Species Of Saxifraga Research Articles

Chemical constituents of five Saxifraga species and their virucidal activities

The chemical constituents of Saxifraga stolonifera, S. fortunei, S. nipponica, S. cortusifolia, and S. rebunshirensis were investigated for structure and virucidal activity. In addition to the Saxifraga species-derived tannins, 30 compounds (1−30) were isolated from the five species. 5-Hydroxy-4-methoxy-3-O-(6-O-caffeoyl)-β-D-glucopyranosyl benzoic acid (1) and kaempferol 3-O-β-d-xylopyranosyl-(1 → 2)-β-D-xylopyranoside (2) were identified as undescribed compounds. Although 1 was isolated as the (E)-isomer of its caffeoyl moiety, under light it became over time a mixture of the (Z)-isomer (1a) and (E)-isomer (1). Kaempferol 3-O-β-d-xylopyranosyl-(1 → 2)-β-D-glucopyranoside (3) was an analog of 2 and a known compound, whose NMR assignment was reconsidered and described. 6-Isopropyl-5,5-dimethyldihydropyrimidine-2,4(1H,3H)-dione (30) was expected to be a racemic mixture based on its optical rotation and X-ray crystallography data. The virucidal activities of the isolated compounds (1–30) against influenza A virus, severe acute respiratory syndrome coronavirus 2, feline calicivirus, and murine norovirus were evaluated to identify the presence of compounds other than the Saxifraga species-derived tannins, which exhibited virucidal activities. Although the isolated compound activities were relatively weak compared to those of Saxifraga species-derived tannins, the potential virucidal activity of the compounds with galloyl groups was confirmed.

A New Species of Saxifraga from Tibet

No Abstract.

Comparative Analysis of Chloroplast Genomes within Saxifraga (Saxifragaceae) Takes Insights into Their Genomic Evolution and Adaption to the High-Elevation Environment

Saxifraga species are widely distributed in alpine and arctic regions in the Northern hemisphere. Highly morphological diversity within this genus brings great difficulties for species identification, and their typical highland living properties make it interesting how they adapt to the extreme environment. Here, we newly generated the chloroplast (cp) genomes of two Saxifraga species and compared them with another five Saxifraga cp genomes to understand the characteristics of cp genomes and their potential roles in highland adaptation. The genome size, structure, gene content, GC content, and codon usage pattern were found to be highly similar. Cp genomes ranged from 146,549 bp to 151,066 bp in length, most of which comprised 130 predicted genes. Yet, due to the expansion of IR regions, the second copy of rps19 in Saxifraga stolonifera was uniquely kept. Through sequence divergence analysis, we identified seven hypervariable regions and detected some signatures of regularity associated with genetic distance. We also identified 52 to 89 SSRs and some long repeats among seven Saxifraga species. Both ML and BI phylogenetic analyses confirmed that seven Saxifraga species formed a monophyletic clade in the Saxifragaceae family, and their intragenus relationship was also well supported. Additionally, the ndhI and ycf1 genes were considered under positive selection in species inhabiting relatively high altitudes. Given the conditions of intense light and low CO2 concentration in the highland, the products of these two genes might participate in the adaptation to the extreme environment.

Recruitment Traits Could Influence Species' Geographical Range: A Case Study in the Genus Saxifraga L.

The reasons why some species occur widespread, while related species have restricted geographical ranges have been attributed to habitat specialization or ecological niche breadth. For species in the genus Saxifraga, habitat specialization alone cannot explain the distributional differences observed. We hypothesize that recruitment traits (i.e., germination, emergence, and survival) may account for differences in geographical ranges and that early life stages correlate to survival. We studied recruitment responses in 13 widespread and 12 narrow-ranged Saxifraga species in the laboratory and common garden experiments using seeds collected from 79 populations in the European Alps. We found that in the laboratory cold temperature led to higher germination percentages compared with warm temperature for both distribution groups. This represents an exception to the general assumption that alpine species require warm cues for germination. In warm laboratory temperatures, widespread species germinated better than narrow-ranged species, indicating a greater tolerance of warm temperatures for the former. Subsequent to germination, recruitment traits between the two distribution groups were lower or null in the common garden, suggesting that the impact of recruitment on species’ geographical ranges occurs at the earliest life stage. Mean time to emergence of narrow-ranged species showed lower variability than that of widespread species. Consistently, intraspecific variation of mean annual temperatures between seed collection sites was lower for narrow-ranged species, indicating a close relationship between home sites and emergence time. Emergence percentage was a strong predictor of survival only for widespread species, underlining that seed and seedling functional traits differ between distribution groups, which require further research. Our results support the view that early life stages are critical to population dynamics and thus can influence species’ geographical ranges. The wider responses to climatic conditions in widespread species may have facilitated their spread across the Alps. Our results also suggest that all Saxifraga species face a considerable threat from climate warming due to their overall cold-adapted recruitment niche.

