Eight microsatellite markers for sympatric alpine shrubs,Phyllodoce aleutica and P. caerulea (Ericaceae)

Phyllodoce aleutica and Phyllodoce caerulea are sympatric alpine‐snowbed plants in northern Japan. They compete for pollinators (bumblebees) each other and the competitive situation varies depending on snow conditions. We isolated and characterized eight microsatellite loci in these species. Additionally, one of 13 primers developed for Rhododendron metternichii was available in these species. The expected heterozygosity of these nine markers ranged from 0.06 to 0.93 in P. aleutica and from 0.09 to 0.96 in P. caerulea. These markers may be useful to reveal the mating system evolution, patterns of pollen flow and the process of natural hybridization in these Phyllodoce species.     

A hybrid zone dominated by fertile F1s of two alpine shrub species, Phyllodoce caerulea and Phyllodoce aleutica, along a snowmelt gradient

In alpine ecosystems, the steep environmental gradients produced by the difference in snowmelt timing create a dynamic selective regime for alpine plants. As these gradients directly alter flowering phenology, they can affect pollen-mediated gene flow among populations of single and related species. In northern Japan, we found a hybrid zone dominated by fertile F(1)s of two alpine shrub species, Phyllodoce caerulea and P. aleutica, along a snowmelt gradient. Seed germination confirmed the fertility of F(1) hybrid, making the rarity and absence of backcross and F(2) plants puzzling. The long-term clonal perpetuation of F(1) hybrids (at least a few thousand years ago) contributes the maintenance of this unique hybrid zone. The distribution patterns of chloroplast DNA haplotypes suggest that F(1) formation might be caused by directional pollen flow between parental species along the snowmelt gradient. Based on these results, we discuss the ecological and evolutionary significance of this unique hybrid zone.     

Variations in bumble bee preference and pollen limitation among neighboring populations: comparisons between Phyllodoce caerulea and Phyllodoce aleutica (Ericaceae) along snowmelt gradients

Two alpine-snowbed shrubs, Phyllodoce caerulea and P. aleutica (Ericaceae), co-occur in locales in northern Japan with early to late snowmelt, but they have different mating systems. Phyllodoce caerulea is an obligate outcrosser in any population, whereas the selfing ability of P. aleutica is highly variable among neighboring populations along snowmelt gradients: it shows high self-compatibility in early to middle snowmelt populations but low self-compatibility in late snowmelt populations. We investigated the relationships between pollinator availability and mating systems of these species along three snowmelt gradients. Relative abundance of flowers and nectar standing crop of P. caerulea decreased from early to late snowmelt plots. Bumble bees preferred P. caerulea to P. aleutica in early and middle snowmelt plots, while their preference shifted to P. aleutica in late snowmelt plots. Pollen limitation was severe in P. aleutica in early to middle snowmelt plots but it was severe in P. caerulea in late snowmelt plots. Seed-set success under natural conditions of P. aleutica was higher than that of P. caerulea in all plots. Thus, we infer that the selfing ability of P. aleutica under pollinator limitation acts as a reproductive assurance. We conclude that the interaction through pollination between the sympatric species is strong enough to cause a phenotypic change in mating system even within a local area.     

Clarification of the genetic component of hybrids between Phyllodoce caerulea and Phyllodoce aleutica (Ericaceae) in Hokkaido,northern Japan

Natural hybridization provides great opportunities to understand the interaction of genetics and ecology in determining species boundaries. We examined the genetic relationships of Phyllodoce taxa and revealed that most hybrids were fertile F₁s and an extremely small number of backcross and no F₂ plants were established in natural conditions. Because this trend was irrespective of regions, we conclude that negative endogenous selection may act after the germination of F₁ seeds and prevent the establishment of later‐generation hybrids. Based on these results, we discuss the ecological and evolutionary significance of natural hybridization in Phyllodoce taxa.     

Flowering phenology influences seed production and outcrossing rate in populations of an alpine snowbed shrub, Phyllodoce aleutica: effects of pollinators and self-incompatibility

Background and Aims

Because of differences in snowmelt time, the reproductive phenologies of alpine plants are highly variable among local populations, and there is large variation in seed set across populations. Temporal variation in pollinator availability during the season may be a major factor affecting not only seed production but also outcrossing rate of alpine plants.

Methods

Among local populations of Phyllodoce aleutica that experience different snowmelt regimes, flowering phenology, pollinator availability, seed-set rate, and outcrossing rate were compared with reference to the mating system (self-compatibility or heterospecific compatibility with a co-occurring congeneric species).

