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Variation in fruits in Episcieae. A, Fleshy display capsule in Nematanthus albus. B, Fleshy display capsule in Drymonia chiribogana. C, Indehiscent fleshy berry in Paradrymonia metamorphophylla. D, Semifleshy capsule in Paradrymonia ciliosa. E, Dry capsule in Episcia xantha. F, Semifleshy capsule in Episcia lilacina. G, Dry capsule showing splash-cup seed dispersal in Lembocarpus amoenus. H, Semifleshy capsule in Nautilocalyx panamensis. (Voucher specimens at US: B, J. L. Clark 7358; C, J. L. Clark 9874; D, J. L. Clark 6791; E, J. L. Clark 11323; F, J. L. Clark 1259; G, J. L. Clark 8841; H, J. L. Clark 12735. Photos: A by Jiri R. Haager, B–G by John L. Clark, and H by James F. Di Loreto.)  

Variation in fruits in Episcieae. A, Fleshy display capsule in Nematanthus albus. B, Fleshy display capsule in Drymonia chiribogana. C, Indehiscent fleshy berry in Paradrymonia metamorphophylla. D, Semifleshy capsule in Paradrymonia ciliosa. E, Dry capsule in Episcia xantha. F, Semifleshy capsule in Episcia lilacina. G, Dry capsule showing splash-cup seed dispersal in Lembocarpus amoenus. H, Semifleshy capsule in Nautilocalyx panamensis. (Voucher specimens at US: B, J. L. Clark 7358; C, J. L. Clark 9874; D, J. L. Clark 6791; E, J. L. Clark 11323; F, J. L. Clark 1259; G, J. L. Clark 8841; H, J. L. Clark 12735. Photos: A by Jiri R. Haager, B–G by John L. Clark, and H by James F. Di Loreto.)  

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The Gesneriaceae is a family known for convergent evolution of complex floral forms. As a result, defining genera and resolving evolutionary relationships among such genera using morphological data alone has been challenging and often does not accurately reflect monophyletic lineages. The tribe Episcieae is the most diverse within Neotropical Gesne...

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... The analyses by Clark et al. (2006), Clark et al. (2012), and Mora and Clark (2016) pointed out the need for taxonomic reorganization for the clade composed of Paradrymonia, Nautilocalyx, and Chrysothemis since these are probably non-monophyletic genera, and the morphology of the pollen grains of their species can be used, together with other data, to assist the taxonomy of the group. Thus, other studies involving the palynology of Columneinae may clarify the taxonomic relationships of the subtribe. ...
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... In recent decades, recorded diversity in the subfamily has grown considerably as new taxa are being described continuously [e.g., 58,[64][65][66] especially in Vietnam [55][56][57][58][66][67][68][69][70][71]. However, molecular phylogenetic studies in the past decade also showed that traditional taxonomy of Gesneriaceae had utilized traits that are homoplasious [72][73][74][75][76]. Consequently, infrafamilial classification and generic delimitation of Gesneriaceae have undergone drastic changes in the past two decades [63,77]. ...
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... The Gesneriaceae family has colonized a great diversity of habitats and developed specialized plant-animal interactions that have led to a great diversity of floral morphologies as well ( Figure 1). These traits were first used for phylogenetic species classification, but because they highly converged in different Gesneriaceae lineages, the early taxonomy of the family was complex and contradictory [6][7][8]. ...
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... Most study species have fleshy fruits ( Table 1) that are presumably consumed by understory birds, as reported for M. rubescens (Kessler-Rios & Kattan, 2012) and closely related species (Loiselle & Blake, 1999), as well as for Psychotria, which is related to Palicourea and similar in habit and habitats (Loiselle et al., 1995;Loiselle & Blake, 1993;Theim et al., 2014). Drymonia have fleshy capsules, often termed "display-capsules" in understory Gesneriaceae, because of their presumed role in animal attraction (Clark et al., 2012). The limited reports suggest these are also consumed by understory birds, as well as frugivorous bats and monkeys (Wiehler, 1983). ...
... The capsular fruits of M. tomentosa probably have wind-or gravity-dispersed seeds, as in many understory Melastomataceae with the same type of fruit (Renner, 1989), potentially making this species the most limited in seed dispersal. All six species have mechanisms to reduce selfing, either via marked herkogamy (i.e., spatial separation of stigma and anthers via style elongation) in the Melastomataceae (Renner, 1989), protandry (i.e., temporal separation of male and female phases, with anthers releasing pollen and dying-off before stigma is receptive) in the Gesneriaceae (Clark et al., 2012;Wiehler, 1983), or distyly (i.e., polymorphism in style length within a population in which flowers in one individual are monomorphic) in the Rubiaceae (Bawa & Beach, 1983). Self-incompatibility (via intramorph incompatibility) is common in the Rubiaceae (Bawa & Beach, 1983), but less clear in the Gesneriaceae and Melastomataceae, where both selfincompatibility and self-compatibility have been documented (Ramírez-Aguirre et al., 2016;Renner, 1989). ...
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... To date, phylogenetic inference in this plant group has mainly relied on plastid markers (e.g., atpB-rbcL, psbA-trnH, trnL-trnF, ndhF) and few multi-copy nuclear ribosomal regions such as ITS, and to a lower extent, low-copy nuclear genes such as GLUTAMINE SYNTHETASE (ncpGS) and CYCLOEDIA (CYC) (reviewed in Möller and Clark, 2013;Roalson and Roberts, 2016). Phylogenetic hypotheses derived from these genetic markers provided the framework to redefine the generic and tribal boundaries and to develop a new formal classification of the family (Zimmer et al., 2002;Perret et al., 2003;Roalson et al., 2005;Clark et al., 2006;Möller et al., 2009Möller et al., , 2011Clark et al., 2012;Weber et al., 2013). The analyses of these sequence data using supermatrix approaches also provided large scale phylogenetic hypotheses for the entire family (768 species; Roalson and Roberts, 2016) and the Gesnerioideae subfamily (583 species; Serrano-Serrano et al., 2017). ...
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Gesneriaceae (ca. 3400 species) is a pantropical plant family with a wide range of growth form and floral morphology that are associated with repeated adaptations to different environments and pollinators. Although Gesneriaceae systematics has been largely improved by the use of Sanger sequencing data, our understanding of the evolutionary history of the group is still far from complete due to the limited number of informative characters provided by this type of data. To overcome this limitation, we developed here a Gesneriaceae-specific gene capture kit targeting 830 single-copy loci (776,754 bp in total), including 279 genes from the Universal Angiosperms-353 kit. With an average of 557,600 reads and 87.8% gene recovery, our target capture was successful across the family Gesneriaceae and also in other families of Lamiales. From our bait set, we selected the most informative 418 loci to resolve phylogenetic relationships across the entire Gesneriaceae family using maximum likelihood and coalescent-based methods. Upon testing the phylogenetic performance of our baits on 78 taxa representing 20 out of 24 subtribes within the family, we showed that our data provided high support for the phylogenetic relationships among the major lineages, and were able to provide high resolution within more recent radiations. Overall, the molecular resources we developed here open new perspectives for the study of Gesneriaceae phylogeny at different taxonomical levels and the identification of the factors underlying the diversification of this plant group.