Molecular Phylogeny of the Neotropical Genus Paradrymonia(Gesneriaceae),

Reexamination of Generic Concepts and the Resurrection of Trichodrymonia and

Centrosolenia
Author(s): M. Marcela Mora and John L. Clark
Source: Systematic Botany, 41(1):82-104.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364416X690561

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Systematic Botany (2016), 41(1): pp. 82–104
© Copyright 2016 by the American Society of Plant Taxonomists
DOI 10.1600/036364416X690561
Date of publication February 11, 2016

Molecular Phylogeny of the Neotropical Genus Paradrymonia (Gesneriaceae),

Reexamination of Generic Concepts and the Resurrection of Trichodrymonia
and Centrosolenia

M. Marcela Mora1,2,4 and John L. Clark1,3

1
The University of Alabama, Department of Biological Sciences, Tuscaloosa, AL 35487, U.S.A.
2
Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166, U.S.A.
3
The Lawrenceville School, Science Department, 2500 Main Street, Lawrenceville, NJ 08648, U.S.A.
4
Author for correspondence: ([email protected])

Communicating Editor: Michael J. Moore

Abstract—Paradrymonia Hanst. (Gesneriaceae) with 40 currently recognized species is a genus of facultative epiphytic herbs that inhabits
the forest understory. The genus occurs throughout the Neotropics except for southeastern Brazil and the Caribbean. Monophyly of neotropi-
cal Paradrymonia and species relationships within the genus and closely related genera were evaluated based on molecular sequence data
from the nuclear ribosomal (nrDNA) internal and external transcribed spacer regions (ITS and ETS), and the plastid (cpDNA) psbA-trnH
intergenic spacer. Traditional Paradrymonia is resolved as polyphyletic and includes taxa in three clades. The clade that contains the type spe-
cies is recognized as Paradrymonia sensu stricto (s. s.) and includes three species. The Paradrymonia s. s. clade is sister to a clade that includes
species of Nautilocalyx, Chrysothemis, and most of the species traditionally recognized as Paradrymonia. Morphological characters useful for
circumscribing generic boundaries include bearded anthers, rosette habit, presence of leaf anisophylly, and lanceolate calyx lobe shape. Our
results support the re-circumscription of Paradrymonia and the resurrection of two previously recognized genera, Centrosolenia and
Trichodrymonia, in order to accommodate the remaining ex-Paradrymonia species. Forty-three new combinations (11 in Centrosolenia, four in
Chrysothemis, and 28 in Trichodrymonia) are made. The genus Nautilocalyx is moderately supported as lacking monophyly, but broader taxon
sampling is necessary before new generic circumscriptions are provided.
Keywords—Chrysothemis, new combinations, plastid DNA, Columneinae, Gesnerieae, Nautilocalyx.

Gesneriaceae is a mostly tropical or subtropical plant family the family with 26 genera and over 700 species (Clark et al.
with 150–160 genera and ca. 3200 species (Burtt and Wiehler 2006, 2012; Clark, 2009). The Columneinae is distinguished
1995; Weber 2004; Skog and Boggan 2006; Weber et al. 2013). from other subtribes of the tribe Gesnerieae by its nodal anatomy
Pre-phylogenetic or traditional classifications had divided the and chromosome number. All members of the Columneinae
family into three subfamilies (Gesnerioideae, Cyrtandroideae, have a distinctive three-trace trilacunar nodal anatomy with
and Coronantheroideae) (Wiehler 1983; Burtt and Wiehler split lateral bundles, in contrast to the conventional one-trace
1995). Phylogenetic-based studies support the recognition of unilacunar nodal anatomy present in members of the other
two subfamilies (Gesnerioideae and Cyrtandroideae) (Smith tribes of Gesnerioideae (Metcalfe 1950; Wiehler 1978). Most
et al. 1997; Mayer et al. 2003; Weber 2004; Möller et al. 2009; subtribes in the Gesnerieae have a base chromosome number
Woo et al. 2011). More recently (Weber et al. 2013), a third of x = 13 or 14 (Wiehler 1983; Skog 1984; Burtt and Wiehler
subfamily (Sanangoideae) was added to accommodate the 1995; Smith et al. 2004). The Columneinae is unique in the
monotypic genus and subfamily that includes Sanango tribe because it has a base chromosome number of x = 9 (or
racemosum (Ruiz & Pav.) Barringer. Traditional classifications x = 8 in Codonanthe (Mart.) Hanst. and Nematanthus Schrad.)
based on morphological data (e.g. Hanstein 1854; Fritsch (Wiehler 1978; Smith and Carroll 1997).
1893–1894; Wiehler 1983) have been shown to be artificial. All genera in the Columneinae have been extensively eval-
Recent molecular-based studies have redefined tribes to uated (i.e. inclusion of generic type species and/or broad
reflect monophyletic groups (Smith et al. 1997; Zimmer et al. sampling) in a phylogenetic context except for Paradrymonia
2002; Roalson et al. 2005; Woo et al. 2011; Weber et al. 2013). Hanst., Nautilocalyx Linden ex Hanst., and Chrysothemis Decne.
Weber et al. (2013) proposed a new comprehensive formal Recent phylogenetic studies have resulted in major shifts that
classification for the family based on molecular phylogenetic have redefined most of the genera in the Columneinae so
studies for New and Old World Gesneriaceae that resulted that they represent monophyletic groups (Clark et al. 2006,
in a re-circumscription of the traditionally recognized tribes. 2012). Although Paradrymonia, Nautilocalyx, and Chrysothemis
For the neotropical Gesneriaceae the number of tribes was form a clade within Columneinae (Clark et al. 2012; Perret
reduced from nine to the following five tribes: Beslerieae, et al. 2012), the relationships and generic delimitations within
Coronanthereae, Gesnerieae, Napeantheae, and Titanotricheae this clade have been problematic because material for generic
(Weber et al. 2013). Gesnerieae, the largest tribe in the family, type species was not available, and taxon sampling was
includes what was previously classified in the following tribes: mostly limited to Andean species. For this study we con-
Gesnerieae, Gloxinieae, Episcieae, Sinningiae, and Sphaerorrhizeae ducted field expeditions to Central and South America to
(Weber et al. 2013). Coronanthereae (sensu Weber et al. 2013), increase taxon sampling and include generic type species
previously recognized as subfamily Coronantheroideae by that were previously unavailable.
Wiehler (1983) and Burtt and Wiehler (1995), is supported in Many genera in the subtribe Columneinae (tribe Episcieae
Gesnerioideae based on recent phylogenetic studies as the sensu Wiehler 1983) have at least one synapomorphy or dis-
sister clade to all other New World Gesneriaceae (Woo et al. tinctive morphological trait. For example, Columnea L. has
2011; Perret et al. 2012; Weber et al. 2013). an indehiscent fleshy berry, Glossoloma Hanst. has resupinate
Columneinae (tribe Episcieae sensu Wiehler 1983) is the flowers, Drymonia Mart. has poricidal anther dehiscence, and
largest and most diverse subtribe in the Gesnerioideae and in Episcia has stolons.

82
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 83

In contrast, Paradrymonia, Nautilocalyx, and Chrysothemis cooperi and S. incarnata were selected as outgroups based on previous phy-
have traditionally been defined by a combination of sym- logenies that strongly support the subtribe Ligeriinae (tribe Sinningieae
sensu Wiehler 1983) as the sister group to Columneinae (Perret et al.
plesiomorphic characters. Paradrymonia has been historically 2003; Zimmer et al. 2002).
recognized by its epiphytic rosette forming habit, aniso- DNA extraction, amplification and sequencing — DNA was isolated
phyllous leaves, campanulate corollas, and semi-fleshy bivalved from fresh leaves dried in silica gel. Extractions of DNA were carried
dehiscent capsules; Nautilocalyx by an obligate terrestrial out using a Qiagen DNeasy plant mini kit following the manufacturer’s
recommendations (Qiagen, Germantown, Maryland).
habit and isophyllous leaves; and Chrysothemis by an obli-
The internal (ITS) and external transcribed spacers (ETS) of the
gate terrestrial habit and connate calyx lobes. nuclear ribosomal cistron, as well as the plastid psbA-trnH intergenic
The species conventionally assigned to the genus Paradrymonia spacer, were sequenced for this study. The ITS region has been shown to
are widely distributed throughout the Neotropics, except be phylogenetically useful for analyses at the species level because it
southeastern Brazil and the Caribbean. The genus is espe- is variable and highly repeated in the plant genome allowing for easy
detection, amplification and sequencing (Baldwin et al. 1995). The ETS
cially diverse in northwestern South America with centers region was chosen because it shares the same favorable features of
of diversity in Colombia and Ecuador. Unlike other genera ITS and it has been shown to provide more variable and parsimony-
in the Gesneriaceae, which are abundant in mid-elevation informative characters than ITS (Baldwin and Markos 1998; Linder
montane forests (800–1,500 m) (Perret et al. 2012), most et al. 2000; Markos and Baldwin 2001, 2002; Vander Stappen et al.
2003). Additionally, the combination of data from both regions has
species in Paradrymonia s. l. occur in lowland rainforests
increased resolution and support (Markos and Baldwin 2001; Vander
(< 800 m). One exception is P. metamorphophylla, which occurs Stappen et al. 2003). The psbA-trnH intergenic spacer is considered
in montane cloud forests up to 2,500 m. Most species of among the most variable regions in the angiosperm plastid genome
Paradrymonia have narrow geographic ranges and grow in and it has been used in population genetic studies, in species-level
primary wet to moist forests on moss-covered rocks, wet phylogenies, and in DNA barcoding (Kress et al. 2005; Štorchová and
Olson 2007; Yao et al. 2010).
logs, or edges of waterfalls and streambanks. The following primers were used to amplify these loci: (1) ITS4 and
Paradrymonia is currently recognized with 40 species (Weber ITS 5 (Baldwin and Markos 1998); (2) 18S-ETS (Roalson et al. 2003)
2004; Skog and Boggan 2007). It is the fourth largest genus and ETS-B (Beardsley and Olmstead 2002); and (3) psbA-trnH (Sang
in Columneinae after Columnea (200 + species.), Drymonia et al. 1997). Promega Taq DNA polymerase kits were used to amplify
DNA in accordance with the manufacturer’s instructions (Promega,
(100 + spp.), and Nautilocalyx (ca. 60 spp.) (Clark 2009). The Madison, Wisconsin).
generic definition of Paradrymonia has been historically con- The PCR reaction conditions for amplification of the ITS and ETS
fusing because taxonomists have at some time recognized regions were as follows: (1) initial denaturizing at 95°C; (2) 34 cycles of
17 (45%) of the described species as homotypic synonyms in 1 min at 95°C, 1 min at 48°C, 1 min at 72°C; and (3) a 7-min extension
at 72°C. The PCR reaction conditions for amplification of the psbA-trnH
other genera. Paradrymonia was first described by Hanstein
spacer were as follows: (1) initial denaturizing at 95°C; (2) 30 cycles
(1854) with only one species, P. glabra (Benth.) Hanst. The con- of 1 min at 95°C, 30 s at 52°C, 30 s at 72°C; and (3) followed by a
cept of Paradrymonia as a genus was short-lived, as Hanstein 7-min extension.
(1865) himself reduced it to a synonym of Episcia Mart. More The PCR reactions were cleaned of unincorporated nucleotides and
than 100 yr later, Wiehler (1973) re-established Paradrymonia excess PCR primers using a PEG precipitation protocol prior to sequenc-
ing. Sequencing reactions were performed with ABI Prism BigDye termi-
by transferring four species that were previously recognized nator cycle sequencing ready reaction kits (PE Biosystems, Foster City,
in Episcia (section Paradrymonia (Hanst.) Leeuwenb.). Wiehler California) and were cleaned using standard ethanol/sodium acetate
(1973) justified the re-circumscription and generic status by precipitation. Sequencing was carried out at on an ABI 3731 genetic
the presence of a rosette habit and oblong-lanceolate leaves. analyzer. Sequence chromatograms were examined and edited using
Sequencher version 3.1.1 (Gene Codes Corporation 1998). GenBank
The taxonomic challenges and non-monophyly of Paradrymonia
accessions with their corresponding voucher information are provided in
are well-established in the literature (Smith and Carroll 1997; Appendix 1. Sequence alignments of all the datasets for the phylogenetic
Clark and Zimmer 2003; Clark et al. 2006, 2012). Paradrymonia analyses are available from TreeBASE (study number 14733).
was recovered as monophyletic by Smith (2000); however, Sequence alignment — Sequences were initially aligned using ClustalW
this study only included a limited sampling of three spe- version 2.0 (Larkin et al. 2007) with default settings and subsequently
manually adjusted using Se-Al version 2.0a11 (Rambaut 2002), a multiple
cies. All other studies (Smith and Carroll 1997; Clark and sequence editor. Automated sequence alignments for the nuclear regions
Zimmer 2003; Clark et al. 2006, 2012) do not support the (ITS and ETS) required few manual adjustments. In contrast, alignments
monophyly of Paradrymonia. The primary objectives of this for the plastid marker, psbA-trnH, required extensive manual editing.
study were to use additional molecular sequence data from Inversions for six species were detected for the psbA-trnH dataset.
Each inverted region was replaced with its reverse complement, such
an expanded taxon sampling to: (1) re-circumscribe a mono-
that sequence homology was maximized across the alignment (Whitlock
phyletic Paradrymonia; (2) identify well-supported clades et al. 2010). Inversions are known to be frequent for the psbA-trnH region
among the Paradrymonia segregates; (3) increase taxon sam- (Sang et al. 1997; Tate and Simpson 2003; Clark et al. 2006; Kårehed et al.
pling relative to previous studies to resolve relationships 2008; Borsch and Quandt 2009) and they appear to be common in non-
among Paradrymonia, Chrysothemis, Nautilocalyx, and closely coding plastid regions associated with stem-loops in the RNA secondary
structures (Kelchner 2000; Štorchová and Olson 2007). The most common
related congeners; (4) to determine the placement of these explanation for these hairpin-associated inversions is a single mutational
genera within the subtribe Columneinae; and (5) evaluate event (Kelchner 2000; Quandt et al. 2003; Kim and Lee 2005). Additionally,
diagnostic morphological characters of well-supported clades inversions were scored as a presence/absence character. Three regions
amongst currently recognized Paradrymonia species. with a combined total of 146 characters were especially variable and
difficult to align and were therefore excluded from the analyses.
Phylogenetic Analyses — Data were analyzed separately (ITS, ETS,
psbA-trnH) for parsimony analyses, and in combination under maximum
Materials and Methods parsimony (MP), maximum likelihood (ML) (Felsenstein 1973), and
Bayesian (Yang and Rannala 1997) methods.
Taxon Sampling — A total of 80 ingroup species belonging to 15 out of No incongruence was detected from analyses of individual markers
22 genera of the subtribe Columneinae were sampled for the phyloge- based on the evaluation of strongly supported nodes in the tree topolo-
netic analyses (Appendix 1). Paradrymonia, Nautilocalyx, and Drymonia gies. Incongruence among data partitions was evaluated by comparison
were represented with 27, 23, and four species respectively. The remaining of bootstrap support for clades (>75%) and posterior probability values
13 genera were represented with one to three species each. Sinningia greater than 0.95. Since there were no strongly supported nodes in conflict
84 SYSTEMATIC BOTANY [Volume 41

