Contrasting levels of β-diversity and underlying phylogenetic trends indicate different paths to chemical diversity in highland and lowland willow species

Diverse specialised metabolites contributed to the success of vascular plants in colonising most terrestrial habitats. Understanding how distinct aspects of chemical diversity arise through heterogeneous environmental pressures can help us understand the effects of abiotic and biotic stress on plant evolution and community assembly. We examined highland and lowland willow species within a phylogenetic framework to test for trends in their chemical α-diversity (richness) and β-diversity (variation among species sympatric in elevation). We show that differences in chemistry among willows growing at different elevations occur mainly through shifts in chemical β-diversity and due to convergence or divergence among species sharing their elevation level. We also detect contrasting phylogenetic trends in concentration and α-diversity of metabolites in highland and lowland willow species. The resulting elevational patterns contribute to the chemical diversity of willows and suggest that variable selective pressure across ecological gradients may, more generally, underpin complex changes in plant chemistry.     

Contrasting patterns of elevational zonation for birds and mammals in the Andes of southeastern Peru

Abstract. To determine the generality of avian diversity patterns, we investigated patterns of elevational zonation shown by birds and mammals along the eastern slope of the Andes Mountains in southeastern Peru. The strong environmental gradient sampled, entirely within Peru's Manu National Park and Biosphere Reserve, supports highly diverse faunas. Elevational distributions of 901 bird species, 129 bat species, and twenty-eight species of native mice exhibit contrasting patterns in species richness, species composition, and species turnover. Birds and bats showed smooth declines of species richness with elevation, whereas the richness of mouse assemblages was unrelated to elevation. For all three groups, the greatest differences were between lowland and highland faunas, although cutoff points for this contrast varied among groups (≈ 500 m for birds, 750 m for bats, and 1000 m for mice). Differences in composition also separated bird and bat faunas on either side of c. 1400 m (the boundary between montance forest and cloud forest); for mice, this faunal transition may take place nearer to 2000 m. Bird and bat faunas lacked the more discrete zonations suggested for mouse assemblages, as indicated by elevational range profiles and nested subset analyses. Distinct highland assemblages are apparent in two-dimensional histograms of range limits of birds and mice, but not for bats. Highland bat species occupy broader elevational ranges than lowland bat species, but for both birds and mice, species at intermediate elevations had the broadest amplitudes. Finally, clumping of range maxima and minima along the gradient identified zones of pronounced species turnover in each group, but these were generally not strongly associated with the locations of ecotones. Differences in zonation of these groups appear to reflect their different biological attributes and phylogenetic histories. Such differences obviously complicate discussions of ‘general’ diversity patterns, and limit the usefulness of birds to forecast or predict diversity patterns in other more poorly known groups—other groups may show elevated diversity and endemism in areas where avian diversity patterns appear unremarkable. The pronounced contrasts between bats and mice, and the generally intermediate character of avian patterns, suggest that future analyses might profitably partition birds into finer, more homogeneous groups of historically and/or ecologically similar species. Group differences in zonation may ultimately prove explicable with information on both species-abundance patterns and resource distributions.     

The linyphiid fauna of eastern Africa (Araneae: Linyphiidae)—distribution patterns, diversity and endemism

Distributions, endemism and diversity among East African linyphiids are analysed and discussed in relation to other forest organisms and the environmental history of eastern African. A total of 231 species are reported from eastern Africa, of which 14 are confined to the Afroalpine region and 114 species to moist forests. Only 12 of the latter are widely distributed. The rest are only known from one or two localities. Information on habitats and distributions of all species is tabulated. Few species are shared between East African mountains and there are no detectable gradients of species diversity between mountains. There is, however, a gradient of decreasing species diversity from high latitudes to the Equator. Vicariance patterns are demonstrated for Elgonia, Ophrynia and Callitrichia in the Eastern Arc Mountains, Tanzania—areas that also hold the highest degree of endemism (> 80% on individual mountains) among linyphiids. The many endemic species on nearby mountains suggest that intermontane dispersal (ballooning) is rare or non-existent. There is no evidence for a distinction between highland and lowland linyphiid faunas, but altitudinal segregation of single species is demonstrated. The question of the reality of highland and lowland faunas cannot be solved by studying the altitudinal distribution of single species. Phylogenetic relationships must be taken into consideration to determine where sister-groups/species are located (lowland or highland).     

