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Saxifraga giraldiana Engler is a common subalpine and alpine plant belonging to Saxifragaceae. However, the genetic diversity of this species has remained to be explored. In this study, we have assembled and characterized the complete chloroplast genome of S. giraldiana, filling this knowledge gap and uncovering its genetic composition. The chloroplast genome is 147,267 bp long and contains 131 genes, including 85 protein-coding genes, 38 tRNA genes, and eight rRNA genes. Furthermore, we have performed a phylogenetic analysis of 19 representative species within Saxifraga. As a result, we have found that S. giraldiana, together with S. implicans and S. stellariifolia, forms a monophyletic group. These findings have implications for the conservation and utilization of S. giraldiana.

Abstract Background Successive stamen movement is a complex plant behavior involving successive uplift of stamens and pollen release, which plays a role in reducing sexual interference, increasing pollen deposition and promoting pollen export. Although reported from several taxa, studies on whether the movement can be influenced by abiotic and biotic factors are scarce. Methods In this study, we here for the first time described a pattern of successive stamen movement in Saxifraga candelabrum (Saxifragaceae). We then compared the rates of stamen movement in S. candelabrum under different weather and varying pollinator visits. Pollen packaging and presentation schedule of S. candelabrum were also investigated. Results The results showed that the number of stamens bent per day in sunny days was significantly higher than overcast and rain. Flowers that receive more pollinator visits (control treatment) had significantly higher number of stamen movement than those that received fewer (removal treatment) and none (bagging treatment). Throughout the staminate phase of a flower, there was a progressive increase in both pollen quantity of individual stamens and pollen presentation during each day. Conclusion Our research demonstrates that successive stamen movement in S. candelabrum was accelerated by favorable weather and increased pollinator visits, which may promote pollen export. Moreover, incremental pollen packaging is likely an adaptation to seasonal regularity in variations of sex ratio resulting from protandry.

Abstract Background Accurate species delimitation is fundamental for testing evolutionary theory and provides essential implications for conservation management. The arctic-alpine genus Saxifraga L. (Saxifragaceae) is taxonomically complex and many species have not been critically assessed. The taxonomic and phylogenetic status of Saxifraga lancangensis Y.Y.Qian, considered as a synonym of Saxifraga mengtzeana Engl. & Irmsch. in previous studies, is re-evaluated in light of new evidence presented here. Results Evidence from morphological comparison and sequencing of plastid genome indicate that S. lancangensis belongs to Saxifraga sect. Irregulares Haw., and is closely related to Saxifraga geifolia Balf.f., and S. mengtzeana. However, S. lancangensis can be diagnosed by its petals with red and clawless base, leaf blade orbicular and leaf margin shallowly dentate. Conclusions The morphological and molecular evidence support the resurrection of S. lancangensis as a distinct species. An updated morphological description based on protologue and fresh material, diagnostic characters, and original photographs of the resurrected species are presented.

Background: Successive stamen movement is a complex plant behavior involving successive uplift of stamens and pollen release, which plays a role in reducing sexual interference, increasing pollen deposition and promoting pollen export. Although reported from several taxa, studies on whether the movement can be influenced by abiotic and biotic factors are scarce. Methods: In this study, we here for the first time described a pattern of successive stamen movement in Saxifraga candelabrum (Saxifragaceae). We then compared the rates of stamen movement in S. candelabrum under different weather and varying pollinator visits. Pollen packaging and presentation schedule of S. candelabrum were also investigated. Results: The results showed that the number of stamens bent per day in sunny days was significantly higher than overcast and rain. Flowers that receive more pollinator visits (control treatment) had significantly higher number of stamen movement than those that received fewer (removal treatment) and none (bagging treatment). Throughout the staminate phase of a flower, there was a progressive increase in both pollen quantity of individual stamens and pollen presentation during each day. Conclusion: Our research demonstrates that successive stamen movement in S. candelabrum was accelerated by favorable weather and increased pollinator visits, which may promote pollen export. Moreover, incremental pollen packaging is likely an adaptation to seasonal regularity in variations of sex ratio resulting from protandry. [ABSTRACT FROM AUTHOR]

Background: Accurate species delimitation is fundamental for testing evolutionary theory and provides essential implications for conservation management. The arctic-alpine genus Saxifraga L. (Saxifragaceae) is taxonomically complex and many species have not been critically assessed. The taxonomic and phylogenetic status of Saxifraga lancangensis Y.Y.Qian, considered as a synonym of Saxifraga mengtzeana Engl. & Irmsch. in previous studies, is re-evaluated in light of new evidence presented here. Results: Evidence from morphological comparison and sequencing of plastid genome indicate that S. lancangensis belongs to Saxifraga sect. Irregulares Haw., and is closely related to Saxifraga geifolia Balf.f., and S. mengtzeana. However, S. lancangensis can be diagnosed by its petals with red and clawless base, leaf blade orbicular and leaf margin shallowly dentate. Conclusions: The morphological and molecular evidence support the resurrection of S. lancangensis as a distinct species. An updated morphological description based on protologue and fresh material, diagnostic characters, and original photographs of the resurrected species are presented. [ABSTRACT FROM AUTHOR]

Chrysosplenium guangxiense H.G.Ye & Gui C.Zhang was first described as a new species in 1994 but later synonymized in the Flora of China treatment with C. glossophyllum H.Hara. Plastid genomes and nrDNA sequences were used to infer the phylogenetic relationships of selected taxa in Chrysosplenium. Our phylogenetic analyses revealed that C. guangxiense belongs to sect. Alternifolia, is closely related to Chrysosplenium Hydrocotylifolium H.Lév. & Vaniot but distant from C. glossophyllum. Morphologically, C. guangxiense could be easily distinguished from C. glossophyllum by having robust rhizomes, basal leaves with a long cuneate base and fewer teeth in the margin, curled sepal margins, and red, larger seeds. It could also be easily distinguished from C. hydrocotylifolium by possessing long elliptic leaves and a long cuneate leaf base. Along with the phylogenetic studies, the complete plastid genome of C. guangxiense was also reported. The plastid genome was 154,004 bp in length and comprised two inverted repeats (IRs) of 28,120 bp, separated by a large single-copy of 80,646 bp and a small single-copy of 17,118 bp. A total of 111 functional genes were discovered, comprising 78 protein-coding genes, 29 tRNA genes, and four rRNA genes. Based on assessment of morphological and molecular data Chrysosplenium guangxiense H.G.Ye & Gui C.Zhang is resurrected from C. glossophyllum H.Hara at species level. A global conservation assessment classifies C. guangxiense as Vulnerable (VU). [ABSTRACT FROM AUTHOR]

Bergenia ciliata (Haw.) Sternb. is an important herb predominantly found in the Indian Himalayan Region. It is widely used in medicines, healthcare systems, cosmetics, fodder, and ornamental purposes. The Illumina sequencing and de novo transcriptome assembly were carried out in B. ciliata to develop and identify simple sequence repeat markers. A total of 18 226 simple sequence repeats (SSRs) were identified wherein di-nucleotides were found to be abundant (47.88%), followed by mono-nucleotide (35.03%) and tri-nucleotide (15.88%) repeats. A total of 11 839 EST-SSR primers were designed, of which 96 primer pairs were commercially synthesized. Finally, 17 primer pairs revealed clear, distinct polymorphic bands, and these primers were validated with 40 diverse B. ciliata accessions. The present study revealed moderate level of genetic diversity (Ho = 0.389, He = 0.542, and PIC = 0.513). Furthermore, the transcriptome data and EST-SSR markers generated during the present investigation could be an important genetic resource for functional genomics, population studies, and conservation genetics of the genus Bergenia. [ABSTRACT FROM AUTHOR]

Societal Impact Statement Many mountain plants persist in small, isolated patches on the verge of extinction. Observational methods of monitoring these populations, such as recording the number of flowering stems, do not indicate the number of genetically distinct individuals, which is crucial information for conserving small populations. Here, the rate of clonal reproduction and number of genetic individuals were measured in the threatened Saxifraga hirculus in Scotland. These methods showed that population size is a poor proxy for genotype diversity and identified highly diverse small populations that may otherwise have been overlooked. This highlights the necessity of using genetic data to ensure the successful conservation of threatened plants. Summary Habitat fragmentation and loss increase the isolation of plant populations, increasing the occurrence of within population reproduction, and the potential for negative genetic effects, such as inbreeding depression and loss of genetic diversity. We use the European protected Marsh Saxifrage (Saxifraga hirculus) in Scotland as an example for declining perennial plants and the genetic resources they encapsulate. S. hirculus has declined due to agricultural intensification, drainage, industrial afforestation and grazing. The species can spread by seed or vegetatively through the production of rhizomes. Flowering is rare though due to grazing, which limits sexual reproduction and gene flow. An almost complete genetic inventory of Scottish populations was done using 11 microsatellite markers. Furthermore, archived DNA samples were used to document temporal genetic changes. We showed that clonal growth is predominant in some populations and genetic diversity (HS and allelic richness) is relatively high. However, the number of genetically distinct individuals (genets) per population is extremely low (3–34). Archived DNA samples showed that some populations consist of the same few genets with no evidence for turnover. Thus, while clonal growth may have helped the species to persist, there is limited creation of new gene combinations. Our findings highlight that reducing grazing pressure and increasing gene flow will be essential to rescue this species from its evolutionary dead end. We demonstrate the benefits of genetic monitoring for determining census population sizes and thus effective plant management and conservation. This work further sets out a strategy for moving this species towards demographic and genetic sustainability.

Societal Impact Statement: Many mountain plants persist in small, isolated patches on the verge of extinction. Observational methods of monitoring these populations, such as recording the number of flowering stems, do not indicate the number of genetically distinct individuals, which is crucial information for conserving small populations. Here, the rate of clonal reproduction and number of genetic individuals were measured in the threatened Saxifraga hirculus in Scotland. These methods showed that population size is a poor proxy for genotype diversity and identified highly diverse small populations that may otherwise have been overlooked. This highlights the necessity of using genetic data to ensure the successful conservation of threatened plants. Summary: Habitat fragmentation and loss increase the isolation of plant populations, increasing the occurrence of within population reproduction, and the potential for negative genetic effects, such as inbreeding depression and loss of genetic diversity. We use the European protected Marsh Saxifrage (Saxifraga hirculus) in Scotland as an example for declining perennial plants and the genetic resources they encapsulate. S. hirculus has declined due to agricultural intensification, drainage, industrial afforestation and grazing. The species can spread by seed or vegetatively through the production of rhizomes. Flowering is rare though due to grazing, which limits sexual reproduction and gene flow.An almost complete genetic inventory of Scottish populations was done using 11 microsatellite markers. Furthermore, archived DNA samples were used to document temporal genetic changes.We showed that clonal growth is predominant in some populations and genetic diversity (HS and allelic richness) is relatively high. However, the number of genetically distinct individuals (genets) per population is extremely low (3–34). Archived DNA samples showed that some populations consist of the same few genets with no evidence for turnover. Thus, while clonal growth may have helped the species to persist, there is limited creation of new gene combinations.Our findings highlight that reducing grazing pressure and increasing gene flow will be essential to rescue this species from its evolutionary dead end. We demonstrate the benefits of genetic monitoring for determining census population sizes and thus effective plant management and conservation. This work further sets out a strategy for moving this species towards demographic and genetic sustainability. [ABSTRACT FROM AUTHOR]

AbstractBergenia purpurascens (Hook. f. et Thoms.) Engl. is one species of traditional Chinese medicinal plant. This is the first publication of its complete chloroplast (cp) genome. The whole cp genome has 157,246 base pairs in length with 132 annotated genes, of which were 87 protein-coding genes, 37 tRNAs, and 8 rRNAs. According to the phylogenetic study, B. purpurascens and Bergenia scopulosa T. P. Wang. 1974 had a sister relationship. This genomic data and conclusions from B. purpurascens phylogenetic research will provide useful information and throw light on more in-depth investigations of the systematics and evolutionary patterns of Saxifragaceae.

AbstractThe perennial herb Heuchera micrantha (Saxifragaceae) is a popular ornamental plant. However, the plastome sequence of H. micrantha has not been reported yet. Here, we assembled the complete plastome of H. micrantha using Illumina high-throughput pair-end sequencing. The plastome is a circular DNA molecule of 155,469 bp, comprising a pair of inverted repeat (IR, 25,654 bp) regions, a small single copy (SSC, 18,050 bp) region, and a large single copy (LSC, 86,111 bp) region. It encodes 129 genes, of which 84 are protein-coding genes, 37 are transfer RNAs, and eight are rRNAs. The total GC content is 37.8%. Phylogenetic analysis shows that H. micrantha, together with three other Heuchera species is clustered with Tiarella cordifolia. This complete plastome is beneficial for future genetic research on the Heuchera group.

Bergenia purpurascens (Hook. f. et Thoms.) Engl. is one species of traditional Chinese medicinal plant. This is the first publication of its complete chloroplast (cp) genome. The whole cp genome has 157,246 base pairs in length with 132 annotated genes, of which were 87 protein-coding genes, 37 tRNAs, and 8 rRNAs. According to the phylogenetic study, B. purpurascens and Bergenia scopulosa T. P. Wang. 1974 had a sister relationship. This genomic data and conclusions from B. purpurascens phylogenetic research will provide useful information and throw light on more in-depth investigations of the systematics and evolutionary patterns of Saxifragaceae. [ABSTRACT FROM AUTHOR]

Micranthes octopetala (Nakai) Y.I.Kim & Y.D. Kim et al. 2015, which belongs to the family Saxifragaceae, is a perennial herb endemic to Korea. M. octopetala was originally treated as a synonym of M. manchuriensis. However, in 2015, molecular phylogenetic analysis confirmed that M. octopetala is an independent species. In this study, the plastid genome of M. octopetala was sequenced for the first time, and the taxonomic position of this species was identified. The complete plastid genome of M. octopetala has a total length of 149 751 bp (large single copy: 83 083 bp; small single copy: 17 196 bp; inverted repeat: 24 736 bp), containing 130 genes, including 79 CDS, 30 tRNAs, and 4 rRNAs. Moreover, the absence of intron in the rpl2 gene, which is a common feature of Saxifragaceae, was confirmed. Phylogenetic analysis based on 79 protein-coding genes from 21 species revealed that M. octopetala belongs to the genus Micranthes, being a sister to other Micranthes species. The plastid genome of M. octopetala obtained in this study provides fundamental information for future studies on the genus Micranthes.

A novel bibenzyl derivative, hydrangchin (1) and ten known compounds (2–11) were isolated from the stems of Hydrangea chinensis Maxim. (Saxifragaceae). The structure of the novel bibenzyl derivative was characterized on the basis of spectral analysis. [ABSTRACT FROM AUTHOR]

Micranthes octopetala (Nakai) Y.I.Kim & Y.D. Kim et al. 2015, which belongs to the family Saxifragaceae, is a perennial herb endemic to Korea. M. octopetala was originally treated as a synonym of M. manchuriensis. However, in 2015, molecular phylogenetic analysis confirmed that M. octopetala is an independent species. In this study, the plastid genome of M. octopetala was sequenced for the first time, and the taxonomic position of this species was identified. The complete plastid genome of M. octopetala has a total length of 149 751 bp (large single copy: 83 083 bp; small single copy: 17 196 bp; inverted repeat: 24 736 bp), containing 130 genes, including 79 CDS, 30 tRNAs, and 4 rRNAs. Moreover, the absence of intron in the rpl2 gene, which is a common feature of Saxifragaceae, was confirmed. Phylogenetic analysis based on 79 protein-coding genes from 21 species revealed that M. octopetala belongs to the genus Micranthes, being a sister to other Micranthes species. The plastid genome of M. octopetala obtained in this study provides fundamental information for future studies on the genus Micranthes. [ABSTRACT FROM AUTHOR]

Functional traits fall along a continuum from resource conservative to acquisitive and are powerful predictors of the ecological settings necessary for a species to persist and establish. As a consequence, a major problem that functional trait analysis could address is understanding the ecological contexts necessary for the persistence of polyploid plants, because early generation polyploids, or "neopolyploids," are at a high extinction risk. Because neopolyploidy could increase nutrient limitation, growth strategies should shift to accommodate the increased need for resources, but this prediction is untested. To address this gap, we compared the functional trait responses of diploids, synthetic neotetraploids, and naturally occurring tetraploids of Heuchera cylindrica, an herbaceous perennial plant, to nutrient manipulations in a greenhouse experiment. We found strong support for the hypothesis that neotetraploidy increases nutrient requirements, as evidenced by reduced productivity and increased tissue concentrations of nitrogen and phosphorus in neotetraploids. We also found that the repeated formation of independent origins of neotetraploidy led to differing responses to nutrient supply, but neotetraploidy generally shifted functional traits to be more resource acquisitive and inefficient. Taken together, our results suggest that shifts in functional trait responses may constrain the ability of neopolyploids to establish in nutrient‐poor habitats. [ABSTRACT FROM AUTHOR]

