Reproducing under a warming climate: long winter flowering and extended flower longevity in the only Mediterranean and maritime Primula

Under the pressure of global warming, general expectations of species migration and evolution of adaptive traits should always be confirmed with species‐specific studies. Within this framework, some species can be used as study systems to predict possible consequences of global warming also on other relatives. Unlike its mountain congeneric, Primula palinuri Petagn. has endured all the climatic fluctuations since the Pleistocene, while surviving on Mediterranean coastal cliffs. The aim of this work was to investigate the possible evolution of reproductive biological and ecological traits in P. palinuri adaptation to a warmer environment. Data showed that flowering starts in mid‐winter; single flowers remain open for over a month, changing from pendulous to erect. The number of insects visiting flowers of P. palinuri increases during the flowering season, and pollination reduces flower longevity. Overall, the best pollen performances, in terms of viability and germinability, occur at winter temperatures, while pollinator activity prolongs flowering until spring. Moreover, extended longevity of single flowers optimises reproductive success. Both phenotypic plasticity and selective processes might have occurred in P. palinuri. However, we found that reproductive traits of the only Mediterranean Primula remain more associated with cold mountain habitats than warm coastal cliffs. Given the rapid trend of climate warming, migration and new adaptive processes in P. palinuri are unlikely. Response to past climate warming of P. palinuri provides useful indications for future scenarios in other Primula species.     

Plasticity in functional traits in the context of climate change: a case study of the subalpine forb Boechera stricta (Brassicaceae)

Environmental variation often induces shifts in functional traits, yet we know little about whether plasticity will reduce extinction risks under climate change. As climate change proceeds, phenotypic plasticity could enable species with limited dispersal capacity to persist in situ, and migrating populations of other species to establish in new sites at higher elevations or latitudes. Alternatively, climate change could induce maladaptive plasticity, reducing fitness, and potentially stalling adaptation and migration. Here, we quantified plasticity in life history, foliar morphology, and ecophysiology in Boechera stricta (Brassicaceae), a perennial forb native to the Rocky Mountains. In this region, warming winters are reducing snowpack and warming springs are advancing the timing of snow melt. We hypothesized that traits that were historically advantageous in hot and dry, low‐elevation locations will be favored at higher elevation sites due to climate change. To test this hypothesis, we quantified trait variation in natural populations across an elevational gradient. We then estimated plasticity and genetic variation in common gardens at two elevations. Finally, we tested whether climatic manipulations induce plasticity, with the prediction that plants exposed to early snow removal would resemble individuals from lower elevation populations. In natural populations, foliar morphology and ecophysiology varied with elevation in the predicted directions. In the common gardens, trait plasticity was generally concordant with phenotypic clines from the natural populations. Experimental snow removal advanced flowering phenology by 7 days, which is similar in magnitude to flowering time shifts over 2–3 decades of climate change. Therefore, snow manipulations in this system can be used to predict eco‐evolutionary responses to global change. Snow removal also altered foliar morphology, but in unexpected ways. Extensive plasticity could buffer against immediate fitness declines due to changing climates.     

Flowering phenology and reproductive fitness along a mountain slope: maladaptive responses to transplantation to a warmer climate in Campanula thyrsoides

In many biomes, global warming has resulted in advanced and longer growing seasons, which has often led to earlier flowering in plant taxa. Elevational gradients are ideal to study the effects of global warming as they allow transplantation of plants from their original cooler higher elevations down to elevations with a prospective climate. We transplanted plants from ten populations of the European alpine monocarpic herb species Campanula thyrsoides L. to three sites along a steep mountain slope (600, 1,235 and 1,850 m above sea level) in the Swiss Alps and asked whether reproductive phenology adjusts plastically to elevation and if these responses were adaptive, i.e. increased the fitness of plants. We further assessed current genetic differentiation in phenotypic traits and whether any such origin effects were due to adaptation to climatic conditions of origin. Our results showed that transplantation to lower elevations caused strong shifts in phenology, with plants starting growth and flowering earlier than plants placed at higher elevations. However, compared to flower production at high elevation, number of flowers per plant decreased 21 % at mid- and 61 % at low elevation. The shift in phenology thus came with a high cost in fitness, and we suggest that phenology is maladaptive when C. thyrsoides faces temperature conditions deviating from its natural amplitude. We conclude that the frequently reported phenological shift in plant species as a response to global warming may include heavy fitness costs that may hamper species survival.     