Parapatry and niche complementarity of Peruvian Desert geckos (Phyllodactylus): the ambiguous role of competition

The Sechura Desert of Peru is among the most arid, barren regions of South America. Four species of nocturnal geckos (Phyllodactylus) are parapatric in part of the desert. By comparing niche associations of these species in allopatry and parapatry, I attempt to determine whether the observed parapatric distributions and niche dimension complementarity are related to competition — as is frequently assumed. While parapatry suggests a role for competition, distributional patterns can alternatively be related to adaptations of geckos to different physical environments (sandy desert and rocky foothill) that abut in the study area. Niche complementarity might also be a result of competition, but potentially contradictory evidence suggests that niche complementarity might instead be the result of adaptations developed in allopatry and having no relationship to competition. The ambiguity of these interpretations sets limits on the significance of this kind of evidence: in the absence of attempts to falsify alternative explanations, observations of parapatry or of niche dimension complementarity do not demonstrate conclusively the impact of competition as a force structuring communities.     

Examining variation in the leaf mass per area of dominant species across two contrasting tropical gradients in light of community assembly

Understanding variation in key functional traits across gradients in high diversity systems and the ecology of community changes along gradients in these systems is crucial in light of conservation and climate change. We examined inter‐ and intraspecific variation in leaf mass per area (LMA) of sun and shade leaves along a 3330‐m elevation gradient in Peru, and in sun leaves across a forest–savanna vegetation gradient in Brazil. We also compared LMA variance ratios (T‐statistics metrics) to null models to explore internal (i.e., abiotic) and environmental filtering on community structure along the gradients. Community‐weighted LMA increased with decreasing forest cover in Brazil, likely due to increased light availability and water stress, and increased with elevation in Peru, consistent with the leaf economic spectrum strategy expected in colder, less productive environments. A very high species turnover was observed along both environmental gradients, and consequently, the first source of variation in LMA was species turnover. Variation in LMA at the genus or family levels was greater in Peru than in Brazil. Using dominant trees to examine possible filters on community assembly, we found that in Brazil, internal filtering was strongest in the forest, while environmental filtering was observed in the dry savanna. In Peru, internal filtering was observed along 80% of the gradient, perhaps due to variation in taxa or interspecific competition. Environmental filtering was observed at cloud zone edges and in lowlands, possibly due to water and nutrient availability, respectively. These results related to variation in LMA indicate that biodiversity in species rich tropical assemblages may be structured by differential niche‐based processes. In the future, specific mechanisms generating these patterns of variation in leaf functional traits across tropical environmental gradients should be explored.     

Ecological correlates of the distribution limits of two poeciliid species along a salinity gradient

Identifying the environmental factors responsible for the formation of a species' distribution limit is challenging because organisms interact in complex ways with their environments. However, the use of statistical niche models in combination with the analysis of phenotypic variation along environmental gradients can help to reduce such complexity and identify a subset of candidate factors. In the present study, we used such approaches to describe and identify factors responsible for the parapatric distribution of two closely‐related livebearer fish species along a salinity gradient in the lowlands of Trinidad, West Indies. The downstream distribution limits of Poecilia reticulata were strongly correlated with the brackish–freshwater interface. We did not observe significant phenotypic variation in life‐history traits for this species when comparing marginal with more central populations, suggesting that abrupt changes in conditions at the brackish–freshwater interface limit its distribution. By contrast, Poecilia picta was present across a wide range of salinities, although it gradually disappeared from upstream freshwater localities. In addition, P. picta populations exhibited an increase in offspring size in localities where they coexist with P. reticulata, suggesting a role for interspecific competition. The parapatric distribution of these two species, suggests that P. reticulata distributions are limited by an abiotic factor (salinity), whereas P. picta is limited by a biotic factor (interspecific competition). Similar parapatric patterns have been previously described in other systems, suggesting they might be a common pattern in nature. © 2013 The Linnean Society of London     

Growth characteristics of populations of Tibetan origin in Nepal.

Previous growth studies of highland-dwelling populations in the ecologically diverse areas of Peru and Ethiopia have yielded highly varied results: the retarded growth of the Peruvian sample was attributed to the effects of hypoxia, while the increased height and weight of the highland Ethiopian sample could be traced to better health conditions in the highland village than in the lowland village studied. In an attempt to provide a basis for evaluating studies of growth at high altitude, the present study compared Sherpa children living in the Everest region of Nepal with Tibetan children living in Kathmandu. It was found that: (1) the growth of Sherpa and Tibetan children is considerably retarded compared to other high altitude populations; (2) despite conditions favorable for optimum growth among the Tibetans, their growth resembled that of the Sherpas and (3) increased chest circumference, which seems to reflect a developmental acclimatization to hypoxia among Peruvian high-altitude natives, was not seen among the Sherpas.     