Key Results

Flowering occurred sequentially among populations reflecting snowmelt time from mid-July to late August. The visit frequency of bumble-bees increased substantially in late July when workers appeared. Both seed set and outcrossing rate increased as flowering season progressed. Although flowers were self-compatible and heterospecific compatible, the mixed-pollination experiment revealed that fertilization with conspecific, outcrossing pollen took priority over selfing and hybridization, indicating a cryptic self-incompatibility. In early snowmelt populations, seed production was pollen-limited and autogamous selfing was common. However, genetic analyses revealed that selfed progenies did not contribute to the maintenance of populations due to late-acting inbreeding depression.

Conclusions

Large variations in seed-set and outcrossing rates among populations were caused by the timing of pollinator availability during the season and the cryptic self-incompatibility of this species. Despite the intensive pollen limitation in part of the early season, reproductive assurance by autogamous selfing was not evident. Under fluctuating conditions of pollinator availability and flowering structures, P. aleutica maintained the genetic composition by conspecific outcrossing.Key words: Alpine snowbed, autogamy, bumble-bee, cryptic self-incompatibility, flowering phenology, mixed pollination, outcrossing rate, Phyllodoce aleutica, pollination success, seasonality, self-pollination     

Performance of two alpine plant species along environmental gradients in an alpine meadow ecosystem in central Tibet

Understanding how biotic interactions and abiotic conditions affect plant performance is important for predicting changes in ecosystem function and services in variable environments. We tested how performances of Astragalus rigidulus and Potentilla fruticosa change along gradients of biotic interactions (represented by plant species richness, abundance of the dominant plant species Kobresia pygmaea, and herbivory intensity) and abiotic conditions (represented by elevation, aspect, and slope steepness) across a semi-arid landscape in central Tibet. Redundancy analyses showed that the biotic variables explained 30 and 39 % of the variation in overall performance of A. rigidulus (P = 0.03) and P. fruticosa (P = 0.01), respectively. Abiotic variables did not contribute significantly to variation among A. rigidulus populations. Plant size decreased with species richness in both species and was larger on south- rather than north-facing slopes. Reproductive effort for both species was significantly negatively related to the abundance of K. pygmaea and both species had larger reproductive effort on south- rather than north- and west-facing slopes. The proportion of biomass allocated to sexual reproduction in P. fruticosa was negatively correlated with K. pygmaea abundance and herbivory intensity. The population density of P. fruticosa was positively related to elevation, species richness, and K. pygmaea abundance. We conclude that plant performance at a local scale was more strongly related to biotic than abiotic conditions, but different components of plant performance responded differently to predictor variables and the responses were species-specific. These findings have important implications for rangeland management under changing environmental conditions.     

Competition and evolution along environmental gradients: patterns, boundaries and sympatric divergence

The present study examined how competitive interactions and environmental conditions generate species boundaries and determine species distributions. A spatially explicit, quantitative genetic, two-species competition model was used to manipulate the strengths of competition, gene flow and local adaptation along environmental gradients. This allowed us to assess the long-term persistence of each species and whether the ranges they inhabited had boundaries in space or were unlimited. We found that a species boundary arises along less steep environmental gradients when the strength of stabilizing selection and diversifying selection are similar. We also found that a species boundary may arise along shallow environmental gradients if interspecific competition is more intense than intraspecific, which relaxes previous requirements for steep gradients for generating range limits. We determined an analytical form for the critical environmental gradient as a function of ecological and genetic parameters at which a species boundary is expected to arise by competition. Results suggest an alternative to resource competition as an explanation for phenotypic divergence between sympatric competitors. Competitors sharing a trait that is under stabilizing selection along an environmental gradient may segregate spatially and evolve in different regions, with phenotypic sympatric divergence reflecting the resulting clines. Along various types of environmental gradients, variation in stabilizing selection intensities could lead to contrasting patterns in the distribution of species. For stabilizing selection strengths in accord with field data estimates, this study predicts that the level of sympatric character divergence would be limited along environmental gradients.     