The effectiveness of pre‐ and post‐zygotic barriers in avoiding hybridization between two snapdragons (Antirrhinum L.: Plantaginaceae)

Reproductive barriers play an important role in the maintenance of species boundaries. However, to date, few studies have provided a detailed analysis of reproductive isolation barriers between species or examined their importance in maintaining species identity. This is the first detailed study into pre‐ and post‐zygotic reproductive isolation barriers in Antirrhinum, based on a mixed population with two species that rarely co‐occur. The study revealed that pollinator constancy and preference and poor hybrid seed viability were the most important reproductive isolating mechanisms. Reproductive isolation was practically complete by both pre‐ and post‐zygotic barriers. Average pre‐zygotic isolation was greater than post‐zygotic isolation, in accordance with the trend observed in flowering plants in which reproductive isolation is principally caused by pre‐zygotic mechanisms. However, average post‐zygotic isolation was also high, in contrast to what was expected among Antirrhinum spp. This case highlights the importance of quantifying the reproductive isolation barriers thoroughly to understand how and why species boundaries are maintained. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 159–172.     

Selection on flowering time in Mediterranean high-mountain plants under global warming

Under climate warming, plants will undergo novel selective pressures to adjust reproductive timing. Adjustment between reproductive phenology and environment is expected to be higher in arctic and alpine habitats because the growing season is considerably short. As early- and late-flowering species reproduce under very different environmental conditions, selective pressures on flowering phenology and potential effects of climate change are likely to differ between them. However, there is no agreement on the magnitude of the benefits and costs of early- vs. late-flowering species under a global warming scenario. In spite of its relevance, phenotypic selection on flowering phenology has rarely been explored in alpine plants and never in Mediterranean high mountain species, where selective pressures are very different due to the summer drought imposed over the short growth season. We hypothesized that late-flowering plants in Mediterranean mountains should present stronger selective pressures towards early onset of reproduction than early-flowering species, because less water is available in the soil as growing season progresses. We performed selection analyses on flowering onset and duration in two high mountain species of contrasting phenology. Since phenotypic selection can be highly context-dependent, we studied several populations of each species for 2 years, covering their local altitudinal ranges and their different microhabitats. Surrogates of biotic selective agents, like fruitset for pollinators and flower and fruit loss for flower and seed predators, were included in the analysis. Differences between the early- and the late-flowering species were less than expected. A consistent negative correlational selection of flowering onset and duration was found affecting plant fitness, i.e., plants that bloomed earlier flowered for longer periods improving plant fitness. Nevertheless, the late-flowering species may experience higher risks under climate warming because in extremely warm and dry years the earlier season does not bring about a longer flowering duration due to summer drought.     

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta‐analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho‐ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ¹³C). We found LMA, Narea, Nmass and δ¹³C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ¹³C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross‐species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.     

Failure of sexual reproduction found in micropropagated critically endangered plants prior to reintroduction: a cautionary tale

Micropropagation is a useful technique for ex situ multiplication and restoration of critically endangered plant species, but the sexual reproductive behaviour of micropropagated plants is seldom evaluated prior to reintroduction. We examined the critically endangered species Rulingia sp. ‘Trigwell Bridge’, with only three remaining plants known in the wild, as a model case to examine this issue. Abnormalities in micropropagated plants of this species related to four floral traits (lengths of sepals, petals and anthers and width of anthers). The number of pollen grains per flower of abnormal individuals was lower than in plants with apparently normal flowers (wild types), but not significantly so (P = 0.068). Pollen viability for the abnormal plant (0.87 ± 0.26%) was significantly lower than for the plants exhibiting wild‐type floral morphology (45.42 ± 4.47%). Experimental manipulations were used to examine the mating behaviour of normal and abnormal plants. The results showed that both male and female reproductive failure was linked to individuals exhibiting abnormal flowering attributes. Such aberrant reproductive performance in a micropropagated rare species predicates caution when using micropropagated plants in reintroduction programmes, highlighting the importance of screening for reproductive normality prior to release of micropropagated plants (especially for critically endangered species where reliance on in vitro propagation methods is often a necessity). © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 278–284.     

