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The significance of prezygotic isolation for speciation has been recognized at least since the Modern Synthesis. However, fundamental questions remain. For example, how are genetic associations between traits that contribute to prezygotic isolation maintained? What is the source of genetic variation underlying the evolution of these traits? And how do prezygotic barriers affect patterns of gene flow? We address these questions by reviewing genetic features shared across plants and animals that influence prezygotic isolation. Emerging technologies increasingly enable the identification and functional characterization of the genes involved, allowing us to test established theoretical expectations. Embedding these genes in their developmental context will allow further predictions about what constrains the evolution of prezygotic isolation. Ongoing improvements in statistical and computational tools will reveal how pre- and postzygotic isolation may differ in how they influence gene flow across the genome. Finally, we highlight opportunities for progress by combining theory with appropriate data. 

Abstract Different populations of plant species can adapt to their local pollinators and diverge in floral traits accordingly. Floral traits are subject to pollinator‐driven natural selection to enhance plant reproductive success. Studies on temperate plant systems have shown pollinator‐driven selection results in floral trait variation along elevational gradients, but studies in tropical systems are lacking. We analyzed floral traits and pollinator assemblages in the Neotropical bee‐pollinated taxonCostus guanaiensisvar.tarmicusacross four sites along a steep elevational gradient in Peru. We found variations in floral traits of size, color, and reward, and in the pollinator assemblage along the elevational gradient. We examined our results considering two hypotheses, (1) local adaptation to different bee assemblages, and (2) the early stages of an evolutionary shift to a new pollinator functional group (hummingbirds). We found some evidence consistent with the adaptation ofC. guanaiensisvar.tarmicusto the local bee fauna along the studied elevational gradient. Corolla width across sites was associated with bee thorax width of the local most frequent pollinator. However, we could not rule out the possibility of the beginning of a bee‐to‐hummingbird pollination shift in the highest‐studied site. Our study is one of the few geographic‐scale analyses of floral trait and pollinator assemblage variation in tropical plant species. Our results broaden our understanding of plant‐pollinator interactions beyond temperate systems by showing substantial intraspecific divergence in both floral traits and pollinator assemblages across geographic space in a tropical plant species. 

Abstract— Costus flammulus is a new herbaceous species endemic to montane cloud forests of the volcanic cordilleras in northern Costa Rica. Costus flammulus has been mistaken for C. wilsonii , but phylogenetic evidence demonstrates that it is closely related to the widespread lowland species C. pulverulentus . Here, we use an integrated framework of species concepts to evaluate whether C. flammulus and C. pulverulentus are distinct species. First, we re-evaluate prior phylogenetic analyses to assess whether C. flammulus bifurcated from or budded off from within C. pulverulentus and whether C. flammulus is monophyletic. We then compare phenotypic traits to determine which diagnostic vegetative and inflorescence traits can be used to identify species in herbarium specimens and examine whether floral traits may confer floral isolation. We compare pollinator assemblages to examine whether pollinator specificity may contribute to reproductive isolation. Finally, we model species distributions and climatic niche overlap to assess ecogeographic isolation. We found that C. flammulus is a monophyletic species phenotypically, ecologically, and geographically distinct from C. pulverulentus and may have speciated as a peripheral isolate at the high elevation range edge of C. pulverulentus . Several lines of evidence, such as C. pulverulentus paraphyly, range size asymmetry, and C. flammulus’ nested distribution and vegetative traits, suggest that C. flammulus budded off from a C. pulverulentus ‐like progenitor species, evolving to tolerate a colder and more seasonal montane environment. 

The spiral gingers (Costus L.) are a pantropical genus of herbaceous perennial monocots; the Neotropical clade of Costus radiated rapidly in the past few million years into over 60 species. The Neotropical spiral gingers have a rich history of evolutionary and ecological research that can motivate and inform modern genetic investigations. Here, we present the first two chromosome-level genome assemblies in the genus, for C. pulverulentus and C. lasius, and briefly compare their synteny. We assembled the C. pulverulentus genome from a combination of short-read data, Chicago and Dovetail Hi-C chromatin-proximity sequencing, and alignment with a linkage map. We assembled the C. lasius genome with Pacific Biosciences HiFi long reads and alignment to the C. pulverulentus genome. These two assemblies are the first published genomes for non-cultivated tropical plants. These genomes solidify the spiral gingers as a model system and will facilitate research on the poorly understood genetic basis of tropical plant diversification. 