Global patterns of species richness of the holarctic alpine herbSaxifraga: the role of temperature and habitat heterogeneity

AbstractThe effects of contemporary climate, habitat heterogeneity and long-term climate change on species richness are well studied for woody plants in forest ecosystems, but poorly understood for herbaceous plants, especially in alpine–arctic ecosystems. Here, we aim to test if the previously proposed hypothesis based on the richness–environment relationship could explain the variation in richness patterns of the typical alpine–arctic herbaceous genus Saxifraga.Using a newly compiled distribution database of 437 Saxifraga species, we estimated the species richness patterns for all species, narrow- and wide-ranged species. We used generalized linear models and simultaneous autoregressive models to evaluate the effects of contemporary climate, habitat heterogeneity and historical climate on species richness patterns. Partial regressions were used to determine the independent and shared effects of different variables. Four widely used models were tested to identify their predictive power in explaining patterns of species richness.We found that temperature was negatively correlated with the richness patterns of all and wide-ranged species, and that was the most important environmental factor, indicating a strong conservatism of its ancestral temperate niche. Habitat heterogeneity and long-term climate change were the best predictors of the spatial variation of narrow-ranged species richness. Overall, the combined model containing five predictors can explain ca. 40%–50% of the variation in species richness. We further argued that additional evolutionary and biogeographical processes might have also played an essential role in shaping the Saxifraga diversity patterns and should be considered in future studies.

Comparative analysis of spatial genetic structures in sympatric populations of two riparian plants, Saxifraga acerifolia and Saxifraga fortunei.

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. 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. 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. 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.

The Role of Hybridisation in the Making of the Species-Rich Arctic-Alpine Genus Saxifraga (Saxifragaceae)

Evolutionary processes fuelling rapid species diversification are not yet fully understood, although their major contribution to overall patterns of plant biodiversity is well established. Hybridisation is among the least understood of these processes, despite its multifaceted role in speciation processes being widely accepted. Species of the large arctic-alpine genus Saxifraga are notorious for their ability to hybridise; however, the overall role of hybridisation and polyploidisation for the diversification of this genus remains unknown. Here, we provide a comprehensive genus-wide review of hybridisation accounts and ploidy levels. We find that the sections of Saxifraga vary greatly in their propensity to hybridise. The majority of natural hybridisation accounts are from recent localised events (n = 71). Hybridisation hotspots were located in the Pyrenees and the European Alps, thus contrasting with the overall distribution of species richness in the genus. Hybrids or hybrid populations are often short-lived in Saxifraga due to a multitude of reproductive barriers, most commonly low F1 hybrid fertility. However, these barriers are not always fully effective, allowing for backcrossing and the formation of hybrid swarms. In addition, we find that the incidence of polyploidy varies widely across different sections of Saxifraga, with species-rich sections Porphyrion and Saxifraga showing divergent polyploidy proportions. Overall, we show that hybridisation and polyploidisation played differential roles in the diversification of this large genus. Nevertheless, a significant proportion of species are yet to be scrutinised, particularly among the Asian Saxifraga species, illustrating the need for systematic further study to fully unravel the role of hybridisation during the evolution of Saxifraga.