Landscape genetics of alpine-snowbed plants: comparisons along geographic and snowmelt gradients

The genetic structure of three snowbed-herb species (Peucedanum multivittatum, Veronica stelleri, and Gentiana nipponica) was analyzed using allozymes across nine populations arranged as a matrix of three snowmelt gradients x three geographic locations within 3 km in the Taisetsu Mountains, northern Japan. Phenologically asynchronous populations are packed within a local area in alpine snowbeds, because flowering season of alpine plants depends strongly on the timing of snowmelt. Moderate genetic differentiation was detected among local populations in every species (FST=0.03-0.07). There was a significant correlation between the geographic distance and genetic distance in the P. multivittatum populations, but not in the V. stelleri and G. nipponica populations. On the other hand, a significant correlation between the phenological distance caused by snowmelt timing and genetic distance was detected in the V. stelleri and G. nipponica populations, but not in the P. multivittatum populations. The snowmelt gradient or geographic separation influenced hierarchical genetic structure of these species moderately (FRT <0.04). Restriction of gene flow due to phenological separation and possible differential selection along the snowmelt gradient may produce genetic clines at microgeographic scale in these species.     

Seasonal changes in pollen limitation and femaleness along the snowmelt gradient in a distylous alpine herb,Primula modesta

Flowering phenology of alpine plants is strongly determined by the timing of snowmelt, and the conditions of pollination of widely distributed plants vary greatly during their flowering season. We examined the reproductive success of the distylous alpine herb, Primula modesta, along the snowmelt gradient under natural conditions, and compared it with the result of artificial pollination experiments. In addition, the compositions and visit frequencies of pollinators to the flower of P. modesta were examined during the flowering period. The pin and thrum plants of P. modesta growing at the same site have an equal ability to produce seeds if a sufficient amount of legitimate pollen grains are deposited on the stigma surface. However, under natural conditions, their seed‐set success was often (even if not always) restricted by pollen limitation, and the functional gender of the pin and thrum plants biased to the female and male, respectively, associated with their growing sites. These variations were not ascribed to resource limitation nor biased morph ratio but to the seasonal changes in pollination situations, a replacement of pollinator types from long‐ to short‐tongued pollinators resulted in unidirectional pollen transfer from long stamens (thrum plants) to long styles (pin plants). The functional gender specialization may enhance the evolution of dioecy from heterostyly, but the severe pollen limitation may cause the breakdown of heterostyly into homostyly. To consider the evolutionary pathway of heterostylous plants, an accumulation of the empirical data is required demonstrating how phenological synchrony between plants and pollinators is decided and to what degree this relationship is stable over years, along with estimates of selection and gene flow in individual plants.     

Direct and indirect control by snow cover over decomposition in alpine tundra along a snowmelt gradient

We assessed direct and indirect effects of snow cover on litter decomposition and litter nitrogen release in alpine tundra. Direct effects are driven by the direct influence of snow cover on edaphoclimatic conditions, whereas indirect effects result from the filtering effect of snow cover on species’ abundance and traits. We compared the in situ decomposition of leaf litter from four dominant plant species (two graminoids, two shrubs) at early and late snowmelt locations using a two-year litter-bag experiment. A seasonal experiment was also performed to estimate the relative importance of winter and summer decomposition. We found that growth form (graminoids vs. shrubs) are the main determinants of decomposition rate. Direct effect of snow cover exerted only a secondary influence. Whatever the species, early snowmelt locations showed consistently reduced decomposition rates and delayed final stages of N mineralization. This lower decomposition rate was associated with freezing soil temperatures during winter. The results suggest that a reduced snow cover may have a weak and immediate direct effect on litter decomposition rates and N availability in alpine tundra. A much larger impact on nutrient cycling is likely to be mediated by longer term changes in the relative abundance of lignin-rich dwarf shrubs.     

Biomass and diversity of dry alpine plant communities along altitudinal gradients in the Himalayas

A non-linear relationship between phytodiversity and altitude has widely been reported, but the relationship between phytomass and altitude remains little understood. We examined the phytomass and diversity of vascular plants along altitudinal gradients on the dry alpine rangelands of Ladakh, western Himalaya. We used generalized linear and generalized additive models to assess the relationship between these vegetation parameters and altitude. We found a hump-shaped relationship between aboveground phytomass and altitude. We suspect that this is engendered by low rainfall and trampling/excessive grazing at lower slopes by domestic livestock, and low temperature and low nutrient levels at higher slopes. We also found a unimodal relationship between plant species-richness and altitude at a single mountain as well as at the scale of entire Ladakh. The species-richness at the single mountain peaked between 5,000 and 5,200 m, while it peaked between 3,500 and 4,000 m at entire Ladakh level. Perhaps biotic factors such as grazing and precipitation are, respectively, important in generating this pattern at the single mountain and entire Ladakh.     