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In the Flora of China account of Saxifraga mengtzeana Engl. & Irmsch., eight synonyms were attributed to it and one variant, recognized as Saxifraga epiphylla Gornall & Ohba, was split from it. This study reevaluates the taxonomic status of some of the synonyms and of the segregated species in light of new evidence presented here. Morphological and molecular evidence demonstrate that collections from northwestern Yunnan and Sichuan are genetically differentiated from those in southeastern Yunnan and neighboring Guangxi. Observations in the field and in cultivation show that the peltate petiole attachment diagnostic of S. mengtzeana var. peltifolia Engl. & Irmsch. is developmentally labile. Similar observations combined with molecular data show that viviparous phenotypes, formerly treated as S. epiphylla, although largely under genetic control, occur sporadically throughout the ranges of both northern and southern taxa. Collections from northwestern Yunnan and Sichuan are best recognized as Saxifraga geifolia Balf.f., whereas those from southeastern Yunnan and neighboring Guangxi are S. mengtzeana. Peltate‐leaved variants of the latter are given no status and are relegated to complete synonymy. Viviparous phenotypes of S. mengtzeana and S. geifolia are recognized at the rank of forma. [ABSTRACT FROM AUTHOR]

The current distribution of most species results from ecological niche, past distribution, and migrations during glacial–interglacial periods and in situ evolution. Here, we disentangle the colonization history of Saxifraga longifolia Lapeyr., a limestone plant abundant in the Pyrenees and rare in other Iberian mountains and the African Atlas. Our working hypothesis is that the current distribution results from the shrinkage of a more extensive distribution in previous cold periods. We sampled 160 individuals of 32 populations across the whole distribution range and sequenced four DNA regions (rpl32‐trnL, rps16‐trnQ, trnS‐trnG, and ITS). Ecological conditions were modeled to identify factors promoting high genetic diversity and long‐term persistence areas for S. longifolia. In addition, we inferred phylogenetic relationships, phylogeographic divergence, genetic diversity, and migration routes. Seven plastid haplotypes were found, of which six occur in the Pyrenees and one in the High Atlas (Morocco). Discrete phylogeographic analysis (DPA) estimated migration routes predominantly from the Pyrenees to the other areas. Colonization events to those areas appear to have taken place recently given that the rest of the Iberian mountains do not harbor exclusive haplotypes. Iberian–Northern African distribution was inferred to be the result of long‐distance dispersal because the split between Iberian and High Atlas haplotypes is estimated to have taken place in the last 4 million years ago when the Strait of Gibraltar was already open. Migrations from the Pyrenees to the south may have been favored by a corridor of predominant limestone rocks along Eastern Iberia, followed by successful overcoming the Strait of Gibraltar to reach northern Africa. [ABSTRACT FROM AUTHOR]

Understanding the ranges of rare and endangered species is central to conserving biodiversity in the Anthropocene. Species distribution models (SDMs) have become a common and powerful tool for analyzing species–environment relationships across geographic space. Although evaluating the distribution of rare species is integral to their conservation, this can be difficult when limited distribution data are available. Community science platforms, such as iNaturalist, have emerged as alternative sources for species occurrence data. Although these observations are often thought to be of lower quality than those of natural history collections, they may have potential for improving SDMs for species with few occurrence records from collections. Here, we investigate the utility of iNaturalist data for developing SDMs for a rare high‐elevation plant, Telesonix jamesii. Because methods for modeling rare species are limited in the literature, five different modeling techniques were considered, including profile methods, statistical models, and machine learning algorithms. The inclusion of iNaturalist data doubled the number of usable records for T. jamesii. We found that a random forest (RF) model using ensemble training data performed the highest of any model (area under curve = 0.98). We then compared the performance of RF models that use only natural history training data and those that use a combination of natural history (herbarium specimens) and iNaturalist training data. All models heavily relied on climate data (mean temperature of driest quarter, and precipitation of the warmest quarter), indicating that this species is under threat as climate continues to change. Validation datasets affected model fits as well. Models using only herbarium data performed slightly poorer when evaluated with cross‐validation than when validated externally with iNaturalist data. This study can serve as a model for future SDM studies of species with similar data limitations. [ABSTRACT FROM AUTHOR]

A major challenge in evolutionary biology is to understand how complex traits of multiple functions have diversified and codiversified across interacting lineages and geographic ranges. We evaluate intra- and interspecific variation in floral scent, which is a complex trait of documented importance for mutualistic and antagonistic interactions between plants, pollinators, and herbivores. We performed a large-scale, phylogenetically structured study of an entire plant genus (Lithophragma, Saxifragaceae), of which several species are coevolving with specialized pollinating floral parasites of the moth genus Greya (Prodoxidae). We sampled 94 Lithophragma populations distributed across all 12 recognized Lithophragma species and subspecies, and four populations of related saxifragaceous species. Our results reveal an unusually high diversity of floral volatiles among populations, species, and clades within the genus. Moreover, we found unexpectedly major changes at each of these levels in the biosynthetic pathways used by local populations in their floral scents. Finally, we detected significant, but variable, genus- and species-level patterns of ecological convergence in the floral scent signal, including an impact of the presence and absence of two pollinating Greya moth species. We propose that one potential key to understanding floral scent variation in this hypervariable genus is its geographically diverse interactions with the obligate specialized Greya moths and, in some species and sites, more generalized copollinators.