The Importance of Including Soil Properties When Disentangling the Drivers of Species Richness: The Case of the Alpine Genus Saxifraga L. in China

Despite the numerous studies on the large-scale patterns of species richness, the spatial variation and determinants of species richness for alpine plant are still an outstanding question and critical to future biodiversity conservation. The genus Saxifraga is a typical alpine plant group with high species richness in the Himalaya-Hengduan mountain regions, China. We performed simple regression models and variance partitioning to assess the importance of different factors, especially soil-related ones, in driving Saxifraga richness patterns. The results showed that environmental energy, habitat heterogeneity, and soil heterogeneity together dominated the spatial variation of species richness. The coarse fragments volume of soil, elevation range, and soil heterogeneity, are positively related to Saxifraga richness. Soil slightly outperforms habitat heterogeneity in predicting the spatial variation of Saxifraga species richness with an explanatory power of 39.3% and 36.6%, respectively. Environmental energy, such as the maximum temperature of the warmest quarter, is negatively correlated with species richness and explains 44.8% of spatial variation of Saxifraga richness. Multiple regression models, including three variables, each representing energy, soil, and habitat heterogeneity, can only explain 53.1% variation of species richness. Variance partitioning outscored 26% of the shared effects of the three variables, while the independent effect of each variable is less than 10%. These results indicated the energy, soil, and habitat heterogeneity together are primary determinants of the spatial variation of Saxifraga species richness. However, there are probably other hidden factors predicting species richness variation due to the low explanatory power of the multiple regression models. Our study emphasizes the significance of soil properties in determining species richness patterns in China, especially for the alpine plant groups. The negative association of species richness with temperature suggests a potential threat of alpine biodiversity loss in HHM from future warming.

Development and application of polymorphic microsatellite markers in Saxifraga genus

In order to develop genomic-SSR markers for species of Saxifraga genus, a mixed plant genomic DNA sample was sequenced based on high-throughput Illumina MiSeq platform. According to genomic sequencing data, SSR loci were identified with MISA software, and then primers were designed with Primer 3 software. A total of 120 pairs of primers were randomly synthesized and amplified in genomic DNA of a few plant samples. Those primers who have yielded polymorphic bands and were considered easy to amplify were identified. After that, transferability of these primers was evaluated, and phylogenetic relationship of 25 species of Saxifraga genus was analyzed with UPGMA (unweighted pair group method analysis). In our results, 587 256 sequences containing SSRs were identified from a total of 1 881 979 combined read pairs obtained in genomic sequencing. Primers were designated to amplify SSRs containing two to six nucleotide repeat units, screened in a small portion of species. Finally, 17 pairs of primers which have produced abundant of polymorphic bands with little problem were amplified in 25 species of Saxifraga genus. A total of 2 687 polymorphic bands were obtained, the average polymorphic rate was 158 bands per pairs of primers. The transferability rate was ranging from 88.0% to 100% across 25 species of Saxifraga. In phylogenetic analysis, the clustering of 25 species based on 17 pairs of SSR primers was different from morphological classification. Our analysis has provided molecular data for genetic relationship of Saxifraga genus, and the transferable and polymorphic SSRs have provided information for genetic diversity research.

Population Genetic Differentiation and Taxonomy of Three Closely Related Species of Saxifraga (Saxifragaceae) from Southern Tibet and the Hengduan Mountains.

The effects of rapid, recent uplift of the Hengduan Mountains on evolution and diversification of young floristic lineages still remain unclear. Here, we investigate diversification of three closely related Saxifraga species with a distribution restricted to the Hengduan Mountains (HM) and southern Tibet, and comment on their taxonomy based on molecular evidence. Three chloroplast DNA fragments (rbcL, trnL-F, trnS-G) and the nuclear ribosomal DNA internal transcribed spacer (ITS) were employed to study genetic structure across 104 individuals from 12 populations of Saxifraga umbellulata, S. pasumensis, and S. banmaensis. Chloroplast DNA (cpDNA) phylogenies revealed two well supported clades, corresponding to S. umbellulata and S. pasumensis plus S. banmaensis. Topology of the ITS phylogeny was largely congruent with that generated from cpDNA haplotypes, but with minor conflicts which might be caused by incomplete lineage sorting. Analyses of molecular variance of both cpDNA and ITS datasets revealed that most variation was held between S. pasumensis s.l. (with S. banmaensis) and S. umbellulata (92.31% for cpDNA; 69.78% for ITS), suggesting a high degree of genetic divergence between them. Molecular clock analysis based on ITS dataset suggested that the divergence between S. pasumensis s.l. and S. umbellulata can be dated to 8.50 Ma, probably a result of vicariant allopatric diversification associated with the uplift events of the HM. Vicariance associated with HM uplifts may also have been responsible for infraspecific differentiation in S. pasumensis. In contrast, infraspecific differentiation in S. umbellulata was most likely triggered by Quaternary glaciations. The much lower levels of gene diversity within populations of S. pasumensis compared with S. umbellulata could have resulted from both range contractions and human collection on account of its putative medicinal properties. Combining evidence from morphology, geographical distributions and molecular phylogenetic data, we recommend that S. banmaensis should be treated as a synonym of S. pasumensis which in turn, and based on the same sources of evidence, should be treated as a separate species rather than as a variety of S. umbellulata.

Evolutionary radiations in the species-rich mountain genus Saxifraga L.

BackgroundA large number of taxa have undergone evolutionary radiations in mountainous areas, rendering alpine systems particularly suitable to study the extrinsic and intrinsic factors that have shaped diversification patterns in plants. The species-rich genus Saxifraga L. is widely distributed throughout the Northern Hemisphere, with high species numbers in the regions adjacent to the Qinghai-Tibet Plateau (QTP) in particular the Hengduan Mountains and the Himalayas. Using a dataset of 297 taxa (representing at least 60% of extant Saxifraga species), we explored the variation of infrageneric diversification rates. In addition, we used state-dependent speciation and extinction models to test the effects of geographic distribution in the Hengduan Mountains and the entire QTP region as well as of two morphological traits (cushion habit and specialized lime-secreting glands, so-called hydathodes) on the diversification of this genus.ResultsWe detected two to three rate shifts across the Saxifraga phylogeny and two of these shifts led to radiations within two large subclades of Saxifraga, sect. Ciliatae Haworth subsect. Hirculoideae Engl. & Irmsch. and sect. Porphyrion Tausch subsect. Kabschia Engl. GEOSSE analyses showed that presence in the Hengduan Mountains had a positive effect on diversification across Saxifraga. Influence of these mountains was strongest in Saxifraga sect. Ciliatae subsect. Hirculoideae given its pronounced distribution there, and thus the radiation in this group can be classified at least partially as geographic. In contrast, the evolution of the cushion life form and lime-secreting hydathodes had positive effects on diversification only in selected Saxifraga sections, including sect. Porphyrion subsect. Kabschia. We therefore argue that radiation in this group was likely adaptive.ConclusionsOur study underlines the complexity of processes and factors underpinning plant radiations: Even in closely related lineages occupying the same life zone, shifts in diversification are not necessarily governed by similar factors. In conclusion, alpine plant radiations result from a complex interaction among geographical settings and/or climatic modifications providing key opportunities for diversification as well as the evolution of key innovations.

Testing the influence of gravity on flower symmetry in five Saxifraga species.

Flower symmetry is considered a species-specific trait and is categorized in asymmetry, actinomorphic symmetry, bisymmetry and zygomorphic symmetry. Here we report on the intra-individual variation of flower symmetry in the genus Saxifraga and the influence of light, gravity and intrinsic factors on the development of flower symmetry. We tested five species-Saxifraga cuneifolia, Saxifraga imparilis, Saxifraga rotundifolia, Saxifraga stolonifera and Saxifraga umbrosa-concerning six flower parameters-angles between petals, petal length, petal pigmentation, angular position of carpels, movement of stamens and (only for S. imparilis and S. stolonifera) the length of the two lower elongated petals in regard to their position towards the stem. Specimens of all species were tested on a vertical clinostat as a gravity compensator, on a horizontal clinostat as a light incidence compensator and on a stationary control. The results show that the angle of incident light has no apparent impact on flower symmetry, whereas gravity affects the angular position of petals in S. cuneifolia and S. umbrosa and the petal colouration in S. rotundifolia. In S. cuneifolia and S. umbrosa, the absence of directional gravity resulted in the development of actinomorphic flowers, whereas the corresponding control flowers were zygomorphic. The development of flowers in S. rotundifolia was not altered by this treatment. The length of the two elongated petals in S. stolonifera and S. imparilis was not affected by gravity, but rather was determined by position of the flower within the inflorescence and resulted in asymmetrical flowers.