the three datasets were combined in a total evidence approach (Kluge The main difference between the topologies of the individ-
1989) as suggested by Seelanan et al. (1997) and Wiens (1998). ual ITS and ETS consensus trees (not shown) is the place-
The MP analyses were performed using the heuristic search option in
PAUP* (Swofford 2003) with the following settings: 100 random addition
ment of the Core Columneinae and Guiana Shield clades.
replications with tree bisection and reconnection (TBR) branch swapping, The individual analysis of ITS resulted in a poorly supported
and MULTREES option in effect with a maximum of 10,000 trees being branch (BS < 50%) that subtends the Guiana Shield and
retained. Bootstrap analysis (Felsenstein 1985) with 1,000 replicates and Paradrymonia alliance clades as sister groups. In contrast,
10 random sequenced additions per replicate was conducted to assess
the ETS and the total evidence analyses result in a strongly
support values for the MP analysis.
The best-fitting model of DNA substitution was determined using supported sister-group relationship (BS = 100%) of the Guiana
the Akaike information criterion (Akaike 1974) under ML settings as Shield clade and the Core Columneinae + Paradrymonia alli-
implemented by jModelTest (Posada, 2008). The best model of sequence ance clades (Fig. 1). In addition, a sister-group relationship
evolution was the general time-reversible model (GTR+I+gamma) with between the Core Columneinae and the Paradrymonia alli-
some sites assumed invariant and with variable sites assumed to follow
a gamma distribution. The following parameter estimates were obtained:
ance clades is strongly supported by the MP strict consensus
(1) base frequencies of 0.2569 (A), 0.2172 (C), 0.2298 (G), 0.2960 (T); of the ETS marker (BS = 90%); and moderately to strongly
(2) substitution rates of 0.8679 (A–C), 2.5181 (A–G), 0.9418 (A–T), supported for the combined analysis (MP BS = 78%; ML
0.7413 (C–G), 4.6177 (C–T), 1 (G–T); and (3) a gamma distribution shape BS = 84%; PP = 0.98, Fig. 1).
parameter of 0.5030. A ML heuristic bootstrapping search was per-
The traditionally recognized Paradrymonia is strongly sup-
formed using the default parameters in RAxML 7.2.6 through the CIPRES
portal (Stamatakis et al. 2008) for 100 replicates, repeated 10 times to gen- ported as polyphyletic (i.e. Paradrymonia s. l. or Clade II in
erate 1,000 replicates, starting from random trees. Fig. 1) based on the phylogenetic analyses of the combined
Bayesian posterior probabilities (PP) for the combined dataset were datasets. Currently recognized Paradrymonia species were
calculated using the Bayesian Markov Chain Monte Carlo algorithm recovered as belonging to two clades in the Columneinae (Core
(MCMC) of BEAST v1.8.0 (Drummond et al. 2012) to further investigate
the impact of tree construction method on topology. The species tree Columneinae clade and three lineages in the Paradrymonia
prior was set to a Yule Process following author recommendations. We alliance clade; Figs. 1 & 2). The Paradrymonia alliance clade
ran two independent runs of 50,000,000 generations and sampled every comprises seven strongly supported clades (herein referred
5000 generations. Convergence was evaluated by comparing traces and to as clades I–VII in Figs. 1, 2). Among these clades, three
posterior distributions of parameter estimates using Tracer (Rambaut
and Drummond 2007), ensuring that the log-likelihood of the model had
contain species of Paradrymonia (clades I, II, and VII in
reached a plateau and the effective sampling size (ESS) of each parameter Figs. 1, 2). Clade I (MP BS = 98%; ML BS = 98%; PP = 1)
was at least 200 or greater. The two runs were then combined using includes the type species (P. glabra), as well as P. ciliosa and
LogCombiner after the first 25% of trees were discarded as part of the P. campostyla. This small clade is strongly supported (MP
burn-in phase. The remaining trees were used to calculate posterior proba-
BS = 98%; ML BS = 98%; PP = 1) as the sister group to
bilities in a maximum credibility tree using TreeAnnotator (Drummond
and Rambaut 2007) with a posterior probability threshold of 0.5 and the remaining species of the Paradrymonia alliance clade.
median node. Trees were visualized in FigTree (Rambaut 2009). Clade II is also strongly supported (MP BS = 95%; ML
BS = 90%; PP = 1) and includes Nautilocalyx cordatus
(= Centrosolenia hirsuta), Nautilocalyx coccineus (= C. coccinea),
Results
and Paradrymonia densa (= C. densa). Clade VI (Trichodrymonia
Phylogenetic analyses — The total evidence analyses (MP, clade) contains most of the traditionally recognized species
ML, and Bayesian inference) resulted in trees with similar of “Paradrymonia” and is strongly supported as the sister
topologies. The same strongly supported clades were recov- group to clade VI that contains the generic type species for
ered in all analyses, and differences in topology occurred Nautilocalyx (N. bracteatus).
only in clades with low support. Therefore, support values The topology of the Bayesian maximum clade credibility
(Bootstrap/posterior probability values) were mapped onto (MCC) tree (Fig. 2) is more resolved than the topology of the
the parsimony strict consensus tree (Fig. 1). The MP, ML, MP tree (Fig. 1), but overall there were no strongly sup-
and Bayesian analyses of the combined regions consistently ported branches that are incongruent. Likewise, the ML tree
revealed three highly supported main clades in the subtribe (Fig. S1) is congruent with the MCC tree (Fig. 2). One main
Columneinae (Figs. 1, 2 Supplementary Fig. S1): (1) the difference between the MCC (Fig. 2) tree in contrast to the MP
Guiana Shield clade (MP BS = 92%; ML BS = 97%; PP = 1.0); tree (Fig. 1) is the placement of Centrosolenia poryphyrotricha
(2) the Core Columneinae clade (MP BS = 96%; ML BS = 98%; (=Nautilocalyx porphyrotrichus). Its phylogenetic placement
PP = 0.98); and (3) the Paradrymonia alliance clade (MP BS = in the MCC tree is within Centrosolenia (Fig. 2) and it is
98%; ML BS = 98%; PP = 1.0). unresolved near the base of the Paradrymonia alliance in the
Table 1 presents a summary of statistics for the MP analy- MP tree (Fig. 1). Relatively long branches (Fig. 2) were found
ses of the combined dataset and individual partitions. Among in Cremersia platula (Guiana Shield clade) and Codonanthopsis
the individual partitions, the percentage of informative char- ulei (Core Columneinae clade).
acters was highest in the ETS region (51%), while psbA-trnH Phylogenetic utility of ITS, ETS, and psbA-trnH — ITS and
had the lowest percentage of informative characters (14%, ETS were the most informative regions among the three
Table 1). markers. The nuclear regions provided four times as many
The strict consensus of the MP trees based on the plastid parsimony-informative characters as the psbA-trnH cpDNA
dataset (psbA-trnH) was mostly unresolved (Fig. S2), with marker (Table 1). Although both nuclear regions provided
only two pairs of species that were well-supported (BS MP about the same number of informative characters (ITS = 283;
> 90%) and congruent with the nrDNA datasets. Those ETS = 285), ETS yielded a greater number of phylogenetically
species pairs are: Rhoogeton cyclophyllus + R. viviparus and informative characters per sequence length (Table 1). The
Nautilocalyx sp. Mora 931 + Nautilocalyx sp. Mora 949 (Fig. S2). sequence data from ETS and ITS provided more resolution
In contrast, the individual nuclear markers (e.g. ITS and ETS), and more parsimony-informative characters (Fig. S3) com-
as well as the combined analysis (Fig. 1), provided higher pared to psbA-trnH. Of the three markers, psbA-trnH resulted
resolution and support than the plastid dataset. in the least number of variable and parsimony-informative
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 85

Fig. 1. Parsimony strict consensus tree from the total evidence analysis of three datasets (nrDNA ITS, nrDNA ETS, and cpDNA trnH-psbA) showing
the three main clades within subtribe Columneinae and the principal monophyletic subclades within the Paradrymonia + Nautilocalyx + Chrysothemis
clade (Paradrymonia alliance clade). MP bootstrap values are above the branches, and those derived from maximum likelihood (when nodes are shared)
are below the branches. Only bootstrap values ≥ 50% are shown. Thickened branches indicate ≥ 0.9 Bayesian posterior probability values. An asterisk (*)
following the species name indicates the species is the type species of the genus.
86 SYSTEMATIC BOTANY [Volume 41

Fig. 2. Maximum clade credibility (MCC) tree of the combined dataset (ETS, ITS and psbA-trnH) for 82 taxa obtained from the BEAST analysis.
Node support is given as Bayesian posterior probability (PP): black circles at nodes denote PP = 1.0, grey circles denote 0.95 ≤ PP ≤0.99, white circles
denote 0.90 ≤ PP ≤ 0.94. Unlabelled nodes denote PP < 0.90.
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 87

Table 1. Summary statistics derived from the parsimony analyses of nuclear and plastid markers

Combined Combined nuclear

Statistic ITS ETS nuclear data psbA-trnH and plastid data

Total # of taxa 82 73 82 73 82
Base pairs in the alignment 1–758 759–1313 1–1313 1313–1886 1–1886
Total length (aligned) 758 555 1313 573 1886
Included characters 758 555 1313 470 1795
Number of MP trees 71 29 12 3 25
Tree length (steps) 1436 1193 2711 239 2977
Variable characters (%) 398 (52%) 373 (67%) 751 (57%) 153 (32%) 950 (53%)
Parsimony-informative characters (%) 283 (37%) 285 (51%) 576 (44%) 68 (14%) 638 (35%)
Consistency Index 0.407 0.503 0.443 0.574 0.463
Retention index 0.700 0.735 0.706 0.439 0.704

characters and provided little support for clades within the significant difference is taxon sampling where only 20 spe-
subtribe Columneinae (Fig. S2). cies are shared between the current study and Clark et al.
(2012). It should be noted that the sister-group relationship
of the Core Columneinae + Guiana Shield clades in Clark
Discussion
et al. (2012) is weakly supported in the MP and Bayesian
Comparison with other studies of Columneinae (=Episcieae) analyses (MP BS = 61; PP = 0.77) and strongly supported in
phylogeny—Nuclear markers (ITS and ETS) and the com- the ML analysis (ML BS = 100). In contrast, the sister-group
bined analyses are congruent in supporting the non- relationship in the present study of the Core Columneinae
monophyly of Paradrymonia, as suggested by earlier studies and the Paradrymonia alliance clades is moderately supported
with more limited taxon sampling (Smith and Carroll 1997; in all the analyses (MP BS = 78; MP BS = 84; PP = 0.99). It is
Clark and Zimmer 2003; Clark et al. 2006, 2012). recommended that future studies of the Columneinae include
The placement of the Guiana Shield clade as the sister ETS because it provides more parsimony-informative char-
group to all the members of the subtribe Columneinae is acters relative to previous studies that depended on large
moderately supported in our analyses (BS MP = 78%, BS quantities of cpDNA markers that resulted in relatively few
ML = 84%, PP = 0.99; Figs. 1, 2). The sister-group relation- parsimony-informative characters.
ship of the Guiana Shield clade with the Core Columneinae + Analyses by Perret et al. (2012) with a broader emphasis
Paradrymonia alliance clades is congruent with Clark et al. in the Gesnerioideae placed the Paradrymonia alliance clade
(2006), who used a total evidence analysis based on ITS, as the sister group to all of the remaining Columneinae. One
psbA-trnH, and morphological data to produce a compre- potential reason for this incongruence could be attributed
hensive phylogeny of the subtribe Columneinae (155 spe- to the use of plastid markers in Perret et al. (2012) in con-
cies and 21 out of the 22 genera). However, the support for trast to the use of nuclear markers here. A number of studies
the placement of the Guiana Shield as sister clade to all of have shown that different genomes (e.g. nuclear vs. plastid
the remaining members of Columneinae was low (BS < 50%). markers) could produce incongruent phylogenetic trees due
Clark et al. (2006) included more taxa compared to the to different historic signals or processes such as hybridization,
current study (151 vs. 82), but their focus was on the Core paralogy, incomplete lineage sorting, and scarcity of nucleo-
Columneinae clade. In contrast, we sampled 35 (18%) more tide homologies among others (e.g. Sang et al. 1997; Slovák
species representing the Paradrymonia alliance clade and et al. 2014; Zhang et al. 2015). Likewise, incongruence may
included an additional nrDNA molecular marker (ETS). The also result from errors such as misspecification of models or
addition of ETS and increased taxon sampling resulted in problems during the sequencing and the alignment that are
more parsimony-informative characters and increased sup- difficult to discard (Zhang et al. 2015).
port at deep branches relative to previous studies (Table 1). Alternatively, discordance in tree topologies between the
The utility of ETS significantly increases support by adding two studies might be due to differences in taxon sampling.
285 (51%) parsimony-informative characters compared to The Perret et al. (2012) study lacked taxa from the Guiana
283 (37%) for ITS and 165 (31%) for psbA-trnH (Table 1). Shield and included only four species from the Paradrymonia
In contrast to Clark et al. (2006), the placement of the alliance clade (Chrysothemis melittifolia, Chrysothemis pulchella,
Paradrymonia alliance and Guiana Shield clades in the pres- Trichodrymonia macrophylla, and Nautilocalyx aeneus). As a
ent study are not congruent with Clark et al. (2012). The result of limited taxon sampling, comparisons at the level of
different topology between these two studies could be attrib- Columneinae phylogeny in the present study are of limited
uted to the use of different loci. Clark et al. (2012) used four value with Perret et al. (2012).
plastid and four nuclear loci for the analysis of the subtribe Analyses presented support the sister-group relationship
Columneinae with ITS as the only mutual marker in both between the Paradrymonia alliance clade and the Columneinae
studies. Interestingly, many of cpDNA markers in Clark et al. core clade (Figs. 1, 2). These two clades are more diverse than
(2012) were limited in parsimony-informative characters. For the Guiana Shield clade and have centers of diversity in the
example, Clark et al. (2012) generated 8,714 base pairs from Amazon, northern Andes, and Central America.
14 loci for the Core Columneinae clade and only 383 (4.3%) The phylogenetic utility of plastid and nuclear DNA
were parsimony-informative (Table 2 in Clark et al. 2012). markers — The alignment of the psbA-trnH region was par-
In contrast, in the present study the utility of ETS alone ticularly challenging due to the presence of numerous inver-
accounted for 1300 + base pairs and resulted in 285 (51%) sions and indels. Three regions with a total of 146 characters
parsimony-informative characters (Table 1). An additional were especially variable and difficult to align and were
88 SYSTEMATIC BOTANY [Volume 41

