Phenotypic variation seems not to be associated with the genetic profile in Zygopetalum (Orchidaceae): a case study of a high-elevation rocky complex.

BACKGROUND: Hybridization associated with polyploidy studies is rare in the tropics. The genus Zygopetalum (Orchidaceae) was investigated here as a case study of Neotropical plants. In the rocky highlands of the Ibitipoca State Park (ISP), southeast Brazil, individuals with intermediate colors and forms between the species Z. maculatum and Z. triste were commonly identified. METHODS AND RESULTS: Chromosomal analysis and DNA quantity showed a uniform population. Regardless of the aspects related to the color and shape of floral structures, all individuals showed 2n = 96 chromosomes and an average of 14.05 pg of DNA. Irregularities in meiosis associated with chromosome number and C value suggest the occurrence of polyploidy. The genetic distance estimated using ISSR molecular markers revealed the existence of genetic variability not related to morphological clusters. Morphometric measurements of the flower pieces revealed that Z. maculatum shows higher variation than Z. triste although lacking a defined circumscription. CONCLUSION: The observed variation can be explained by the polyploid and phenotypic plasticity resulting from the interaction of the genotypes with the heterogeneous environments observed in this habitat.

Anatomical, histochemical, and developmental approaches reveal the long-term functioning of the floral nectary in Tocoyena formosa (Rubiaceae).

Tocoyena formosa has a persistent floral nectary that continues producing nectar throughout flower and fruit development. This plant also presents an intriguing non-anthetic nectary derived from early-developing floral buds with premature abscised corolla. In this study, we characterize the structure, morphological changes, and functioning of T. formosa floral nectary at different developmental stages. We subdivided the nectary into four categories based on the floral and fruit development stage at which nectar production started: (i) non-anthetic nectary; (ii) anthetic nectary, which follows the regular floral development; (iii) pericarpial nectary, derived from pollinated flowers following fruit development; and (iv) post-anthetic nectary that results from non-pollinated flowers after anthesis. The nectary has a uniseriate epidermis with stomata, nectariferous parenchyma, and vascular bundles, with a predominating phloem at the periphery. The non-anthetic nectary presents immature tissues that release the exudate. The nectary progressively becomes more rigid as the flower and fruit develop. The main nectary changes during flower and fruit development comprised the thickening of the cuticle and epidermal cell walls, formation of cuticular epithelium, and an increase in the abundance of calcium oxalate crystals and phenolic cells near the vascular bundles. Projections of the outer periclinal walls toward the cuticle in the post-anthetic nectary suggest nectar reabsorption. The anatomical changes of the nectary allow it to function for an extended period throughout floral and fruit development. Hence, T. formosa nectary is a bivalent secretory structure that plays a crucial role in the reproductive and defensive interactions of this plant species.

High floral disparity without pollinator shifts in buzz-bee-pollinated Melastomataceae.

Shifts among functional pollinator groups are commonly regarded as sources of floral morphological diversity (disparity) through the formation of distinct pollination syndromes. While pollination syndromes may be used for predicting pollinators, their predictive accuracy remains debated, and they are rarely used to test whether floral disparity is indeed associated with pollinator shifts. We apply classification models trained and validated on 44 functional floral traits across 252 species with empirical pollinator observations and then use the validated models to predict pollinators for 159 species lacking observations. In addition, we employ multivariate statistics and phylogenetic comparative analyses to test whether pollinator shifts are the main source of floral disparity in Melastomataceae. We find strong support for four well-differentiated pollination syndromes ('buzz-bee', 'nectar-foraging vertebrate', 'food-body-foraging vertebrate', 'generalist'). While pollinator shifts add significantly to floral disparity, we find that the most species-rich 'buzz-bee' pollination syndrome is most disparate, indicating that high floral disparity may evolve without pollinator shifts. Also, relatively species-poor clades and geographic areas contributed substantially to total disparity. Finally, our results show that machine-learning approaches are a powerful tool for evaluating the predictive accuracy of the pollination syndrome concept as well as for predicting pollinators where observations are missing.

