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• Although the evolution of the selfing syndrome often involves reductions in floral size, pollen and nectar, few studies of selfing syndrome divergence have examined nectar. We investigate whether nectar traits have evolved independently of other floral size traits in the selfing syndrome, whether nectar traits diverged due to drift or selection, and the extent to which quantitative trait locus (QTL) analyses predict genetic correlations. • We use F5 recombinant inbred lines (RILs) generated from a cross between Ipomoea cordatotriloba and Ipomoea lacunosa. We calculate genetic correlations to identify evolutionary modules, test whether trait divergence was due to selection, identify QTLs and perform correlation analyses to evaluate how well QTL properties reflect genetic correlations. • Nectar and floral size traits form separate evolutionary modules. Selection has acted to reduce nectar traits in the selfing I. lacunosa. Genetic correlations predicted from QTL properties are consistent with observed genetic correlations. • Changes in floral traits associated with the selfing syndrome reflect independent evolution of at least two evolutionary modules: nectar and floral size traits. We also demonstrate directional selection on nectar traits, which is likely to be independent of selection on floral size traits. Our study also supports the expected mechanistic link between QTL properties and genetic correlations.

Background Floral morphs are characterized differentiations in reciprocal positions of sexual organs and ancillary floral traits in heterostylous plants. However, it remains unclear how differential floral morphs ensure reproductive success between morphs using the same pollinator. Results Measurements of floral traits in white-flowered Tirpitzia sinensis with long corolla tubes indicated that it is typically distylous, long-styled (L-) morph producing more but smaller pollen grains per flower than short-styled (S-) morph. Both morphs secreted more nectar volume at night than in the day and the sugar composition was rich in sucrose, potentially adaptive to pollination by hawkmoths ( Macroglossum spp.) which were active at dusk. A bumblebee species functioned as the nectar robber in both morphs and a honeybee as the pollen feeder in the S-morph. The L-morph secreted more nectar volume but relatively lower sucrose/hexose ratio than the S-morph. Floral visitation rate by hawkmoths was higher but its pollination efficiency was lower in the S-morph than the L-morph. Hand pollination treatments indicated self-incompatibility in T. sinensis and seed set of open-pollinated flowers did not differ between morphs. Conclusions Our findings suggest that the two morphs differ with respect to traits relevant to pollination. The L-morph, with its exserted stigma, has more pollen grains per anther and a greater volume of nectar, which may prolong the foraging time and increase the pollination efficiency of hawkmoths. The S-morph has a higher sucrose/hexose ratio in its nectar which can be more attractive to hawkmoths and increase the visit rates. Ancillary polymorphic floral traits between two morphs are adaptive to hawkmoth and ensure reproductive success in distylous plant T. sinensis .

Although floral and extrafloral nectar traits are important for plant reproduction and defense, we know little about their genetic basis. Only a handful of studies have quantified heritable variation for nectar traits, primarily in controlled environments that minimize environmental variation. Most such studies have reported strong genetic influences, with heritabilities often >0.35. However, because nectar traits are often very responsive to environmental variation, even substantial amounts of genetic variation may be swamped out in the field. Environmental variation deserves to be studied in its own right, including exploration of genotype x environment interaction for nectar traits. Most genetic studies of nectar have focused on production rate and concentration, whereas we know almost nothing about the heritability of other important traits such as production patterns, sugar ratios, amino acid composition, taste, and scent. Likewise, almost nothing is known about the heritability of extrafloral nectar traits. Important progress on all of these fronts can be made using simple experimental designs to quantify environmental effects, genotype x environment interactions, clonal repeatability, and correlated traits. There is great promise in molecular approaches, but their use will not obviate the need for more quantitative genetic studies in the field and greenhouse.

Abstract Background Floral nectar is an important determinant of plant–pollinator interactions and an integral component of pollination syndromes, suggesting it is under pollinator-mediated selection. However, compared to floral display traits, we know little about the evolutionary ecology of nectar. Combining a literature review with a meta-analysis approach, we summarize the evidence for heritable variation in nectar traits and link this variation to pollinator response and plant fitness. We further review associations between nectar traits and floral signals and discuss them in the context of honest signalling and targets of selection. Scope Although nectar is strongly influenced by environmental factors, heritable variation in nectar production rate has been documented in several populations (mean h2 = 0.31). Almost nothing is known about heritability of other nectar traits, such as sugar and amino acid concentrations. Only a handful of studies have quantified selection on nectar traits, and few find statistically significant selection. Pollinator responses to nectar traits indicate they may drive selection, but studies tying pollinator preferences to plant fitness are lacking. So far, only one study conclusively identified pollinators as selective agents on a nectar trait, and the role of microbes, herbivores, nectar robbers and abiotic factors in nectar evolution is largely hypothetical. Finally, there is a trend for positive correlations among floral cues and nectar traits, indicating honest signalling of rewards. Conclusions Important progress can be made by studies that quantify current selection on nectar in natural populations, as well as experimental approaches that identify the target traits and selective agents involved. Signal–reward associations suggest that correlational selection may shape evolution of nectar traits, and studies exploring these more complex forms of natural selection are needed. Many questions about nectar evolution remain unanswered, making this a field ripe for future research.