Effects of disturbance on plant regrowth along snow pack gradients in alpine habitats

Human disturbance in alpine habitats is expected to increase, and improved knowledge of short-term recovery after disturbance events is necessary to interpret vegetation responses and formulate planning and mitigation efforts. The ability of a plant community to return to its original state after a disturbance (community resilience) depends on species composition and environmental conditions. The aim of this study is to analyze initial short-term effects of disturbance in alpine plant communities in contrasting climates (oceanic vs. continental; central Norway). We used a nested block-design to examine vegetative regrowth and seedling recruitment after experimental perturbation. Three plant community types along the snow pack gradient were exposed to (1) no disturbance, (2) clipping, and (3) clipping and uprooting. Slow vegetative regrowth and low seedling establishment rates were found in dry alpine ridges and late-melting oceanic snowbed communities. Leeside habitats with intermediate snow conditions were found more resilient. The difference was related to growth form and species diversity. Woody species, which dominated in ridges and oceanic snowbeds, showed the most negative response to disturbance. Species-rich plant communities dominated by graminoids and herbs showed higher rates of regrowth. Species richness seems to cause resilience to the plant communities through higher response diversity. Plant communities at the extreme ends of abiotic gradients, ridges and late-melting snowbeds, will be most sensitive to both disturbance and environmental change. In an up-scaled human-used landscape disturbance effects will be amplified and further limit recovery to a pre-disturbance state.     

Warming effects on shoot developmental growth and biomass production in sympatric evergreen alpine dwarf shrubs Empetrum nigrum and Loiseleuria procumbens

Effects of experimental warming on shoot developmental growth and biomass production were preliminarily investigated in two evergreen dwarf shrubs Empetrum nigrum and Loiseleuria procumbens, using the International Tundra Experiments open-top chamber (OTC) method, in the Tateyama Range, central Japan. An OTC was installed over shrub (E. nigrum and L. procumbens) -dominated vegetation and over shrub-forb (such as Anemone narcissiflora var. nipponica and Solidago virga-aurea ssp. leiocarpa) mixed vegetation, and stem samples of the evergreen shrubs were obtained at 26 months after installing the OTC. The OTC increased the daily mean temperature by 0.1°C to 1.8°C, on average, during the growing season. Shoot developmental growth and biomass production were considerably different between species of different vegetation types. The boreal species E. nigrum generally showed better growth inside the OTC than the arctic and subarctic species L. procumbens. Both species showed significantly larger shoot elongation and biomass production inside the OTC over shrub-dominated vegetation, whereas smaller or reduced growth was detected inside the OTC over shrub-forb mixed vegetation. The variations of growth responses to warming between species of different vegetation types are discussed, especially in relation to interspecific competition under a simulated environmental change.     

Evergreen alpine shrubs have high freezing resistance in spring,irrespective of snowmelt timing and exposure to frost: an investigation from the Snowy Mountains,Australia

Over winter, alpine plants are protected from low-temperature extremes by a blanket of snow. Climate change predictions indicate an overall reduction in snowpack and an earlier thaw; a situation which could expose the tips of shrubs which extend above the snowpack to freezing events in early spring, and cause foliar frost damage during the onset of physiological activity. We assessed the photosynthetic responses of freezing-damaged shrub leaves from an assay of freezing temperatures in the Snowy Mountains in south-eastern Australia, using chlorophyll fluorometery ex situ. We sampled leaves that were exposed early during the spring thaw and leaves that were buried in snow for up to two extra weeks, from four evergreen shrub species at monthly intervals following the period of snowmelt. Freezing resistance (estimated from LT₅₀) was poorest at the earliest spring sampling time, in both exposed above-snow and protected below-snow foliage in all species. Protected foliage in early spring had lower freezing resistance than exposed foliage, but not significantly so. By the third sampling time, freezing resistance was significantly better in the lower protected foliage (LT₅₀ of ? 14) compared with the upper exposed foliage (LT₅₀ of ? 10) in one species. Over the course of spring, freezing resistance improved significantly in all species, with LT₅₀ values of between ? 10 and ? 15 °C by the third sampling time, which is lower than the minimum air temperatures recorded at that time (> ? 5 °C). The results indicate that the dominant evergreen shrub species in this area may only be susceptible to freezing events very early in spring, before a period of frost-hardening occurs after snowmelt. Later in spring, these alpine shrubs appear frost hardy, thus further perpetuating the positive feedbacks surrounding shrub expansion in alpine areas.     