therefore excluded from the analyses. Many regions of indels endemic genera except for Lampadaria and Christopheria. While
were similar in unrelated taxa (i.e. species from different the placement of Lampadaria remains uncertain, Christopheria
genera), indicating a high level of homoplasy. Sang et al. (as Episcia xantha Leeuwenb. in Clark et al. 2012) was strongly
(1997) also noticed considerable homoplasy of indel characters supported as nesting in the Guiana Shield clade. Pagothyra
for psbA-trnH within Paeonia (Paeoniaceae). In the present consists of a single species, P. maculata, a climbing vine with
study, the remaining portion of the alignment included in the isophyllous leaves and bell-shaped yellow flowers mottled
analysis was rather conserved and therefore of limited value with bright red spots. This taxon was first placed in the genus
in resolving relationships. Like Shaw et al. (2005), we con- Episcia by Hooker (1890) and then transferred to Paradrymonia
clude that psbA-trnH provides limited information relative to by Wiehler (1978). Recently, this taxon was recognized as a
other markers and its use in future phylogenetic studies of monotypic genus, based on the unique apomorphy of the
the Gesneriaceae is not highly recommended. ventral corolla lobe occluding the throat like a trap door
Likewise, other phylogenetic studies of the Gesneriaceae (Smith and Clark 2013). A recent analysis (Clark et al. 2012)
have shown that the phylogenetic signal of psbA-trnH and supports a sister-taxon relationship between Christopheria
other plastid markers are less informative than nuclear non- xantha and Pagothyra maculata.
coding sequences. For example, in Clark et al. (2012), ITS Core Columneinae clade — The Core Columneinae clade
yielded at least five times the percentage of phylogenetically includes Drymonia longifolia Poepp., which had been recog-
informative characters as any single non-coding plastid region nized as a member of Paradrymonia (Wiehler 1973) but was
used in the analysis. Some regions like the trnL-F spacer and originally described in Drymonia by Poeppig (1840). Wiehler
the trnL intron had 2% or fewer parsimony-informative char- (1973) made the combination in Paradrymonia because he
acters per length of sequence (vs. 20% for ITS). Although noted the presence of longitudinal slits instead of poricidal
Clark et al. (2012) included 14 markers, 10 of which were anther dehiscence. The presence of poricidal anther dehis-
from cpDNA non-coding regions, the combined regions cence in Drymonia longifolia was observed in the early stage
yielded less than 5% informative characters. In contrast, the of anthesis and then developed into longitudinal slits as the
present study yielded 35% of parsimony-informative char- flower matured (Clark, pers. obs.). The shift from poricidal
acters using only three markers. The data presented here to longitudinal anther dehiscence has been observed for
and in other studies support that when analyzed separately, other species of Drymonia (Clark et al. 2006) and therefore
cpDNA noncoding sequences result in poorly supported phy- makes it challenging to assign a definitive character state for
logenies and limited resolution within the subtribe Columneinae. anther dehiscence. Furthermore, the presence of isophyllous
Although some relationships remain to be resolved, in the leaves, leafy calyx, and succulent display capsule are further
present study the combination of ITS and ETS provided suf- evidence that this taxon does not belong in Paradrymonia.
ficient characters for robust results and resolution amongst Results presented here and in previous studies (Zimmer
closely related taxa. Therefore, the use of ETS for future analy- et al. 2002; Clark and Zimmer 2003; Clark et al. 2006, 2012)
ses in the Gesneriaceae is encouraged. strongly support the placement of Drymonia longifolia as a
Phylogenetic implications for the taxonomy of Columneinae member of Drymonia.
and Paradrymonia s. l.—MP, ML, and Bayesian analyses for Paradrymonia alliance clade — The Paradrymonia alliance
the combined ITS, ETS, and psbA-trnH datasets recovered clade is strongly supported (Fig. 1) in this study as well as
similar phylogenies for subtribe Columneinae. Our results in previous analyses (Zimmer et al. 2002; Clark and Zimmer
strongly support that Paradrymonia must be substantially 2003; Clark et al. 2006, 2012). Our results show that this
re-circumscribed to reflect the phylogenetic results presented clade comprises seven moderately to strongly supported lin-
here (i.e. monophyletic genera). Formal nomenclatural changes eages (clades I–VII in Fig. 1). Species relationships within
regarding these findings are provided in the taxonomic section. these clades varied by analytical approach usually when
Below we describe the clades presented in Figs. 1 and 2. the nodes were not strongly supported (BS values < 80%,
Guiana Shield clade — The Guiana Shield clade includes PP < 0.8) or were unresolved.
taxa that are endemic to that region in northeastern South Paradrymonia s. s. (subclade I)—Paradrymonia s. s. is
America. This region is defined by the underlying geologi- strongly supported as monophyletic (BS = 98, PP = 1) and
cal formation known as the Guiana Shield and extends includes the generic type species, Paradrymonia glabra
from western Colombia to eastern Brazil (States of Amapá, (Figs. 3A, B), as well as P. ciliosa (Figs. 3C, D) and
Pará, Roraima, and Amazonas), Venezuela (States of Delta P. campostyla. Interestingly, P. glabra was recognized by
Amacuro, Bolívar, and Amazonas), Guyana, Suriname, and Wiehler (1973) as a synonym of P. ciliosa. Extensive fieldwork
French Guiana (Hollowell and Reynolds 2005). Interestingly, and an ongoing monographic revision of Paradrymonia sug-
this region has a surprisingly large number of endemic genera gest that these two taxa are not conspecific based on morpho-
of Gesneriaceae (Weber 2004). The restricted distribution and logical and geographical data. Both species have creeping
endemism of the Columneinae in this clade could be the stems, clustered leaves, elongate and coriaceous leaf blades,
outcome of geology, topography, climatic factors (Kelloff and fimbriate corolla lobes, and bearded anthers (Fig. 3F). Char-
Funk 2004; Antonelli et al. 2009; Antonelli and Sanmartín acters that differentiate Paradrymonia glabra from P. ciliosa
2011), geographic barriers to dispersal (Avise 2000), and low include the following: P. glabra has white, straight corollas
intrinsic dispersal ability (Croteau 2010), among others. Out covered with white trichomes (Fig. 3A), while P. ciliosa has
of the eight genera endemic to this region, seven belong to cream-white, angulated corollas covered with red trichomes
the Columneinae: Lembocarpus Leeuwenb., Rhoogeton Leeuwenb., (Fig. 3C). Additionally, P. glabra is endemic to the Venezuelan
Lampadaria Feuillet & L. E. Skog, Cremersia Feuillet & L. E. Andes while P. ciliosa is a widespread taxon that is distributed
Skog (Feuillet and Skog 2003), and the recently described from Nicaragua to northeastern Brazil.
monotypic genera Christopheria and Pagothyra (Smith and As defined here, Paradrymonia s. s. includes species with
Clark, 2013). The results presented here sampled all of the anisophyllous leaves, succulent, large elliptic-oblanceolate leaf
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 89

Fig. 3. Paradrymonia clade (Paradrymonia s. s. in Paradrymonia clade from Fig. 1). A. Lateral flower of Paradrymonia glabra showing straight
(non-angulate corolla). B. Front view of P. glabra showing fimbriations on lower corolla lobe. C. Lateral flower of P. ciliosa showing angulate
corolla. D. Flower of P. ciliosa showing fimbriations on lower corolla lobe. E. Leaves of P. glabra. F. Bearded anthers of P. glabra. Images from
John L. Clark. A, B, and D from J. L. Clark et al. 11331. B and C from J. L. Clark et al. 12490.
90 SYSTEMATIC BOTANY [Volume 41

blades with decurrent bases (Fig. 3E), petioles longer than In addition to the species sampled here, it is predicted that
the inflorescence, flowers with calyx lobes linear to lanceolate other unsampled species also belong to Centrosolenia because
(Fig. 3), corolla lobes crenate to fimbriate with elongate fim- they share a suite of characters such as similar habit (terres-
briations on lower lobe (Figs. 3B, D), and anthers with an trial herb), obovate to oblanceolate leaf shape, membranous
apical tuft of trichomes (i.e. “bearded anthers”; Fig. 3F). leaf texture, crenate margins, oblong calyx lobes, and tubular
The presence of bearded anthers in Paradrymonia s. s. is a corollas. These species are from the Venezuelan Guiana Shield
distinctive synapomorphy that has not been previously rec- (e.g. Nautilocalyx chimantensis, N. crenatus, N. orinocensis, and
ognized at the generic level and bearded anthers are not N. ruber) and the Amazon basin (e.g. N. paujiensis, N. pusillus,
known to occur in other groups of Gesneriaceae. Although N. roseus, and N. vestitus). The present molecular analysis in
only two species included in the phylogeny have bearded conjunction with morphological characters and geographic
anthers, other species such as Paradrymonia buchtienii, P. lutea, distribution provide strong support for the Centrosolenia clade.
P. yatua, and P. tepui also have this distinctive character. It is The genus Centrosolenia is re-established here with 12 new
predicted that these species belong to the same clade because combinations (see taxonomic treatment below).
they present bearded anthers and are vegetatively similar to Pictus clade (subclade iii)—This subclade includes
other members of Paradrymonia s. s. Nautilocalyx pictus and a recently discovered, undescribed
The sister taxon to Paradrymonia glabra and P. ciliosa is species from Peru (M. M. Mora 949). Both species have in
Paradrymonia campostyla, which lacks bearded anthers. In common the presence of flowers in axillary fascicles, acumi-
addition to lacking bearded anthers it also differs by the nate linear-lanceolate bracts, elongate calyx lobes, creeping
presence of elongate climbing stems with isophyllous leaves habit, elongate internodes, isophyllous leaves, and bullate
(vs. anisophyllous leaves and erect shoots), small oblong- leaf blades with crenate to crenulate margins. Taxonomic
elliptic to oblong-ovate leaf blades with subcordate bases changes are not recommended for this clade until addi-
(vs. oblanceolate with decurrent bases), and petioles shorter tional sampling includes more species of Nautilocalyx.
than the flowers (vs. petioles longer than the flowers). We Chrysothemis clade (subclade IV)—This clade includes
refrain from recognizing Paradrymonia campostyla as a dif- Chrysothemis pulchella (the generic type species of Chrysothemis),
ferent genus even though it is morphologically distinct C. friedrichsthaliana, Nautilocalyx melittifolius (=Chrysothemis
from the core Paradrymonia clade based on the characters melittifolia), and N. panamensis (=Chrysothemis panamensis)
described above. (Figs. 1, 2). All members of this clade are obligate terres-
Centrosolenia (subclade II)—The Centrosolenia clade trial herbs that occasionally form tubers.
includes three species (Figs. 1, 2). Centrosolenia is an avail- Chrysothemis has been traditionally distinguished from
able name that includes Centrosolenia hirsuta, which is a other species in the subtribe Columneinae by the presence of
synonym of Nautilocalyx cordatus and the generic type spe- connate calyx lobes, which make the calyx look like a cup
cies for Centrosolenia Benth. The name Centrosolenia Benth. with irregular or five-pointed rim (Fig. 4E). These calyces
predates Nautilocalyx Linden ex Hanst., but the latter was have been described as “water calyces” because they often
conserved by Feuillet and Skog (1990). retain water and secrete liquid that allow the immature buds
Morphological characters that are shared among species to develop under an aqueous solution that protects the buds
in this clade include: (1) the presence of decumbent stems from floral herbivores (Burtt and Woods 1975; Endress 1996;
that root and branch at the nodes; (2) variegated leaf blades Carlson and Harms 2007). Studies of C. friedrichsthaliana by
(especially along the midrib) (Fig. 4A) with asymmetrical Carlson and Harms (2007) suggest that the water calyx helps
bases and anastomosed tertiary venation; and (3) elongate protect immature flower buds from a small species of moth
narrow-tubular corollas (Fig. 4A). Some immature individ- (Alucita sp.) that is a floral herbivore.
uals of Paradrymonia densa have variegated leaves that lose The present phylogenetic analysis supports the expansion
their variegation at maturity. An important diagnostic char- of Chrysothemis to include N. panamensis, and N. melittifolius.
acter for Centrosolenia that distinguishes it from subclade V We also include new combinations for N. adenosiphon and
(Amazonian “Nautilocalyx”) and subclade VI (Nautilocalyx N. colonensis in Chrysothemis based on the study of herbarium
s. s.) is the presence of oblong-lanceolate to oblong-spathulate specimens. Although these species have rather relatively free
calyx lobes. The species of Nautilocalyx s. l. in subclades calyx lobes, they share with all other species an obligate ter-
(V and VI) have calyx lobes that range from broadly ovate restrial habit with underground tubers and succulent stems.
and leaf-like (Amazonian “Nautilocalyx”, Fig. 4B) to linear- Other diagnostic characters for the Chrysothemis clade include
lanceolate (Nautilocalyx s. s., Figs. 4C, D). The ML and the following: (1) elliptic leaves with crenate-dentate to
Bayesian analyses result in placing Nautilocalyx porphyrotrichus dentate-serrate margins; (2) slightly asymmetrical (oblique)
into Centrosolenia, but support is moderately low or weak leaf bases; (3) cross-venulate to reticulate veins; and (4) semi-
(ML BS = 72; PP = 0.63; Figs. 2, S1). It is interesting to note succulent fully reflexed capsules when opened.
that Leeuwenberg (1958) recognized six subsections in a Amazonian “Nautilocalyx” clade (subclade v)—This
classification of the genus Episcia, and included Nautilocalyx clade includes most of the species traditionally placed in
porphyrotrichus as Episcia porphyrotricha in section Episcia, Nautilocalyx. The species in this clade are obligate terrestrial
subsection Centrosolenia. This species has bullate varie- herbs with similarities to members of the Nautilocalyx s. s.
gated leaves with white or yellow on the midrib, and red clade such as isophyllous leaves and succulent stems. An
narrow-tubular corollas, characters common in other spe- important distinction between the Amazon “Nautilocalyx”
cies in the Centrosolenia clade. For the reasons mentioned clade and the Nautilocalyx s. s. clade is that the calyx lobes
above, we propose to transfer Nautilocalyx porpyrothrichus are overlapping at the base or folded in a conduplicate
to Centrosolenia. Interestingly, all of the species in the manner such that each lobe is appressed to an adjacent
Centrosolenia clade represented in the phylogeny are from the lobe and folded lengthwise with the margin curved upward
Guiana Shield. (Fig. 4B). The calyx lobes are also leaf-like and cover the basal
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 91

Fig. 4. Morphological characters in selected species from the Paradrymonia clade (from Fig. 1). A. Centrosolenia hirsuta (= Nautilocalyx cordatus,
Centrosolenia clade) showing variegated leaves and elongate tubular corollas. B. Front view of Nautilocalyx ecuadoranus (Amazonian “Nautilocalyx”
clade in Fig. 1) showing ovate calyx lobes and corolla tube longitudinally sulcate with blunt ridges on both, the dorsal and ventral sides. C. Nautilocalyx
colombianus (Nautilocalyx s. s. clade in Fig. 1) showing trumpet-shaped corolla and lanceolate calyx lobes. D. Nautilocalyx erytranthus showing trumpet-
shaped corolla with reflexed lobes and lanceolate calyx lobes (Nautilocalyx s. s. clade in Fig. 1). E. Chrysothemis friedrichsthaliana (Chrysothemis clade)
showing flower with fused calyx lobes. F. Trichodymonia pedunculata (= Paradrymonia pedunculata; Trichodrymonia clade) showing rosette habit and
elongated leaves with decurrent base. G–H. Flowers showing absence of fimbriations on corolla lobes. G. Frontal view of the salverform corolla in
Trichodrymonia ulei (= P. ulei). H. Frontal view of the trumpet-shaped corolla in Trichodrymonia aurea (= P. aurea). I. Lateral view of the hypocyrtoid
corolla of T. hypocyrta (= P. hypocyrta). Images from R. Myrh (A) and J. L. Clark (B–I).
92 SYSTEMATIC BOTANY [Volume 41