Abstract In the Flora of China account of Saxifraga mengtzeana Engl. & Irmsch., eight synonyms were attributed to it and one variant, recognized as Saxifraga epiphylla Gornall & Ohba, was split from it. This study reevaluates the taxonomic status of some of the synonyms and of the segregated species in light of new evidence presented here. Morphological and molecular evidence demonstrate that collections from northwestern Yunnan and Sichuan are genetically differentiated from those in southeastern Yunnan and neighboring Guangxi. Observations in the field and in cultivation show that the peltate petiole attachment diagnostic of S. mengtzeana var. peltifolia Engl. & Irmsch. is developmentally labile. Similar observations combined with molecular data show that viviparous phenotypes, formerly treated as S. epiphylla, although largely under genetic control, occur sporadically throughout the ranges of both northern and southern taxa. Collections from northwestern Yunnan and Sichuan are best recognized as Saxifraga geifolia Balf.f., whereas those from southeastern Yunnan and neighboring Guangxi are S. mengtzeana. Peltate‐leaved variants of the latter are given no status and are relegated to complete synonymy. Viviparous phenotypes of S. mengtzeana and S. geifolia are recognized at the rank of forma.

Tertiary relicts often show evolutionary stasis in morphology and ecology and have been hypothesized to retain stable population sizes in refugia. However, recent studies have reported that some relicts evolutionarily shifted their physiology, ecology, and morphology and experienced various patterns of demography. To understand the historical survival of relict plants, a multidimensional study investigating the evolution of ecological and morphological traits as well as population demographic history is needed. The genus Tanakaea (Saxifragaceae) comprises two species in China and Japan. These species share most vegetative characteristics and are sometimes treated as a single species. The distribution pattern is relictual, as the populations are confined to small areas in mesic warm temperate forests less influenced by Quaternary glacial climates. Focusing on the relictual plant group, this study tested the hypotheses of evolutionary stasis and population stability in long‐term refugia. Genetic analyses using plastome sequences and genome‐wide single‐nucleotide polymorphisms revealed divergence of the two species approximately 6.8 million years ago and strong genetic differentiation of the regional populations. Demographic analysis revealed that almost all populations retained stable population sizes during glacial–interglacial climate changes, supporting the traditional view. However, morphological assessments revealed a simultaneous shift in breeding systems (from hermaphrodite to dioecy/non‐clonal to clonal reproduction) in Japanese species and intraspecific differentiation of leaf traits. Therefore, the relict species do not show evolutionary stasis in every aspect. Changes in reproductive characteristics may have contributed to their long‐term in situ survival. [ABSTRACT FROM AUTHOR]

Peltoboykinia tellimoides, which is distributed in Japan and China, is the type species of Peltoboykinia. In this study, we sequenced and assembled the complete chloroplast (cp) genome of P. tellimoides and reconstructed the phylogeny of Saxifragaceae based on the whole cp genome sequences. The cp genome of P. tellimoides was 156,274 bp in length, comprising a pair of inverted repeat regions (25,099 bp) separated by a large single copy region (88,109 bp) and a small single copy region (17,967 bp). The genome encoded 112 unique genes consisting of 78 different protein-coding genes, 30 transfer RNA and four ribosomal RNA genes, with 16 duplicated genes in the inverted repeats. Phylogenetic analysis indicated that P. tellimoides together with three Chrysosplenium species formed a high support clade, which was sister to Micranthes melanocentra.

Background and aimsMany plant-pollinator interactions are mediated by floral scents that can vary among species, among populations within species and even among individuals within populations. This variation could be innate and unaffected by the environment, but, because many floral volatiles have amino-acid precursors, scent variation also could be affected by differences in nutrient availability among environments. In plants that have coevolved with specific pollinators, natural selection is likely to favour low phenotypic plasticity in floral scent even under different conditions of nutrient availability if particular scents or scent combinations are important for attracting local pollinators.MethodsClonal pairs of multiple seed-families of two Lithophragma bolanderi (Saxifragaceae) populations were subjected to a high and a low nutrient treatment. These plants are pollinated primarily by host-specific Greya moths. It was evaluated how nutrient treatment affected variation in floral scent relative to other vegetative and reproductive traits.Key resultsFloral scent strength (the per-flower emission rate) and composition were unaffected by nutrient treatment, but low-nutrient plants produced fewer and lighter leaves, fewer scapes and fewer flowers than high-nutrient plants. The results held in both populations, which differed greatly in the number and composition of floral scents produced.ConclusionsThe results reveal a strong genetic component both to scent composition and emission level, and partly contrasts with the only previous study that has assessed the susceptibility of floral volatile signals to variation in the abundance of nutrients. These results, and the tight coevolutionary relationship between Lithophragma plants and their specialized Greya moth pollinators, indicate that reproductive traits important to coevolving interactions, such as the floral scent of L. bolanderi, may be locally specialized and more canalized than other traits important for plant fitness.

Closely related species often have similar traits and sometimes interact with the same species. A crucial problem in evolutionary ecology is therefore to understand how coevolving species diverge when they interact with a set of closely related species from another lineage rather than with a single species. We evaluated geographic differences in the floral morphology of all woodland star plant species (Lithophragma, Saxifragaceae) that are pollinated by Greya (Prodoxidae) moths. Flowers of each woodland star species differed depending on whether plants interact locally with one, two, or no pollinating moth species. Plants of one species grown in six different environments showed few differences in floral traits, suggesting that the geographic differences are not due significantly to trait plasticity. Greya moth populations also showed significant geographic divergence in morphology, depending on the local host and on whether the moth species co-occurred locally. Divergence in the plants and the moths involved shifts in combinations of partially correlated traits, rather than any one trait. The results indicate that the geographic mosaic of coevolution can be amplified as coevolving lineages diversify into separate species and come together in different combinations in different ecosystems.

Chrysosplenium nudicaule Bunge, Tibetan name ‘Yajima,’ growing in the highlands of China is a perennial herb belonging to the genus Chrysosplenium Saxifragaceae. As a traditional Chinese medicine, it has been used to treat digestive diseases for hundreds of years. The complete chloroplast genome of Chrysosplenium nudicaule is 152,775 bp in length and comprises two inverted repeats (IR, 25,962 bp), a large single-copy region (LSC, 83,533 bp), and a small single-copy region (SSC, 17,318 bp). It harbors 112 genes, including 78 protein-coding genes, four ribosomal RNA genes, and 30 transfer RNA genes. In addition, the rpl32 gene was deleted. The GC content of the whole chloroplast genome is 37.54%. This chloroplast genome resource will be useful for study on the evolution and genetic diversity of C. nudicaule in the future.

Saxifraga florulenta is a monocarpic species endemic to SW Alps, whose reproductive biology is still unknown. Its knowledge may provide useful information on how the species may face the global warming. In this study, we investigated the plant–pollinator interactions and the mating system of the plant. We described the phenology of the inflorescences, we evaluated the type of pollinators, we verified self and cross-pollination and lastly we quantified the reproductive success of the plants. In spite of the low visitation rate of pollinators, S. florulenta shows high fruit set and seed set. The plant exhibits a prolonged flowering period and has a definite inflorescence, characterized by a sequential blooming bearing both female- or male-phase flowers at the same time; in addition, flowers show a delayed maturation and movement of stamens. These floral and inflorescence traits lead to a mixed reproduction mode which guarantees reproductive success by adopting the "best of two worlds" strategy. The species maximizes its resource investment by having inflorescence size and number of flowers directly related to plant size, whilst fruit and seed set as well as seeds per flower did not show any statistically significant relationship with plant size. [ABSTRACT FROM AUTHOR]

Saxifraga viridiflora, a remarkable new species of the genus Saxifraga sect. Irregulares (Saxifragaceae) from Guangxi, is described and illustrated herein. This new species morphologically differs from all known S. sect. Irregulares taxa by its greenish petals, verruculose sepals, and thick leathery leaf blade abaxially scarlet with white spots. [ABSTRACT FROM AUTHOR]

As mentioned in the texts such as Vedas, Charaka Samhita and Sushruta Samhita, Bergenia ciliata is a known to be the ancient and medicinal plant belonging to the family Saxifragaceae. It can be found as a shrub or an herb, while some are trees or vines. The family includes about 80 genera, 1250 species and has biological properties that are possessed by the different plant parts including antibacterial, antiviral, antioxidant, anti-ulcer, anti-diabetic, analgesic, anti-plasmodial, antitumor, antitussive activity, antineoplastic, diuretic and anti-inflammatory properties. Many of its members grow in rocky places. Bergenia ciliata itself shows that the plant originate between rocks and appears to break them or that it possesses lithotroptic property. It's different plant parts are reported to have different medicinal properties. There are three species of Bergenia found in India Bergenia ligulata, Bergenia ciliata, Bergenia stracheyi. The rhizomes of these plants are used in the indigenous system of medicines. [ABSTRACT FROM AUTHOR]

Abstract Background Chrysosplenium L. (Saxifragaceae) is a genus of plants widely distributed in Northern Hemisphere and usually found in moist, shaded valleys and mountain slopes. This genus is ideal for studying plant adaptation to low light conditions. Although some progress has been made in the systematics and biogeography of Chrysosplenium, its chloroplast genome evolution remains to be investigated. Results To fill this gap, we sequenced the chloroplast genomes of six Chrysosplenium species and analyzed their genome structure, GC content, and nucleotide diversity. Moreover, we performed a phylogenetic analysis and calculated non-synonymous (Ka) /synonymous (Ks) substitution ratios using the combined protein-coding genes of 29 species within Saxifragales and two additional species as outgroups, as well as a pair-wise estimation for each gene within Chrysosplenium. Compared with the outgroups in Saxifragaceae, the six Chrysosplenium chloroplast genomes had lower GC contents; they also had conserved boundary regions and gene contents, as only the rpl32 gene was lost in four of the Chrysosplenium chloroplast genomes. Phylogenetic analyses suggested that the Chrysosplenium separated to two major clades (the opposite group and the alternate group). The selection pressure estimation (Ka/Ks ratios) of genes in the Chrysosplenium species showed that matK and ycf2 were subjected to positive selection. Conclusion This study provides genetic resources for exploring the phylogeny of Chrysosplenium and sheds light on plant adaptation to low light conditions. The lower average GC content and the lacking gene of rpl32 indicated selective pressure in their unique habitats. Different from results previously reported, our selective pressure estimation suggested that the genes related to photosynthesis (such as ycf2) were under positive selection at sites in the coding region.

Abstract Background Plastid gene loss and pseudogenization has been widely documented in parasitic and mycoheterotrophic plants, which have relaxed selective constraints on photosynthetic function. More enigmatic are sporadic reports of pseudogenization and loss of important photosynthesis genes in lineages thought to be fully photosynthetic. Here we report the complete plastid genome of Saniculiphyllum guangxiense, a critically endangered and phylogenetically isolated plant lineage, along with genomic evidence of reduced chloroplast function. We also report 22 additional plastid genomes representing the diversity of its containing clade Saxifragales, characterizing gene content and placing variation in a broader phylogenetic context. Results We find that the plastid genome of Saniculiphyllum has experienced pseudogenization of five genes of the ndh complex (ndhA, ndhB, ndhD, ndhF, and ndhK), previously reported in flowering plants with an aquatic habit, as well as the surprising pseudogenization of two genes more central to photosynthesis (ccsA and cemA), contrasting with strong phylogenetic conservatism of plastid gene content in all other sampled Saxifragales. These genes participate in photooxidative protection, cytochrome synthesis, and carbon uptake. Nuclear paralogs exist for all seven plastid pseudogenes, yet these are also unlikely to be functional. Conclusions Saniculiphyllum appears to represent the greatest degree of plastid gene loss observed to date in any fully photosynthetic lineage, perhaps related to its extreme habitat specialization, yet plastid genome length, structure, and substitution rate are within the variation previously reported for photosynthetic plants. These results highlight the increasingly appreciated dynamism of plastid genomes, otherwise highly conserved across a billion years of green plant evolution, in plants with highly specialized life history traits.

Chrysospleniumguangxiense H.G.Ye & Gui C.Zhang was first described as a new species in 1994 but later synonymized in the Flora of China treatment with C.glossophyllum H.Hara. Plastid genomes and nrDNA sequences were used to infer the phylogenetic relationships of selected taxa in Chrysosplenium . Our phylogenetic analyses revealed that C.guangxiense belongs to sect. Alternifolia, is closely related to Chrysospleniumhydrocotylifolium H.Lév. & Vaniot but distant from C.glossophyllum . Morphologically, C.guangxiense could be easily distinguished from C.glossophyllum by having robust rhizomes, basal leaves with a long cuneate base and fewer teeth in the margin, curled sepal margins, and red, larger seeds. It could also be easily distinguished from C.hydrocotylifolium by possessing long elliptic leaves and a long cuneate leaf base. Along with the phylogenetic studies, the complete plastid genome of C.guangxiense was also reported. The plastid genome was 154,004 bp in length and comprised two inverted repeats (IRs) of 28,120 bp, separated by a large single-copy of 80,646 bp and a small single-copy of 17,118 bp. A total of 111 functional genes were discovered, comprising 78 protein-coding genes, 29 tRNA genes, and four rRNA genes. Based on assessment of morphological and molecular data Chrysospleniumguangxiense H.G.Ye & Gui C.Zhang is resurrected from C.glossophyllum H.Hara at species level. A global conservation assessment classifies C.guangxiense as Vulnerable (VU)., Competing Interests: The authors have declared that no competing interests exist., (Long-Fei Fu, Tian-Ge Yang, De-Qing Lan, Xi-Tang Chen, Hong Liu.)

Leaf venation is an important trait used for specifying and grouping plants. In the present study leaf venation types of 30 Saxifraga L. species, found in Indian Western Himalaya, were studied. Out of these, leaf venation patterns of 20 species were not previously studied. In total, five major types of leaf venations viz. acrodomous (perfect-basal; perfect-suprabasal; imperfect-suprabasal), camptodromous (eucamptodromous), campylodromous, hyphodromous, and palinactinodromous – were recorded. Section Micranthes showed palinactinodromous; section Ciliatae showed mainly acrodromous; section Mesogyne showed palinactinodromous; and sect. Porphyrion showed acrodromous, eucamptodromous and hyphodromous types of venations. Palinactinodromous venation type was found to be the ancestral type. Further molecular study is needed to be done to solve the taxonomic complexity of the genus. [ABSTRACT FROM AUTHOR]

In the present study leaf venation types of 30 Saxifraga L. species, found in Indian Western Himalaya, were studied. Out of these, leaf venation patterns of 20 species were not previously studied. In total, five major types of leaf venations- acrodomous (perfect-basal; perfect-suprabasal; imperfect-suprabasal), camptodromous (eucamptodromous), campylodromous, hyphodromous, and palinactinodromous – were recorded. Section Micranthes show palinactinodromous; section Ciliatae show mainly acrodromous; section Mesogyne show palinactinodromous; and sect. Porphyrion show acrodromous, eucamptodromous and hyphodromous types of venation. Palinactinodromous appear as the ancestral venation type.

Chrysosplenium sangzhiense Hong Liu, a new species from Hunan, China, is described and illustrated. The phylogenetic analysis revealed that the new species belongs to subgen. Chrysosplenium and is closely related to C. grayanum, C. nepalense and C. sinicum. The chromosome number of the new species is 2n = 46, indicating a novel basic number x = 23 in Chrysosplenium that is different from other species. This also suggests that C. sangzhiense is probably an allopolyploid derivative of a species with x = 11 and one with x = 12. Morphologically, C. sangzhiense can be easily distinguished from C. grayanum, C. nepalense, C. sinicum and C. cavaleriei, a species not included in our phylogenetic analysis by a suite of characters relating to the sterile shoots, basal leaves, cauline leaves, flowering stem, sepals, disc, capsule and seed. A global conservation assessment is performed, and classifies C. sangzhiense as Least Concern (LC). [ABSTRACT FROM AUTHOR]

Premise: The genetic structure between plant populations is facilitated by the spatial population arrangement and limited dispersal of seed and pollen. Saxifraga acerifolia, a local endemic species in Japan, is a habitat specialist that is confined to waterfalls in riparian environments. Its sister species, Saxifraga fortunei, is a generalist that is widely distributed along riverbanks. Here, we examined sympatric populations of the two Saxifraga species to test whether the differences in habitat preference and colonization process influenced regional and local genetic structures. Methods: To reveal genetic structures, we examined chloroplast microsatellite variations and genome‐wide nucleotide polymorphisms obtained by genotyping by sequencing. We also estimated the gene flow among and within populations and performed landscape genetic analyses to evaluate seed and pollen movement and the extent of genetic isolation related to geographic distance and/or habitat differences. Results: We found strong genetic structure in the specialist S. acerifolia, even on a small spatial scale (<1 km part); each population on a different waterfall in one river system had a completely different predominant haplotype. By contrast, the generalist S. fortunei showed no clear genetic differentiation. Conclusions: Our findings suggest that the level of genetic isolation was increased in S. acerifolia by the spatially fragmented habitat and limited seed and pollen dispersal over waterfalls. Habitat differentiation between the sister taxa could have contributed to the different patterns of gene flow and then shaped the contrasting genetic structures. [ABSTRACT FROM AUTHOR]

The family Saxifragaceae, the current composition of which is one of the great surprises of molecular systematics, has been subject to massive improvements in the knowledge of phylogenetic relationships. Nevertheless, developments from phylogenomic efforts have yet to be mobilized to inform biogeography and taxonomy. Here, we use a recent order‐level phylogeny for Saxifragaceae and related families covering 72% of species with a set of new analyses to assess habitat evolution and biogeography. Our results suggest a North American origin of the family in cold alpine habitats, followed by rapid recent evolution of diverse habitat tolerances. We also combine these recent phylogenomic results and a synthesis of the literature to improve generic limits and tribal classification of Saxifragaceae. We recognize 40 genera in 10 tribes, with 14 new combinations, and elevate one subtribe as well as describing four new taxa at the tribal level. Finally, we synthesize information about biogeography and morphology for the family. [ABSTRACT FROM AUTHOR]

Peltoboykinia tellimoides, which is distributed in Japan and China, is the type species of Peltoboykinia. In this study, we sequenced and assembled the complete chloroplast (cp) genome of P. tellimoides and reconstructed the phylogeny of Saxifragaceae based on the whole cp genome sequences. The cp genome of P. tellimoides was 156,274 bp in length, comprising a pair of inverted repeat regions (25,099 bp) separated by a large single copy region (88,109 bp) and a small single copy region (17,967 bp). The genome encoded 112 unique genes consisting of 78 different protein-coding genes, 30 transfer RNA and four ribosomal RNA genes, with 16 duplicated genes in the inverted repeats. Phylogenetic analysis indicated that P. tellimoides together with three Chrysosplenium species formed a high support clade, which was sister to Micranthes melanocentra. [ABSTRACT FROM AUTHOR]

Chemical defenses, repellents, and attractants are important shapers of species interactions. Chemical attractants could contribute to the divergence of coevolving plant-insect interactions, if pollinators are especially responsive to signals from the local plant species. We experimentally investigated patterns of daily floral scent production in three Lithophragma species (Saxifragaceae) that are geographically isolated and tested how scent divergence affects attraction of their major pollinator-the floral parasitic moth Greya politella (Prodoxidae). These moths oviposit through the corolla while simultaneously pollinating the flower with pollen adhering to the abdomen. The complex and species-specific floral scent profiles were emitted in higher amounts during the day, when these day-flying moths are active. There was minimal divergence found in petal color, which is another potential floral attractant. Female moths responded most strongly to scent from their local host species in olfactometer bioassays, and were more likely to oviposit in, and thereby pollinate, their local host species in no-choice trials. The results suggest that floral scent is an important attractant in this interaction. Local specialization in the pollinator response to a highly specific plant chemistry, thus, has the potential to contribute importantly to patterns of interaction specificity among coevolving plants and highly specialized pollinators.

Mountains are among the most biodiverse places on Earth, and plant lineages that inhabit them have some of the highest speciation rates ever recorded. Plant diversity within the alpine zone - the elevation above which trees cannot grow-contributes significantly to overall diversity within mountain systems, but the origins of alpine plant diversity are poorly understood. Here, we quantify the processes that generate alpine plant diversity and their changing dynamics through time in Saxifraga (Saxifragaceae), an angiosperm genus that occurs predominantly in mountain systems. We present a time-calibrated molecular phylogenetic tree for the genus that is inferred from 329 low-copy nuclear loci and incorporates 73% (407) of known species. We show that upslope biome shifts into the alpine zone are considerably more prevalent than dispersal of alpine specialists between regions, and that the rate of upslope biome shifts increased markedly in the last 5 Myr, a timeframe concordant with a cooling and fluctuating climate that is likely to have increased the extent of the alpine zone. Furthermore, alpine zone specialists have lower speciation rates than generalists that occur inside and outside the alpine zone, and major speciation rate increases within Saxifraga significantly pre-date increased rates of upslope biome shifts. Specialisation to the alpine zone is not therefore associated with speciation rate increases. Taken together, this study presents a quantified and broad scale perspective of processes underpinning alpine plant diversity., (© 2024. The Author(s).)

Saxifraga L. is the largest genus in Saxifragaceae and a characteristic component of the herbaceous flora of the temperate and alpine mountains in the Northern Hemisphere. Section Irregulares is a small group of 15–20 species, representing one of the early‐diverged lineages in the genus characterized with unique zygomorphic flowers. We used both nuclear internal transcribed spacer and chloroplast DNA regions (psbA‐trnH, trnL‐F, and matK) to reconstruct its species relationships, estimate divergence times, and infer its historical biogeography. Our phylogenetic results corroborate the monophyly of sect. Irregulares and its sister relationship to sect. Heterisia from North America. The section was well resolved into two lineages corresponding to their morphological features and biogeographic distributions. One represents ser. Stoloniferae including S. stolonifera Curtis and S. nipponica Makino with long‐creeping stolons/rhizomes and small petals with spots and the other comprises the remaining taxa (ser. Rufescentes) which lack long‐creeping rhizomes. Spots on leaves (abaxially spotted vs. abaxially without spots) and spots on petals (without spots vs. with spots) are inferred to be phylogenetically informative within ser. Rufescentes. Divergence time estimates and ancestral area analysis suggested a western North American origin of sect. Irregulares with migration into East Asia by way of the Bering land bridge in the Middle Oligocene. The development of drying and desertification belt in the late Miocene could have played an important role in the subsequent restriction and separation of the north and south lineages within eastern Asia. [ABSTRACT FROM AUTHOR]