Abstract The spiral gingers (Costus L.) are a pantropical genus of herbaceous perennial monocots; the Neotropical clade of Costus radiated rapidly in the past few million years into over 60 species. The Neotropical spiral gingers have a rich history of evolutionary and ecological research that can motivate and inform modern genetic investigations. Here, we present the first 2 chromosome-level genome assemblies in the genus, for C. pulverulentus and C. lasius, and briefly compare their synteny. We assembled the C. pulverulentus genome from a combination of short-read data, Chicago and Dovetail Hi-C chromatin-proximity sequencing, and alignment with a linkage map. We annotated the genome by mapping a C. pulverulentus transcriptome and querying mapped transcripts against a protein database. We assembled the C. lasius genome with Pacific Biosciences HiFi long reads and alignment to the C. pulverulentus genome. These 2 assemblies are the first published genomes for non-cultivated tropical plants. These genomes solidify the spiral gingers as a model system and will facilitate research on the poorly understood genetic basis of tropical plant diversification. 

1. The evolution of hummingbird pollination is common across angiosperm lineages throughout the Americas, presenting an opportunity to examine convergence in both traits and environments to better understand how complex phenotypes arise. We examine multiple independent shifts from bee to hummingbird pollination in the Neotropical spiral gingers (Costus) and use our data to address several common explanations for the prevalence of bee to bird pollination transitions. 2. We use floral traits of species with observed pollinators to predict pollinators of unobserved species and reconstruct ancestral pollination states on a well-resolved phylogeny. We examine whether independent transitions evolve towards the same phenotypic optimum and whether shifts to hummingbird pollination are associated with high elevation or climatic niche. 3. Traits predicting hummingbird pollination include small flower size, brightly-colored floral bracts, and the absence of nectar guides. We find many shifts to hummingbird pollination and no reversals, a single shared phenotypic optimum across hummingbird flowers, and no association between pollination and elevation or climatic niche. 4. Costus presents surprising findings compared to other plant clades. Hummingbird flowers are consistently smaller than bee flowers and primary flower colors are not predictive of pollinators. Moreover, hummingbird pollination shows no association with high elevation, as found in other tropical plants. 

Costus flammulus is a new herbaceous species endemic to montane cloud forests of the volcanic cordilleras in northern Costa Rica. Costus flammulus has been mistaken for C. wilsonii, but phylogenetic evidence demonstrates that it is closely related to the widespread lowland species C. pulverulentus. Here, we used an integrated framework of species concepts to evaluate whether C. flammulus and C. pulverulentus are distinct species. First, we re-evaluate prior phylogenetic analyses to assess whether C. flammulus bifurcated from or budded off from within C. pulverulentus and whether C. flammulus is monophyletic. We then compare phenotypic traits to determine which diagnostic vegetative and inflorescence traits can be used to identify species in herbarium specimens and examine whether floral traits may confer floral isolation. We compare pollinator assemblages to examine whether pollinator specificity may contribute to reproductive isolation. Finally, we model species distributions and climatic niche overlap to assess ecogeographic isolation. We found that C. flammulus is a monophyletic species phenotypically, ecologically, and geographically distinct from C. pulverulentus and may have speciated as a peripheral isolate at the high elevation range edge of C. pulverulentus. Several lines of evidence, such as C. pulverulentus paraphyly, range size asymmetry, and C. flammulus’ nested distribution and vegetative traits, suggest that C. flammulus budded off from a C. pulverulentus‐like progenitor species, evolving to tolerate a colder and more seasonal montane environment. 

Abstract The study of plant functional traits and variation among and within species can help illuminate functional coordination and trade-offs in key processes that allow plants to grow, reproduce and survive. We studied 20 leaf, above-ground stem, below-ground stem and fine-root traits of 17 Costus species from forests in Costa Rica and Panama to answer the following questions: (i) Do congeneric species show above-ground and below-ground trait coordination and trade-offs consistent with theory of resource acquisition and conservation? (ii) Is there correlated evolution among traits? (iii) Given the diversity of habitats over which Costus occurs, what is the relative contribution of site and species to trait variation? We performed a principal components analysis (PCA) to assess for the existence of a spectrum of trait variation and found that the first two PCs accounted for 21.4 % and 17.8 % of the total trait variation, respectively, with the first axis of variation being consistent with a continuum of resource-acquisitive and resource-conservative traits in water acquisition and use, and the second axis of variation being related to the leaf economics spectrum. Stomatal conductance was negatively related to both above-ground stem and rhizome specific density, and these relationships became stronger after accounting for evolutionary relatedness, indicating correlated evolution. Despite elevation and climatic differences among sites, high trait variation was ascribed to individuals rather than to sites. We conclude that Costus species present trait coordination and trade-offs that allow species to be categorized as having a resource-acquisitive or resource-conservative functional strategy, consistent with a whole-plant functional strategy with evident coordination and trade-offs between above-ground and below-ground function. Our results also show that herbaceous species and species with rhizomes tend to agree with trade-offs found in more species-rich comparisons.