gibbosity of the corolla tube. Most species in this clade have to the Paradrymonia s. s. clade such as rosette habit (Fig. 4F),
tubular white or cream corollas. The corolla tube is usually anisophyllous leaf pairs, and elongated oblanceolate leaf
3 × or longer than the calyx lobes and is typically longitudi- blades. An important difference between Trichodrymonia and
nally sulcate with blunt ridges on both the dorsal and ven- Paradrymonia is the absence of fimbriations on the corolla
tral surfaces (Fig. 4B). The Nautilocalyx s. s. clade has calyx lobes in the former (Figs. 4G–I) and presence in the latter
lobes that are nearly free, linear-lanceolate, and the corolla (Figs. 4A–F). Additionally, bearded anthers are present
spur is exposed (Figs. 4C, D). in Paradrymonia (Fig. 3) and absent in Trichodrymonia. The
Nautilocalyx s. s. clade (subclade vi)—This study is following characters define Trichodrymonia: (1) facultative
the first to include Nautilocalyx bracteatus, which is the type epiphytic herbs; (2) leaves clustered in an apical rosette
species for Nautilocalyx. This species is endemic to Colombia (Fig. 4F); (3) maroon sulcate petioles; (4) corollas salverform
and occurs in the Departments of Antioquia, Chocó, and (Fig. 4G) to trumpet-shaped (Fig. 4H), and occasionally
Santander. The inclusion of this species in the analysis is cru- hypocyrtoid (Fig. 4I); (5) anthers glabrous with longitudinal
cial for the circumscription of Nautilocalyx. The genus was dehiscence; (6) leaf pairs usually anisophyllous; and (7) fruits
established by Hanstein (1854) in an introduction on genera a semi-fleshy bivalved dehiscent capsule.
of Gesneriaceae in the New World, but 11 yr later he reduced
it to a synonym of Episcia (Hanstein 1865). Bentham (1876) Conclusions
then divided the genus Episcia into six sections including sect.
Nautilocalyx. Sprague (1912) resurrected the genus on the basis Analyses of relationships among traditionally recognized
of the position of the ovules on one or both surfaces of the Paradrymonia provide the most resolved phylogeny available
placenta. However, Wiehler (1978) found Sprague’s character to for the genus. Paradrymonia as previously defined is taxonom-
be inconsistent and variable among some members of Episcieae. ically evaluated and recircumscribed. New combinations are
The Nautilocalyx s. s. clade includes N. antioquensis, provided that are consistent with the recognition of mono-
N. bracteatus, N. colombianus (Fig. 4C), and the recently phyletic genera based on the present phylogeny and inferred
described species of Nautilocalyx erytranthus (Fig. 4D). This morphological synapomorphies. Paradrymonia is reduced to
eight species (See taxonomic treatment), while the remain-
clade is defined by a funnel- or trumpet-shaped corolla with
ing ex-Paradrymonia species are placed in other genera.
spreading lobes, narrow lanceolate to linear-lanceolate calyx
Chrysothemis is monophyletic, but the results presented here
lobes with the dorsal lobe greatly reduced and recurved to
strongly support a broader circumscription that includes
accommodate the gibbosity at the base of the corolla, and a
some species previously recognized in Nautilocalyx. Our results
corolla tube that is less than 2 × longer than the calyx (vs. 3 ×
also show that Nautilocalyx is clearly paraphyletic, but the rec-
longer than the calyx in the Amazonian “Nautilocalyx” clade).
ognition of additional new genera is not recommended until
Most of the species in the Nautilocalyx s. s. clade are dis-
more comprehensive taxon sampling is included, as well as
tributed in the Chocó biogeographic region in Colombia and
additional loci. The Nautilocalyx s. s. clade comprises species
Ecuador and Andes and inter-Andean valleys of Colombia.
mostly from Central America, northwestern South America
One exception is N. erytranthus (Fig. 4D), which occurs to (the Chocó Biogeographic region) and the northern Andes.
the east of the Andes in the Amazon basin of Colombia and Nautilocalyx s. s. is sister to Trichodrymonia and its species
northern Ecuador. mainly differ from the ones comprising the Amazonian
Trichodrymonia clade—The Trichodrymonia clade is com- “Nautilocalyx” clade by the presence of infundibuliform or
prised of most of the traditionally recognized species of trumpet-shaped corollas (vs. tubular) and linear calyces
Paradrymonia and includes the generic type species Trichodrymonia (vs. leafy and ovate). Future studies with increased taxon
congesta Oerst. (=Paradrymonia congesta (Oerst.) Wiehler). The sampling may support the segregation of the Amazonian
Trichodrymonia clade is a strongly supported monophyletic “Nautilocalyx” as a genus. Centrosolenia is resurrected based on
group (MP BS = 88; ML BS = 90; PP = 0.94) and is the sister molecular and morphological evidence. Nautilocalyx needs fur-
clade to Nautilocalyx s. s. (MP BS = 72; ML BS = 78 PP = 1.0). ther taxon sampling before taxonomic changes are proposed.
A major challenge for this project was locating the generic
type species of Trichodrymonia because it was only known from
the type specimen collected by Liebmann in 1842 in Chinantla, Taxonomic Treatment
Mexico. After more than a 150-yr hiatus, the first author In the present treatment an identification key to major
re-discovered this species on a 2010 collecting expedition to the clades and genera in the Paradrymonia alliance is provided.
type locality. The inclusion of Trichodrymonia congesta in the Trichodrymonia and Centrosolenia are resurrected to accommo-
molecular phylogeny is necessary to unambiguously assign a date species that were previously in Paradrymonia. Descriptions
genus name to this clade and to re-establish Trichodrymonia. for Chrysothemis, Paradrymonia s. s. and for the resurrected
To account for the phylogenetic relationships and to genera Trichodrymonia and Centrosolenia are included. An
accommodate this large segregate of Paradrymonia s. s., here index of names, including the new combinations, with the
we resurrect the genus Trichodrymonia and make 25 new com- accepted names in bold, is provided to facilitate referencing
binations (see taxonomic treatment below). The species in this currently recognized species circumscriptions with previous
clade share many vegetative characters with those belonging treatments (Appendix 2).

Key to the genera of the PARADRYMONIA alliance

1. Stem shorter than leaves or with leaves clustered at apex forming a rosette . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Leaves equal to subequal in a pair, usually variegated in young individuals, smaller leaf shaped like the larger one,
not deciduous, leaf base cuneate, rounded or subcordate; margin crenate; calyx lobes oblong-spatulate to
oblong-lanceolate, rounded or bluntly acute at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centrosolenia
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 93

2. Leaves strongly unequal in a pair, never variegated, smaller leaf usually stipule-like or deciduous, leaf blade typically long
decurrent on the petiole; margin serrate to serrulate, calyx lobes usually lanceolate to linear-lanceolate, acuminate at apex . . . . . . . . . . . . . 3
3. Ventral lobe of corolla entire, anthers glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichodrymonia
3. Ventral lobe of corolla fimbriate, anthers bearded with an apical tuft of elongate trichomes . . . . . . . . . . . . . . . . . . . . . . . . . . . Paradrymonia
1. Stem longer than leaves or with leaves evenly distributed along the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Leaves strongly unequal in a pair, the smaller one usually stipule-like or deciduous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichodrymonia
4. Leaves equal to moderately unequal in a pair, if unequal, the smaller leaf shaped like the larger one . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Stem with adventitious roots along the nodes and internodes, ventral lobe of the corolla with margin
long-fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paradrymonia barbata
5. Stem with adventitious roots absent or present only at some nodes; ventral lobe of the corolla with margin
entire to crenate, serrate or short-fimbriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Corolla narrow-tubular and elongate, tube about 5 × longer than broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centrosolenia
6. Corolla infundibuliform, trumpet shaped or salver-form, usually with tube less than 4 × longer than broad . . . . . . . . . . . . . . . . . . . . . . . 7
7. Leaf blade coriaceous with inconspicuous tertiary venation above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paradrymonia campostyla
7. Leaf blade usually thin with conspicuous tertiary venation above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Plants terrestrial, epiphytic, lithophytic or vining; stem decumbent, creeping or repent on the
substrate; calyx lobes free or nearly free, usually narrow, linear-lanceolate, oblanceolate,
never overlapping or conduplicate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Stems usually more than 1 m long, flowers usually several per leaf axil; calyx lobes lanceolate
to oblanceolate, corolla lobes less than 1 cm long (Amazon Basin and Guianas) . . . . . . . . . . . . . . . . . . . . . . . Pictus clade
9. Stems less than 1 m long, flowers usually 5 or fewer per leaf axil; infundibuliform to salverform
calyx lobes linear-lanceolate to ovate-lanceolate; corolla lobes 1 cm or longer (Central America,
Colombian inter-Andean valleys and Amazon Basin) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nautilocalyx s. s.
8. Plants terrestrial; stem predominantly erect, free-standing; calyx lobes connate into a cup or if free,
relatively wide, ovate-lanceolate, overlapping at the base or with margins folded in a conduplicate
manner such that each lobe is appressed to an adjacent lobe and folded lengthwise with the margin
curved upward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Plants forming tubers, leaf blade usually ovoid-elliptic to elliptic, usually rounded or subcordate
at base and ± oblique, sometimes decurrent into the petiole; leaf margin crenate to crenate-dentate;
calyx lobes nearly free or usually connate into a tube, calyx lobes green, red or yellow; corolla yellow,
orange, purplish or white, sometimes with the limb with red striations . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrysothemis
10. Plants never forming tubers, leaf blade usually oblanceolate, and attenuate at base,
decurrent into the petiole; leaf margin serrate; calyx lobes connate only at the very
base, green or tinged with purple; corolla cream-white or light yellow, sometimes
with the limb with spots wine-red to maroon-red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amazonian “Nautilocalyx” clade

Centrosolenia Benth., emend. M.M. Mora & J.L. Clark predominantly inhabit the slopes of the high tepuis (table
mountains), the summit areas of low elevation tepuis, or
Centrosolenia Benth., London J. Bot. 5: 362. 1846.—TYPE:
the understory of the forested high plains between 300
Centrosolenia hirsuta Benth.
and 1500 m.
Herbs, terrestrial or saxicolous; stems elongate, terete,
1. Centrosolenia bryogeton (Leeuwenb.) M. M. Mora & J. L.
decumbent, rooting and branching at the nodes. Leaves
Clark, comb. nov. Episcia bryogeton Leeuwenb., Acta Bot.
opposite, subequal to unequal in a pair; petioles usually
Neerl. 7: 312, 400. 1958. Nautilocalyx bryogeton (Leeuwenb.)
hirsute or densely villous; leaf blades broadly oblong or
Wiehler, Selbyana 5: 30. 1978.—TYPE: GUYANA. Cuyuni-
ovate-oblong, apex broadly rounded, shortly acute or rarely
Mazaruni: Kurupung River, near Makreba Falls, A.S.
subobtuse, base asymmetrical, rounded to subcordate, usually
Pinkus 12 (holotype: NY!; isotype: US!).
variegated and rugose or bullate above, scabrous or sparsely
pubescent, hairs aggregated in the center of the vein-areoles; 2. Centrosolenia chimantensis (L. E. Skog & Steyerm.)
margin shallowly crenate-dentate to serrate, lateral nerves M. M. Mora & J. L. Clark, comb. nov. Nautilocalyx
anastomosing near margin, conspicuously reticulated at chimantensis L. E. Skog & Steyerm., Novon 1: 217. 1991.—
least in the lower surface. Inflorescences axillary, cymose, TYPE: VENEZUELA. Bolívar: Chimantá Massif, common
2–8-flowered; pedicels short, densely hirsute to villous. Calyx along shaded wet bluffs and by waterfall along SW-
lobes nearly free, spathulate-oblong to lanceolate, apex obtuse facing sandstone bluffs and Chimantá-tepuí (Torono-
or rounded, shallowly 2–3 repand-dentate in upper one-third. tepuí), near southern corner, 1700 m, 19–20 May 1953,
Corolla oblique in calyx, narrowly tubular, gibbous basally J.A. Steyermark 75473 (holotype: US!; isotypes: F!, VEN).
on upper surface with slight ovate-oblong spur, villous
3. Centrosolenia coccinea (Feuillet & L. E. Skog) M. M.
throughout, white, violet, or deep red, corolla lobes orbicular,
Mora & J. L. Clark, comb. nov. Nautilocalyx coccineus
nearly equal with upper two lobes slightly reduced. Androe-
Feuillet & L. E. Skog, Brittonia 54: 352. 2003 (“2002”).—
cium with stamens adnate just above the base of the corolla
TYPE: GUYANA. Potaro-Siparuni: upper Potaro River
tube, filaments coiling after anthesis, anthers suborbicular or
Region, upper slopes of Mt. Wokomung, 5°05′N,
reniform, coherent in pairs. Disc a single dorsal nectariferous
59°50′W, 1540–1600 m, 11 Jul 1989, B.M. Boom & G.J.
gland. Gynoecium with narrowly ovoid ovary, pilose-sericeous,
Samuels 9186 (holotype: US!; isotype: NY!).
stigma stomatomorphic. Fruit a semi-succulent bivalved capsule.
Distribution and habitat — Centrosolenia currently includes 4. Centrosolenia crenata (Feuillet) M. M. Mora & J. L. Clark,
15 species that are endemic to the Guiana Shield. Most of comb. nov. Nautilocalyx crenatus Feuillet, J. Bot. Res. Inst.
the species are found growing on shady banks of rivers Texas 2: 825. 2008.—TYPE: VENEZUELA. Amazonas:
and streams, in crevices or on wet mossy rocks, or in the Cerro Yapacana, alrededores del campamento a lo largo
understory of the rainforests. In the Guiana region, they del río en las faldas en la parte suroeste, 3°45′N, 66°45′W,
94 SYSTEMATIC BOTANY [Volume 41

825 m, 4 May 1970 (fl), J. A. Steyermark & G. Bunting Yutajé, 9 km NW of settlement of Yutajé, along stream
103068 (holotype: US!; isotypes: NY!, VEN!). on plateau north of unnamed 1760 m peak, 5°41'N,
66°10′W, 1050–1300 m, 7 Mar 1987 (fl), R.L. Liesner &
5. Centrosolenia densa (C. H. Wright) Sprague, Bull. Misc.
B.K. Holst 21728 (holotype: US!; isotypes: MO!, VEN!).
Inform. Kew 1912: 87. 1912. Episcia densa C. H. Wright,
Bull. Misc. Inform. Kew 1895: 17. 1895. Paradrymonia 13. Centrosolenia vestita (Feuillet) M. M. Mora & J. L. Clark,
densa (C. H. Wright) Wiehler, Selbyana 5: 50. 1978.— comb. nov. Nautilocalyx vestitus Feuillet, J. Bot. Res.
TYPE: GUYANA. River Masouria, G.S. Jenman 2414 Inst. Texas 2: 833. 2008.—TYPE: VENEZUELA. Bolívar:
(holotype: K!). Cumbre del Cerro Guaiquinima, a lo largo del affluente
del Río Carapo, 1 km río arriba del Salto Szczerbanari,
6. Centrosolenia hirsuta Benth., London J. Bot. 5: 362. 5°44′N, 63°41′W, 730–750 m, 23–24 May 1978 (fl), J. A.
1846. Episcia hirsuta (Benth.) Hanst., Linnaea 34: 350. Steyermark, P. Berry, G. C. K. Dunsterville & E. Dunsterville
1865 (“1865-1866”), non Nautilocalyx hirsutus (Sprague) 117244 (holotype: VEN!).
Sprague (1912).—Type: Venezuela. Amazonas: banks
of the Río Paramu [“Parama” in the protologue], R.H. Chrysothemis Decne., emend. M. M. Mora & J. L. Clark
Schomburgk s.n. (holotype: K!). The type label says Chrysothemis Decne., Rev. Hort. (Paris) ser. 3.3: 242. 1849.—
“British Guiana” but this is an error based on expedi- TYPE: Chrysothemis pulchella Donn ex Sims
tion itineraries that were published by Rivière (2006)
and van Dam (2002). Herbs, terrestrial; tubers usually present; stems succulent,
subquadrangular, erect, branched or not, mostly with spread-
Episcia cordata Gleason, Bull. Torrey Bot. Club 58: 466. 1931. ing adventitious roots at the base. Leaves opposite, subequal
Nautilocalyx cordatus (Gleason) L.E. Skog in L. E. Skog & in a pair; petiole flattened above, maroon or green, often
Steyerm., Novon 1: 217. 1991.—TYPE: VENEZUELA. winged along the margin; leaf blades elliptic or oblong-
Amazonas: Tate 878 (holotype: NY!, isotypes: K!, US!). elliptic to oblong-lanceolate, apex acuminate, base cuneate
7. Centrosolenia orinocensis (Feuillet) M. M. Mora & J. L. and decurrent onto petiole or acute to rounded or subcordate
Clark, comb. nov. Nautilocalyx orinocensis Feuillet, J. at base and ± oblique, margins crenate or crenate-serrate,
Bot. Res. Inst. Texas 2: 833. 2008.—TYPE: VENEZUELA. dark green above, paler green or purplish below, usually
Amazonas: Upper Orinoco river, Sierra Guaharibo, near with margins red or purple; ends of primary veins arcuate,
Raudal de los Guaharibos, slopes of “Mt. Rimbaud,” secondary veins reticulate. Inflorescence axillary, with or
light growth near the top, 30 Jul 1951 (fl), L.C. Croizat without peduncles, sometimes an umbellate or rarely com-
429 (holotype: NY!). pound umbellate, sometimes reduced to a single axillary
flower, shorter than the subtended leaf; bracts, oblong,
8. Centrosolenia paujiensis (Feuillet) M. M. Mora & J. L. lanceolate or linear. Calyx lobes green, red or yellow, fre-
Clark, comb. nov. Nautilocalyx paujiensis Feuillet, J. Bot. quently fused for most of their length or sometimes connate
Res. Inst. Texas 2: 827. 2008.—TYPE: VENEZUELA. only at the very base or free; usually fused in an elongate
Bolívar: 17 km E of El Paují by road and 64 km W tube, when free the lobes are ovate. Corolla yellow, orange,
of Santa Elena by road, 4 km N of highway, Río Las purple or white, usually spotted or lined on the limb,
Ahallas, 4°30′N, 61°30′W, 850 m, 28 Oct 1985 (fl), R.L. trumpet-shaped nearly erect in the calyx or sometimes oblique
Liesner 19044 (holotype: US!; isotypes: MO!, VEN). in the calyx, base gibbous, rarely spurred, glabrous or pilose
9. Centrosolenia porphyrotricha (Leeuwenb.) M. M. Mora & outside, inside with glandular hairs in the throat; tube cylin-
J. L. Clark, comb. nov. Episcia porphyrotricha Leeuwenb. drical; limb spreading; lobes 5, subequal, suborbicular, entire.
Acta Bot. Neerl. 7: 311, Figure 26. 1958. Nautilocalyx Androecium with 4 stamens, didynamous, included, con-
porphyrotrichus (Leeuwenb.) Wiehler, Phytologia 27: 308. nate at the base, inserted at the base of the corolla, anthers
1973.—TYPE: GUYANA. Cuyuni Mazaruni: Pakaraima orbicular, dehiscent by a longitudinal slit. Gynoecium with
Mountains, Wenamu River, Jul 1925 (fl), Davenport 7 superior ovary, pubescent, ovoid, style mostly glabrous, erect;
(holotype: K!). stigma bilobed, pubescent with glandular hairs. Disc usually
a single dorsal nectary gland, rarely four. Fruit a globose
10. Centrosolenia pusilla (Feuillet) M. M. Mora & J. L. bivalved capsule. Seeds dark brown, obliquely striate, funiculi
Clark, comb. nov. Nautilocalyx pusillus Feuillet, J. Bot. elongate, white, translucent.
Res. Inst. Texas 2: 827. 2008.—TYPE: VENEZUELA. Distribution and habitat — A genus of nine species native
Bolívar: near El Paují, Río Cabass, waterfall, 4°30′N, to the West Indies, Mexico (Chiapas), Guatemala to Colombia,
61°35′W, 800–900 m, 3 Nov 1985 (fl), R.L. Liesner 19429 Venezuela, Guyana, Surinam, French Guiana and Brazil
(holotype: US!; isotypes: MO!, VEN!). (Amazonas). Growing in shaded areas along roads and
streams of wet forests.
11. Centrosolenia rosea (Feuillet) M. M. Mora & J. L. Clark,
comb. nov. Nautilocalyx roseus Feuillet, J. Bot. Res. Inst. 1. Chrysothemis adenosiphon (Leeuwenb.) M. M. Mora and
Texas 2: 830. 2008.—TYPE: VENEZUELA. Bolívar: N J. L. Clark, comb. nov. Episcia adenosiphon Leeuwenb.
side of Auyan tepui, along banks of Quebrada Honda, Acta Bot. Neerl. 18: 585. 1969. Nautilocalyx adenosiphon
Mar 1969 (fl), G.C.K. Dunsterville & E. Dunsterville s.n. (Leeuwenb.) Wiehler, Selbyana 5: 29. 1978. TYPE:
(holotype: VEN!). Venezuela. Bolívar, Steyermark 88162 (holotype: WAG!;
isotypes: VEN, WAG).
12. Centrosolenia rubra (Feuillet) M. M. Mora & J. L. Clark,
comb. nov. Nautilocalyx ruber Feuillet, J. Bot. Res. Inst. 2. Chrysothemis colonensis (Wiehler) M. M. Mora and
Texas 2: 830. 2008.—TYPE: VENEZUELA. Amazonas: J. L. Clark, comb. nov. Nautilocalyx colonensis Wiehler,
Dept. Atures, 4 km of Río Coro-Coro, W of Serranía de Selbyana 5: 89. 1978. TYPE: type collection made from
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 95