Heterostyly.

Scharman and Lenhard introduce heterostyly, a phenomenon where individuals in a plant population produce flowers with more than one morphologically distinct form.

Relictithismia kimotsukiensis, a new genus and species of Thismiaceae from southern Japan with discussions on its phylogenetic relationship.

The family Thismiaceae, known as "fairy lanterns" for their urn- or bell-shaped flowers with basally fused tepals, consists of non-photosynthetic flowering monocots mainly in tropical regions, extending into subtropical and temperate areas. Here, we propose a new mycoheterotrophic genus, Relictithismia Suetsugu & Tagane (Thismiaceae), with its monotypic species Relictithismia kimotsukiensis Suetsugu, Yas.Nakam. & Tagane from Kimotsuki Mountains in the Osumi Peninsula, Kagoshima Prefecture, Kyushu Island, southern Japan. Relictithismia resembles Haplothismia Airy Shaw in having a cluster of tuberous roots, a feature previously observed only in this genus within the family Thismiaceae. However, it differs in having solitary flowers (vs. 2-6-flowered pseudo-raceme in Haplothismia), anther thecae largely separated (vs. connate), and the presence of an annulus (vs. absent). Additionally, Relictithismia differs from the geographically overlapping genus Thismia Griff. in its stamen structure and the position of the annulus. In Relictithismia, the stamens lack connectives, and its free filaments arise from the annulus located inside the perianth mouth, while in Thismia, the stamens typically have connate connectives, forming a staminal tube pendulous from the annulus located at the mouth of the floral tube. Our morphological and phylogenetic data indicated that R. kimotsukiensis holds an early-diverging position within the family, situated outside the Old World Thismia clade. This paper offers an extensive description and color photographs of R. kimotsukiensis, complemented by notes on its phylogenetic relationship and evolutionary history.

Two lineages of Lemna aequinoctialis (Araceae, Lemnoideae) based on physiology, morphology, and phylogeny.

Lemna aequinoctialis Welw. is a widely spread species that has diverse physiological and molecular properties. Flower characteristics are important factors in deducing taxonomical status; however, owing to the rarity of flowering observations in Lemna, studying them has been a prolonged challenge. In this study, physiological and morphological analyses were conducted by inducing flowering, and molecular analysis was done based on the two chloroplast DNA loci (matK, atpF-atpH intergeneric spacer) of L. aequinoctialis sensu Landolt (1986) from 70 strains found in 70 localities in Japan, Korea, Thailand, and the US. In total, 752 flowering fronds from 13 strains were observed based on axenic conditions. Two different trends in flower organ development-protogyny and adichogamy-were detected in these strains. Their physiological traits were divided into two groups, showing different morphological features based on frond thickness, root cap, and anther sizes. Molecular analysis showed two lineages corresponding to two physiological groups. These were identified as L. aequinoctialis sensu Beppu et al. (1985) and L. aoukikusa Beppu et Murata based on the description of the nomenclature of L. aoukikusa. These were concluded as independent taxa and can be treated as different species. Furthermore, the distribution of L. aoukikusa is not only limited to Japan.

Parallel evolution of morphological and genomic selfing syndromes accompany the breakdown of heterostyly.

Evolutionary transitions from outcrossing to selfing in flowering plants have convergent morphological and genomic signatures and can involve parallel evolution within related lineages. Adaptive evolution of morphological traits is often assumed to evolve faster than nonadaptive features of the genomic selfing syndrome. We investigated phenotypic and genomic changes associated with transitions from distyly to homostyly in the Primula oreodoxa complex. We determined whether the transition to selfing occurred more than once and investigated stages in the evolution of morphological and genomic selfing syndromes using 22 floral traits and both nuclear and plastid genomic data from 25 populations. Two independent transitions were detected representing an earlier and a more recently derived selfing lineage. The older lineage exhibited classic features of the morphological and genomic selfing syndrome. Although features of both selfing syndromes were less developed in the younger selfing lineage, they exhibited parallel development with the older selfing lineage. This finding contrasts with the prediction that some genomic changes should lag behind adaptive changes to morphological traits. Our findings highlight the value of comparative studies on the timing and extent of transitions from outcrossing to selfing between related lineages for investigating the tempo of morphological and molecular evolution.