Genetic divergence and speciation in lowland and montane peruvian poison frogs

Amazonia is famous for high biodiversity, and the highlands of the transition zone between the Andes and the lowlands of the Amazon basin show particularly high species diversity. Hypotheses proposed to explain the high levels of diversity in the highlands include repeated parapatric speciation across ecological gradients spanning the transition zone, repeated allopatric speciation across geographic barriers between the highlands and lowlands, divergence across geographic barriers within the transition zone, and simple lineage accumulation over long periods of time. In this study, we investigated patterns of divergence in frogs of the genus Epipedobates (family Dendrobatidae) using phylogenetic and biogeographic analyses of divergence in mitochondrial DNA (1778 aligned positions from genes encoding cyt b, 12S and 16S rRNA for 60 Epipedobates and 11 outgroup specimens) and coloration (measured for 18 specimens representing nine species in Epipedobates). The majority of phenotypic and species diversity in the poison frog genus Epipedobates occurs in the transition zone, although two morphologically conserved members of the genus are distributed across the lowlands of the Amazon basin. Phylogenetic analysis reveals that there is a single highland clade derived from an ancestral colonization event in northern Peru by a population of lowland ancestry. Epipedobates trivittatus, a widespread Amazonian species, is a member of the highland clade that reinvaded the lowlands. Comparative analyses of divergence in coloration and mtDNA reveals that divergence in coloration among populations and species in the highlands has been accelerated relative to the lowlands. This suggests a role for selection in the divergence of coloration among populations and species.     

Environmental factors determining the distribution of highland plants at low-altitude algific talus sites

Algific talus is a micro-scale habitat type where highland plants (subalpine and alpine species) are found, disjunct from their typical range, in lowland forests. On algific talus, cold airflows from the interstices between talus fragments create a local microclimate colder than surrounding forests. Despite of the widely-known occurrence of unique vegetation on algific talus, critical environmental factors determining the distribution of highland species in this habitat type are unclear. In order to reveal the environmental factors enabling highland species to inhabit algific talus, we investigated the vegetation and environments of 26 algific talus sites and four reference (non-algific talus) sites in Hokkaido, northern Japan. Several algific talus sites were dominated by highland species, while some algific talus sites and all non-algific talus sites were dominated by lowland species. Community analysis based on detrended correspondence analysis (DCA) and canonical corresponding analysis (CCA) revealed that the algific talus sites dominated by highland species had lower ground temperature, more acidic soil, larger canopy openness, and less diverse vegetation than the sites dominated by lowland species. Highland plants might be maintained under conditions stressful for lowland plants, resulting in less competitive situation. Generalized linear models (GLM), used to evaluate the response of individual highland species to environmental factors, revealed that preferable environmental conditions for highland plants are highly species specific. These results indicate that the maintenance of diverse environments is crucial for the conservation of the unique vegetation and local populations of highland species in algific talus areas.     

The role of climate for the range limits of parapatric European land salamanders

Abrupt range limits of parapatric species may serve as a model system to understand the factors that determine species’ range borders. Theory suggests that parapatric range limits can be caused by abiotic conditions along environmental gradients, biotic interactions or a combination of both. Geographic ranges of the parapatric salamanders, Salamandra salamandra and S. atra, meet in small contact zones in the European Alps and to date, the cause of parapatry and the restricted range of S. atra remain elusive. We combine multivariate approaches and climatic data analysis to explore niche differentiation among the two salamanders with respect to the available climatic environment at their contact zones. Our purpose is to evaluate whether climatic conditions explain the species’ sharp range limits or if biotic interactions may play a role for range delimitation. Analyses were carried out in three contact zones in Switzerland to assess possible geographic variation. Our results indicate that both species occur at localities with different climatic conditions as well as the presence of a strong climatic gradient across the species’ range limits. Although the species’ climatic niches differ moderately (with a wider niche breadth for S. atra), interspecific niche overlap is found. Comparisons among the contact zones confirm geographic variation in the species’ climatic niches as well as in the conditions within the geographically available space. Our results suggest that the change in climatic conditions along the recognized gradient represents a determining factor for species’ range limits within contact zones. However, our analyses of geographic variation in climatic conditions reveal that both salamander species can occur in a much wider range of conditions than observed within contact zones. This finding and the interspecific climatic niche overlap within each contact zone provides indirect evidence that biotic interactions (likely competition) between the two species may also determine their range limits.     