Flowering plant species and their nectar-feeding vertebrates exemplify some of the most remarkable biotic interactions in the Neotropics. In the Brazilian Atlantic Forest, several species of birds (especially hummingbirds), bats, and non-flying mammals, as well as one lizard feed on nectar, often act as pollinators and contribute to seed output of flowering plants. We present a dataset containing information on flowering plants visited by nectar-feeding vertebrates and sampled at 166 localities in the Brazilian Atlantic Forest. This dataset provides information on 1902 unique interactions among 515 species of flowering plants and 129 species of potential vertebrate pollinators and the patterns of species diversity across latitudes. All plant–vertebrate interactions compiled were recorded through direct observations of visits, and no inferences of pollinators based on floral syndromes were included. We also provide information on the most common plant traits used to understand the interactions between flowers and nectar-feeding vertebrates: plant growth form, corolla length, rate of nectar production per hour in bagged flowers, nectar concentration, flower color and shape, time of anthesis, presence or absence of perceptible fragrance by human, and flowering phenology as well as the plant’s threat status by International Union for Conservation of Nature (IUCN) classification. For the vertebrates, status of threat by IUCN classification, body mass, bill or rostrum size areprovided. Information on the frequency of visits and pollen deposition on the vertebrate’s body is provided from the original source when available. The highest number of unique interactions is recorded for birds (1771) followed by bats (110). For plants, Bromeliaceae contains the highest number of unique interactions (606), followed by Fabaceae (242) and Gesneriaceae (104). It is evident that there was geographical bias of the studies throughout the southeast of the Brazilian Atlantic Forest and that most effort was directed to flower–hummingbird interactions. However, it reflects a worldwide tendency of more plants interacting with birds compared with other vertebrate species. The lack of similar protocols among studies to collect basic data limits the comparisons among areas and generalizations. Nevertheless, this dataset represents a notable effort to organize and highlight the importance of vertebrate pollinators in this hotspot of biodiversity on Earth and represents the data currently available. No copyright or proprietary restrictions are associated with the use of this data set. Please cite this data paper when the data are used in publications or scientific events.

Accurate variant calling is critical for identifying the genetic basis of complex traits, yet filters used in variant detection may inadvertently exclude valuable genetic information. In this study, we compare common sequencing depth filters, used to eliminate error‐prone variants associated with repetitive regions and technical issues, with a biologically relevant filtering approach that targets expected Mendelian segregation. The resulting variant sets were evaluated in the context of nectar volume quantitative trait loci (QTL) mapping in sunflower (Helianthus annuus L.). Our previous research failed to detect an interval containing a strong candidate gene for nectar production (HaCWINV2). We removed hard filters and implemented a chi‐square goodness‐of‐fit test to retain variants that segregate according to expected genetic ratios. We demonstrate that biologically relevant filtering retains more significant QTL and candidate genes, including HaCWINV2, while removing variants due to technical errors more effectively, and accounted for 48.55% of nectar production phenotypic variation. In finding nine putative homologs of Arabidopsis genes with nectary function within QTL regions, we demonstrate that this filtering strategy has a higher power of true variant detection in QTL mapping than the commonly used variant depth filtering strategy. Future research will adapt the technique to multiple population contexts, such as genomic selection. Core Ideas Discovering biologically meaningful variants from sequence data requires a careful and critical view of bioinformatic workflows. The use of arbitrary filters can remove significant genomic variation that contributes to the phenotype of interest. Arbitrary filters can also fail to remove variant call errors. A chi‐square filtering strategy based on segregation ratio retained a larger number of valid variants. More candidate regions with putative nectar‐related genes and better statistical support were discovered. Plain Language Summary In genomic research, identifying genetic markers is key to understanding complex traits, but traditional methods for filtering genetic data can sometimes miss important information. In this study, we explored a new data filtering approach and mapped genes related to nectar production in sunflower. We applied a more flexible filtering method that considers how markers are expected to segregate in breeding populations. Our previous work failed to identify an important gene previously hypothesized to be involved in nectar production, likely due to overly strict filtering. Our improved approach identified nine sunflower genes related to nectar production genes in the model species Arabidopsis thaliana, compared to zero genes identified from the previous filtering strategy. This study highlights the value of using flexible, biologically relevant filtering methods, which can lead to better results in plant genomic studies.