The mechanism underlying asymmetric bending of lateral petals in Delphinium (Ranunculaceae).

During the process of flower opening, most petals move downward in the direction of the pedicel (i.e., epinastic movement). In most Delphinium flowers, however, their two lateral petals display a very peculiar movement, the mirrored helical rotation, which requires the twist of the petal stalk. However, in some lineages, their lateral petals also exhibit asymmetric bending that increases the degree of mirrored helical rotation, facilitating the formation of a 3D final shape. Notably, petal asymmetric bending is a novel trait that has not been noticed yet, so its morphological nature, developmental process, and molecular mechanisms remain largely unknown. Here, by using D. anthriscifolium as a model, we determined that petal asymmetric bending was caused by the localized expansion of cell width, accompanied by the specialized array of cell wall nano-structure, on the adaxial epidermis. Digital gene analyses, gene expression, and functional studies revealed that a class I homeodomain-leucine zipper family transcription factor gene, DeanLATE MERISTEM IDENTITY1 (DeanLMI1), contributes to petal asymmetric bending; knockdown of it led to the formation of explanate 2D petals. Specifically, DeanLMI1 promotes cell expansion in width and influences the arrangement of cell wall nano-structure on the localized adaxial epidermis. These results not only provide a comprehensive portrait of petal asymmetric bending for the first time but also shed some new insights into the mechanisms of flower opening and helical movement in plants.

Nectar-mediated avian pollination in Cattleya (Orchidaceae: Laeliinae).

Orchids are notable for the variety of rewards produced for their pollinators. Nevertheless, many species are rewardless and pollinated by food-deceptive strategies. This is the case for Cattleya, a Neotropical genus of more than 100 species supposedly pollinated by food-deception. Here, I studied a member of the clade Sophronitis (Cattleya cernua) in forest areas of southeastern Brazil. The study included analyses of floral morphology, light microscopy of secretory tissues, histochemistry of flower resources, and measurements of nectar volume and concentration. Data on pollinators and mechanisms of pollination were collected in the field by means of film records, while breeding systems were studied using experimental treatments applied to cultivated specimens. The flower traits of Cattleya cernua show strong adaptation to avian pollination. Flowers produce 1-7 µl of a moderately concentrated nectar (17-50%). A secretory tissue at the distal portion of the cuniculus is responsible for nectar secretion. Secreted nectar is stored in the nectar chamber. Pollinaria deposition on the apex of bird beaks allows both short- and long-billed birds to pollinate the flowers. Plants from the studied population were fully self-compatible but pollinator-dependent. Pollinator frequency was higher than in food-deceptive Cattleya. As far as is known, this is the first record of nectar production in Cattleya. The discovery of a reward-producing species (C. cernua) within a genus entirely pollinated by food-deception strategies (Cattleya) provides clues and new insights into understanding the diversification of Laeliinae across the Neotropics.

Angiosperm flowers reached their highest morphological diversity early in their evolutionary history.

Flowers are the complex and highly diverse reproductive structures of angiosperms. Because of their role in sexual reproduction, the evolution of flowers is tightly linked to angiosperm speciation and diversification. Accordingly, the quantification of floral morphological diversity (disparity) among angiosperm subgroups and through time may give important insights into the evolutionary history of angiosperms as a whole. Based on a comprehensive dataset focusing on 30 characters describing floral structure across angiosperms, we used 1201 extant and 121 fossil flowers to measure floral disparity and explore patterns of floral evolution through time and across lineages. We found that angiosperms reached their highest floral disparity in the Early Cretaceous. However, decreasing disparity toward the present likely has not precluded the innovation of other complex traits at other morphological levels, which likely played a key role in the outstanding angiosperm species richness. Angiosperms occupy specific regions of the theoretical morphospace, indicating that only a portion of the possible floral trait combinations is observed in nature. The ANA grade, the magnoliids, and the early-eudicot grade occupy large areas of the morphospace (higher disparity), whereas nested groups occupy narrower regions (lower disparity).

Reproductive isolation between two sympatric bat-pollinated Bauhinia (Leguminosae).

Several barriers contribute to reproductive isolation between plant species, which can be classified as pre- or post-pollination. Understanding the strength of these barriers could clarify the factors that maintain reproductive isolation and thus species integrity. In this study, we quantified reproductive isolation between two bat-pollinated co-occurring Bauhinia species (B. acuruana and B. pentandra) with similar flower morphology. Over the course of 18 months, we assessed reproductive isolation between these two Bauhinia species by quantifying the individual strengths and absolute contributions of five pre- and post- pollination barriers. Our data showed that both species are completely isolated in their reproduction by a combination of several barriers. Although they co-occur in a few populations, we found a high degree of geographic isolation between them. And although their flowering periods overlap, there is a significant difference in flowering peaks. Both species have the same pollinating bats, but the interspecific transfer of pollen between the plant species may be reduced due to the different length of the flower stamens, resulting in different pollen deposition on the bats' bodies. We have documented complete incompatibility between taxa and conclude that pre- and post-pollination barriers are important factors in preventing gene flow, even in contact zones between these two species of Bauhinia. We highlight that our work is the first study to use methods to estimate the strength of reproductive isolation barriers between bat-pollinated species.

The fertility tracks of pollen tube in the ovary of Solanum nigrum by three-dimensional reconstruction.

In this paper, we present an enhanced method for automatically capturing a large number of consecutive paraffin sections using a microscope. Leveraging these microstructural images, we employed three-dimensional visualisation and reconstruction techniques to investigate the dispersal growth process of pollen tube bundles upon entering the ovary of Solanum nigrum. Additionally, we explored their behaviour within different ovules and examined the relationship between the germination rate of seeds and the fertilisation process. Our findings reveal that despite the abundance of Solanum nigrum seeds, only a fraction of them is capable of successful germination. The germination rate of seeds is closely related to whether fertilisation of the ovules and pollen tubes is completed. Due to the limited number of pollen tubes entering the ovary, only a portion of the ovules can be fertilised. The proportion of fertilised ovules positively correlates with the germination rate of the seeds. Through three-dimensional reconstruction, we observed a phenomenon of proximity during the pollination process, wherein ovules closer to the pollen tube bundles are more likely to be fertilised. Furthermore, fertilised ovules exhibited significant changes in morphology and embryo sac structure. The number of fertilised ovules directly impacts the germination rate of wild Solanum nigrum seeds. Although all Solanum nigrum ovules have the potential to develop into seeds, most seeds originating from unfertilised ovules are unable to germinate normally, resulting in an incomplete germination rate of seeds and preventing it from reaching 100%.

Repeated Evolution of Unorthodox Feeding Styles Drives a Negative Correlation between Foot Size and Bill Length in Hummingbirds.

AbstractDifferences among hummingbird species in bill length and shape have rightly been viewed as adaptive in relation to the morphology of the flowers they visit for nectar. In this study we examine functional variation in a behaviorally related but neglected feature: hummingbird feet. We gathered records of hummingbirds clinging by their feet to feed legitimately as pollinators or illegitimately as nectar robbers-"unorthodox" feeding behaviors. We measured key features of bills and feet for 220 species of hummingbirds and compared the 66 known "clinger" species (covering virtually the entire scope of hummingbird body size) with the 144 presumed "non-clinger" species. Once the effects of phylogenetic signal, body size, and elevation above sea level are accounted for statistically, hummingbirds display a surprising but functionally interpretable negative correlation. Clingers with short bills and long hallux (hind-toe) claws have evolved-independently-more than 20 times and in every major clade. Their biomechanically enhanced feet allow them to save energy by clinging to feed legitimately on short-corolla flowers and by stealing nectar from long-corolla flowers. In contrast, long-billed species have shorter hallux claws, as plant species with long-corolla flowers enforce hovering to feed, simply by the way they present their flowers.

Not just females and males: Unravelling the complex sex determinism of the hemp palm, Trachycarpus fortunei.

PREMISE: The ornamental Asian palm Trachycarpus fortunei (Arecaceae: Coryphoideae) is widely planted in temperate regions. In Europe, it has spread outside of gardens, particularly on the southern side of the Alps. Sexual expression in the species is complex, varying from dioecy to polygamy. This study investigated (1) sexual floral development and (2) genetic markers implicated in sex determinism. METHODS: The morphology and anatomy of floral organs at different developmental stages were studied using SEM observations and anatomical section. Sex determinism was explored using a genome-wide association study approach, searching for correlations between 31,000 single-nucleotide polymorphisms and sex affiliation of 122 palms from 21 wild populations. RESULTS: We observed that sexual differentiation appears late in floral development of T. fortunei. Morpho-anatomical characters of flowers conducive to panmixia were observed, such as well-differentiated septal nectaries that are thought to promote cross-pollination. At the molecular level, homozygous and heterozygous allelic systems with closely linked regions were found for sex determinism in individuals with female and "dominant-male" phenotypes, respectively. Through our wide sampling in the southern Alps, the closely linked genetic regions in males suggest that at least fifteen percent of wild palms are the direct offspring of "males" that can also produce fertile pistillate flowers. CONCLUSIONS: Trachycarpus fortunei is a further example of unstable sexual expression found in the family Arecaceae and represents an evolutionary path towards an XY genetic system. Our structural and genetic results may explain the high species dispersal ability in the southern Alps.

The evolution of flower-pollinator trait matching, and why do some alpine gingers appear to be mismatched?

BACKGROUND AND AIMS: Morphological matching between flower and pollinator traits has been documented in diverse plant lineages. Indeed, the matching of corolla tube length and pollinator tongue length has been cited repeatedly as a classic case of coevolution. However, there are many possible evolutionary routes to trait matching. Our aim here is both to review the evolutionary mechanisms of plant-pollinator trait matching and to investigate a specific case of trait matching/mismatching in a genus of alpine gingers. METHODS: Roscoea gingers with long corolla tubes in the western Himalayas have pollinators with correspondingly long tongues, but the match between corolla tube and pollinator tongue lengths is not seen in the eastern Himalayas. Six floral traits were measured, including corolla tube depth, an internal trait controlling pollinator access to nectar. We calculated coefficients of variation and phylogenetically controlled correlation patterns of these traits in six Roscoea species in order to gain possible insights into stabilizing selection and modularization of these traits. KEY RESULTS: The distal (nectar-containing) portion of the corolla tube exhibited lower coefficients of variations than did the basal portion. This is consistent with the hypothesis that pollinators mediate stabilizing selection on the distal, but not basal, portion of the corolla tube. This result, combined with phylogenetic data, suggests that the elevated liquid level of nectar in the distal tube evolved subsequent to dispersal into the eastern Himalayan region and loss of long-tongue pollinators. After accounting for phylogeny, corolla tube length, anther length, style length and labellum width were all intercorrelated. Corolla-tube depth was not part of this covariational module, however, suggesting separate adaptation to short-tongued pollinators. CONCLUSIONS: The reduction in functional corolla tube depth in the Roscoea appears to be related to the loss of long-tongued pollinators associated with dispersal to the eastern Himalayas and pollination by short-tongued pollinators. The apparent mismatch between floral tubes and pollinator tongues is a case of cryptic trait matching between flowers and pollinators, underscoring the importance of combining floral anatomy with pollination ecology in assessing plant-pollinator trait matching.