Dichotrachelus pesarinii sp.n., a missing link between the species from the central and the western southern Alps (Coleoptera: Curculionidae: Cyclominae)

A new species of Dichotrachelus associated with Saxifraga caesia is described, D. pesarinii sp.n., and a phylogenetic analysis is carried out based on morphological and molecular mtCOI data. The new species resolved as the sister group of the D. sulcipennis group, whose members are associated with Saxifraga oppositifolia. This placement suggests that this group differentiated in the central southern Alps, and that the new species represents an evolutionary link between the species from the central and eastern southern Alps and those from the western Alps. The evolution of the association of Dichotrachelus with particular Saxifraga species remains contradictory; scenarios are proposed.

Leaf venation patterns in the genus Saxifraga (Saxifragaceae)

Details of leaf venation in 150 species of Saxifraga L. representing all 13 sections are reported. Three main patterns were observed: palinactinodromous, acrodromous and camptodromous. Species with lobed leaves have palinactinodromous patterns, whereas those with entire leaves are acrodromous or camptodromous. Thus species from sections Heterisia, Irregulares, Cymbalaria, Cotylea and Mesogyne are palinactinodromous; species from sections Saxifraga, Ciliatae, Bronchiales and Trachyphyllum, in which some have lobed and some have entire leaves, are either palinactinodromous or acrodromous, according to the leaf margin; species from sections Ligulatae and Porphyrion, whose hydathodes secrete calcium bicarbonate, have camptodromous venation. Intermediate states occur in section Gymnopera (between palinactinodromous and camptodromous) and in section Porphyrion subsection Oppositifoliae and section Xanthizoon (intermediate between acrodromous and camptodromous). The pattern of distribution across the genus and selected outgroups shows that the ancestral state is likely to be palinactinodromous.

Comparative germination ecology of two altitudinal vicariant Saxifraga species endemic to the north of Spain

Seeds of high-mountain species are thought to germinate rapidly, synchronously and at high percentages after a cold period, with limited dependence on the external environment; yet, empirical evidence only partially supports this behaviour. We performed a comparative study of the germination response of two closely related taxa along an altitude gradient in northern Spain. Seeds from several maternal families of six populations of Saxifraga trifurcata (lowland species) and S.canaliculata (highland species) were subjected to temperature and stratification treatments. Germination percentages and germination rates were analysed using generalised linear mixed modelling and accelerated failure-time modelling. We found that germination percentages and germination rates were high and dependent on incubation temperature in both species. Within species, seeds from higher altitudes had higher germination percentages under all conditions. Cold-wet stratification negatively affected germination success, particularly in the lowland species. Overall, the highland species was less responsive to the experimental treatments and showed more synchronous germination patterns. We conclude that seeds from these two Saxifraga species germinate as efficiently as species from other habitats, but have a narrower germination response, probably due to the stronger selective pressures in their harsh environments. Finally, a cold, wet stratification period is not a prerequisite for the germination of high-mountain S.canaliculata, and its strong negative effect on the germination of its lowland relative S.trifurcata may contribute to the altitudinal segregation of these two species.

SAXIFRAGA MINUTISSIMA, A NEW SPECIES FROM THE GARHWAL HIMALAYA, INDIA, AND ITS IMPLICATIONS FOR THE TAXONOMY OF THE GENUS SAXIFRAGA (SAXIFRAGACEAE)

Saxifraga minutissima D.S.Rawat, a new and extremely small species of Saxifraga (Saxifragaceae), is described from the Garhwal Himalaya, India. It differs from all other species of Saxifraga, except Saxifraga bicuspidata, in having five stamens and lacking petals. It can be distinguished from Saxifraga bicuspidata in having leaves and sepals entire. The finely striate pollen exine pattern of Saxifraga minutissima indicates that the species belongs to Saxifraga section Ciliatae. Its prostrate, axillary leafy shoots and lack of crisped, rufous hairs strongly suggests a place in Saxifraga subsection Serpyllifoliae, where one of its closest relatives may be Saxifraga stella-aurea.

Chromosome studies in four species of Saxifraga L. (Saxifragaceae) collected in Tuva and adjacent regions, Russia

Four species of Saxifraga such as S. aestivalis, S. hieracifolia, S. foliolosa and S. hirculus were collected in Russia and observed their somatic chromosome numbers. Saxifraga aestivalis showed the somatic chromosome number of 2n=28, S. hieracifolia showed that of 2n=114, S. foliolosa showed that of 2n=60, and S. hirculus showed that of 2n=16. These chromosome numbers verifi ed with the previous reports, expect for S. foliolosa. The chromosome numbers of 2n=60 for S. foliolosa was reported here for the fi rst time. Chromosome sizes of four species varied in mean lengths with each others; S. aestivalis and S. foliolosa had small-sized mean chromosomes 1.31 μm and 1.47 μm long respectively, S. hieracifolia had medium sized, mean chromosome 2.75 μm long, and S. hirculus had large sized, mean chromosome 4.4 μm long. Saxifraga aestivalis had a bi-modality and decrease in chromosome size from the largest to the smallest chromosomes, while the other three species had mono-modal or gradual decrease in chromosome size from the largest to the smallest chromosomes.

Systematic significance of pollen nucleus number in the genus Saxifraga (Saxifragaceae)

Pollen cell number was studied in 124 species of Saxifraga and six species of Micranthes. Binucleate pollen is present in all the six species of Micranthes studied and in 96 species of Saxifraga. Trinucleate pollen occurs in 28 species of Saxifraga and is confined to section Ciliatae subsection Hirculoideae, where it is a putative synapomorphy, and to two species of subsection Serpyllifoliae. Trinucleate pollen does not characterise the whole of section Ciliatae as has been thought. The state has probably evolved at least twice, and possibly three times, in the genus. One new nomenclatural combination is made in Micranthes.

Do alpine plants facilitate each other’s pollination? Experiments at a small spatial scale

Short growing seasons, low temperatures, and frequent strong wind classify high mountains as adverse environments, in which pollinator abundance and activity are reduced. In such environments, plants growing in dense stands comprising several species and thus exhibiting larger and more diverse flower displays may profit by attracting more visits from scarce alpine pollinators than do plants that grow alone or in patches only composed of conspecifics. To study whether aggregation of plants increases (facilitation) or decreases (competition) the attraction of pollinators, we measured the rate and numbers with which insects entered experimental plots in the Swiss Alps, and their behaviour at flowers in plots that they entered. The plots contained individuals of the blue-flowering cushion plant Eritrichium nanum, either alone or mixed with white- to yellowish-flowering Saxifraga species. Pollinators were generally rare: in 55% of 236 observation periods, no pollinators were observed. Over 95% of the pollinators were Diptera. The average probability of observing any insect at all was higher in plots that contained some Saxifraga flowers, including mixed plots, than in those containing only E. nanum flowers. However, although insects tended to choose Saxifraga as the first flower visited in mixed plots, in all other regards their visitation of Saxifraga and E. nanum flowers in such plots was statistically indistinguishable. We also detected no effect of floral neighbourhood on the frequencies of potentially geitonogamous visits or of transitions among individual plants of the same or different species. Thus, our study suggests that the presence of Saxifraga may facilitate visitation to E. nanum at larger spatial scales, but gives no evidence for either competition or facilitation at small scales within floral neighbourhoods.

Effects of floral neighborhood on seed set and degree of outbreeding in a high-alpine cushion plant

Plants flowering together may influence each other's pollination and fecundity over a range of physical distances. Their effects on one another can be competitive, neutral, or facilitative. We manipulated the floral neighborhood of the high-alpine cushion plant Eritrichium nanum in the Swiss Alps and measured the effects of co-flowering neighbors on both the number of seeds produced and the degree of inbreeding and outbreeding in the offspring, as deduced from nuclear microsatellite markers. Seed set of E. nanum did not vary significantly with the presence or absence of two Saxifraga species growing as near neighbors, but it was higher in E. nanum cushions growing at low conspecific density than in those growing at high density. In addition, floral neighborhood had no detectable effect on the degree of selfing of E. nanum, but seeds from cushions growing at low conspecific density were more highly outbred than seeds from cushions at high density. Thus, there was no evidence of either competition or facilitation between E. nanum and Saxifraga spp. as mediated by pollinators at the spatial scale of our experimental manipulation. In contrast, the greater fecundity of E. nanum cushions at low density was consistent with reduced intraspecific competition for pollinators and might also represent a beneficial effect of highly outbred seeds as brought about by more long-distance pollinator flights under low-density conditions.