Plant functional traits mediate reproductive phenology and success in response to experimental warming and snow addition in Tibet

Global climate change is predicted to have large impacts on the phenology and reproduction of alpine plants, which will have important implications for plant demography and community interactions, trophic dynamics, ecosystem energy balance, and human livelihoods. In this article we report results of a 3‐year, fully factorial experimental study exploring how warming, snow addition, and their combination affect reproductive phenology, effort, and success of four alpine plant species belonging to three different life forms in a semiarid, alpine meadow ecosystem on the central Tibetan Plateau. Our results indicate that warming and snow addition change reproductive phenology and success, but responses are not uniform across species. Moreover, traits associated with resource acquisition, such as rooting depth and life history (early vs. late flowering), mediate plant phenology, and reproductive responses to changing climatic conditions. Specifically, we found that warming delayed the reproductive phenology and decreased number of inflorescences of Kobresia pygmaea C. B. Clarke, a shallow‐rooted, early‐flowering plant, which may be mainly constrained by upper‐soil moisture availability. Because K. pygmaea is the dominant species in the alpine meadow ecosystem, these results may have important implications for ecosystem dynamics and for pastoralists and wildlife in the region.     

Abundance and habitat segregation in Mediterranean grassland species: the importance of seed weight

Abstract. We analysed the relationship between seed traits (weight, shape and dispersal structures) and the abundance and habitat segregation of Mediterranean grassland species. To take into account possible correlations with other plant traits, the study also includes 5 vegetative traits (growth form, plant longevity, clonality, onset of flowering and plant size) of commonly accepted functional importance. Data were recorded for 85 species from dehesa grasslands in central Spain. Species abundance was measured in upper (dry and less productive, high stress) and lower (moist and more productive, low stress) slope zones in the same area. Habitat segregation was estimated using an index based on the relative frequencies of species in upper and lower slope zones. Multiple regression models were fitted using species, as well as phylogenetically independent contrasts, as data points. Annual small‐seeded species without specialised dispersal structures are over‐represented in dehesa grasslands. Abundance was negatively related to seed weight in upper slope zones. None of the recorded plant traits were related to abundance in the lower slope zones. Habitat segregation was mainly related to seed weight, but also to some vegetative traits. Annual, early flowering and small‐seeded species were relatively more abundant in the upper than the lower slope zones. This pattern is independent of phylogeny. Our results suggest that in dry Mediterranean grasslands, abundance of many species is determined by dispersal (production of numerous small seeds) rather than by competitive ability.     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.     

Exploring the causes of variation in phenology and morphology in Mediterranean geophytes: a genus-wide study of Cyclamen

The phenology and morphology of Mediterranean plants are constrained by drought in summer and cold temperatures in winter. In this study we examine how climatic factors and phylogenetic constraints have shaped variation in the phenology and morphology of 17 species of the genus Cyclamen cultivated in uniform garden conditions. We quantify the extent to which traits differ among subgenera and thus represent conserved traits within evolutionary lineages. We also explore whether leaf, flowering and seed-release phenology are correlated among species, and thus whether variation in flowering phenology results from selection on dispersal phenology. Our results show a significant influence of subgenus membership on leaf and flowering phenology but not on morphological traits or the timing of seed release. Among-species variation in foliage height, leaf size and seed mass (but not in floral traits) is correlated with chromosome number. Leaf traits show that species with a shorter vegetative period have a higher capacity for resource acquisition. Major phenological shifts, i.e. spring vs. autumn flowering and a decoupling of leaf and flower phenology in autumnal flowering species, thus occurred prior to the diversification of species in each subgenus and not as a response to selection on dispersal timing. Leaf and flowering phenology illustrate a gradient of strategies from autumn flowering in the absence of leaves (hysteranthous species) to spring flowering with fully developed foliage (synanthous species). In the former, flowering is uncoupled from resource acquisition by simultaneous photosynthesis, indicative that hysteranthy is a response to temporal unpredictability in the onset of rain after the summer drought. Our results support the idea that whereas leaf development is controlled primarily by moisture availability and secondarily by temperature, flowering is temperature dependent, above a minimum moisture threshold. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145 , 469–484.     