Floral Diversity and Pollination Syndromes in Agave subgenus Manfreda.

The genus Agave is an ecological keystone of American deserts and both culturally and economically important in Mexico. Agave is a large genus of about 250 species. The radiation of Agave is marked by an initial adaptation to desert environments and then a secondary diversification of species associated with pollinator groups, such as hummingbirds and nocturnal moths. Phylogenetic analyses place Agave subgenus Manfreda, or the "herbaceous agaves," in a monophyletic clade that likely evolved in part as an adaptation to novel pollination vectors. Here, we present a morphological and observational study assessing the evolution of floral form in response to pollinator specialization within this understudied group. We found significant visitation by hummingbirds and nocturnal moths to several species within the Agave subgenus Manfreda. These observations also align with our morphological analyses of floral organs and support the evolution of distinct pollination syndromes. We found that not all floral morphology is consistent within a pollination syndrome, suggesting hidden diversity in the evolution of floral phenotypes in Agave. We also characterize the morphological variation between herbarium and live specimens, demonstrating that special consideration needs to be made when combining these types of data. This work identifies the potential for studying the functional evolution of diverse floral forms within Agave and demonstrates the need to further explore ecological and evolutionary relationships to understand pollinator influence on diversification in the genus.

Adaptation to pollination by fungus gnats underlies the evolution of pollination syndrome in the genus Euonymus.

BACKGROUND AND AIMS: Dipteran insects are known pollinators of many angiosperms, but knowledge on how flies affect floral evolution is relatively scarce. Some plants pollinated by fungus gnats share a unique set of floral characters (dark red display, flat shape and short stamens), which differs from any known pollination syndromes. We tested whether this set of floral characters is a pollination syndrome associated with pollination by fungus gnats, using the genus Euonymus as a model. METHODS: The pollinator and floral colour, morphology and scent profile were investigated for ten Euonymus species and Tripterygium regelii as an outgroup. The flower colour was evaluated using bee and fly colour vision models. The evolutionary association between fungus gnat pollination and each plant character was tested using a phylogenetically independent contrast. The ancestral state reconstruction was performed on flower colour, which is associated with fungus gnat pollination, to infer the evolution of pollination in the genus Euonymus. KEY RESULTS: The red-flowered Euonymus species were pollinated predominantly by fungus gnats, whereas the white-flowered species were pollinated by bees, beetles and brachyceran flies. The colour vision analysis suggested that red and white flowers are perceived as different colours by both bees and flies. The floral scents of the fungus gnat-pollinated species were characterized by acetoin, which made up >90 % of the total scent in three species. Phylogenetically independent contrast showed that the evolution of fungus gnat pollination is associated with acquisition of red flowers, short stamens and acetoin emission. CONCLUSIONS: Our results suggest that the observed combination of floral characters is a pollination syndrome associated with the parallel evolution of pollination by fungus gnats. Although the role of the red floral display and acetoin in pollinator attraction remains to be elucidated, our finding underscores the importance of fungus gnats as potential contributors to floral diversification.

The evolution of Ericaceae flowers and their pollination syndromes at a global scale.

PREMISE: Floral evolution in large clades is difficult to study not only because of the number of species involved, but also because they often are geographically widespread and include a diversity of outcrossing pollination systems. The cosmopolitan blueberry family (Ericaceae) is one such example, most notably pollinated by bees and multiple clades of nectarivorous birds. METHODS: We combined data on floral traits, pollination ecology, and geography with a comprehensive phylogeny to examine the structuring of floral diversity across pollination systems and continents. We focused on ornithophilous systems to test the hypothesis that some Old World Ericaceae were pollinated by now-extinct hummingbirds. RESULTS: Despite some support for floral differentiation at a continental scale, we found a large amount of variability within and among landmasses, due to both phylogenetic conservatism and parallel evolution. We found support for floral differentiation in anther and corolla traits across pollination systems, including among different ornithophilous systems. Corolla traits show inconclusive evidence that some Old World Ericaceae were pollinated by hummingbirds, while anther traits show stronger evidence. Some major shifts in floral traits are associated with changes in pollination system, but shifts within bee systems are likely also important. CONCLUSIONS: Studying the floral evolution of large, morphologically diverse, and widespread clades is feasible. We demonstrate that continent-specific radiations have led to widespread parallel evolution of floral morphology. We show that traits outside of the perianth may hold important clues to the ecological history of lineages.