Asymmetric interspecific aggression in New Guinean songbirds that replace one another along an elevational gradient

A common pattern in tropical avifaunas is for closely related species to inhabit largely parapatric elevational distributions such that they replace one another along the elevational gradient. A long‐standing hypothesis for this pattern is that parapatry is maintained by interspecific interference competition mediated by interspecific aggression. However, empirical tests of this hypothesis remain scarce. We used reciprocal playback experiments to measure interspecific aggression in five species‐pairs of New Guinean passerine elevational replacements. We found evidence of interspecific aggression in three species‐pairs. In these three cases, interspecific aggression was asymmetric, with the lower elevation species more aggressive towards the upper elevation species than vice versa. Two patterns suggest that this interspecific aggression is a learned response to the presence of a heterospecific competitor rather than misdirected intraspecific aggression or an evolved response to a competitor. First, when present, interspecific aggression was always strongest at the upper elevation range margin of the lower elevation species (i.e. in the elevational zone in which the two species were found in close proximity and thus interacted with each other), and diminished over very short distances away from this zone. Secondly, the two species‐pairs that did not exhibit interspecific aggression had narrow ‘no man's land’ gaps between their elevational distributions such that heterospecifics did not encounter one another, possibly explaining the lack of interspecific aggression in these examples. Our results support the hypothesis that interspecific aggression is one factor influencing elevational limits in species‐pairs of New Guinean elevational replacements.     

Comparative ecology of a lowland and a subalpine species of Mnium in the northern Rocky Mountains

Factors that set the altitudinal limits of plants have been relatively well explored for many land plant groups, but not for bryophytes. Bryophytes typically represent a significant portion of alpine floras with many species specific to highland systems. Differences between highland and lowland bryophytes have been underinvestigated. In the present study spanning three growing seasons, a subalpine and a lowland moss were both reciprocally planted as apical fragments and transplanted as adults between sites at 1400 m and 2000 m in the Front Ranges of the Rocky Mountains, Alberta. The lowland species, Mnium spinulosum, was less tolerant of conditions at 2000 m than the subalpine species, M. arizonicum, was to conditions at 1400 m. In particular, M. spinulosum had lower establishment from both apical fragments and spores at higher elevation sites. Both species had significantly lower establishment during the abnormally cold growing season of 1999, but fragments of M. arizonicum were better able to adjust their investment in establishment. The effect of a dominant feather moss, Hylocomium splendens, on establishment and transplant health was tested for M. arizonicum. Establishment of M. arizonicum was lower in Hylocomium mats than on bare humus regardless of site elevation suggesting allelopathy; however, stem survival in adult transplants was higher in Hylocomium mats than in Mnium dominated microsites at the higher elevation suggesting facilitation. Competition, rather than a lack of physiological plasticity, probably determines the lower elevation limit of the subalpine moss, while poor establishment ability at low temperatures accounts for the upper elevation limit of the montane moss. This revised version was published online in June 2006 with corrections to the Cover Date.     

Elevational variation in diversity and composition of land-snail faunas in a Tanzanian forest

We sampled terrestrial molluscs at fifteen elevations between 400 and 2000 m in Mwanihana Forest Reserve, Tanzania and recorded 84 taxa. Four diversity measures – species density (D_P), species richness (D₂₀) and observed (S_O) and interpolated (S_I) alpha diversity – were measured; beta diversity and abundance were also investigated. Mean elevational range was 470 m. D_P, D₂₀, alpha diversity and mollusc abundance increased with elevation, but alpha diversity peaked at 1695 m (S_O) or 1500 m (S_I) and declined at higher elevations. Maximum beta diversity was at 1000 m. Soil pH was negatively correlated with elevation and D_P. Cluster analysis divided the sites into lowland (400–900 m) and highland (>1000 m) groups. Axis 1 of a canonical correspondence analysis was associated with altitude and suggested a faunal discontinuity at 1000 m. Variation within the highland (>1000 m) and lowland faunal sets was related to elevation and forest structure respectively. The findings indicate that mollusc diversity peaks at intermediate elevations. This may be related to the combined effects of low rainfall conditions at low elevations and increasing effects of soil leaching at high elevations, both of which may limit mollusc diversity and abundance. Diversity at intermediate altitudes may be further elevated because of faunal mixing of lowland and highland groups.     