greenhouse-grown plant (origin of material: Panama. ex Sims) Mart. ex G. Don, Gen. Syst. 4: 656. 1838.
Colón, Río Escandaloso, tributary of Rio Boquerón, near Tussacia pulchella (Donn ex Sims) Riechenb. ex Walp.,
Mina #2, 20 May 1978, R. Dressler 5817, (PMA)), culti- Repert Bot. Syst. 6: 740. 1847. Skiophila pulchella (Donn
vated at the Marie Selby Botanical Gardens, MSBG live ex Sims) Hanst., Linnaea 26: 207. 1854.— Type: type col-
accession no. W-2573, 24 Aug 1978, H. Wiehler 78137 lection made from cultivated material, “Cult. Hort. Kew,”
(holotype: SEL!; isotypes: BH!, K!, MO!, NY!, PMA!, US!). J. Woodford s.n. (holotype: BM).
3. Chrysothemis dichroa Leeuwenb., Acta Bot. Neerl. 7: Besleria melittifolia sensu Drapiez, non L., Besleria umbellata
331. 1958.—TYPE: COLOMBIA. Vichada, R. Spruce 3612 Herb. Banks ex Sims, nom. nud. pro syn.
(holotype: K!; isotypes: BM!, BR!, P!, W)
Chrysothemis aurantiaca Decne., Rev. Hort. [ser. 3,4] 22: 381.
4. Chrysothemis friedrichsthaliana (Hanst.) H. E. Moore, 1855.— Type: type collection made from cultivated plant
Baileya 2: 87. 1954. Tussacia friedrichsthaliana Hanst., Linnaea (origin of material: Colombia), cultivated at Culta H.R.P.
34: 337. 1865.—TYPE: GUATEMALA or PANAMA, [cultivé à Paris] Serre chaude, Aug 1850 (lectotype
E. Friedrichsthal 3612 (holotype: W; isotypes: F, W). designated by Leeuwenberg, 1958: P!).
5. Chrysothemis kuhlmannii Hoehne, Sellowia 9: 43. Cyrtandromoea minor Ridl., J. Straits Branch Roy. Asiat. Soc.
1958.—TYPE: BRAZIL. Mato Grosso, J. Kuhlmann 2300 49: 20. 1908.—TYPE: MALAYSIA: Sarawak. HT: Anon.
(holotype: R). s.n. (SING).
6. Chrysothemis melittifolia (L.) M. M. Mora and J. L. Tussacia woodsonii C.V. Morton, Ann. Missouri Bot. Gard. 26:
Clark, comb. nov. Besleria melittifolia L. Sp. Pl. 2: 619. 308. 1939.—TYPE: PANAMA: Chiriquí: between Río
1753. Episcia melittifolia (L.) Mart., Nov. Gen. Sp. Pl. Chiriquí and Remedios,15-50 m, 11 July 1938. Woodson,
3: 42. 1829. Skiophila melittifolia (L.) Hanst., Linnaea 26: Allen & Seibert 1195 (holotype: US!; isotypes: GH, MO!,
207, 215, 1853. Alloplectus melittifolius (L.) Mart. ex NY!, US!).
Loud., (“mellitifolia”) Encycl. Pl. (new ed.) : 1402. 1855.
Chrysothemis melittifolia (L.) G. Don, (“melissaefolia”), Chrysothemis villosa (Benth.) Leeuwenb., Acta Bot. Neerl. 7:
Encycl. Pl. (new ed.) : 1402. 1855. Episcia melittifolia var. 338. 1958. Tussacia villosa Benth., London J. Bot. 5: 363.
typica Urb., Symb. Antill. 2 : 354. 1901. Episcia melittifolia 1846.—TYPE: Guyana. Upper Takutu-Upper Essequibo,
f. typica (Urb.) Stehlé, Bull. Soc. Bot. France 109: 31. 1962. Kanuku Mts., R. Schomburgk s.n. (holotype: K!).
Episcia melittifolia f. guadalupensis (DC.) Stehlé, Bull. Chrysothemis semiclausa (Hanst.) Leeuwenb. Acta Bot. Neerl.
Soc. Bot. France 109: 32. 1962. Nautilocalyx melittifolius 7: 338. 1958. Tussacia semiclausa Hanst., Illustr. Hortic.
(L.) Wiehler, Phytologia 27(5): 307. 1973. Nautilocalyx 17: 140, t. 28. 1870.— Type: Brazil. Rio Branco, Wallis
melittifolius var. guadalupensis (DC.) Fournet, comb. s.n. 1865 (holotype: B†?).
illeg., Fl. Ill. Phan. Guad. Mart. 1325. 1978.— Type:
plate 48 in Burmann, Plant Amer. Car. Plumier. 1756 9. Chrysothemis rupestris (Benth.) Leeuwenb. Acta Bot.
as “Besleria pedunculis ramosis.” (neotype, designated Neerl. 7: 336. 1958. Tussacia rupestris Benth., London J.
by Leeuwenberg, 1958: 308). Bot.5: 363.— Type: Guyana: Kanuku Mts., R. Schomburgk
s.n. (holotype: K!; photos: U, WAG).
Besleria guadalupensis DC., Prodr. 7: 538. 1839. Episcia guadalupensis
(DC.) Hanst. Linnaea 34 : 345. 1865.—TYPE: GUADELOUPE. Paradrymonia Hanst., emend. M. M. Mora & J. L. Clark
L’Herminier s.n. (lectotype: G-DC).
Paradrymonia Hanst., Linnaea 26: 207. 1854.— Type:
Chrysothemis venosa Decne., Rev. Hort. [ser. 3, 3] 21 : 242. 1849. Paradrymonia glabra (Benth.) Hanst.
7. Chrysothemis panamensis (Seem.) M. Mora and J. L. Episcia sect. Paradrymonia (Hanst.) Leeuwenb., Meded. Bot.
Clark, comb. nov. Scheeria panamensis Seem. Bot. Voy. Mus. Herb. Rijks Univ. Utrecht 146: 311. 1958.
Herald: 185. 1854. Nautilocalyx panamensis (Seem.) Seem.,
Epiphytes, facultative; stems subwoody, elongate, creeping
Bot. Voy. Herald, Suppl. 250. Figure 26. 1854. Achimenes
or ascending with many adventitious roots. Leaves usually
panamensis (Seem.) Hemsl. in Godm. & Salv., Biol. Cent.-
clustered on a short stem, opposite, rarely equal, usually
Amer., Bot. 2 : 475. 1882. Episcia panamensis (Seem.) C.V.
strongly unequal in a pair, when strongly anisophyllous, the
Morton Acta Bot. Venez. 2(2): 71. 1966.— Type: Panama.
smaller leaf usually sessile, reduced, subulate and early decid-
Seemann 235 (holotype: BM!; isotype: K!).
uous; larger leaf semi-erect and extending beyond the shoot
Drymonia villosa Kunth & Bouché, Index Seminum [Berlin] apex; often rather long–petiolate, petiole winged adaxially
1847: 12. 1847. Episcia villosa (Kunth & Bouché) Hanst., (U-shaped in cross section), stout and succulent; blade of
Linnaea 34: 348. 1865. Nautilocalyx villosus (Kunth & the larger leaf obovate to widely oblanceolate, acuminate
Bouché) Sprague, Bull. Misc. Inform. Kew 1912: 88. at the apex, cuneate at the base or long decurrent on the
1912.— Type: Venezuela, Caracas, Malcato. Gollmer s.n. petiole, coriaceous to succulent, margin serrulate some-
(holotype: B†). times with glandular teeth. Leaves of isophyllous species
(i.e. P. campostyla and P. barbata) rather short–petiolate;
Episcia inclinata Brandegee, Univ. Calif. Publ. Bot. 6: 63. 1914.—
leaf blade lanceolate to rarely oblanceolate, rounded to
TYPE: MEXICO: Chiapas: Finca Mexiquito, Purpus 6851
subcordate at base, coriaceous, hirsute, the margin crenate
(holotype: UC; isotypes: BM, F, MO!, NY!, US!, WAG(2)).
and ciliose. Inflorescences axillary, reduced pair-flowered
8. Chrysothemis pulchella (Donn ex Sims) Decne. Besleria cyme and appearing in fascicles; bracts linear, pedicel hirsute.
pulchella Donn, nom. nud., Besleria pulchella Donn ex Calyx lobes nearly free, hirsute; 4 subequal, linear to lanceo-
Sims, Bot. Mag. 28: pl. 1146. 1808. Episcia pulchella (Donn late, apex acuminate; fifth (ventral) lobe smaller and narrower.
96 SYSTEMATIC BOTANY [Volume 41

Corolla oblique in the calyx, tubular with a broad limb, base Cerro Marahuaca, “Sima Camp,” south–central portion
with well-developed spur, corolla lobes subequal, ventral of forested slopes along eastern branch of Caño Negro,
lobe slightly larger, margins range from entire to crenate, 3°43′N, 65°31′W, 1140 m, 21–24 Feb 1985 (fl), J.
or with fimbriations (on ventral lobe). Androecium with Steyermark & B. Holst 130443 (holotype: US!; isotypes:
stamens adnate just above the base of the corolla tube, MO!, VEN!).
filaments coiling after anthesis, anthers oblong, elongate
Paradrymonia hamata Feuillet, J. Bot. Res. Inst. Texas 3: 585.
trichomes clustered at base (barbate), longitudinally dehis-
2009.— Type: Venezuela. Amazonas. Depto. Río Negro,
cent, coherent in pairs. Nectary reduced to a large double-
Cerro de la Neblina, Río Yatúa, 140–1700 m, 31 Dec
connate dorsal gland, white, glabrous. Gynoecium with
1957 (fl), B. Maguire, J.J. Wurdack &. C.K. Maguire 42563
ovary villous or sericeous. Fruit a semi-fleshy, bivalved,
(holotype: NY!).
dehiscent capsule.
Distribution and habitat — A genus with approximately 5. Paradrymonia glabra (Benth.) Hanst., Linnaea 26: 207.
10 species distributed in the understory of rainforests of 1854. Centrosolenia glabra Benth. Bot. Mag. 76: t. 4552.
Central America, the Amazon basin, and the Guianas. 1850. Episcia glabra (Benth.) Hanst., Linnaea 34(3): 349.
Most species grow on moist rocks or logs in primary and 1865.—TYPE: type collection made from greenhouse-
secondary rainforests. grown plant (origin of material: Venezuela, La Guayra),
cultivated at the Royal Botanic Gardens, Kew, H. Wagener
1. Paradrymonia barbata Feuillet & L. E. Skog, Brittonia
s.n. (holotype: K!; isotype: K!).
54: 356, 2003, Fig. 3.— Type: GUYANA. Cuyuni-Mazaruni
Region: Permanent miner’s campsite near Eping River, 6. Paradrymonia lutea Feuillet, J. Bot. Res. Inst. Texas
E of several diamond pits, 6°00′N, 60°10′W, 122 m, 3: 585. 2009.— Type: Venezuela. Amazonas: Depto. Río
2 Feb 1991(fl), T. McDowell & A. Stobey 3810 (holotype: Negro, Neblina Massif, Canyon Grande, along Río
BRG; isotypes; K! US!). Mawarinuma, ca. 7 km ENE of Puerto Chimo, 0°50–51′
N, 66°02–06′W, 300 m, 9–14 Jul 1984 (fl), G. Davidse & J.S.
2. Paradrymonia buchtienii (Mansf.) Wiehler, Selbyana 5:49.
Miller 27212 (holotype: US!; isotypes: MO!, NY! VEN).
1978. Episcia buchtienii Mansf. Repert. Spec. Nov. Regni
Veg. 38: 25. 1935.— Type: Bolivia. La Paz: Mapiri. Buchtien 7. Paradrymonia tepui Feuillet, J. Bot. Res. Inst. Texas
1344 (holotype: B, destroyed; lectotype designated by 3: 588. 2009.— Type: Venezuela. Amazonas: Depto. Río
Wiehler, 1978: NY!; isolectotypes: HBG! NY! SEL! US!). Negro, Cerro Aracamuni, summit, Proa Camp, in ravines
and near edge of tepui, 01°32′N, 65°49′W, 1400 m, 31 Oct
3. Paradrymonia campostyla (Leeuwenb.) Wiehler, Selbyana
1987 (fl & fr), R.L. Liesner & G. Carnevali 22679 (holotype:
5: 49. 1978. Drymonia campostyla Leeuwenb., Acta Botanica
US!; isotype: MO!).
Neerlandica, 7: 305, 393. 1958.— Type: Surinam. Jonker &
Jonker 625 (holotype: U!; isotype: US!). 8. Paradrymonia yatua Feuillet, J. Bot. Res. Inst. Texas
3: 588. 2009.— Type: Venezuela. Amazonas: Depto. Río
4. Paradrymonia ciliosa (Mart.) Wiehler, Phytologia 27:
Negro, Río Yatúa, at base of Piedra Arauicaua, 100–140 m,
308. 1973. Hypocyrta ciliosa Mart. Nov. Gen. Sp. Pl. 3: 53.
3 Feb 1954 (fl), B. Maguire, J. J. Wurdack & G. S. Bunting
1829. Episcia ciliosa (Mart.) Hanst., Flora Brasiliensis 8(1):
37466 (holotype: US!; isotype: NY!).
403. 1864. Columnea ciliosa (Mart.) Kuntze, Revis. Gen.
Plantarum 2: 472. 1891.— Type: Brazil: Amazonas. Martius, Trichodrymonia Oerst. emend. M. M. Mora & J. L. Clark
Obs. 3117 s.n.; (holotype: M!, photos: U, US, WAG).
Trichodrymonia Oerst., Centralamer. Gesner. 38. 1858.—
Episcia hansteiniana Mansf., Repert. Spec. Nov. Regni Veg. Type: Trichodrymonia congesta Oerst.
38: 25. 1935. Paradrymonia hansteiniana (Mansf.) Wiehler,
Epiphytes, facultative; stems subwoody, usually reduced
Selbyana 5: 54. 1978.— Type: Peru. Loreto, Iquitos.
to basal rosette, with numerous adventitious roots. Leaves
Tessmann 5088 (holotype: B [destroyed]).— Type: Peru.
opposite, equal to strongly unequal in a pair, usually clus-
Loreto. Carretera Oleoducto Secundario entre los
tered during anthesis or widely spread on an elongate shoot
Campamentos Bartra 1 y Bartra 4, Diaz 1401 (neotype,
when young; petioles short or elongate, but always shorter
here designated: US!; isoneotypes: MO!, SEL!).
than the leaf-blade, larger leaf of anisophyllous plants semi-
Centrosolenia decurrens C. V. Morton, Publ. Field Mus. erect and extending beyond the shoot apex; smaller leaf
Nat. Hist., Bot. Ser. 18(4): 1158. 1938. Episcia decurrens sessile, reduced, subulate and early deciduous; the petiole of
(C. V. Morton) Leeuwenb., Acta Bot. Neerl. 8: 53. the larger leaf of a pair well-developed, winged adaxially
1959. Paradrymonia decurrens (C. V. Morton) Wiehler, (U-shaped in cross section), stout and succulent, usually
Phytologia 27:308. 1973.— Type: Costa Rica. Limón: maroon; leaf blades obovate to widely oblanceolate, some-
Finca Montecristo, on Río Reventazón, below El Cairo, times ovate, apex acuminate; base usually cuneate or long
25 m, 18–19 Feb 1926. P.C. Standley & J. Valerio 48589 decurrent along petiole, rarely subcordate; thin to coriaceous;
(holotype: US!). margin dentate to serrulate. Inflorescences many-flowered, in
sessile or short-pedunculate pair-flowerd cymes; bracts linear
Paradrymonia prististoma Wiehler, Phytologia 73: 233. 1992.—
to ovate; pedicels glabrous or hirsute. Calyx lobes nearly
Type: Ecuador. Napo: unfinished road from Tena to
free, linear to lanceolate, apex acuminate, glabrous to hirsute.
Latacunga, along Río Pano, 16–18 km from Tena, 23 Apr
Corolla oblique in the calyx, infundibuliform, trumpet-shaped
1986, H. Wiehler & GRF Expedition 86184 (holotype:
or salverform, rarely hypocyrtoid, spurred at the base.
SEL!: isotype: QCA!).
Androecium with stamens adnate just above the base of the
Paradrymonia glandulosa Feuillet, J. Bot. Res. Inst. Texas 3: 583. corolla tube, filaments coiling after anthesis, anthers oblong,
2009.— Type: Venezuela. Amazonas: Depto. Atabapo, glabrous, longitudinally dehiscent, coherent in pairs. Nectary
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 97

reduced to a large double-connate dorsal gland, white, gla- Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1159. 1938.
brous. Gynoecium with ovary glabrous to sericeous. Fruit a Paradrymonia congesta (Oerst.) Wiehler, Selbyana 5: 50.
semi-fleshy, bivalved, dehiscent capsule, rarely a berry. 1978.— Type: Mexico. Oaxaca: Chinantla, Oct 1842,
Distribution and habitat — A genus of 40 currently rec- F. M. Liebmann 9214 (holotype: C!; isotypes: C!, F!, K!,
ognized species and at least 10 that are new to science. M, US!).
Trichodrymonia is distributed from southern Mexico to Central
8. Trichodrymonia darienensis (Seem.) M. M. Mora & J. L.
America, the Andes and the Amazon basin. Members of this
Clark, comb. nov. Alloplectus darienensis Seem., Bot. Voy.
genus grow predominantly in the understory of rainforests,
Herald: 187. 1854. Episcia dariensis (Seem.) Leeuwenb.,
on wet slopes, stream banks, or on moist rocks or logs.
Acta. Bot. Neerl. 8: 52. 1959. Paradrymonia darienensis
1. Trichodrymonia alata (Kriebel) M. M. Mora & J. L. Clark, (Seem.) Wiehler, (“dariensis”), Selbyana 5: 50. 1978.—
comb. nov. Paradrymonia alata Kriebel, Rhodora 106: Type: Panama. Darién: Cape Corrientes, dark woods,
47. 2004.— Type: Costa Rica. Limón: Cordillera de B. T. Seemann 1058; (holotype: K!; isotypes: BM!, MO!).
Talamanca, 200 m aguas abajo de la confluencia de 9. Trichodrymonia erythropus (Hook. f.) M. M. Mora & J. L.
Quebrada Cañabral con Río Barbilla, 10°00′10″N, Clark, comb. nov. Episcia erythropus Hook. f., Bot. Mag.
83°25′30″W, 100 m, 5 Nov 1988, G. Herrera 2287 (holo- 102: t. 6219. 1876. Paradrymonia erythropus (Hook. f.)
type: INB; isotype: MO!). Wiehler, Selbyana 5: 50. 1978.—TYPE: type collection
2. Trichodrymonia alba (Wiehler) M. M. Mora & J. L. Clark, from cultivated material (origin of material: Colombia),
comb. nov. Paradrymonia alba Wiehler, Selbyana 5: 46, cultivated at Royal Botanic Gardens, Kew, Veitch s.n.
pl. 12A. 1978.— Type: Panama. Bocas del Toro: Filo de (holotype: K!).
Almirante, valley beyond ridge when approaching from 10. Trichodrymonia flava (Wiehler) M. M. Mora & J. L.
Almirante, 22 May 1972, H. Wiehler & R. Dressler 72303 Clark, comb. nov. Paradrymonia flava Wiehler, Selbyana
(holotype: SEL!). 5: 51, pl. 12C. 1978.— Type: Panama: Colón: Río Guanche,
3. Trichodrymonia apicaudata (M. M. Mora & J. L. Clark) near Portobelo, 10 Aug 1971. H. Wiehler & R. Dressler
M. M. Mora & J. L. Clark, comb. nov. Paradrymonia 71158. (holotype: SEL; isotypes: MO!, PMA).
apicaudata M. M. Mora & J. L. Clark, J. Bot. Res. Inst. 11. Trichodrymonia gibbosa (Wiehler) M. M. Mora & J. L.
Texas 6: 66. 2012.— Type: Colombia. Valle: From Clark, comb. nov. Paradrymonia gibbosa Wiehler, Selbyana
Campoalegre into area controlled by Corporación Valle 5: 52, pl. 12D. 1978.— Type: Colombia. Valle: Old road
del Cauca, trail uphill behind last camp (El Chanco), from Cali to Buenaventura, near La Elsa, 30 Apr 1972,
04°00′N, 076°40′W, 400–610 m, 17 Feb 1989, J. F. Smith, H. Wiehler & R. Dressler 7278 (lectotype, designated by
R. Bernal, X. Londoño & W. Devia 1357 (holotype: SEL!; Clark et al. 2003: SEL!).
isotypes: F!, MO!, US!, WIS!).
12. Trichodrymonia gigantea (Wiehler) M. M. Mora & J. L.
4. Trichodrymonia aurea (Wiehler) M. M. Mora & J. L. Clark, comb. nov. Paradrymonia gigantea Wiehler, Selbyana
Clark, comb. nov. Paradrymonia aurea Wiehler, Selbyana 5: 53, pl. 12A. 1978.—TYPE: type collection made from
5: 46, pl 12B. 1978.— Type: Ecuador. Pastaza: road Puyo greenhouse-grown plant (origin of material: Colombia.
to Canelos, 8 km E of Puyo, 21 Apr 1986, H. Wiehler & Valle: old road from Cali to Buenaventura, near La Elsa,
GRF Expedition 86129 (neotype, designated by Clark et al. 30 Apr 1972, H. Wiehler et al. 7236 (SEL)), cultivated at
2003: SEL!; isoneotype: US!). Marie Selby Botanical Gardens, live accession no W-1687,
Paradrymonia fuquaiana Wiehler, Phytologia 73: 232. 1992.— 3 Nov 1976, H. Wiehler et al. 76247 (holotype: SEL!;
Type: Ecuador. Napo: N of Tena, road Hollin-Loreto, isotype: US!).
21 Apr 1986, H. Wiehler & GRF Expedition 95116 (neo- 13. Trichodrymonia hirta (L. E. Skog) M. M. Mora & J. L.
type, designated by Clark et al. 2003: SEL!). Clark, comb. nov. Paradrymonia hirta L. E. Skog, Ann.
5. Trichodrymonia binata (Wiehler) M. M. Mora & J. L. Missouri Bot. Gard. 65: 967. 1979.— Type: Panama.
Clark, comb. nov. Paradrymonia binata Wiehler, Phytologia Darién: Cuasí-Caná trail between Cerro Campamiento
73: 231. 1992.— Type: Ecuador. Esmeraldas: km 5–18 and La Escalera to Páramo, east of Tres Bocas, cloud forest
on road Lita to Alto Tambo, 18 Jan 1987, C. H. Dodson, and mossy forest. Kirkbride & Duke 1293 (holotype: MO!)
A. Hitrz, D. Benzing, C. Luer & J. Luer 16833 (neotype, 14. Trichodrymonia hypocyrta (Wiehler) M. M. Mora &
designated by Clark et al. 2003: SEL!; isoneotype: MO). J. L. Clark, comb. nov. Paradrymonia hypocyrta Wiehler,
Selbyana 2: 82, pl. 25C. 1977.— Type: Ecuador. Los Ríos:
6. Trichodrymonia conferta (C. V. Morton) M. M. Mora &
Montaña de Ila, km 12, road from Patricia Pilar to 24
J. L. Clark, comb. nov. Centrosolenia conferta C. V. Morton,
de Mayo, 540 m, 30 May 1976, C. H. Dodson 6092 (holo-
J. Wash. Acad. Sci. 35(4): 126. 1945. Episcia conferta
type: SEL!).
(C. V. Morton) Leeuwenb., Acta Bot. Neerl. 8: 52. 1959.
Paradrymonia conferta (C. V. Morton) Wiehler, Selbyana 15. Trichodrymonia lacera (Wiehler) M. M. Mora & J. L.
5(1): 50. 197.— Type: Colombia. Antioquia: north of Clark, comb. nov. Paradrymonia lacera Wiehler, Selbyana
Dabeiba, on road to Turbo, 300–350 m, 25 Feb – 1 Mar 7: 343, pl. 5A. 1984.— Type: Ecuador. Cotopaxi: 3 km E
1942, R. D. Metcalf & J. Cuatrecasas 30200 (holotype: of El Palmar, on road from Quevedo to Latacunga, 800 m,
US!; isotype: US!). 5 Apr 1980, C. H. Dodson & A. H. Gentry 10194 (holotype:
SEL!; isotypes: QCA!, QCNE!).
7. Trichodrymonia congesta Oerst., Centralamer. Gesner.
38–39. 1858. Episcia congesta (Oerst.) Hanst. Linnaea 34: 16. Trichodrymonia lineata (C. V. Morton) M. M. Mora &
347. 1865. Centrosolenia congesta (Oerst.) C. V. Morton, J. L. Clark, comb. nov. Centrosolenia lineata C. V. Morton,
98 SYSTEMATIC BOTANY [Volume 41

Ann. Missouri Bot. Gard. 29: 41. 1942. Episcia lineata des plaines de Surubres,” Jul 1890, H Pittier & H.T.
(C. V. Morton) Leeuwenb., Acta Bot. Neerl. 8: 53. 1959. Durand 2654 [Biolley 343] (neotype, designated by Clark
Paradrymonia lineata (C. V. Morton) Wiehler, Phytologia et al. 2003: US!; isotype: BR).
27: 308. 1973.— Type: Panama. Coclé: hills on trail to
23. Trichodrymonia peltata (C. V. Morton) M. M. Mora &
La Mesa, north of El Valle de Anton, 1000 m, 31 Aug
J. L. Clark, comb. nov. Episcia peltata C. V. Morton,
1941, P.H. Allen 2717 (holotype:US!; isotypes: MO!).
J. Wash. Acad. Sci. 35: 131. 1945. Nautilocalyx peltatus
Paradrymonia lurida (C. V. Morton & Raymond) Wiehler, (C. V. Morton) Wiehler, Selbyana 5(1): 39. 1978.— Type:
Phytologia 27: 308. 1973. Episcia lurida C. V. Morton & Colombia. Antioquia: Collected on banks of Río Cauca
Raymond, Baileya 18: 9. 1971.—Type: type collection at Puerto Valdivia, 240–260 m, 17–20 Feb 1942. R.D.
made from greenhouse-grown plant (origin of material: Metcalf & J. Cuatrecasas 30098 (holotype: US!; isotypes:
Costa Rica. Alajuela, Cariblanco, Sarapiqui Valley, cut- F!, GH, UC).
tings then received by the Bailey Hortorium, Ithaca, NY
24. Trichodrymonia peltatifolia (J. L. Clark & M. M. Mora)
through H. E. Moore) cultivated at the Jardin Botanique
M. M. Mora & J. L. Clark, comb. nov. Paradrymonia
de Montreal, live accession no. 2209–60, Aug 1966,
peltatifolia J. L. Clark & M. M. Mora, Novon 23: 18.
C. Horich s.n. (holotype: US!; isotype: MTJB).
2014.— Type: Panama. Colón: Distr. Donoso, helipad
17. Trichodrymonia longipetiolata (Donn. Sm.) M. M. Mora & ZP-P9, 391 m, 08°51′5″N, 080°40′19″ W, 20 Jul. 2011 (fl.),
J. L. Clark, comb. nov. Episcia longipetiolata Donn. Sm. Bot. J. L. Clark & L. Martinez 12550 (holotype: US!; isotypes:
Gaz. 25: 152–153. 1898. Centrosolenia longipetiolata (Donn. E!, K!, MO!, NY!, PMA!, SCZ!, SEL!, UNA!).
Sm.) C.V. Morton, Publ. Field Mus. Nat. Hist., Bot. Ser.
25. Trichodrymonia sastrei (Wiehler) M. M. Mora & J. L.
18 : 1159. 1938. Paradrymonia longipetiolata (Donn. Sm.)
Clark, comb. nov. Paradrymonia sastrei Wiehler, Gesneriana
Wiehler, (“longipedunculata”) Selbyana 5: 54. 1978.—
1: 71, Figure 21. 1995.— Type: Colombia. Amazonas: Río
Type: Costa Rica. Guanacaste: borders of the road to
Igara-Parana (tributary of Río Putumayo), corregimiento
Carrillo, 300 m, Jun 1890, A. Tonduz 2493 (holotype:
La Chorrera, San Antonio, loma Obiraehidi, 9 Jul 1974,
US!; isotypes: BR!, CR, WAG!).
C. Sastre 3608 (holotype: P!).
18. Trichodrymonia macrophylla (Wiehler) M. M. Mora &
26. Trichodrymonia sericea (Wiehler) M. M. Mora & J. L.
J. L. Clark, comb. nov. Paradrymonia macrophylla Wiehler,
Clark, comb. nov. Paradrymonia sericea Wiehler, Selbyana
Selbyana 5: 56, pl. 13B. 1978.— Type: Panama. Coclé:
5: 57, pl 13C. 1978.— Type: type collection made from
hills N of El Valle de Anton, 15 Aug 1971, H. Wiehler &
greenhouse-grown plant (origin of material: Colombia.
R. Dressler 71286 (holotype: SEL!).
Valle: old road from Cali to Buenaventura, below La
19. Trichodrymonia maguirei (Feuillet) M. M. Mora & Elsa, on clay cliff near waterfall, 30 Apr 1972, H. Wiehler
J. L. Clark, comb. nov. Paradrymonia maguirei Feuillet, et al. 7261, sterile collection at SEL), cultivated at the
J. Bot. Res. Inst. Texas 3: 134. 2009.— Type: Venezuela. Marie Selby Botanical Gardens, MSBG live accession no.
Amazonas: Alto Orinoco, Cerro Marahuaca, 1000 m, W-1644, 13 May 1975, H. Wiehler 75270 (holotype: SEL!).
3 May 1949, B. Maguire & B. Maguire, Jr. 29185 (holotype:
27. Trichodrymonia splendens (Freiberg) M. M. Mora &
NY, pro parte: specimen A and material in the pocket).
J. L. Clark, comb. nov. Paradrymonia splendens Freiberg,
20. Trichodrymonia metamorphophylla (Donn. Sm.) M. M. Phyton (Horn, Austria) 37: 136.1997.— Type: Ecuador.
Mora & J. L. Clark, comb. nov. Alloplectus metamorpho- Imbabura: Los Cedros Biological Station, 17 Mar 1996,
phyllus Donn. Sm. Bot. Gaz. 52: 52. 1911. Paradrymonia M. Freiberg 96008 (holotype: QCA; isotype: ULM!).
metamorphophylla (Donn. Sm.) Wiehler, Phytologia 27:
28. Trichodrymonia tylocalyx (Wiehler) M. M. Mora & J. L.
327. 1973.— Type: Panama. Coclé: hills N of El Valle
Clark, comb. nov. Paradrymonia tylocalyx Wiehler, Selbyana
de Anton, 15 Aug 1971, H. Wiehler & R. Dressler 71286
5: 58, pl. 13D. 1978.— Type: type collection from cultivated
(holotype: SEL!; isotype: F!, K!, MO!, NY!, PMA, US!).
material (origin of material: Colombia. Valle: old road
21. Trichodrymonia ommata (L. E. Skog) M. M. Mora & from Cali to Buenaventura, below La Elsa, 30 Apr 1972,
J. L. Clark, comb. nov. Paradrymonia ommata L.E. Skog, H. Wiehler et al. 7253 (SEL)), cultivated at the Marie
Brittonia 30: 324. 1978.— Type: Panama. Bocas Del Toro: Selby Botanical Gardens, MSBG live accession no. W-1691,
Quebrada Huron, 90–120 m, 11 Apr 1968, J. Kirkbride & 13 May 1975, H. Wiehler 75271 (holotype: SEL!).
J. Duke 437 (holotype: MO!; isotypes: REED, SCZ).
29. Trichodrymonia ulei (Wiehler) M. M. Mora & J. L.
22. Trichodrymonia pedunculata (L. E. Skog) M. M. Mora & Clark, comb. nov. Paradrymonia ulei Wiehler, Gesneriana
J. L. Clark, comb. nov. Paradrymonia pedunculata L. E. 1: 71, Figure 22. 1995.— Type: Peru. Loreto: Pongo de
Skog, Brittonia 30: 325. 1978.— Type: Panama. Darién: Cainarachi, below Yurimaguas, Sep 1902, E. Ule 6328
Puerto St. Dorothea, 21 Jul 1962, J. Dwyer 2268 (holo- (holotype: HBG!).
type: MO!; isotype: US!).
Rhoogeton panamensis Wiehler, Phytologia 73: 239. 1992.— Acknowledgments. This research is based on a doctoral dissertation
Type: type collection made from greenhouse-grown in the Department of Biological Sciences at The University of Alabama.
plant (origin of material: Panama. Coclé: El Valle de Committee members are gratefully acknowledged for their feedback and
Anton, La Mesa, 19 Jun 1978, Dressler s. n.), cultivated support during the first author’s tenure as a graduate student. Those
committee members are: Juan Lopez-Bautista, Martha Powell, Leslie Rissler,
at the Gesneriad Research Foundation (GRF), live acces- Laurence E. Skog, and John L. Clark. This research was greatly facilitated
sion no. G-2633, 8 Jul 1985, H. Wiehler 8356 (holotype: by the Aquatic Biology Enhancement Fellowship. Additional financial
GES, not found).— Costa Rica. “Terres, rochers humides support was provided by the Nellie Sleeth Scholarship Endowment
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 99

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Washington, DC: Dept. of Botany, Smithsonian Institution. http://
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based on nrDNA and cpDNA trnL–F and trnE–T spacer region
Slovák, M., J. Kučera, E. Záveská, and P. Vd’ačný. 2014. Dealing with
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discordant genetic signal caused by hybridisation, incomplete line- Zhang, Q., T. S. Feild, and A. Antonelli. 2015. Assessing the impact of
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study of closely related members of the genus Picris subsection graphical history, and phylogenetic diversity. American Journal of
Hieracioides (Compositae). PLoS One 9(9): e104929. Botany 102: 566–580.
Smith, J. F. 2000. Phylogenetic resolution within the tribe Episcieae
(Gesneriaceae): congruence of ITS and ndhF sequences from parsi-
mony and maximum-likelihood analyses. American Journal of Botany
Appendix 1. Taxa included in the analyses, voucher information,
87: 883–897.
locality and GenBank accession numbers for ITS ETS and trnH-psbA.
Smith, J. F. and C. L. Carroll. 1997. A cladistic analysis of the tribe
Unvouchered samples taken from live material grown at the U.S. Botany
Episcieae (Gesneriaceae) based on ndhF sequences: origin of
Research Greenhouses (USBRG) are designated by their accession
morphological characters. Systematic Botany 22: 713–724.
number. Sequences not obtained are designated by—; * indicate generic
Smith, J. F. and J. L. Clark. 2013. Molecular phylogenetic analyses reveal
type species; herbarium acronyms follow Thiers (2015).
undiscovered monospecific genera in the tribe Episcieae (Gesneriaceae).
Systematic Botany 38: 451–463.
Taxon, Voucher, Locality, GenBank accessions: ITS, ETS, psbA-trnH
Smith, J. F., S. B. Draper, L. C. Hileman, and D. A. Baum. 2004. A phyloge-
spacer
netic analysis within tribes Gloxinieae and Gesnerieae (Gesnerioideae:
Gesneriaceae). Systematic Botany 29: 947–958. Ingroup—Alloplectus hispidus (Kunth) Mart., J.L. Clark 7720 (US),
Smith, J. F., J. C. Wolfram, K. D. Brown, C. L. Carroll, and D. S. Denton. Ecuador, DQ211111, KF040256, DQ211219; Alloplectus weirii (Kuntze)
1997. Tribal relationships in the Gesneriaceae: evidence from DNA Wiehler, J.L. Clark 5788 (US), Ecuador, AF543233, KF040257, DQ211192;
sequences of the plastid gene ndhF. Annals of the Missouri Botanical Alsobia dianthiflora (H. E. Moore & R. G. Wilson) Wiehler, J. Hall s. n.
Garden 84: 50–66. (SEL), Cultivated (Costa Rica), DQ211160, KF040258, DQ211303; Alsobia
Sprague, T. A. 1912. The genus Nautilocalyx. Bulletin of Miscellaneous punctata (Lindl.) Hanst. *, J.L. Clark 8851 (US), Cultivated (Mexico),
Information, Royal Botanic Gardens, Kew. 1912: 85–90. DQ211159, KF040259, DQ211302; Chrysothemis friedrichstaliana (Hanst.)
Stamatakis, A., P. Hoover, and J. Rougemont. 2008. A rapid bootstrap H. E. Moore, J.L. Clark 10018 (US), Colombia, KF040179, KF040261,
algorithm for the RAxML web servers. Systematic Biology 57: 758–771. KF040106; Chrysothemis pulchella (Donn. Sm. ex Sims) Decne. *, J.L.
Štorchová, H. and M. S. Olson. 2007. The architecture of the chloroplast Clark 8864 (US), Cultivated (unknown), KF040180, KF040262, DQ211344;
psbA-trnH non-coding region in angiosperms. Plant Systematics and Cobananthus calochlamys (Donn. Sm.) Wiehler *, J.L. Clark 5613 (US),
Evolution 268: 235–256. Cultivated (Guatemala), AF543273, KF040263, DQ211304; Codonanthe
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 101

carnosa (Gardner) Hanst., J.L. Clark 6268 (US), Cultivated (Brazil), KF040281, KF040127; Nautilocalyx sp. Mora 939, M.M. Mora 939 (US),
AF543271, KF040264, DQ211296; Codonanthe gracilis (Mart.) Hanst., J.F. Peru, KF040202, KF040282, KF040128; Nautilocalyx sp. Mora 949, M.M.
Smith 3721 (US), Cultivated (Brazil), KF040183, KF040265, KF040110; Mora 949 (US), Peru, KF040203, KF040283, KF040129; Nautilocalyx sp.
Codonanthopsis anisophylla (Feuillet & L.E. Skog) Chautems & Mat. Mora 970, M.M. Mora 970 (UNA), Ecuador, KF040198, KF040278,
Perret, H.D. Clarke 10413 (US), Guyana, DQ211181, KF040305, DQ211334; KF040124; Nematanthus albus Chautems, J.L. Clark 6266 (US), Culti-
Codonanthopsis ulei Mansf., J.L. Clark 8868 (US), Cultivated (Brazil), vated (Brazil), AF543270, KF040300, DQ211318; Nematanthus corticola
DQ211167, KF040260, DQ211314; Columnea dressleri Wiehler, J.L. Clark Schrad. *, J.L. Clark 6271 (US), Cultivated (Brazil), AF543268, KF040301,
8559 (US), Panama, DQ211117, KF040266, DQ211230; Columnea linearis DQ211316; Pagothyra maculata (Hook. f.) J.F. Sm. & J.L. Clark *, K. Redden
Oerst., J.L. Clark 6274 (US), Cultivated (Costa Rica), AF543240, KF040267, 2231 (US), French Guiana, KF040245, KF040321,—; Paradrymonia aurea
DQ211243; Columnea scandens L., J.L. Clark 8879 (US), Cultivated Wiehler, J.L. Clark 5409 (US), Ecuador, KF040228, KF040306, KF040150;
(unknown), KF040186, KF040268, KF040113; Crantzia cristata (L.) Paradrymonia binata Wiehler, J.L. Clark 8848 (US), Ecuador, KF040229,—,
Scop., J.L. Clark 6546 (US), Martinique, DQ211154, KF040269, DQ211294; DQ211307; Paradrymonia campostyla (Leeuwenb.) Wiehler, J.L. Clark
Cremersia platula Feuillet & L.E. Skog *, J.J. de Granville 14868 (CAY), 8855 (US), Cultivated (French Guiana), DQ211180, KF040307, DQ211333;
French Guiana, DQ211152,—,—; Drymonia killipii Wiehler, J.L. Clark Paradrymonia ciliosa (Mansf.) Wiehler, H.D. Clarke 10239 (US), Guyana,
7521 (US), Ecuador, DQ211136, KF040270, DQ211271; Drymonia lanceolata DQ211182, KF040308, DQ211335; Paradrymonia congesta (Oerst.)
(Hanst.) C.V.Morton, J.L. Clark 8553 (US), Panama, DQ211139, KM079401, Wiehler, M.M. Mora 969 (US), Mexico, KF040232, KF040309, KF040154;
DQ211276; Drymonia longifolia Poepp., J.L. Clark 6262 (US), Ecuador, Paradrymonia densa (C.H. Wright) Wiehler, K. Redden 1060 (US), Guyana,
AF543264, KM079371, DQ211340; Drymonia urceolata Wiehler, J.L. Clark DQ211184, KF040310,—; Paradrymonia erythropus (Hook. f.) Wiehler,
5225 (US), Ecuador, AF543265, KF040273, DQ211289; Episcia cupreata M.M. Mora 805 (US), Colombia, KF040234, KF040311, KF040155;
(Hook.) Hanst., J.L. Clark 8844 (US), Cultivated (Colombia), DQ211165, Paradrymonia flava Wiehler, J.L. Clark 12547 (US), Panama, KF040223,—,
KF040274, DQ211312; Episcia lilacina Hanst., J.L. Clark 8881 (US), Costa KF040146; Paradrymonia flava Wiehler, J.L. Clark 8846 (US), Panama,
Rica, KF040194, KF040275, DQ211309; Glossoloma medusaeum (L.E. Skog) KF040235,—, KF040156; Paradrymonia gibbosa Wiehler, M.M. Mora 832
J.L.Clark, J.L. Clark 4973 (US), Ecuador, AF543223, KF040276, DQ211200; (US), Colombia, KF040236, KF040312, KF040157; Paradrymonia gigantea
Glossoloma tetragonum Hanst., J.L. Clark 8547 (US), Panama, DQ211104, Wiehler, M.M. Mora 846 (US), Colombia, KF040237, KF040313, KF040158;
KF040277, DQ211207; Lembocarpus amoenus Leeuwenb. *, J.L. Clark 8841 Paradrymonia glabra (Benth.) Hanst., J.L. Clark 10075 (UNA), Cultivated
(US), Cultivated (French Guiana), DQ211172,—, DQ211323; Nautilocalyx (Venezuela), KF040238, KF040314, KF040159; Paradrymonia hypocyrta
antioquensis Wiehler, M.M. Mora 806 (US), Colombia, KF040205, KF040285, Wiehler, J.L. Clark 10010 (US), Ecuador, KF040240, KF040316, KF040160;
KF040130; Nautilocalyx bracteatus (Planch.) Sprague, M.M. Mora 800 (US), Paradrymonia lacera Wiehler, J.L. Clark 11942 (UNA), Cultivated
Colombia, KF040206, KF040286, KF040131; Nautilocalyx bullatus (Lem.) (Ecuador), KF040241, KF040317, KF040161; Paradrymonia macrophylla
Sprague, M.M. Mora 971 (UNA), Cultivated (Peru), KF040207, KF040287, Wiehler, J.L. Clark 8545 (US), Panama, DQ211174, KF040319, DQ211325;
KF040132; Nautilocalyx coccineus Feuillet & L.E. Skog, H.D. Clarke 10295 Paradrymonia macrophylla Wiehler, M.M. Mora 811 (US), Ecuador,
(US), Guyana, DQ211185, KF040288, DQ211338; Nautilocalyx colombianus KF040244, KF040320, KF040164; Paradrymonia metamorphophylla (Donn.
Wiehler, J.L. Clark 12454 (US), Panama, KF040209, KF040289, KF040134; Sm.) Wiehler, J.L. Clark 6028 (US), Ecuador, DQ211178, KF040322,
Nautilocalyx cordatus (Gleason) L.E. Skog, K. Redden 2359 (US), Guyana, DQ211330; Paradrymonia pedunculata L.E. Skog, USBRG 1994-184
KF040210, KF040290, KF040135; Nautilocalyx erytranthus J.L. Clark & (US), Cultivated (Costa Rica & Panama), DQ211179, KF040323, DQ211332;
M.M. Mora, J.L. Clark 8268 (US), Ecuador, KF040199, KF040279, KF040125; Paradrymonia peltatifolia J.L. Clark & M.M. Mora, J.L. Clark 12550 (US),
Nautilocalyx forgetii (Sprague) Sprague, J.L. Clark 8847 (US), Culti- Panama, KF040248, KF040324, KF040167; Paradrymonia sericea Wiehler,
vated (Peru), KF040212, KF040292, KF040137; Nautilocalyx glandulifer M.M. Mora 838 (US), Colombia, KF040249, KF040325, KF040168;
Wiehler, J.L. Clark 10012 (US), Ecuador, KF040211, KF040291, KF040136; Paradrymonia splendens M.Freiberg, J.L. Clark 7351 (US), Ecuador,
Nautilocalyx hirsutus (Sprague) Sprague, M.M. Mora 950 (US), Peru, DQ211173,—,—; Paradrymonia ulei Wiehler, M.M. Mora 866 (US), Peru,
KF040204, KF040284,—; Nautilocalyx melittifolius (L.) Wiehler, J.L. Clark KF040251, KF040326, KF040169; Paradrymonia sp. Mora 816, M.M. Mora
6540 (US), Martinique, GQ344532, KF040293, DQ211326; Nautilocalyx 816 (US), Colombia, KF040224, KF040302, KF040147; Paradrymonia
pallidus (Sprague) Sprague, J.L. Clark 9352 (US), Ecuador, KF040214, sp. Mora 823, M.M. Mora 823 (US), Colombia, KF040242, KF040318,
KF040294, KF040139; Nautilocalyx panamensis (Seem.) Seem., J.L. Clark KF040162; Paradrymonia sp. Mora 845, M.M. Mora 845 (US), Colombia,
12735 (US), Panama, KF040215, KF040295,—; Nautilocalyx pemphidius KF040239, KF040315,—; Paradrymonia sp. Mora 842, M.M. Mora 842
L.E. Skog, D. Bell 324 (US), Venezuela, DQ211176, KF040296, DQ211328; (US), Colombia, KF040225, KF040303, KF040148; Paradrymonia sp. Mora
Nautilocalyx pictus (W. Hook.) Sprague, H.D. Clarke 9974 (US), Guyana, 895, M.M. Mora 895 (US), Colombia, KF040226, KF040304,—; Rhoogeton
DQ211188,—, DQ211342; Nautilocalyx porphyrotrichus (Leeuwenb.) cyclophyllus Leeuwenb. *, H.D. Clarke 10350 (US), Guyana, DQ211163,
Wiehler, J.L. Clark 10449 (UNA), Cultivated (unknown), KF040218, KF040327, DQ211310; Rhoogeton viviparus Leeuwenb., H.D. Clarke 9255
KF040297,—; Nautilocalyx punctatus Wiehler, K. Redden 3407 (US), (US), Guyana, DQ211164, KF040328, DQ211311.
Venezuela, KF040219, KF040298, KF040142; Nautilocalyx whitei Rusby, J.L. Outgroup—Sinningia cooperi (Paxt.) Wiehler, J.L. Clark 8857 (US),
Clark 6793 (US), Bolivia, DQ211189, KF040299, DQ211343; Nautilocalyx Cultivated (Brazil), DQ211097, KF040329, DQ211299; Sinningia incarnata
sp. Mora 931, M.M. Mora 931 (US), Peru, KF040200, KF040280, KF040126; (Aubl.) D.L. Denham, J.L. Clark 8849 (US), Cultivated (Colombia),
Nautilocalyx sp. Mora 933, M.M. Mora 933 (US), Peru, KF040201, JQ953785, KF040330, DQ211300.
102 SYSTEMATIC BOTANY [Volume 41

Appendix 2. Index of names for Centrosolenia, Chrysothemis, Paradrymonia and Trichodrymonia (Accepted names in bold).

Alloplectus darienensis Seem. Trichodrymonia darienensis

Alloplectus metamorphophyllus Donn. Sm. Trichodrymonia metamorphophylla
Besleria melittifolia L. Chrysothemis melitifolia
Besleria pulchella Donn ex Sims Chrysothemis pulchella
Besleria pulchella Donn, nom. nud. Chrysothemis pulchella
Centrosolenia bryogeton (Leeuwenb.) M. M. Mora & J. L. Clark
Centrosolenia chimantensis (L. E. Skog & Steyermark) M. M. Mora & J. L. Clark
Centrosolenia coccinea (Feuillet & L. E. Skog) M. M. Mora & J. L. Clark
Centrosolenia conferta C. V. Morton Trichodrymonia conferta
Centrosolenia congesta (Oerst.) C. V. Morton Trichodrymonia congesta
Centrosolenia crenata (Feuillet) M. M. Mora & J. L. Clark
Centrosolenia decurrens C. V. Morton Paradrymonia ciliosa
Centrosolenia densa (C. H. Wright) Sprague
Centrosolenia glabra Benth. Paradrymonia glabra
Centrosolenia hirsuta Benth
Centrosolenia lineata C.V. Morton Trichodrymonia lineata
Centrosolenia orinocensis (Feuillet) M. M. Mora & J. L. Clark
Centrosolenia paujiensis (Feuillet) M. M. Mora & J. L. Clark
Centrosolenia porphyrotricha (Leeuwenb.) M. M. Mora & J. L. Clark
Centrosolenia pusilla (Feuillet) M. M. Mora & J. L. Clark
Centrosolenia rosea (Feuillet) M. M. Mora & J. L. Clark
Centrosolenia rubra (Feuillet) M. M. Mora & J. L. Clark
Centrosolenia vestita (Feuillet) M. M. Mora & J. L. Clark
Chrysothemis adenosiphon (Leeuwenb.) M. M. Mora and J. L. Clark
Chrysothemis colonensis (Wiehler) M. M. Mora and J. L. Clark
Chrysothemis dichroa Leeuwenb.
Chrysothemis friedrichsthaliana (Hanst.) H. E. Moore
Chrysothemis kuhlmannii Hoehne
Chrysothemis melitifolia (L.) M. M. Mora and J. L. Clark
Chrysothemis panamensis (Seem.) M. M. Mora and J. L. Clark
Chrysothemis pulchella (Donn ex Sims) Decne.
Chrysothemis rupestris (Benth.) Leeuwenb.
Chrysothemis semiclausa (Hanst.) Leeuwenb.
Columnea ciliosa (Mart.) Kuntze Paradrymonia ciliosa
Drymonia campostyla Leeuwenb. Paradrymonia campostyla
Episcia adenosiphon Leeuwenb. Chrysothemis adenosiphon
Episcia bryogeton Leeuwenb. Centrosolenia bryogeton
Episcia buchtienii Mansf. Paradrymonia buchtienii
Episcia ciliosa (Mart.) Hanst. Paradrymonia ciliosa
Episcia conferta (C.V. Morton) Leeuwenb. Trichodrymonia conferta
Episcia congesta (Oerst.) Hanst. Trichodrymonia congesta
Episcia cordata Gleason Centrosolenia hirsuta
Episcia dariensis (Seem.) Leeuwenb. Trichodrymonia darienensis
Episcia decurrens (C. V. Morton) Leeuwenb. Paradrymonia ciliosa
Episcia densa C. H. Wright Centrosolenia densa
Episcia erythropus Hook. f. Trichodrymonia erythropus
Episcia glabra (Benth.) Hanst. Paradrymonia glabra
Episcia hansteiniana Mansf. Paradrymonia ciliosa
Episcia hirsuta (Benth.) Hanst. Centrosolenia hirsuta
Episcia lineata (C. V. Morton) Leeuwenb. Trichodrymonia lineata
Episcia longipetiolata Donn. Sm. Trichodrymonia longipetiolata
Episcia lurida C. V. Morton & Raymond Trichodrymonia lineata
Episcia peltata C. V. Morton Trichodrymonia peltata
Episcia porphyrotricha Leeuwenb. Centrosolenia porphyrotricha
Episcia pulchella (Donn ex Sims) Mart. ex G. Don Chrysothemis pulchella
Hypocyrta ciliosa Mart. Paradrymonia ciliosa
Nautilocalyx bryogeton (Leeuwenb.) Wiehler Centrosolenia bryogeton
Nautilocalyx chimantensis L. E. Skog & Steyermark Centrosolenia chimantensis
Nautilocalyx coccineus Feuillet & L. E. Skog Centrosolenia coccinea
Nautilocalyx colonensis Wiehler Chrysothemis colonensis
Nautilocalyx cordatus (Gleason) L. E. Skog Centrosolenia hirsuta
Nautilocalyx crenatus Feuillet Centrosolenia crenata
Nautilocalyx orinocensis Feuillet Centrosolenia orinocensis
Nautilocalyx panamensis (Seem.) Seem. Chrysothemis panamensis
Nautilocalyx paujiensis Feuillet Centrosolenia paujiensis
Nautilocalyx porphyrotrichus (Leeuwenb.) Wiehler Centrosolenia porphyrotricha
Nautilocalyx pusillus Feuillet Centrosolenia pusilla
Nautilocalyx roseus Feuillet Centrosolenia rosea
Nautilocalyx ruber Feuillet Centrosolenia rubra
Nautilocalyx vestitus Feuillet Centrosolenia vestita
Nautilocalyx villosus (Kunth & Bouché) Sprague Chrysothemis panamensis
Paradrymonia alata Kriebel Trichodrymonia alata
(Continued)
2016] MORA AND CLARK: PHYLOGENY OF PARADRYMONIA (GESNERIACEAE) 103

Appendix 2. (Continued).

Paradrymonia alba Wiehler Trichodrymonia alba

Paradrymonia apicaudata M. M. Mora & J. L. Clark Trichodrymonia apicaudata
Paradrymonia aurea Wiehler Trichodrymonia aurea
Paradrymonia barbata Feuillet & L. E. Skog
Paradrymonia binata Wiehler Trichodrymonia binata
Paradrymonia buchtienii (Mansf.) Wiehler
Paradrymonia campostyla (Leeuwenb.) Wiehler
Paradrymonia ciliosa (Mart.) Wiehler
Paradrymonia conferta (C.V. Morton) Wiehler Trichodrymonia conferta
Paradrymonia congesta (Oerst.) Wiehler Trichodrymonia congesta
Paradrymonia darienensis (Seem.) Wiehler Trichodrymonia darienensis
Paradrymonia decurrens (C. V. Morton) Wiehler Paradrymonia ciliosa
Paradrymonia densa (C.H. Wright) Wiehler Centrosolenia densa
Paradrymonia erythropus (Hook. f.) Wiehler Trichodrymonia erythropus
Paradrymonia flava Wiehler Trichodrymonia flava
Paradrymonia fuquaiana Wiehler Trichodrymonia aurea
Paradrymonia gibbosa Wiehler Trichodrymonia gibbosa
Paradrymonia gigantea Wiehler Trichodrymonia gigantea
Paradrymonia glabra (Benth.) Hanst.
Paradrymonia glandulosa Feuillet Paradrymonia ciliosa
Paradrymonia hamata Feuillet Paradrymonia ciliosa
Paradrymonia hansteiniana (Mansf.) Wiehler Paradrymonia ciliosa
Paradrymonia hirta L.E. Skog Trichodrymonia hirta
Paradrymonia hypocyrta Wiehler Trichodrymonia hypocyrta
Paradrymonia lacera Wiehler Trichodrymonia lacera
Paradrymonia lineata (C. V. Morton) Wiehler Trichodrymonia lineata
Paradrymonia longipetiolata (Donn. Sm.) Wiehler Trichodrymonia longipetiolata
Paradrymonia lurida (C. V. Morton & Raymond) Wiehler Trichodrymonia lineata
Paradrymonia lutea Feuillet
Paradrymonia macrophylla Wiehler Trichodrymonia macrophylla
Paradrymonia maguirei Feuillet Trichodrymonia maguirei
Paradrymonia metamorphophylla (Donn. Sm.) Wiehler Trichodrymonia metamorphophylla
Paradrymonia ommata L. E. Skog Trichodrymonia ommata
Paradrymonia pedunculata L. E. Skog Trichodrymonia pedunculata
Paradrymonia peltatifolia J. L. Clark & M. M. Mora Trichodrymonia peltatifolia
Paradrymonia prististoma Wiehler Paradrymonia ciliosa
Paradrymonia sastrei Wiehler Trichodrymonia sastrei
Paradrymonia sericea Wiehler Trichodrymonia sericea
Paradrymonia splendens Freiberg Trichodrymonia splendens
Paradrymonia tepui Feuillet
Paradrymonia tylocalyx Wiehler Trichodrymonia tylocalyx
Paradrymonia ulei Wiehler Trichodrymonia ulei
Paradrymonia yatua Feuillet
Rhoogeton panamensis Wiehler Trichodrymonia pedunculata
Scheeria panamensis Seem. Chrysothemis panamensis
Skiophila melittifolia (L.) Hanst. Chrysothemis melittifolia
Skiophila pulchella (Donn ex Sims) Hanst. Chrysothemis pulchella
Trichodrymonia alata (Kriebel) M.M. Mora & J.L. Clark
Trichodrymonia alba (Wiehler) M.M. Mora & J.L. Clark
Trichodrymonia apicaudata (M. M. Mora & J. L. Clark) M. M. Mora & J. L. Clark
Trichodrymonia aurea (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia binata (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia conferta (C.V. Morton) M. M. Mora & J. L. Clark
Trichodrymonia congesta Oerst.
Trichodrymonia darienensis (Seem.) M. M. Mora & J. L. Clark
Trichodrymonia erythropus (Hook. f.) M. M. Mora & J. L. Clark
Trichodrymonia flava (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia gibbosa (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia gigantea (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia hirta (L. E. Skog) M. M. Mora & J. L. Clark
Trichodrymonia hypocyrta (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia lacera (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia lineata (C. V. Morton) M. M. Mora & J. L. Clark
Trichodrymonia longipetiolata (Donn. Sm.) M. M. Mora & J. L. Clark
Trichodrymonia macrophylla (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia maguirei (Feuillet) M. M. Mora & J. L. Clark
Trichodrymonia metamorphophylla (Donn. Sm.) M. M. Mora & J. L. Clark
Trichodrymonia ommata (L. E. Skog) M. M. Mora & J. L. Clark
Trichodrymonia pedunculata (L. E. Skog) M. M. Mora & J. L. Clark
Trichodrymonia peltata (C. V. Morton) M. M. Mora & J. L. Clark
Trichodrymonia peltatifolia (J. L. Clark & M. M. Mora) M. M. Mora & J. L. Clark
Trichodrymonia sastrei (Wiehler) M. M. Mora & J. L. Clark
(Continued)
104 SYSTEMATIC BOTANY [Volume 41

Appendix 2. (Continued).

Trichodrymonia sericea (Wiehler) M. M. Mora & J. L. Clark

Trichodrymonia splendens (Freiberg) M. M. Mora & J. L. Clark
Trichodrymonia tylocalyx (Wiehler) M. M. Mora & J. L. Clark
Trichodrymonia ulei (Wiehler) M. M. Mora & J. L. Clark
Tussacia friedrichsthaliana Hanst. Chrysothemis friedrichsthaliana
Tussacia pulchella (Donn ex Sims) Riechenb. ex Walp. Chrysothemis pulchella
Tussacia rupestris Benth. Chrysothemis rupestris
Tussacia semiclausa Hanst. Chrysothemis semiclausa
Tussacia woodsonii C.V. Morton Chrysothemis pulchella