Nitrogen addition affects floral and vegetative traits, reproduction, and pollinator performance in Capsicum annuum L.

BACKGROUND AND AIMS: It has been demonstrated that nitrogen (N) addition alters flower morphology, floral rewards and pollinator performance. However, little is known about the effects of N addition on plant reproduction, including fruit set and seed set during selfing and outcrossing, floral and vegetative traits, and pollinator performance. We hypothesized that N addition would influence fruit set, seed set in selfed and outcrossed flowers, the relationship between vegetative and flower traits, and pollinator performance. METHODS: A 2-year pot experiment was conducted in which Capsicum annuum was exposed to three levels of relatively short-term N supply, i.e. 0 g m-2 (no N addition, as a control), 4 g m-2 (4N) and 16 g m-2 (16N), which are equivalent to about 0-, 1- and 4-fold of the peak local N deposition. We measured flower rewards, flower morphology, flowering phenology, as well as pollinator visitation rate, fruit set and seed set by self- and outcross-fertilization of C. annuum. RESULTS: The four levels of N addition increased plant biomass, biomass allocation to flowers, flower size, stigma-anther separation, nectar production and pollen production, resulting in an increase in pollinator visitation and fruit set. Nevertheless, the control and 16 levels of N addition reduced plant biomass, biomass allocation to flowers, flower size and stigma-anther separation, and nectar and pollen production, and consequently decreased pollinator visitation and fruit set. Exclusion of pollinators and hand-pollination experiments revealed that low levels of N addition were associated with high seed set in outcrossed flowers; however, this trend was reversed in flowers grown in the control and 16N treatments. CONCLUSION: Our results suggest that an optimal level of 4N can enhance the correlation between flower traits, pollinator performance and plant reproduction. Our findings cast new light on the underlying mechanisms of plant-pollinator interactions and plant adaptation to nitrogen deposition.

Profile of a flower: How rates of morphological evolution drive floral diversification in Ericales and angiosperms.

PREMISE: Recent studies of floral disparity in the asterid order Ericales have shown that flowers vary strongly among families and that disparity is unequally distributed between the three flower modules (perianth, androecium, gynoecium). However, it remains unknown whether these patterns are driven by heterogeneous rates of morphological evolution or other factors. METHODS: Here, we compiled a data set of 33 floral characters scored for 414 species of Ericales sampled from 346 genera and all 22 families. We conducted ancestral state reconstructions using an equal-rates Markov model for each character. We estimated rates of morphological evolution for Ericales and for a separate angiosperm-wide data set of 19 characters and 792 species, creating "rate profiles" for Ericales, angiosperms, and major angiosperm subclades. We compared morphological rates among flower modules within each data set separately and between data sets, and we compared rates among angiosperm subclades using the angiosperm data set. RESULTS: The androecium exhibits the highest evolutionary rates across most characters, whereas most perianth and gynoecium characters evolve more slowly in both Ericales and angiosperms. Both high and low rates of morphological evolution can result in high floral disparity in Ericales. Analyses of an angiosperm-wide floral data set reveal that this pattern appears to be conserved across most major angiosperm clades. CONCLUSIONS: Elevated rates of morphological evolution in the androecium of Ericales may explain the higher disparity reported for this floral module. Comparing rates of morphological evolution through rate profiles proves to be a powerful tool in understanding floral evolution.