Interactions with animal pollinators have helped shape the stunning diversity of flower morphologies across the angiosperms. A common evolutionary consequence of these interactions is that some flowers have converged on suites of traits, or pollination syndromes, that attract and reward specific pollinator groups. Determining the genetic basis of these floral pollination syndromes can help us understand the processes that contributed to the diversification of the angiosperms. Here, we characterize the genetic architecture of a bee-to-hummingbird pollination shift in Aquilegia (columbine) using QTL mapping of 17 floral traits encompassing color, nectar composition, and organ morphology. In this system, we find that the genetic architectures underlying differences in floral color are quite complex, and we identify several likely candidate genes involved in anthocyanin and carotenoid floral pigmentation. Most morphological and nectar traits also have complex genetic underpinnings; however, one of the key floral morphological phenotypes, nectar spur curvature, is shaped by a single locus of large effect.

Evolution of complex phenotypes depends on the adaptive importance of individual traits, and the developmental changes required to modify traits. Floral syndromes are complex adaptations to pollinators that include color, nectar, and shape variation. Hummingbird-adapted flowers have evolved a remarkable number of times from bee-adapted ancestors in Penstemon, and previous work demonstrates that color over shape better distinguishes bee from hummingbird syndromes. Here, we examined the relative importance of nectar volume and nectary development in defining Penstemon pollination syndromes. We tested the evolutionary association of nectar volume and nectary area with pollination syndrome across 19 Penstemon species. In selected species, we assessed cellular-level processes shaping nectary size. Within a segregating population from an intersyndrome cross, we assessed trait correlations between nectar volume, nectary area, and the size of stamens on which nectaries develop. Nectar volume and nectary area displayed an evolutionary association with pollination syndrome. These traits were correlated within a genetic cross, suggesting a mechanistic link. Nectary area evolution involves parallel processes of cell expansion and proliferation. Our results demonstrate that changes to nectary patterning are an important contributor to pollination syndrome diversity and provide further evidence that repeated origins of hummingbird adaptation involve parallel developmental processes in Penstemon.

In the formation of species, adaptation by natural selection generates distinct combinations of traits that function well together. The maintenance of adaptive trait combinations in the face of gene flow depends on the strength and nature of selection acting on the underlying genetic loci. Floral pollination syndromes exemplify the evolution of trait combinations adaptive for particular pollinators. The North American wildflower genus Penstemon displays remarkable floral syndrome convergence, with at least 20 separate lineages that have evolved from ancestral bee pollination syndrome (wide blue-purple flowers that present a landing platform for bees and small amounts of nectar) to hummingbird pollination syndrome (bright red narrowly tubular flowers offering copious nectar). Related taxa that differ in floral syndrome offer an attractive opportunity to examine the genomic basis of complex trait divergence. In this study, we characterized genomic divergence among 229 individuals from a Penstemon species complex that includes both bee and hummingbird floral syndromes. Field plants are easily classified into species based on phenotypic differences and hybrids displaying intermediate floral syndromes are rare. Despite unambiguous phenotypic differences, genome-wide differentiation between species is minimal. Hummingbird-adapted populations are more genetically similar to nearby bee-adapted populations than to geographically distant hummingbird-adapted populations, in terms of genome-wide d XY . However, a small number of genetic loci are strongly differentiated between species. These approximately 20 “species-diagnostic loci,” which appear to have nearly fixed differences between pollination syndromes, are sprinkled throughout the genome in high recombination regions. Several map closely to previously established floral trait quantitative trait loci (QTLs). The striking difference between the diagnostic loci and the genome as whole suggests strong selection to maintain distinct combinations of traits, but with sufficient gene flow to homogenize the genomic background. A surprisingly small number of alleles confer phenotypic differences that form the basis of species identity in this species complex.

Plant–pollinator interactions are mediated by floral signals and by the quantity and quality of floral rewards. Biotic and abiotic disturbances can influence plant reproductive success through both direct effects on plant performance and indirect effects on pollinator attraction. In this study, we examined the effects of drought on buckwheat (Fagopyrum esculentum Moensch), a globally cultivated plant that is prone to drought stress, dependent on insect pollinators for reproduction, and increasingly utilized in on‐farm conservation. Between drought‐stressed and control plants, we compared: nectar quantity and chemical composition, pollen quantity, floral volatile emissions, visits by both managed and wild pollinators, and plant reproductive success. Drought‐stressed plants produced significantly fewer flowers and less nectar per flower, though pollen quantity per flower was unaffected. Nectar from drought‐stressed plants had a lower proportion of sucrose relative to total sugars, though overall sugar concentration was unaffected. Significantly fewer bumble bees, honey bees, and flies were recorded on drought‐stressed plants. While there was no significant difference in the quantity of total floral volatile emissions, volatile compositions differed, with drought‐stressed plants having higher emissions of (Z)‐3‐hexenol, isobutyraldehyde, 2‐methylbutanal, and 3‐methylbutanal. Finally, drought stress had negative effects on seed set and total seed mass per plant. Our results show that drought stress can have significant effects on floral traits and pollinator attraction, reducing plant reproductive success, and the nectar resources available to pollinators. Thus, the potential value of this plant in pollinator conservation and as a honey plant may be reduced under drought stress.