Morphological and phenological differences in Nothofagus pumilio from contrasting elevations: Evidence from a common garden

Abstract We tested the hypothesis that contrasting elevations select distinct growth patterns and vegetative phenology in Nothofagus pumilio, a winter deciduous tree that dominates mountain forests of Patagonia. Analysis of saplings maintained under common‐garden conditions for 4 years showed a significant decrease in shoot annual growth, leaf size, and a delay in bud‐break, and leaf expansion with increased elevation of their site of origin. Rapid gain in height seems to be advantageous at low elevation in such light‐demanding species. Lower stature high‐elevation plants have wider branching angles and greater branching ratios (number of branches/number of internodes) than low‐elevation plants. Compact growth at high elevation may be related to strong winds and irradiance. Plants from different elevations had distinct growth patterns during the common‐garden experiment. This could be of importance in Mediterranean‐climate areas characterized by highly unpredictable precipitation regimes. Also, liberation of growth‐suppressed seedlings may follow different environmental signals in low‐ and high‐habitats, which might explain such time‐dependent responses to optimal conditions under cultivation. While these greenhouse‐grown N. pumilio saplings showed heritable differences in plant architectural traits and leafing phenology, it was not clear how the genotypes characteristic of particular elevations would respond to longer growing seasons such as those predicted under global warming.     

Family affiliation,sex ratio and sporophyte frequency in unisexual mosses

Patterns of sex expression and sex ratios are key features of the life histories of organisms. Bryophytes are the only haploid‐dominant land plants. In contrast with seed plants, more than half of bryophyte species are dioecious, with rare sexual expression and sporophyte formation and a commonly female‐biased sex ratio. We asked whether variation in sex expression, sex ratio and sporophyte frequency in ten dioecious pleurocarpous wetland mosses of two different families was best explained by assuming that character states evolved: (1) in ancestors within the respective families or (2) at the species level as a response to recent habitat conditions. Lasso regression shrinkage identified relationships between family membership and sex ratio and sporophyte frequency, whereas environmental conditions were not correlated with any investigated reproductive trait. Sex ratio and sporophyte frequency were correlated with each other. Our results suggest that ancestry is more important than the current environment in explaining reproductive patterns at and above the species level in the studied wetland mosses, and that mechanisms controlling sex ratio and sporophyte frequency are phylogenetically conserved. Obviously, ancestry should be considered in the study of reproductive character state variation in plants. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 163–172.     

Dissecting components of flowering pattern: size effects on female fitness

Flowering synchrony is essential for plant reproductive success, especially in the case of small‐sized populations of self‐incompatible species. Closely related to synchrony, flowering intensity influences pollinator attraction and pollinator movements. Thus, a high flowering intensity may increase pollinator attraction but, at the same time, may also increase the probability of geitonogamous pollinations. Depending on the mating system, the female fitness of plants in small populations may be affected by both the positive effects of higher flowering synchrony and pollinator attraction and the negative effects of geitonogamous pollinations induced by a high flowering intensity. It was hypothesized that different‐sized plants in a population would show contrasting flowering patterns, resulting in differences in pollinator behaviour. These influences could result in differences in mating and female reproductive success. This hypothesis was tested by studying the flowering pattern of Erodium paularense (Geraniaceae), a rare and endangered endemic of central Spain. The temporal distribution of flower production was explored throughout the reproductive season, and the probability of xenogamy and geitonogamy and their relationship to plant size and fitness components were calculated. The analysis of this partially self‐compatible species showed diverse flowering patterns related to different plant sizes. Small plants produced a larger number of seeds per fruit in spite of having lower values of flowering synchrony. By contrast, large plants produced a larger number of seeds from geitonogamous pollinations. The effect of different flower displays and outcrossing rates on seed set varied throughout the season in the different groups. Our findings highlight the relevance of individual plant size‐dependent phenology on female reproductive success and, in particular, on the relationship between flowering synchrony and fitness. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156 , 227–236.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Plant reproductive strategies vary under low and high pollinator densities