Condition-specific competition governs the geographic distribution and diversity of ectoparasites

Understanding how environmental parameters interact to govern species distributions is a shared goal of ecology and biogeography. Biotic and abiotic conditions can change distributions by affecting the nature of interspecific interactions. Although documented in free-living systems, this context dependency has been neglected in parasite interactions. We investigated the influence of condition-specific competition on the specificity of two species of feather lice (Phthiraptera: Ischnocera) that share a host, the mourning dove (Zenaida macroura). We show that relative humidity restricts the range of one species, Columbicola macrourae3 (i.e., the C. macrourae lineage found on mourning doves), to the more humid eastern United States. The second species, Columbicola baculoides, an arid-adapted species, is restricted to drier regions of the western United States by C. macrourae3, which outcompetes it in experiments. Thus, arid conditions in the West provide C. baculoides with a climatic refuge from the competitively superior C. macrourae3, effectively doubling parasite diversity on one host species. These results support the hypothesis that abiotic factors can determine species distributions on the stressful end of an environmental gradient while interspecific competition governs distributions at the benign end. The balance between these factors is subject to change as environmental conditions change, even if the host distribution remains unaffected.     

Infection with Batrachochytrium dendrobatidis is common in tropical lowland habitats: Implications for amphibian conservation

Numerous species of amphibians declined in Central America during the 1980s and 1990s. These declines mostly affected highland stream amphibians and have been primarily linked to chytridiomycosis, a deadly disease caused by the chytrid fungus Batrachochytrium dendrobatidis (Bd). Since then, the majority of field studies on Bd in the Tropics have been conducted in midland and highland environments (>800 m) mainly because the environmental conditions of mountain ranges match the range of ideal abiotic conditions for Bd in the laboratory. This unbalanced sampling has led researchers to largely overlook host–pathogen dynamics in lowlands, where other amphibian species declined during the same period. We conducted a survey testing for Bd in 47 species (n = 348) in four lowland sites in Costa Rica to identify local host–pathogen dynamics and to describe the abiotic environment of these sites. We detected Bd in three sampling sites and 70% of the surveyed species. We found evidence that lowland study sites exhibit enzootic dynamics with low infection intensity and moderate to high prevalence (55% overall prevalence). Additionally, we found evidence that every study site represents an independent climatic zone, where local climatic differences may explain variations in Bd disease dynamics. We recommend more detection surveys across lowlands and other sites that have been historically considered unsuitable for Bd occurrence. These data can be used to identify sites for potential disease outbreaks and amphibian rediscoveries.     

Different trends of neighboring populations of Lesser Kestrel: Effects of climate and other environmental conditions

The sensitivity of population trends to the climate and environment is generally considered a species-specific trait. However, evidence that populations may show different responses to the climate and environmental conditions is growing. Whether this differential sensitivity may arise even among neighboring populations remains elusive. We compared the trends of two neighboring populations of the Lesser Kestrel Falco naumanni, using data from a 12-year survey of 158 colonies in Sicily, Italy; the two populations inhabiting a lowland and an highland area, respectively. Population trends were modeled through the TRIM algorithms implemented in R (package rtrim). A reversed U-shaped population trend was observed in the lowland, while the highland population showed oscillations around a stable trend. Sahel rainfall 2 years before each annual survey significantly affected population variation in the lowland, while rainfall in March and an index of primary productivity in the breeding areas affected population variation in the highland. This suggests that the population in the lowland may be limited mainly by winter survival in Sahel, because the lowland may be an optimal breeding area for this species. In contrast, the highland population, which occupies a different part of the climatic niche of the species, may be limited mainly by reproductive output, because rainfall in March and the primary productivity in May could represent prey availability immediately before and during the breeding months. Overall, our findings suggest that population-specific environmental sensitivity might occur even over small (<100 km) geographical scales, highlighting the need for population-specific conservation strategies.     

