PP242460, PP242701, PP245674, —, PP246025, PP243879, PP241257, PP241864, PP243405, PP245553, PP247954, PP247355, OR885201, PP246996, PP241014, PP246383, PP249130, PP247119, PP248655, —, —, PP244362, PP246505, PP248071, PP249363, PP246140, PP244000, PP241982, PP243184, PP244602, PP246751, PP247594, PP247835, PP249486, PP249248, PP248888, PP249009, PP248768; C. P. Bove et al. 2292 (R), PP242944, PP245433, PP246263, PP244244, PP242340, PP242824, PP242220, PP243519, PP241377, PP245909, PP244841, PP241136, PP244124, PP243066, PP242104, PP245196, PP245315, PP246629, PP244723, PP241746, PP243637, PP244484, PP241623, PP242583, PP245794, PP243300, PP245078, PP248306, PP247474, PP247716, PP248427, PP247238, PP246875, PP243758, PP241499, PP242461, PP242702, PP245675, PP244960, PP246026, PP243880, PP241258, PP241865, PP243406, PP245554, PP247955, PP247356, OR885202, PP246997, PP241015, PP246384, PP249131, PP247120, PP248656, PP248543, PP248189, PP244363, PP246506, PP248072, PP249364, PP246141, PP244001, PP241983, PP243185, PP244603, PP246752, PP247595, PP247836, PP249487, PP249249, PP248889, PP249010, PP248769. L. sp. 3: C. P. Bove and C. T. Philbrick 2413 (MICH, R), PP242941, PP245430, PP246260, PP244241, PP242337, PP242821, PP242218, PP243516, PP241374, PP245906, PP244838, PP241133, PP244121, PP243063, PP242101, PP245193, PP245312, PP246626, PP244720, PP241743, PP243634, PP244481, PP241620, PP242580, PP245791, PP243297, PP245075, PP248303, PP247471, PP247713, PP248424, PP247235, PP246872, PP243755, PP241496, PP242458, PP242699, PP245672, PP244958, PP246023, PP243877, PP241255, PP241862, PP243403, PP245551, PP247952, PP247353, OR885199, PP246994, PP241012, PP246381, PP249128, PP247117, PP248653, PP248541, PP248187, PP244360, PP246503, PP248069, PP249361, PP246138, PP243998, PP241980, PP243182, PP244600, PP246749, PP247592, PP247833, PP249484, PP249246, PP248886, PP249007, PP248766. L. wilsonii C.P.Bove & C.T.Philbrick: C. P. Bove and C. T. Philbrick 2474 (R), PP242940, —, PP246259, PP244240, PP242336, PP242820, PP242217, PP243515, PP241373, PP245905, PP244837, PP241132, PP244120, PP243062, PP242100, PP245192, PP245311, PP246625, PP244719, PP241742, PP243633, PP244480, PP241619, PP242579, PP245790, PP243296, PP245074, PP248302, PP247470, PP247712, PP248423, PP247234, PP246871, PP243754, PP241495, PP242457, PP242698, PP245671, PP244957, PP246022, PP243876, PP241254, PP241861, PP243402, PP245550, PP247951, PP247352, OR885198, PP246993, PP241011, PP246380, PP249127, PP247116, PP248652, PP248540, PP248186, PP244359, PP246502, PP248068, PP249360, PP246137, PP243997, PP241979, PP243181, PP244599, PP246748, PP247591, PP247832, PP249483, PP249245, PP248885, PP249006, PP248765.
Macropodiella Engl. M. heteromorpha (Baill.) C.Cusset: J. Ghogue et al. gahr24 (YA, Z/ZT), PP242948, PP245437, PP246267, PP244248, PP242344, PP242828, PP242224, PP243523, PP241381, PP245913, PP244845, PP241140, PP244128, PP243070, PP242108, PP245200, PP245319, PP246633, PP244727, PP241750, PP243641, PP244488, PP241627, PP242587, PP245798, —, PP245082, PP248310, PP247478, PP247720, PP248431, PP247242, PP246879, PP243762, PP241503, PP242465, PP242706, PP245679, PP244964, PP246030, PP243884, PP241262, PP241869, —, PP245558, PP247959, PP247360, OR885206, PP247001, PP241019, PP246388, PP249135, PP247124, PP248659, PP248547, PP248192, PP244367, PP246510, PP248076, PP249368, PP246145, PP244005, PP241987, PP243189, PP244607, PP246756, PP247599, PP247840, PP249491, —, PP248893, PP249014, PP248773.