Long‐term variation in the population density of honey bees Apis mellifera across landscapes has been shown to correlate with variation in the floral traits of plant populations in these landscapes, suggesting that variations in pollinator population density and foraging rates can drive floral trait evolution of their host plants. However, it remained to be determined whether this variation in plant traits is associated with adaptive variation in plant reproductive strategies under conditions of high and low pollinator densities. Here we conducted a reciprocal transplant experiment to examine how this variation in floral traits, under conditions of either high and low pollinator density, impacted seed production in the Tibetan lotus Saussurea nigrescens. In 2014 and 2015, we recorded the floral traits, pollinator visitation rates, and seed production of S. nigrescens populations grown in both home sites and foreign sites, where sites varied in honey bee population density. Our results demonstrated that the floral traits reflected those of their original population, regardless of their current location. However, seed production varied with both population origin and transplant site. Seed number was positively correlated with flower abundance in the pollinator‐rich sites, but with nectar production in the pollinator‐poor sites. Pollinator visitation rate was also positively correlated with flower number at pollinator‐rich sites, and with nectar volume at pollinator‐poor sites. Overall, the local genotype had higher seed production than nonlocal genotypes in home sites. However, when pollen is hand‐supplemented, plants from pollinator‐rich populations had higher seed production than plants from pollinator‐poor populations, regardless of whether they were transplanted to pollinator‐rich or ‐poor sites. These results suggest that the plant genotypic differences primarily drive variation in pollinator attraction, and this ultimately drives variation in seed: ovule ratio. Thus, our results suggest that flowering plant species use different reproductive strategies to respond to high or low pollinator densities.     

Plant population differentiation and climate change: responses of grassland species along an elevational gradient

Mountain ecosystems are particularly susceptible to climate change. Characterizing intraspecific variation of alpine plants along elevational gradients is crucial for estimating their vulnerability to predicted changes. Environmental conditions vary with elevation, which might influence plastic responses and affect selection pressures that lead to local adaptation. Thus, local adaptation and phenotypic plasticity among low and high elevation plant populations in response to climate, soil and other factors associated with elevational gradients might underlie different responses of these populations to climate warming. Using a transplant experiment along an elevational gradient, we investigated reproductive phenology, growth and reproduction of the nutrient‐poor grassland species Ranunculus bulbosus, Trifolium montanum and Briza media. Seeds were collected from low and high elevation source populations across the Swiss Alps and grown in nine common gardens at three different elevations with two different soil depths. Despite genetic differentiation in some traits, the results revealed no indication of local adaptation to the elevation of population origin. Reproductive phenology was advanced at lower elevation in low and high elevation populations of all three species. Growth and reproduction of T. montanum and B. media were hardly affected by garden elevation and soil depth. In R. bulbosus, however, growth decreased and reproductive investment increased at higher elevation. Furthermore, soil depth influenced growth and reproduction of low elevation R. bulbosus populations. We found no evidence for local adaptation to elevation of origin and hardly any differences in the responses of low and high elevation populations. However, the consistent advanced reproductive phenology observed in all three species shows that they have the potential to plastically respond to environmental variation. We conclude that populations might not be forced to migrate to higher elevations as a consequence of climate warming, as plasticity will buffer the detrimental effects of climate change in the three investigated nutrient‐poor grassland species.     

Experimental evidence for specialized bird pollination in the endangered South African orchid Satyrium rhodanthum and analysis of associated floral traits

The bird pollination syndrome is characterized by red, unscented flowers with dilute nectar in long nectar tubes. However, the extent to which plants with such traits actually depend on birds for seed production is seldom determined experimentally, and traits such as colour and scent production are often assessed only subjectively. We documented bird pollination and quantified floral traits in the critically endangered Satyrium rhodanthum (Orchidaceae) from mistbelt grasslands in the summer‐rainfall region of South Africa. Direct observations and motion trigger camera footage revealed amethyst sunbirds as the only pollinators, despite the presence of other potential pollinators. Experimental exclusion of sunbirds significantly reduced pollination and fruit set to near zero. Pollination success in naturally pollinated plants was close to 100% in one year, and fruit set varied from 23 to 64% in other years. Pollen transfer efficiency was 5.8%, which is lower than in related insect‐pollinated species, probably due to a tendency of birds to wipe pollinaria from their beak. Flowers of S. rhodanthum only reflect light in the red range of the spectrum, and they produce only a few aliphatic and monoterpene scent compounds at comparatively low emission rates. Nectar volume and sugar concentration varied between 2.7 and 3.7 μL and 23.7 and 25.9%, respectively. We conclude that S. rhodanthum is highly specialized for pollination by sunbirds. Colour, scent and nectar characteristics differ from insect‐pollinated Satyrium species and are consistent with those expected for bird‐pollinated flowers, and may contribute to lack of visitation by other potential long‐tongued pollinators. Habitat loss probably underlies the critically endangered conservation status of S. rhodanthum, but the specialization for pollination by a single bird species means that reproduction in this orchid is vulnerable to losses in surrounding communities of plants that subsidize the energetic requirements of sunbirds. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 177 , 141–150.     

Summer aridity rather than management shapes fitness‐related functional traits of the threatened mountain plant Arnica montana

Semi‐natural mountain grasslands are increasingly exposed to environmental stress under climate change. However, which are the environmental factors that limit plants in these grasslands? Also, is the present management effective against these changes? Fitness‐related functional traits may offer a way to detect changes in performance and provide new insights into their vulnerability to climate change. We investigated changes in performance and variability of functional traits of the mountain grassland target species Arnica montana along a climate gradient in Central German low mountain ranges. This gradient represents at its lower end climate conditions that are expected at its upper end under future climate change. We measured vegetative, generative, and physiological traits to account for multiple ways of plant responses to the environment. Using mixed effects and multivariate models, we evaluated changes in trait values among individuals as well as the variability of their populations in order to assess performance under changing summer aridity and different management regimes. Fitness‐related performance of most traits showed strongly positive associations with reduced summer aridity at higher elevations, while only specific leaf area and leaf dry matter content showed no association. This suggests a higher performance level at less arid montane sites and that the physiological traits are less sensitive to this climate change factor. The coefficient of variation of almost all traits declined steadily with decreasing site aridity. We suggest that this reduced variability indicates a lower environmental stress level for A. montana toward its environmental optimum at montane elevations, especially because the trait performance increased simultaneously. Surprisingly, management factors and habitat characteristics had only low influence on both trait performance and variability. In summary, summer aridity had a stronger effect to shape the trait performance and variability of A. montana under increased environmental stress than management and other habitat characteristics.     

Assessing Intraspecific Variation in Effective Dispersal Along an Altitudinal Gradient: A Test in Two Mediterranean High-Mountain Plants

Background

Plant recruitment depends among other factors on environmental conditions and their variation at different spatial scales. Characterizing dispersal in contrasting environments may thus be necessary to understand natural intraspecific variation in the processes underlying recruitment. Silene ciliata and Armeria caespitosa are two representative species of cryophilic pastures above the tree line in Mediterranean high mountains. No explicit estimations of dispersal kernels have been made so far for these or other high-mountain plants. Such data could help to predict their dispersal and recruitment patterns in a context of changing environments under ongoing global warming.

Methods

We used an inverse modelling approach to analyse effective seed dispersal patterns in five populations of both Silene ciliata and Armeria caespitosa along an altitudinal gradient in Sierra de Guadarrama (Madrid, Spain). We considered four commonly employed two-dimensional seedling dispersal kernels exponential-power, 2Dt, WALD and log-normal.

Key Results

No single kernel function provided the best fit across all populations, although estimated mean dispersal distances were short (<1 m) in all cases. S. ciliata did not exhibit significant among-population variation in mean dispersal distance, whereas significant differences in mean dispersal distance were found in A. caespitosa. Both S. ciliata and A. caespitosa exhibited among-population variation in the fecundity parameter and lacked significant variation in kernel shape.

Conclusions

This study illustrates the complexity of intraspecific variation in the processes underlying recruitment, showing that effective dispersal kernels can remain relatively invariant across populations within particular species, even if there are strong variations in demographic structure and/or physical environment among populations, while the invariant dispersal assumption may not hold for other species in the same environment. Our results call for a case-by-case analysis in a wider range of plant taxa and environments to assess the prevalence and magnitude of intraspecific dispersal variation.