Growth-ring variations of dwarf shrubs reflect regional climate signals in alpine environments rather than topoclimatic differences

Aim Our main aim is to determine if ring‐width variations in Empetrum hermaphroditum reflect regional or local topoclimate signals in an alpine environment. In the case that topoclimate provides the dominant signal, a secondary aim is to link these to spatial distribution patterns of different vegetation types. Location The study area is situated in the middle alpine belt in the Vågåmo region, Central Norwegian Scandes. Sampling sites cover different topoclimates: ridges, north‐facing slopes and south‐facing slopes. Methods We constructed ring‐width chronologies of E. hermaphroditum for each type of microsite for the common period 1951–2004. Climate data were prepared on an hourly, daily and growing‐season time scale. Climate–growth relationships were evaluated using bivariate correlations and regression tree methods for continuous time‐series analyses. In addition, extreme growth anomalies (pointer years) were compared with the climate conditions in those years. The impact of water supply on wood anatomy was determined by correlating the conductive area (percentage of vessel per growth ring) with a running mean (sum) of 10‐day intervals for temperature and precipitation. Results This study indicates that mean summer (June–August) temperatures determine the width of the growth rings of E. hermaphroditum irrespective of topoclimate. The length of the growing season, which is the most differentiating climatic factor between microsites, does not substantially alter the anatomical ring structure. Microsite differences in mean growth rates are attributed to the higher frequency of warm days. Extremely warm days limit ring‐width development at south‐facing slopes, while plants at ridges and north‐facing slopes still benefit from higher temperatures. As a consequence, pointer years are not developed synchronously at all microsites. Vessel formation is affected by available moisture, especially in the later part of the growing season. Main conclusions Topoclimate induces slight modifications of annual growth‐ring increments of E. hermaphroditum at different microsites. In contrast to the distribution patterns of vegetation types that are determined by snow cover, growth‐ring variations are related to summer temperature conditions, and the prominent regional climate signal is still reflected at all microsites. This offers the opportunity to reconstruct climatic change in alpine regions from dwarf shrub ring‐width chronologies.     

Arthropod community composition along snowmelt gradients in snowbeds in the Snowy Mountains of south‐eastern Australia

Environmental gradients drive variation in community composition across a range of spatial scales. In alpine regions, areas of long‐lasting snow (‘snow patches’) create snowmelt gradients that drive considerable change in vegetation structure and composition over small spatial scales. This study examined whether there is parallel variation in arthropod communities using snowmelt gradients in the Australian Alps. Mites (Acarina) were the most common arthropods in snow patches, followed by springtails while, among the insects, the orders Hymenoptera (primarily Formicidae), Diptera, Coleoptera (primarily Carabidae) and Hemiptera (primarily Cicadellidae) dominated. Along the snowmelt gradient, arthropod assemblages changed from having equal proportions of predators and herbivores in early‐melting zones to being predator‐dominated in late‐melting zones, particularly early in the growing season. This followed a transition in vegetation cover and composition and was driven by higher numbers of predacious carabid beetles in later‐melting zones. Overall, however, our results suggest that snowbed arthropod communities in the Australian alpine zone are more sensitive to short‐term effects, such as time since snowmelt, than to differences in vegetation structure and composition or long‐term patterns of snowmelt. Continued advancement of snowmelt timing due to warmer spring temperatures is therefore likely to have more impact on the seasonality of snowbed arthropod communities than on the overall community composition.     

Evidence for divergence between sympatric stone charr and Dolly Varden along unique environmental gradients in Kamchatka

Here, we describe the biological and ecological differences between partially anadromous Dolly Varden and riverine stone charr distributed in the Kamchatka River middle course. Endemic stone charr, being the ambush predator, is defined by the accelerated growth and prolonged lifespan, robust body, and large mouth, as well as specific marble coloration since an early age. The significant restriction in gene flow between the stone charr and benthos-eating Dolly Varden was supported by an allelic distribution of eight microsatellite loci. Herewith, the identity of mitochondrial DNA control region and intron sequences of growth hormone and metallothionein genes confirms a relatively recent diversification of the groups within the single basin. We suggest the key role of the specific environment for reproduction isolation of stone charr. It occupies coniferous taiga zone with dark-colored water, whereas Dolly Varden spawns uppercourse in the zone of stunted, windblown forest. Conifer litter decomposition products could drive natural selection of stone charr via metabolism acceleration and resistance to toxicants. The metabolic difference could promote piscivory and development of the specific morphological peculiarities of the stone charr. Therefore, we present the first evidence in favor of trophic-based sympatric diversification in a river basin at high latitudes.