Marathrum Bonpl. M. foeniculaceum Bonpl.: C. T. Philbrick and Ramey 6284 (EAP), PP242953, PP245442, PP246272, PP244253, PP242349, PP242833, PP242229, PP243528, PP241386, PP245918, PP244850, PP241145, PP244133, PP243075, PP242113, PP245205, PP245324, PP246638, PP244732, PP241755, PP243646, PP244493, PP241632, PP242592, PP245803, PP243306, PP245087, PP248315, PP247483, PP247725, PP248436, PP247247, PP246884, PP243767, PP241508, PP242470, PP242711, PP245684, PP244969, PP246035, PP243889, PP241267, PP241874, PP243411, PP245563, PP247964, PP247365, OR885211, PP247006, PP241024, PP246393, PP249140, PP247129, PP248664, PP248552, PP248197, PP244372, PP246515, PP248081, PP249373, PP246150, PP244010, PP241992, PP243194, PP244612, PP246761, PP247604, PP247845, PP249496, PP249257, PP248898, PP249019, PP248778. M. plumosum (Novelo & C.T.Philbrick) C.T.Philbrick & C.P.Bove: C. T. Philbrick 6264 (MICH), PP242954, PP245443, PP246273, PP244254, PP242350, PP242834, PP242230, PP243529, PP241387, PP245919, PP244851, PP241146, PP244134, PP243076, PP242114, PP245206, PP245325, PP246639, PP244733, PP241756, PP243647, PP244494, PP241633, PP242593, PP245804, PP243307, PP245088, PP248316, PP247484, PP247726, PP248437, PP247248, PP246885, PP243768, PP241509, PP242471, PP242712, PP245685, PP244970, PP246036, PP243890, PP241268, PP241875, PP243412, PP245564, PP247965, PP247366, OR885212, PP247007, PP241025, PP246394, PP249141, PP247130, PP248665, PP248553, PP248198, PP244373, PP246516, PP248082, PP249374, PP246151, PP244011, PP241993, PP243195, PP244613, PP246762, PP247605, PP247846, PP249497, PP249258, PP248899, PP249020, PP248779. M. utile Tul.: A. M. Bedoya AMB497 (ANDES), MN165814; C. T. Philbrick et al. 5886 (MICH, VEN), PP242955, PP245444, PP246274, PP244255, PP242351, PP242835, PP242231, PP243530, PP241388, PP245920, PP244852, PP241147, PP244135, PP243077, PP242115, PP245207, PP245326, PP246640, PP244734, PP241757, PP243648, PP244495, PP241634, PP242594, PP245805, PP243308, PP245089, PP248317, PP247485, PP247727, PP248438, PP247249, PP246886, PP243769, PP241510, PP242472, PP242713, PP245686, PP244971, PP246037, PP243891, PP241269, PP241876, PP243413, PP245565, PP247966, PP247367, OR885213, PP247008, PP241026, PP246395, PP249142, PP247131, PP248666, PP248554, PP248199, PP244374, PP246517, PP248083, PP249375, PP246152, PP244012, PP241994, PP243196, PP244614, PP246763, PP247606, PP247847, PP249498, PP249259, PP248900, PP249021, PP248780.
Monandriella Engl. M. linearifolia Engl.: J. Ghogue gho1663 (YA, Z/ ZT), PP242956, —, —, —, —, —, —, —, —, —, —, PP241148, —, —, —, —, —, PP246641, —, —, —, —, PP241635, —, —, —, —, PP248318, —, —, —, —, —, PP243770, PP241511, PP242473, —, PP245687, —, —, —, —, —, —, —, —, —, OR885214, PP247009, —, —, —, —, —, —, —, —, PP246518, —, —, —, PP244013, —, —, —, PP246764, —, —, —, PP249260, —, —, —.
Mourera Aubl. M. alcicornis (Tul.) P.Royen: C. P. Bove and C. T. Philbrick 1870 (MICH, R), PP242959, PP245447, PP246277, PP244258, PP242354, PP242838, PP242234, PP243533, PP241391, PP245923, PP244855, PP241151, PP244138, PP243080, PP242118, PP245210, PP245329, PP246644, PP244737, PP241760, PP243651, PP244498, PP241638, PP242597, PP245808, PP243311, PP245092, PP248321, PP247488, PP247730, PP248441, PP247252, PP246889, PP243773, PP241514, PP242476, PP242716, PP245690, PP244974, PP246040, PP243894, PP241272, PP241879, PP243416, PP245568, PP247969, PP247370, OR885217, PP247012, PP241029, PP246398, PP249145, PP247134, PP248669, PP248557, PP248202, PP244377, PP246521, PP248086, PP249378, PP246155, PP244016, PP241997, PP243199, PP244617, PP246767, PP247609, PP247850, PP249501, PP249263, PP248903, PP249024, PP248783; M. aspera (Raeusch.) Tul.: C. P. Bove et al. 2260 (MICH, R), PP242960, PP245448, PP246278, PP244259, PP242355, PP242839, PP242235, PP243534, PP241392, PP245924, PP244856, PP241152, PP244139, PP243081, PP242119, PP245211, PP245330, PP246645, PP244738, PP241761, PP243652, PP244499, PP241639, PP242598, PP245809, —, PP245093, PP248322, PP247489, PP247731, PP248442, PP247253, PP246890, PP243774, PP241515, PP242477, PP242717, PP245691, PP244975, PP246041, PP243895, PP241273, PP241880, PP243417, PP245569, PP247970, PP247371, OR885218, PP247013, PP241030, PP246399, PP249146, PP247135, PP248670, PP248558, PP248203, PP244378, PP246522, PP248087, PP249379, PP246156, PP244017, PP241998, PP243200, PP244618, PP246768, PP247610, PP247851, PP249502, PP249264, PP248904, PP249025, PP248784. M. elegans (Tul.) Baillon: C. P. Bove and C. T. Philbrick 1899 (MICH, R), PP242961, PP245449, PP246279, PP244260, PP242356, PP242840, PP242236, PP243535, PP241393, PP245925, PP244857, PP241153, PP244140, PP243082, PP242120, PP245212, PP245331, PP246646, PP244739, PP241762, PP243653, PP244500, PP241640, PP242599, PP245810, PP243312, PP245094, PP248323, PP247490, PP247732, PP248443, PP247254, PP246891, PP243775, PP241516, PP242478, PP242718, PP245692, PP244976, PP246042, PP243896, PP241274, PP241881, PP243418, PP245570, PP247971, PP247372, OR885219, PP247014, PP241031, PP246400, PP249147, PP247136, PP248671, PP248559, PP248204, PP244379, PP246523, PP248088, PP249380, PP246157, PP244018, PP241999, PP243201, PP244619, PP246769, PP247611, PP247852, PP249503, PP249265, PP248905, PP249026, PP248785. M. fluviatilis Aubl.: C. P. Bove et al. 1955 (MICH, R), PP242962, PP245450, PP246280, PP244261, PP242357, PP242841, PP242237, PP243536, PP241394, PP245926, PP244858, PP241154, PP244141, PP243083, PP242121, PP245213, PP245332, PP246647, PP244740, PP241763, PP243654, PP244501, PP241641, PP242600, PP245811, PP243313, PP245095, PP248324, PP247491, PP247733, PP248444, PP247255, PP246892, PP243776, PP241517, PP242479, PP242719, PP245693, PP244977, PP246043, PP243897, PP241275, PP241882, PP243419, PP245571, PP247972, PP247373, OR885220, PP247015, PP241032, PP246401, PP249148, PP247137, PP248672, PP248560, PP248205, PP244380, PP246524, PP248089, PP249381, PP246158, PP244019, PP242000, PP243202, PP244620, PP246770, PP247612, PP247853, PP249504, PP249266, PP248906, PP249027, PP248786.
Noveloa C.T.Philbrick. N. coulteriana (Tul.) C.T.Philbrick: C. T. Philbrick 6329 (MICH, UAT), PP242964, PP245452, PP246282, PP244263, PP242359, PP242843, PP242239, PP243538, PP241396, PP245928, PP244860, PP241156, PP244143, PP243085, PP242123, PP245215, PP245334, PP246649, PP244742, PP241765, PP243656, PP244503, PP241643, PP242602, PP245813, PP243315, PP245097, PP248326, PP247493, PP247735, PP248446, PP247257, PP246894, PP243778, PP241519, PP242481, PP242721, PP245695, PP244979, PP246045, PP243899, PP241277, PP241884, PP243421, PP245573, PP247974, PP247375, OR885222, PP247017, PP241034, PP246403, PP249150, PP247139, PP248674, PP248562, PP248207, PP244382, PP246525, PP248091, PP249383, PP246160, PP244021, PP242002, PP243204, PP244622, PP246772, PP247614, PP247855, PP249506, PP249268, PP248908, PP249029, PP248788; C. T. Philbrick et al. 6270 (MICH), PP242963, PP245451, PP246281, PP244262, PP242358, PP242842, PP242238, PP243537, PP241395, PP245927, PP244859, PP241155, PP244142, PP243084, PP242122, PP245214, PP245333, PP246648, PP244741, PP241764, PP243655, PP244502, PP241642, PP242601, PP245812, PP243314, PP245096, PP248325, PP247492, PP247734, PP248445, PP247256, PP246893, PP243777, PP241518, PP242480, PP242720, PP245694, PP244978, PP246044, PP243898, PP241276, PP241883, PP243420, PP245572, PP247973, PP247374, OR885221, PP247016, PP241033, PP246402, PP249149, PP247138, PP248673, PP248561, PP248206, PP244381, —, PP248090, PP249382, PP246159, PP244020, PP242001, PP243203, PP244621, PP246771, PP247613, PP247854, PP249505, PP249267, PP248907, PP249028, PP248787. N. longifolia (Novelo & C.T.Philbrick) C.T.Philbrick: C. T. Philbrick 6340 (MICH, UAT), PP242965, PP245453, PP246283, PP244264, PP242360, PP242844, PP242240, PP243539, PP241397, PP245929, PP244861, PP241157, PP244144, PP243086, PP242124, PP245216, PP245335, PP246650, PP244743, PP241766, PP243657, PP244504, PP241644, PP242603, PP245814, PP243316, PP245098, PP248327, PP247494, PP247736, PP248447, PP247258, PP246895, PP243779, PP241520, PP242482, PP242722, PP245696, PP244980, PP246046, PP243900, PP241278, PP241885, PP243422, PP245574, PP247975, PP247376, OR885223, PP247018, PP241035, PP246404, PP249151, PP247140, PP248675, PP248563, PP248208, PP244383, PP246526, PP248092, PP249384, PP246161, PP244022, PP242003, PP243205, PP244623, PP246773, PP247615, PP247856, PP249507, PP249269, PP248909, PP249030, PP248789.
Oserya Tul & Wedd. O. perpusilla (Went) P.Royen: C. P. Bove and C. T. Philbrick 2448 (MICH, R), PP242966, PP245454, PP246284, PP244265, PP242361, PP242845, PP242241, PP243540, PP241398, PP245930, PP244862, PP241158, PP244145, PP243087, PP242125, PP245217, PP245336, PP246651, PP244744, PP241767, PP243658, PP244505, PP241645, PP242604, PP245815, PP243317, PP245099, PP248328, PP247495, PP247737, PP248448, PP247259, PP246896, PP243780, PP241521, PP242483, PP242723, PP245697, PP244981, PP246047, PP243901, PP241279, PP241886, PP243423, PP245575, PP247976, PP247377, OR885224, PP247019, PP241036, PP246405, PP249152, PP247141, PP248676, PP248564, PP248209, PP244384, PP246527, PP248093, PP249387, PP246162, PP244023, PP242004, PP243206, PP244624, PP246774, PP247616, PP247857, PP249508, —, PP248910, PP249031, PP248790; C. P. Bove et al. 1946 (MICH, R), PP242967, PP245455, PP246285, PP244266, PP242362, PP242846, PP242242, PP243541, PP241399, PP245931, PP244863, PP241159, PP244146, PP243088, PP242126, PP245218, PP245337, PP246652, PP244745, PP241768, —, PP244506, PP241646, PP242605, PP245816, PP243318, PP245100, PP248329, PP247496, PP247738, PP248449, PP247260, PP246897, PP243781, PP241522, PP242484, PP242724, PP245698, PP244982, PP246048, PP243902, PP241280, PP241887, PP243424, PP245576, PP247977, PP247378, OR885225, PP247020, PP241037, PP246406, PP249153, PP247142, PP248677, PP248565, PP248210, PP244385, PP246528, PP248094, PP249385, PP246163, PP244024, PP242005, PP243207, PP244625, PP246775, PP247617, PP247858, PP249509, —, PP248911, PP249032, PP248791; C. P. Bove et al. 2327 (R), PP242971, PP245459, PP246288, PP244270, PP242366, PP242850, PP242245, PP243545, PP241403, PP245935, PP244867, PP241163, PP244150, PP243092, PP242130, PP245222, PP245341, PP246656, PP244749, PP241772, PP243661, PP244510, PP241650, PP242609, PP245820, PP243322, PP245104, PP248333, PP247500, PP247742, PP248453, PP247264, PP246901, PP243785, PP241526, PP242488, PP242728, PP245702, PP244986, PP246052, PP243906, PP241284, PP241891, PP243428, PP245580, PP247981, PP247382, OR885229, PP247024, PP241041, PP246410, PP249157, PP247146, PP248680, PP248568, PP248214, PP244389, PP246532, PP248098, PP249390, PP246167, PP244028, PP242009, PP243211, PP244629, PP246779, PP247621, PP247862, PP249513, —, PP248915, PP249036, PP248795; C. T. Philbrick 6190 (BBS, MICH), PP242969, PP245457, PP246286, PP244268, PP242364, PP242848, PP242243, PP243543, PP241401, PP245933, PP244865, PP241161, PP244148, PP243090, PP242128, PP245220, PP245339, PP246654, PP244747, PP241770, PP243659, PP244508, PP241648, PP242607, PP245818, PP243320, PP245102, PP248331, PP247498, PP247740, PP248451, PP247262, PP246899, PP243783, PP241524, PP242486, PP242726, PP245700, PP244984, PP246050, PP243904, PP241282, PP241889, PP243426, PP245578, PP247979, PP247380, OR885227, PP247022, PP241039, PP246408, PP249155, PP247144, PP248679, PP248566, PP248212, PP244387, PP246530, PP248096, PP249388, PP246165, PP244026, PP242007, PP243209, PP244627, PP246777, PP247619, PP247860, PP249511, —, PP248913, PP249034, PP248793; C. T. Philbrick et al. 6021 (MICH, VEN), PP242968, PP245456, —, PP244267, PP242363, PP242847, —, PP243542, PP241400, PP245932, PP244864, PP241160, PP244147, PP243089, PP242127, PP245219, PP245338, PP246653, PP244746, PP241769, —, PP244507, PP241647, PP242606, PP245817, PP243319, PP245101, PP248330, PP247497, PP247739, PP248450, PP247261, PP246898, PP243782, PP241523, PP242485, PP242725, PP245699, PP244983, PP246049, PP243903, PP241281, PP241888, PP243425, PP245577, PP247978, PP247379, OR885226, PP247021, PP241038, PP246407, PP249154, PP247143, PP248678, —, PP248211, PP244386, PP246529, PP248095, PP249386, PP246164, PP244025, PP242006, PP243208, PP244626, PP246776, PP247618, PP247859, PP249510, —, PP248912, PP249033, PP248792. O. pilgeri (Mildbr.) C.T. Philbrick & C.P. Bove: C. P. Bove et al. 2520 (MICH, R), PP242902, PP245391, PP246220, PP244201, PP242297, PP242781, PP242178, PP243477, PP241334, PP245867, PP244799, PP241093, PP244081, PP243024, PP242061, PP245153, PP245274, PP246586, PP244680, PP241703, PP243594, PP244441, PP241580, PP242541, PP245753, PP243261, PP245036, PP248263, PP247431, PP247673, PP248386, PP247196, PP246832, PP243715, PP241456, PP242418, PP242659, PP245632, PP244918, PP245984, PP243838, PP241216, PP241823, PP243365, PP245512, PP247914, PP247313, OR885159, PP246954, PP240972, PP246341, PP249089, PP247078, PP248617, PP248502, PP248150, PP244321, PP246463, PP248029, PP249321, PP246098, PP243958, PP241940, PP243144, PP244561, PP246709, PP247553, PP247793, PP249444, PP249209, PP248847, PP248967, PP248726.
Paracladopus M.Kato. P. chantaburiensis Koi & M.Kato: S. Koi et al. tkf24 (TNS), PP242972, PP245460, PP246289, PP244271, PP242367, PP242851, PP242246, —, PP241404, PP245936, PP244868, PP241164, PP244151, PP243093, PP242131, PP245223, PP245342, PP246657, PP244750, PP241773, PP243662, PP244511, PP241651, PP242610, —, —, PP245105, PP248334, PP247501, PP247743, PP248454, PP247265, PP246902, PP243786, PP241527, PP242489, PP242729, PP245703, PP244987, —, PP243907, PP241285, PP241892, PP243429, PP245581, PP247982, PP247383, OR885230, PP247025, PP241042, PP246411, PP249158, PP247147, —, PP248569, PP248215, PP244390, PP246533, PP248099, PP249391, PP246168, PP244029, PP242010, PP243212, PP244630, PP246780, PP247622, PP247863, PP249514, PP249271, PP248916, PP249037, PP248796.
Podostemum Michx. P. ceratophyllum Michx.: B. Ruhfel and Z. Xi 128 (A), JX662415, JX663820, JX663738, JX664438, JX662930, JX665023, JX662296, JX664827, JX662893, JX661961, JX663649, JX664316, JX661833, JX664358, JX664639, JX662770, JX663427, JX662818, JX662082, JX662728, JX664553, JX664107, JX663934, JX663008, JX662449, JX662482, JX663558, JX664226, JX662972, JX663121, JX664181, JX664674, JX663298, JX662213, JX662596, JX662336, JX664510, JX663856, JX663213, JX663889, JX664022, JX661920, JX662254, JX661794, JX662639, JX664597, JX664143, JX664067, JX663981, JX664271, JX663384, JX662558, JX662857, JX663455, —, JX663050, JX662039, JX663495, JX664958, JX662683, JX664790, JX663601, JX661876, JX663530, JX663085, JX662377, JX663168, JX663781, JX664870, JX664913, JX663340, JX664475, JX663254. P. comatum Hicken: C. T. Philbrick et al. 5343 (MICH, R), —, PP245461, PP246290, PP244272, —, PP242852, PP242247, PP243546, —, PP245937, PP244869, PP241165, —, PP243094, —, PP245224, —, PP246658, —, —, PP243663, —, PP241652, —, —, —, —, PP248335, PP247502, —, PP248455, —, —, PP243787, PP241528, PP242490, PP242730, —, —, —, PP243908, —, —, PP243430, —, —, PP247384, OR885231, PP247026, —, PP246412, PP249159, PP247148, PP248681, —, PP248216, PP244391, PP246534, PP248100, PP249392, PP246169, PP244030, —, PP243213, PP244631, PP246781, —, PP247864, PP249515, PP249272, PP248917, PP249038, PP248797. P. ovatum C.T.Philbrick & Novelo: Bove et al. 1124 (R), PP242974, PP245463, PP246292, PP244274, PP242369, PP242854, PP242249, PP243548, PP241406, PP245939, PP244871, PP241167, PP244153, PP243096, PP242133, PP245226, PP245344, PP246660, PP244752, PP241775, PP243665, PP244513, PP241654, PP242612, PP245822, PP243324, PP245107, PP248337, PP247504, PP247745, PP248457, PP247267, PP246904, PP243789, PP241530, PP242492, PP242732, PP245705, PP244989, PP246054, PP243910, PP241287, PP241894, PP243432, PP245583, PP247984, PP247386, OR885233, PP247028, PP241044, PP246414, PP249161, PP247150, PP248683, PP248571, PP248218, PP244393, PP246536, PP248102, PP249394, PP246171, PP244032, PP242012, PP243215, PP244633, PP246783, PP247624, PP247866, PP249517, PP249274, PP248919, PP249040, PP248799. P. scaturiginum (Mart.) C.T.Philbrick & Novelo: C. P. Bove and C. T. Philbrick 2310 (MICH, R), PP242975, PP245464, PP246293, PP244275, PP242370, PP242855, PP242250, PP243549, PP241407, PP245940, PP244872, PP241168, PP244154, PP243097, PP242134, PP245227, PP245345, PP246661, PP244753, PP241776, PP243666, PP244514, PP241655, PP242613, PP245823, PP243325, PP245108, PP248338, PP247505, PP247746, PP248458, PP247268, PP246905, PP243790, PP241531, PP242493, PP242733, PP245706, PP244990, PP246055, PP243911, PP241288, PP241895, PP243433, PP245584, PP247985, PP247387, OR885234, PP247029, PP241045, PP246415, PP249162, PP247151, PP248684, PP248572, PP248219, PP244394, PP246537, PP248103, PP249395, PP246172, PP244033, PP242013, PP243216, PP244634, PP246784, PP247625, PP247867, PP249518, PP249275, PP248920, PP249041, PP248800. P. weddellianum (Tul.) C.T.Philbrick & Novelo: C. P. Bove 1130 (R), PP242976, PP245465, PP246294, —, PP242371, PP242856, PP242251, —, PP241408, —, PP244873, PP241169, PP244155, PP243098, —, PP245228, PP245346, PP246662, —, PP241777, PP243667, PP244515, PP241656, —, —, —, —, PP248339, PP247506, PP247747, —, —, PP246906, PP243791, PP241532, PP242494, PP242734, PP245707, —, —, —, PP241289, —, —, PP245585, —, —, OR885235, PP247030, PP241046, PP246416, PP249163, —, —, —, —, —, PP246538, PP248104, PP249396, —, PP244034, PP242014, PP243217, —, PP246785, PP247626, PP247868, PP249519, PP249276, PP248921, PP249042, PP248801.
Polypleurum (Tul.) Warm. P. stylosum (Wight) J.B.Hall: M. Kato & N. Katayama sl103 (TNS), PP242977, PP245466, PP246295, PP244276, PP242372, —, PP242252, —, PP241409, —, —, PP241170, —, PP243099, —, PP245229, —, PP246663, PP244754, PP241778, PP243668, PP244516, PP241657, —, —, —, —, PP248340, PP247507, PP247748, —, —, PP246907, PP243792, PP241533, PP242495, PP242735, PP245708, PP244991, —, PP243912, —, —, —, PP245586, —, —, OR885236, PP247031, PP241047, PP246417, —, PP247152, —, —, —, PP244395, PP246539, PP248105, PP249397, PP246173, PP244035, PP242015, —, —, PP246786, PP247627, PP247869, PP249520, PP249277, —, PP249043, PP248802.
Rhyncholacis Tul. R. apiculata P.Royen: C. P. Bove and C. T. Philbrick 2466 (MICH, R), PP242983, PP245472, PP246301, PP244282, PP242378, PP242862, PP242258, PP243555, PP241415, PP245946, PP244879, PP241176, PP244161, PP243105, PP242140, PP245235, PP245352, PP246669, PP244760, PP241784, PP243674, PP244522, PP241663, PP242619, PP245829, PP243331, PP245114, PP248346, PP247513, PP247754, PP248464, PP247274, PP246913, PP243798, PP241539, PP242501, PP242741, PP245714, PP244997, PP246061, PP243918, PP241295, PP241901, PP243439, PP245592, PP247991, PP247393, OR885242, PP247037, PP241053, PP246423, PP249169, PP247158, PP248690, PP248578, PP248225, PP244401, PP246545, PP248111, PP249403, PP246179, PP244041, PP242021, PP243223, PP244640, PP246792, PP247633, PP247875, PP249526, PP249283, PP248927, PP249049, PP248808. R. applanata K.I. Goebel.: C. T. Philbrick et al. 6046 (MICH, VEN), PP242982, PP245471, PP246300, PP244281, PP242377, PP242861, PP242257, PP243554, PP241414, PP245945, PP244878, PP241175, PP244160, PP243104, PP242139, PP245234, PP245351, PP246668, PP244759, PP241783, PP243673, PP244521, PP241662, PP242618, PP245828, PP243330, PP245113, PP248345, PP247512, PP247753, PP248463, PP247273, PP246912, PP243797, PP241538, PP242500, PP242740, PP245713, PP244996, PP246060, PP243917, PP241294, PP241900, PP243438, PP245591, PP247990, PP247392, OR885241, PP247036, PP241052, PP246422, PP249168, PP247157, PP248689, PP248577, PP248224, PP244400, PP246544, PP248110, PP249402, PP246178, PP244040, PP242020, PP243222, PP244639, PP246791, PP247632, PP247874, PP249525, PP249282, PP248926, PP249048, PP248807. R. hydrocichorium Tul.: C. P. Bove and C. T. Philbrick 2440 (MICH, R), PP242984, PP245473, PP246302, PP244283, PP242379, PP242863, PP242259, PP243556, PP241416, PP245947, PP244880, PP241177, PP244162, PP243106, PP242141, PP245236, PP245353, PP246670, PP244761, PP241785, PP243675, PP244523, PP241664, PP242620, PP245830, PP243332, PP245115, PP248347, PP247514, PP247755, PP248465, PP247275, PP246914, PP243799, PP241540, PP242502, PP242742, PP245715, PP244998, PP246062, PP243919, PP241296, PP241902, PP243440, PP245593, PP247992, PP247394, OR885243, PP247038, PP241054, PP246424, PP249170, PP247159, PP248691, PP248579, PP248226, PP244402, PP246546, PP248112, PP249404, PP246180, PP244042, PP242022, PP243224, PP244641, PP246793, PP247634, PP247876, PP249527, PP249284, PP248928, PP249050, PP248809. R. jenmanii Engl. forma laciniata P. Royen: C. T. Philbrick et al. 6052 (MICH, VEN), PP242978, PP245467, PP246296, PP244277, PP242373, PP242857, PP242253, PP243550, PP241410, PP245941, PP244874, PP241171, PP244156, PP243100, PP242135, PP245230, PP245347, PP246664, PP244755, PP241779, PP243669, PP244517, PP241658, PP242614, PP245824, PP243326, PP245109, PP248341, PP247508, PP247749, PP248459, PP247269, PP246908, PP243793, PP241534, PP242496, PP242736, PP245709, PP244992, PP246056, PP243913, PP241290, PP241896, PP243434, PP245587, PP247986, PP247388, OR885237, PP247032, PP241048, PP246418, PP249164, PP247153, PP248685, PP248573, PP248220, PP244396, PP246540, PP248106, PP249398, PP246174, PP244036, PP242016, PP243218, PP244635, PP246787, PP247628, PP247870, PP249521, PP249278, PP248922, PP249044, PP248803; C. T. Philbrick et al. 6052A (MICH, VEN), PP242979, PP245468, PP246297, PP244278, PP242374, PP242858, PP242254, PP243551, PP241411, PP245942, PP244875, PP241172, PP244157, PP243101, PP242136, PP245231, PP245348, PP246665, PP244756, PP241780, PP243670, PP244518, PP241659, PP242615, PP245825, PP243327, PP245110, PP248342, PP247509, PP247750, PP248460, PP247270, PP246909, PP243794, PP241535, PP242497, PP242737, PP245710, PP244993, PP246057, PP243914, PP241291, PP241897, PP243435, PP245588, PP247987, PP247389, OR885238, PP247033, PP241049, PP246419, PP249165, PP247154, PP248686, PP248574, PP248221, PP244397, PP246541, PP248107, PP249399, PP246175, PP244037, PP242017, PP243219, PP244636, PP246788, PP247629, PP247871, PP249522, PP249279, PP248923, PP249045, PP248804; C. T. Philbrick et al. 6053 (MICH, VEN), PP242980, PP245469, PP246298, PP244279, PP242375, PP242859, PP242255, PP243552, PP241412, PP245943, PP244876, PP241173, PP244158, PP243102, PP242137, PP245232, PP245349, PP246666, PP244757, PP241781, PP243671, PP244519, PP241660, PP242616, PP245826, PP243328, PP245111, PP248343, PP247510, PP247751, PP248461, PP247271, PP246910, PP243795, PP241536, PP242498, PP242738, PP245711, PP244994, PP246058, PP243915, PP241292, PP241898, PP243436, PP245589, PP247988, PP247390, OR885239, PP247034, PP241050, PP246420, PP249166, PP247155, PP248687, PP248575, PP248222, PP244398, PP246542, PP248108, PP249400, PP246176, PP244038, PP242018, PP243220, PP244637, PP246789, PP247630, PP247872, PP249523, PP249280, PP248924, PP249046, PP248805; C. T. Philbrick et al. 6060 (MICH, R), PP242981, PP245470, PP246299, PP244280, PP242376, PP242860, PP242256, PP243553, PP241413, PP245944, PP244877, PP241174, PP244159, PP243103, PP242138, PP245233, PP245350, PP246667, PP244758, PP241782, PP243672, PP244520, PP241661, PP242617, PP245827, PP243329, PP245112, PP248344, PP247511, PP247752, PP248462, PP247272, PP246911, PP243796, PP241537, PP242499, PP242739, PP245712, PP244995, PP246059, PP243916, PP241293, PP241899, PP243437, PP245590, PP247989, PP247391, OR885240, PP247035, PP241051, PP246421, PP249167, PP247156, PP248688, PP248576, PP248223, PP244399, PP246543, PP248109, PP249401, PP246177, PP244039, PP242019, PP243221, PP244638, PP246790, PP247631, PP247873, PP249524, PP249281, PP248925, PP249047, PP248806. R. paulana C.T. Philbrick & C.P. Bove: C. P. Bove and C. T. Philbrick 2465 (MICH, R), PP242986, PP245475, PP246304, PP244285, PP242381, PP242865, PP242261, PP243558, PP241418, PP245949, PP244882, PP241179, PP244164, PP243108, PP242143, PP245238, PP245355, PP246672, PP244763, PP241787, PP243677, PP244525, PP241666, PP242622, PP245832, PP243334, PP245117, PP248349, PP247516, PP247757, PP248467, PP247277, PP246916, PP243801, PP241542, PP242504, PP242744, PP245717, PP245000, PP246064, PP243921, PP241298, PP241904, PP243442, PP245595, PP247994, PP247396, OR885245, PP247040, PP241056, PP246426, PP249172, PP247161, PP248693, PP248581, PP248228, PP244404, PP246548, PP248114, PP249406, PP246182, PP244044, PP242024, PP243226, PP244643, PP246795, PP247636, PP247878, PP249529, PP249286, PP248930, PP249052, PP248811. R. sp.: C. P. Bove and C. T. Philbrick 2421 (MICH, R), PP242985, PP245474, PP246303, PP244284, PP242380, PP242864, PP242260, PP243557, PP241417, PP245948, PP244881, PP241178, PP244163, PP243107, PP242142, PP245237, PP245354, PP246671, PP244762, PP241786, PP243676, PP244524, PP241665, PP242621, PP245831, PP243333, PP245116, PP248348, PP247515, PP247756, PP248466, PP247276, PP246915, PP243800, PP241541, PP242503, PP242743, PP245716, PP244999, PP246063, PP243920, PP241297, PP241903, PP243441, PP245594, PP247993, PP247395, OR885244, PP247039, PP241055, PP246425, PP249171, PP247160, PP248692, PP248580, PP248227, PP244403, PP246547, PP248113, PP249405, PP246181, PP244043, PP242023, PP243225, PP244642, PP246794, PP247635, PP247877, PP249528, PP249285, PP248929, PP249051, PP248810.
Stonesia G.Taylor. S. ghoguei E.Pfeifer & Rutish.: J. Ghogue gho1665 (YA, Z/ZT), PP242987, PP245476, PP246305, PP244286, PP242382, PP242866, PP242262, PP243559, PP241419, PP245950, PP244883, PP241180, PP244165, PP243109, PP242144, PP245239, PP245356, PP246673, PP244764, PP241788, PP243678, PP244526, PP241667, PP242623, PP245833, —, PP245118, PP248350, PP247517, PP247758, —, PP247278, PP246917, PP243802, PP241543, PP242505, PP242745, PP245718, PP245001, PP246065, PP243922, PP241299, PP241905, PP243443, PP245596, PP247995, PP247397, OR885246, PP247041, PP241057, PP246427, PP249173, PP247162, PP248694, PP248582, PP248229, PP244405, PP246549, PP248115, PP249407, PP246183, PP244045, PP242025, PP243227, PP244644, PP246796, PP247637, PP247879, PP249530, PP249287, PP248931, PP249053, PP248812.
Terniopsis H.C.Chao. T. brevis M.Kato: S. Koi et al. tkf25 (TNS), PP242988, PP245477, —, —, —, —, —, —, PP241420, —, —, PP241181, —, —, —, —, —, —, —, —, PP243679, —, —, —, —, —, —, PP248351, PP247518, —, —, —, PP246918, PP243803, PP241544, PP242506, —, —, —, —, —, —, —, —, —, —, —, OR885247, PP247042, —, —, —, —, —, —, —, —, PP246550, —, PP249408, PP246184, PP244046, PP242026, —, —, —, —, —, —, PP249288, —, —, —.
Thelethylax C.Cusset. T. minutiflora (Tul.) C.Cusset: N. Rakotonirina nr532 (MO), PP242989, PP245478, PP246306, PP244287, PP242383, PP242867, PP242263, PP243560, PP241421, PP245951, PP244884, PP241182, PP244166, PP243110, PP242145, PP245240, PP245357, PP246674, PP244765, PP241789, PP243680, PP244527, PP241668, PP242624, PP245834, PP243335, PP245119, PP248352, PP247519, PP247759, PP248468, PP247279, PP246919, PP243804, PP241545, PP242507, PP242746, PP245719, PP245002, PP246066, PP243923, PP241300, PP241906, PP243444, PP245597, PP247996, PP247398, OR885248, PP247043, PP241058, PP246428, PP249174, PP247163, PP248695, PP248583, PP248230, PP244406, PP246551, PP248116, PP249409, PP246185, PP244047, PP242027, PP243228, PP244645, PP246797, PP247638, PP247880, PP249531, PP249289, PP248932, PP249054, PP248813.
Tristicha Thouars. T. trifaria (Bory ex Willd.) Spreng.: A. Mesterhazy am128 (Z), MN165816; B. Ruhfel 19 (A), PP242990, PP245479, PP246307, PP244288, PP242384, PP242868, PP242264, PP243561, PP241422, —, PP244885, PP241183, PP244167, PP243111, PP242146, PP245241, PP245358, PP246675, PP244766, PP241790, PP243681, PP244528, PP241669, PP242625, PP245835, PP243336, PP245120, PP248353, PP247520, PP247760, PP248469, PP247280, PP246920, PP243805, PP241546, PP242508, PP242747, PP245720, PP245003, —, PP243924, PP241301, PP241907, PP243445, PP245598, PP247997, PP247399, OR885249, PP247044, PP241059, PP246429, PP249175, PP247164, PP248696, PP248584, PP248231, PP244407, PP246552, PP248117, PP249410, PP246186, PP244048, PP242028, PP243229, PP244646, PP246798, PP247639, PP247881, PP249532, PP249290, PP248933, PP249055, PP248814; C. P. Bove and C. T. Philbrick 1867 (R), PP242991, PP245480, PP246308, PP244289, PP242385, PP242869, PP242265, PP243562, PP241423, PP245952, PP244886, PP241184, PP244168, PP243112, PP242147, PP245242, PP245359, PP246676, PP244767, PP241791, PP243682, PP244529, PP241670, PP242626, PP245836, PP243337, PP245121, PP248354, PP247521, PP247761, PP248470, PP247281, PP246921, PP243806, PP241547, PP242509, PP242748, PP245721, PP245004, PP246067, PP243925, PP241302, PP241908, PP243446, PP245599, PP247998, PP247400, OR885250, PP247045, PP241060, PP246430, PP249176, PP247165, PP248697, PP248585, PP248232, PP244408, PP246553, PP248118, PP249411, PP246187, PP244049, PP242029, PP243230, PP244647, PP246799, PP247640, PP247882, PP249533, PP249291, PP248934, PP249056, PP248815.
Weddellina Tul. W. squamulosa Tul.: C. P. Bove et al. 2350 (MICH, R), PP242993, PP245482, PP246310, PP244291, PP242387, PP242871, PP242266, PP243564, PP241425, PP245954, PP244888, PP241186, PP244170, PP243114, PP242149, PP245244, PP245361, PP246678, PP244769, PP241793, PP243684, PP244531, PP241672, PP242628, PP245838, —, PP245123, PP248356, PP247523, PP247763, PP248472, PP247283, PP246923, PP243808, PP241549, PP242511, PP242750, PP245723, PP245006, PP246069, PP243927, PP241304, PP241910, PP243448, PP245601, PP248000, PP247402, OR885252, PP247047, PP241062, PP246432, PP249178, PP247167, —, PP248587, PP248234, PP244410, PP246555, PP248120, PP249413, PP246189, PP244051, PP242031, PP243232, PP244649, PP246801, PP247642, PP247884, PP249535, —, PP248936, PP249058, PP248817; C. T. Philbrick et al. 5827 (R), PP242992, PP245481, PP246309, PP244290, PP242386, PP242870, —, PP243563, PP241424, PP245953, PP244887, PP241185, PP244169, PP243113, PP242148, PP245243, PP245360, PP246677, PP244768, PP241792, PP243683, PP244530, PP241671, PP242627, PP245837, —, PP245122, PP248355, PP247522, PP247762, PP248471, PP247282, PP246922, PP243807, PP241548, PP242510, PP242749, PP245722, PP245005, PP246068, PP243926, PP241303, PP241909, PP243447, PP245600, PP247999, PP247401, OR885251, PP247046, PP241061, PP246431, PP249177, PP247166, —, PP248586, PP248233, PP244409, PP246554, PP248119, PP249412, PP246188, PP244050, PP242030, PP243231, PP244648, PP246800, PP247641, PP247883, PP249534, —, PP248935, PP249057, PP248816.
Zeylanidium (Tul.) Engl. Z. subulatum (Gardner) C.Cusset: M. Kato & N. Katayama sl102 (TNS), PP242994, PP245483, PP246311, PP244292, PP242388, PP242872, PP242267, —, PP241426, PP245955, PP244889, PP241187, PP244171, PP243115, —, PP245245, —, PP246679, PP244770, PP241794, PP243685, PP244532, PP241673, PP242629, —, —, —, PP248357, PP247524, PP247764, —, PP247284, PP246924, PP243809, PP241550, PP242512, PP242751, PP245724, PP245007, —, PP243928, —, —, —, PP245602, —, —, OR885253, PP247048, PP241063, PP246433, PP249179, PP247168, —, PP248588, —, PP244411, PP246556, PP248121, PP249414, PP246190, PP244052, PP242032, —, PP244650, PP246802, PP247643, PP247885, PP249536, PP249292, PP248937, PP249059, PP248818.
Table S1.
Sequencing platform, number of reads, coding region assembly information, and proportion of missing data for each sample.
Table S1 (Continued).
Table S1 (Continued).
Table S1 (Continued).
Table S1 (Continued).
Table S1 (Continued).
Fig. S1.
Heatmap plot showing proportion of the target gene length recovered for each accession. Columns represent plastid genes and rows represent samples of Podostemaceae.
Table S2.
Dataset Characteristics. Best fit models of sequence evolution were chosen by AICc using ModelFinder as implemented in IQ-TREE 1.6.12.
(Continued).
Fig. S2.
Phylogram of the maximum likelihood phylogeny of Podostemaceae inferred from a concatenated dataset of 73 protein-coding genes of the plastid genome. Voucher collection numbers are given after each species name. Branch lengths are in units of estimated substitutions per site.
Fig. S3.
Maximum likelihood phylogeny of Podostemaceae inferred from from a concatenated dataset of 73 protein-coding genes of the plastid genome. Voucher collection numbers are given after each species name. SH-aLRT (left) and ultra-fast bootstrap support (right) values are indicated above the branches.
Fig. S4.
PhyParts conflict analysis with no bootstrap support cutoff, showing 73 gene trees mapped against the maximum likelihood phylogeny from the concatenated analysis. Numbers above branches equal the number of genes concordant with that bipartition, while the numbers below each branch represent the number of conflicting genes. Pie charts at nodes represent the proportion of gene tree concordance (blue slices), the proportion of the most common conflicting bipartition (green), the proportion of all other conflicting bipartitions (red), and the proportion that are uninformative (grey).
Fig. S5.
PhyParts Conflict analysis with a 70% bootstrap support cutoff, showing 73 gene trees mapped against the maximum likelihood phylogeny from the concatenated analysis. Numbers above branches equal the number of genes concordant with that bipartition, while the numbers below each branch represent the number of conflicting genes. Pie charts at nodes represent the proportion of gene tree concordance (blue slices), the proportion of the most common conflicting bipartition (green), the proportion of all other conflicting bipartitions (red), and the proportion that are uninformative given a 70% bootstrap support cutoff (grey).