Adaptive divergence in Darwin's small ground finch (Geospiza fuliginosa): divergent selection along a cline

We examine here, in a single year (2005), phenotypic divergence along a 560‐m elevation gradient in Darwin's small ground finch (Geospiza fuliginosa) in the Galápagos Islands. In this sample, four composite measures of phenotypic traits showed significant differences along the 18‐km geographical cline extending from lowlands to highlands. Compared with lowland birds, highland birds had larger and more pointed beaks, and thicker tarsi, but smaller feet and claws. Finches in an intervening agricultural zone had predominantly intermediate trait values. In a second, mark–recapture study we analyse selection on morphological traits among birds recaptured across years (2000–2005) in lowland and highland habitats. Birds were more likely to survive in the highlands and during the wet season, as well as if they had large beaks and bodies. In addition, highland birds exhibited higher survival rates if they had small feet and pointed beaks – attributes common to highland birds as a whole. Lowland birds were more likely to survive if they possessed the opposite traits. Selection therefore reinforced existing morphological divergence, which appears to reflect local adaptation to differing resources during the predominantly drought‐ridden conditions that characterized the 5‐year study. Alternative explanations – including genetic drift, matching habitat choice, deformation by parasites, and the effects of wear – received little or no support. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 45–59.     

Competition and disturbance affect elevational distribution of two congeneric conifers

Climatic change will affect elevational vegetation distribution because vegetation distribution is related to thermal conditions. However, how elevational species distributions are determined by biotic and abiotic factors is not clear. The long‐term plot census along an elevational gradient is indispensable to clarify mechanisms of elevational distribution of tree species. Two congeneric conifers, the less shade‐tolerant Abies veitchii and shade‐tolerant A. mariesii, dominate at low and high elevations, respectively, in the subalpine zone in Japan. This study investigated the population dynamics of the two species at three elevations (low, middle, high) for 13 years to examine why the two species dominated the different elevations from the viewpoints of competition and disturbance. This study showed that growth and survival rates were not highest at the most dominant elevations for each species. At the high elevation where A. mariesii dominated and small disturbances frequently occurred, the recruitment rate of A. mariesii was highest among the three elevations and that of A. veitchii was largely decreased by tree competition. However, A. veitchii was dominant earlier than A. mariesii at the low elevation after large disturbances by the high growth rate of individual trees. Therefore, A. mariesii was superior to A. veitchii at the high elevation because of its high recruitment rate and large reduction of recruitment of A. veitchii due to competition, while A. veitchii was superior to A. mariesii at the low elevation after large disturbances because of higher growth rate than A. mariesii. It is suggested that the elevational distributions of the two species were determined by elevational changes in population dynamics in relation to competition and disturbance. Long‐term observational studies of forest dynamics among various elevations are indispensable to predict the effects of climatic change on vegetation distribution.     

Incorporating biotic factors in species distribution modeling: are interactions with soil microbes important?

It is increasingly recognized that species distributions are driven by both abiotic factors and biotic interactions. Despite much recent work incorporating competition, predation, and mutualism into species distribution models (SDMs), the focus has been confined to aboveground macroscopic interactions. Biotic interactions between plants and soil microbial communities are understudied as potentially important drivers of plant distributions. Some soil bacteria promote plant growth by cycling nutrients, while others are pathogenic; thus they have a high potential for influencing plant occurrence. We investigated the influence of soil bacterial clades on the distributions of bryophytes and 12 vascular plant species in a high elevation talus‐field ecosystem in the Rocky Mountain Front Range, Colorado, USA. We used an information‐theoretic criterion (AICc) modeling approach to compare SDMs with the following different sets of predictors: abiotic variables, abiotic variables and other plant abundances, abiotic variables and soil bacteria clade relative abundances, and a full model with abiotic factors, plant abundances, and bacteria relative abundances. We predicted that bacteria would influence plant distributions both positively and negatively, and that these interactions would improve prediction of plant species distributions. We found that inclusion of either plant or bacteria biotic predictors generally improved the fit, deviance explained, and predictive power of the SDMs, and for the majority of the species, adding information on both other plants and bacteria yielded the best model. Interactions between the modeled species and biotic predictors were both positive and negative, suggesting the presence of competition, parasitism, and facilitation. While our results indicate that plant–plant co‐occurrences are a stronger driver of plant distributions than plant–bacteria co‐occurrences, they also show that bacteria can explain parts of plant distributions that remain unexplained by abiotic and plant predictors. Our results provide further support for including biotic factors in SDMs, and suggest that belowground factors be considered as well.     

Counterbalancing effects of competition for resources and facilitation against grazing in alpine snowbed communities

Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species‐specific and/or dependent on ontogeny, and whether the balance of positive and negative plant–plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species‐specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants.