Biotic interactions boost spatial models of species richness

Biotic interactions are known to affect the composition of species assemblages via several mechanisms, such as competition and facilitation. However, most spatial models of species richness do not explicitly consider inter‐specific interactions. Here, we test whether incorporating biotic interactions into high‐resolution models alters predictions of species richness as hypothesised. We included key biotic variables (cover of three dominant arctic‐alpine plant species) into two methodologically divergent species richness modelling frameworks – stacked species distribution models (SSDM) and macroecological models (MEM) – for three ecologically and evolutionary distinct taxonomic groups (vascular plants, bryophytes and lichens). Predictions from models including biotic interactions were compared to the predictions of models based on climatic and abiotic data only. Including plant–plant interactions consistently and significantly lowered bias in species richness predictions and increased predictive power for independent evaluation data when compared to the conventional climatic and abiotic data based models. Improvements in predictions were constant irrespective of the modelling framework or taxonomic group used. The global biodiversity crisis necessitates accurate predictions of how changes in biotic and abiotic conditions will potentially affect species richness patterns. Here, we demonstrate that models of the spatial distribution of species richness can be improved by incorporating biotic interactions, and thus that these key predictor factors must be accounted for in biodiversity forecasts.     

Context‐dependent biotic interactions control plant abundance across altitudinal environmental gradients

Many biotic interactions influence community structure, yet most distribution models for plants have focused on plant competition or used only abiotic variables to predict plant abundance. Furthermore, biotic interactions are commonly context‐dependent across abiotic gradients. For example, plant–plant interactions can grade from competition to facilitation over temperature gradients. We used a hierarchical Bayesian framework to predict the abundances of 12 plant species across a mountain landscape and test hypotheses on the context‐dependency of biotic interactions over abiotic gradients. We combined field‐based estimates of six biotic interactions (foliar herbivory and pathogen damage, fungal root colonization, fossorial mammal disturbance, plant cover and plant diversity) with abiotic data on climate and soil depth, nutrients and moisture. All biotic interactions were significantly context‐dependent along temperature gradients. Results supported the stress gradient hypothesis: as abiotic stress increased, the strength or direction of the relationship between biotic variables and plant abundance generally switched from negative (suggesting suppressed plant abundance) to positive (suggesting facilitation/mutualism). For half of the species, plant cover was the best predictor of abundance, suggesting that the prior focus on plant–plant interactions is well‐justified. Explicitly incorporating the context‐dependency of biotic interactions generated novel hypotheses about drivers of plant abundance across abiotic gradients and may improve the accuracy of niche models.     

Incorporating biotic interactions in the distribution models of African wild silk moths (Gonometa species,Lasiocampidae) using different representations of modelled host tree distributions

Biotic interactions influence species niches and may thus shape distributions. Nevertheless, species distribution modelling has traditionally relied exclusively on environmental factors to predict species distributions, while biotic interactions have only seldom been incorporated into models. This study tested the ability of incorporating biotic interactions, in the form of host plant distributions, to increase model performance for two host‐dependent lepidopterans of economic interest, namely the African silk moth species, Gonometa postica and Gonometa rufobrunnea (Lasiocampidae). Both species are dependent on a small number of host tree species for the completion of their life cycle. We thus expected the host plant distribution to be an important predictor of Gonometa distributions. Model performance of a species distribution model trained only on abiotic predictors was compared to four species distribution models that additionally incorporated biotic interactions in the form of four different representations of host plant distributions as predictors. We found that incorporating the moth–host plant interactions improved G. rufobrunnea model performance for all representations of host plant distribution, while for G. postica model performance only improved for one representation of host plant distribution. The best performing representation of host plant distribution differed for the two Gonometa species. While these results suggest that incorporating biotic interactions into species distribution models can improve model performance, there is inconsistency in which representation of the host tree distribution best improves predictions. Therefore, the ability of biotic interactions to improve species distribution models may be context‐specific, even for species which have obligatory interactions with other organisms.     

Individual flowering phenology shapes plant–pollinator interactions across ecological scales affecting plant reproduction

The balance of pollination competition and facilitation among co-flowering plants and abiotic resource availability can modify plant species and individual reproduction. Floral resource succession and spatial heterogeneity modulate plant–pollinator interactions across ecological scales (individual plant, local assemblage, and interaction network of agroecological infrastructure across the farm). Intraspecific variation in flowering phenology can modulate the precise level of spatio-temporal heterogeneity in floral resources, pollen donor density, and pollinator interactions that a plant individual is exposed to, thereby affecting reproduction. We tested how abiotic resources and multi-scale plant–pollinator interactions affected individual plant seed set modulated by intraspecific variation in flowering phenology and spatio-temporal floral heterogeneity arising from agroecological infrastructure. We transplanted two focal insect-pollinated plant species (Cyanus segetum and Centaurea jacea, n = 288) into agroecological infrastructure (10 sown wildflower and six legume–grass strips) across a farm-scale experiment (125 ha). We applied an individual-based phenologically explicit approach to match precisely the flowering period of plant individuals to the concomitant level of spatio-temporal heterogeneity in plant–pollinator interactions, potential pollen donors, floral resources, and abiotic conditions (temperature, water, and nitrogen). Individual plant attractiveness, assemblage floral density, and conspecific pollen donor density (C. jacea) improved seed set. Network linkage density increased focal species seed set and modified the effect of local assemblage richness and abundance on C. segetum. Mutual dependence on pollinators in networks increased C. segetum seed set, while C. jacea seed set was greatest where both specialization on pollinators and mutual dependence was high. Abiotic conditions were of little or no importance to seed set. Intra- and interspecific plant–pollinator interactions respond to spatio-temporal heterogeneity arising from agroecological management affecting wild plant species reproduction. The interplay of pollinator interactions within and between ecological scales affecting seed set implies a co-occurrence of pollinator-mediated facilitative and competitive interactions among plant species and individuals.     

Time after time: flowering phenology and biotic interactions

The role of biotic interactions in shaping plant flowering phenology has long been controversial; plastic responses to the abiotic environment, limited precision of biological clocks and inconsistency of selection pressures have generally been emphasized to explain phenological variation. However, part of this variation is heritable and selection analyses show that biotic interactions can modulate selection on flowering phenology. Our review of the literature indicates that pollinators tend to favour peak or earlier flowering, whereas pre-dispersal seed predators tend to favour off-peak or later flowering. However, effects strongly vary among study systems. To understand such variation, future studies should address the impact of mutualist and antagonist dispersal ability, ecological specialization, and habitat and plant population characteristics. Here, we outline future directions to study how such interactions shape flowering phenology.     

Biotic interactions drive species occurrence and richness in dynamic beach environments

Biotic interactions are predicted to have the strongest influence on species assemblages in extreme environments. We therefore test the hypothesis that in abiotically-severe beaches plant–plant interactions, specifically facilitation, are important relative to abiotic conditions. This hypothesis is tested by assessing the influence of dominant vascular plant species on the fine-scale occurrence and richness of vascular and cryptogam species using a unique dataset of boreal beaches along the Finnish Baltic Sea, characterized by strong post-glacial land uplift and large environmental gradients. We studied three different levels of vegetation patterns across a broad geographical scale; individual species, functional groups and the entire community. Results showed that dominant vascular species strongly drive species occurrence and richness in dynamic beach environments, with some species having an influence similar to that of key abiotic variables. In contrast to expectations, facilitative effects did not dominate in these harsh environments. Instead, the outcomes of biotic interactions were species-specific, and also differed between vascular and cryptogam species, with the former group most strongly influenced by a pioneer species and the latter by a late succession generalist. Our study highlights the importance of incorporating biotic interaction effects into models of multiple vegetation properties and cautions against overly simplistic generalizations to describe relatively idiosyncratic interaction effects.     

Reproductive phenology of Melastomataceae species with contrasting reproductive systems: contemporary and historical drivers

• Flowering and fruiting are key events in the life history of plants, and both are critical to their reproductive success. Besides the role of evolutionary history, plant reproductive phenology is regulated by abiotic factors and shaped by biotic interactions with pollinators and seed dispersers. In Melastomataceae, a dominant Neotropical family, the reproductive systems vary from allogamous with biotic pollination to apomictic, and seed dispersal varies from dry (self‐dispersed) to fleshy (animal‐dispersed) fruits. Such variety in reproductive strategies is likely to affect flowering and fruiting phenologies.
• In this study, we described the reproductive phenology of 81 Melastomataceae species occurring in two biodiversity hotspots: the Atlantic rain forest and the campo rupestre. We aim to disentangle the role of abiotic and biotic factors defining flowering and fruiting times of Melastomataceae species, considering the contrasting breeding and seed dispersal systems, and their evolutionary history.
• In both vegetation types, pollinator‐dependent species had higher flowering seasonality than pollinator‐independent ones. Flowering patterns presented phylogenetic signal regardless of vegetation type. Fruiting of fleshy‐fruited species was seasonal in campo rupestre but not in Atlantic rain forest; the fruiting of dry‐fruited species was also not seasonal in both vegetation types. Fruiting showed a low phylogenetic signal, probably because the influence of environment and dispersal agents on fruiting time is stronger than the phylogenetic affinity.
• Considering these ecophylogenetic patterns, our results indicate that flowering may be shaped by the different reproductive strategies of Melastomataceae lineages, while fruiting patterns may be governed mainly by the seed dispersal strategy and flowering time, with less phylogenetic influence.

     

Species‐specific responses to combined water stress and increasing temperatures in two bee‐pollinated congeners (Echium,Boraginaceae)

1. Water stress and increasing temperatures are two main constraints faced by plants in the context of climate change. These constraints affect plant physiology and morphology, including phenology, floral traits, and nectar rewards, thus altering plant–pollinator interactions.
2. We compared the abiotic stress responses of two bee‐pollinated Boraginaceae species, Echium plantagineum, an annual, and Echium vulgare, a biennial. Plants were grown for 5 weeks during their flowering period under two watering regimes (well‐watered and water‐stressed) and three temperature regimes (21, 24, 27°C).
3. We measured physiological traits linked to photosynthesis (chlorophyll content, stomatal conductance, and water use efficiency), and vegetative (leaf number and growth rate) and floral (e.g., flower number, phenology, floral morphology, and nectar production) traits.
4. The physiological and morphological traits of both species were affected by the water and temperature stresses, although the effects were greater for the annual species. Both stresses negatively affected floral traits, accelerating flower phenology, decreasing flower size, and, for the annual species, decreasing nectar rewards. In both species, the number of flowers was reduced by 22%–45% under water stress, limiting the total amount of floral rewards.
5. Under water stress and increasing temperatures, which mimic the effects of climate change, floral traits and resources of bee‐pollinated species are affected and can lead to disruptions of pollination and reproductive success.

     

Contrasting effects of biotic interactions on richness and distribution of vascular plants,bryophytes and lichens in an arctic–alpine landscape

Biotic interactions may strongly affect the distribution of individual species and the resulting patterns of species richness. However, the impacts can vary depending on the species or taxa examined, suggesting that the influences of interactions on species distributions and diversity are not always straightforward and can be taxon-contingent. The aim of this study was therefore to examine how the importance of biotic interactions varies within a community. We incorporated three biotic predictors (cover of the dominant vascular species) into two correlative species richness modelling frameworks to predict spatial variation in the number of vascular plants, bryophytes and lichens in arctic–alpine Fennoscandia, in N Europe. In addition, predictions based on single-species distribution models were used to determine the nature of the impact (negative vs. positive outcome) of the three dominant species on individual vascular plant, bryophyte and lichen species. Our results suggest that biotic variables can be as important as abiotic variables, but their relative contributions in explaining the richness of sub-dominant species vary among dominant species, species group and the modelling framework implemented. Similarly, the impacts of biotic interactions on individual species varied among the three species groups and dominant species, with the observed patterns partly reflecting species’ biogeographic range. Our study provides additional support for the importance of biotic interactions in modifying arctic–alpine biodiversity patterns and highlights that the impacts of interactions are not constant across taxa or biotic drivers. The influence of biotic interactions, including the taxon contingency and range-based impacts, should therefore be accounted for when developing biodiversity forecasts.     

Unravelling the effects of temperature, latitude and local environment on the reproduction of forest herbs

Aim To investigate the effect of temperature, latitude and local environment on the reproductive traits of widespread perennial forest herbs to better understand the potential impacts of rising temperatures on their population dynamics and colonization capacities. Location Six regions along a latitudinal gradient from France to Sweden. Methods Within each region, we collected data from three to five populations of up to six species. For each species, several variables were recorded in each region (temperature, latitude) and population (local abiotic and biotic environmental variables), and seed production and germination were estimated. Resource investment in reproduction (RIR) was quantified as seed number × seed mass, while germinable seed output (GSO) was expressed as seed number × germination percentage. We performed linear regression and mixed effect models to investigate the effects of temperature (growing degree hours), latitude and local abiotic and biotic environment on RIR and GSO. Results Temperature and latitude explained most of the variation in RIR and GSO for early flowering species with a northerly distribution range edge (Anemone nemorosa, Paris quadrifolia and Oxalis acetosella). Reproduction of the more southerly distributed species (Brachypodium sylvaticum, Circaea lutetiana and Primula elatior), in contrast, was independent of temperature/latitude. In the late summer species, B. sylvaticum and C. lutetiana, variation in RIR and GSO was best explained by local environmental variables, while none of the investigated variables appeared to be related to reproduction in P. elatior. Main conclusions We showed that reproduction of only two early flowering, northerly distributed species was related to temperature. This suggests that the potential reproductive response of forest herbs to climate warming partly depends on their phenology and distribution, but also that the response is to some extent species dependent. These findings should be taken into account when predictions about future shifts in distribution range are made.     

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.     

Phenotypic selection on flowering phenology and pollination efficiency traits between Primula populations with different pollinator assemblages

Floral traits have largely been attributed to phenotypic selection in plant–pollinator interactions. However, the strength of this link has rarely been ascertained with real pollinators. We conducted pollinator observations and estimated selection through female fitness on flowering phenology and floral traits between two Primula secundiflora populations. We quantified pollinator‐mediated selection by subtracting estimates of selection gradients of plants receiving supplemental hand pollination from those of plants receiving open pollination. There was net directional selection for an earlier flowering start date at populations where the dominant pollinators were syrphid flies, and flowering phenology was also subjected to stabilized quadratic selection. However, a later flowering start date was significantly selected at populations where the dominant pollinators were legitimate (normal pollination through the corolla tube entrance) and illegitimate bumblebees (abnormal pollination through nectar robbing hole which located at the corolla tube), and flowering phenology was subjected to disruptive quadratic selection. Wider corolla tube entrance diameter was selected at both populations. Furthermore, the strength of net directional selection on flowering start date and corolla tube entrance diameter was stronger at the population where the dominant pollinators were syrphid flies. Pollinator‐mediated selection explained most of the between‐population variations in the net directional selection on flowering phenology and corolla tube entrance diameter. Our results suggested the important influence of pollinator‐mediated selection on floral evolution. Variations in pollinator assemblages not only resulted in variation in the direction of selection but also the strength of selection on floral traits.     

When resources don't rescue: flowering phenology and species interactions affect compensation to herbivory in Ipomopsis aggregata

The ability of plants to tolerate, or compensate for, herbivore damage is highly variable and has been the subject of much research. Although many plants can compensate for herbivore damage, and some even overcompensate, we cannot yet generalize about the conditions that promote a positive response to damage. Here, we asked how abiotic resources (i.e. plant nutrient status) coupled with biotic interactions – i.e. subsequent interactions with pollinators, seed predators and nectar robbing bumble bees – affect the compensatory ability of Ipomopsis aggregata, a monocarpic herb that has been the subject of much previous debate. We hypothesized that compensation to herbivore damage in I. aggregata (Polemoniaceae) would depend first on plants having an ample supply of resources and, second, on the outcome of subsequent interactions with mutualist pollinators and enemy pre‐dispersal seed predators and nectar robbing bumble bees. We used a fully‐factorial experiment in which plants were watered, fertilized or left as unmanipulated controls, crossed with clipping to simulate herbivore damage to the apical meristem. Resource addition enhanced both male and female components of fitness, but resource enhancement did not provide the means for plants to fully compensate for simulated herbivory. Clipped plants produced significantly more inflorescences, but at the expense of a delay in flowering and fewer total flowers. Clipping significantly reduced losses to dipteran pre‐dispersal seed predators by delaying flowering time, but early flowering plants produced higher numbers of seeds despite incurring higher rates of predation. Clipped plants incurred a higher risk to nectar robbers in one of two years. Overall, clipped plants suffered severe reductions (a nearly 50% reduction in total seed set) in female success, but clipping combined with nutrient addition enhanced male function through increases in per‐flower pollen production. However, because clipped plants produced significantly fewer flowers than unclipped plants, whole‐plant pollen production was significantly reduced by clipping. Pollinator visitation and nectar robbing were variable between clipping treatments and between years and (nectar robbing) among sites. Our results demonstrate that the variability in plant response to herbivory can, at least in part, be driven by plant interactions with mutualists and enemies. Thus, accounting for such interactions and their variability is important to fully understanding plant compensation for herbivore damage and will likely go far to explain variation in plant response that appears to be independent of resources.     

Sugar landscapes and pollinator‐mediated interactions in plant communities

Pollinator‐mediated interactions between plants can play an important role for the dynamics of plant communities. Pollination services depend on the abundance and the foraging behaviour of pollinators, which in turn respond to the availability and distribution of floral resources (notably nectar sugar). However, it is still insufficiently understood how the ‘sugar landscapes’ provided by flowering plant communities shape pollinator‐mediated interactions between multiple plant species and across different spatial scales. A better understanding of pollinator‐mediated interactions requires an integrative approach that quantifies different aspects of sugar landscapes and investigates their relative importance for pollinator behaviour and plant reproductive success. In this study, we quantified such sugar landscapes from individual‐based maps of Protea shrub communities in the Cape Floristic Region, South Africa. The 27 study sites of 4 ha each jointly comprise 127 993 individuals of 19 species. We analysed how rates of visitation by key bird pollinators and the seed set of plants respond to different aspects of sugar landscapes: the distribution of nectar sugar amounts, as well as their quality, taxonomic purity and phenology. We found that pollinator visitation rates strongly depended on phenological variation of site‐scale sugar amounts. The seed set of focal plants increased with nectar sugar amounts of conspecific neighbours and with site‐scale sugar amounts. Seed set increased particularly strongly if site‐scale sugar amounts were provided by plants that offer less sugar per inflorescence. These combined effects of the amount, quality, purity and phenological variation of nectar sugar show that nectar sugar is a common interaction currency that determines how multiple plant species interact via shared pollinators. The responses of pollinator‐mediated interactions to different aspects of this interaction currency alter conditions for species coexistence in Protea communities and may cause community‐level Allee effects that promote extinction cascades.     

Later flowering is associated with a compressed flowering season and reduced reproductive output in an early season floral resource

Climate change‐induced shifts in flowering phenology can expose plants to novel biotic and abiotic environments, potentially leading to decreased temporal overlap with pollinators and exposure to conditions that negatively affect fruit and seed set. We explored the relationship between flowering phenology and reproductive output in the common shrub pointleaf manzanita Arctostaphylos pungens in a lower montane habitat in southeastern Arizona, USA. Contrary to the pattern of progressively earlier flowering observed in many species, long‐term records show that A. pungens flowering onset is shifting later and the flowering season is being compressed. This species can thus provide unusual insight into the effects of altered phenology. To determine the consequences of among‐ and within‐plant variation in flowering time, we documented individual flowering schedules and followed the fates of flowers on over 50 plants throughout two seasons (2012 and 2013). We also measured visitation rates by potential pollinators in 2012, as well as both fruit mass and seeds per fruit of flowers produced at different times. Fruit set was positively related to visitation rate but declined with later dates of flower production in both years. Total fruit production per plant was positively influenced by flowering duration, which declined with later flowering onset, as did fruit mass. Individual flowering schedules were consistent between years, suggesting that plants that begin flowering late have lower reproductive output each year. These patterns suggest that if pointleaf manzanita flowering continues to shift later, its flowering season may continue to become shorter, compressing floral resource availability for pollinators and leading to reduced reproductive output. These results reveal the negative effects of delayed phenology on reproductive output in a long‐lived plant. They highlight the value of using natural variation in flowering time, in combination with long‐term data, to anticipate the consequences of phenological shifts.     

Linking macroecology and community ecology: refining predictions of species distributions using biotic interaction networks

Macroecological models for predicting species distributions usually only include abiotic environmental conditions as explanatory variables, despite knowledge from community ecology that all species are linked to other species through biotic interactions. This disconnect is largely due to the different spatial scales considered by the two sub‐disciplines: macroecologists study patterns at large extents and coarse resolutions, while community ecologists focus on small extents and fine resolutions. A general framework for including biotic interactions in macroecological models would help bridge this divide, as it would allow for rigorous testing of the role that biotic interactions play in determining species ranges. Here, we present an approach that combines species distribution models with Bayesian networks, which enables the direct and indirect effects of biotic interactions to be modelled as propagating conditional dependencies among species’ presences. We show that including biotic interactions in distribution models for species from a California grassland community results in better range predictions across the western USA. This new approach will be important for improving estimates of species distributions and their dynamics under environmental change.     

Variable flowering phenology and pollinator use in a community suggest future phenological mismatch

Recent anthropogenic climate change is strongly associated with average shifts toward earlier seasonal timing of activity (phenology) in temperate-zone species. Shifts in phenology have the potential to alter ecological interactions, to the detriment of one or more interacting species. Recent models predict that detrimental phenological mismatch may increasingly occur between plants and their pollinators. One way to test this prediction is to examine data from ecological communities that experience large annual weather fluctuations. Taking this approach, we analyzed interactions over a four-year period among 132 plant species and 665 pollinating insect species within a Mediterranean community. For each plant species we recorded onset and duration of flowering and number of pollinator species. Flowering onset varied among years, and a year of earlier flowering of a species tended to be a year of fewer species pollinating its flowers. This relationship was attributable principally to early-flowering species, suggesting that shifts toward earlier phenology driven by climate change may reduce pollination services due to phenological mismatch. Earlier flowering onset of a species also was associated with prolonged flowering duration, but it is not certain that this will counterbalance any negative effects of lower pollinator species richness on plant reproductive success. Among plants with different life histories, annuals were more severely affected by flowering–pollinator mismatches than perennials. Specialized plant species (those attracting a smaller number of pollinator species) did not experience disproportionate interannual fluctuations in phenology. Thus they do not appear to be faced with disproportionate fluctuations in pollinator species richness, contrary to the expectation that specialists are at greatest risk of losing mutualistic interactions because of climate change.     

Biotic and abiotic variables show little redundancy in explaining tree species distributions

Abiotic factors such as climate and soil determine the species fundamental niche, which is further constrained by biotic interactions such as interspecific competition. To parameterize this realized niche, species distribution models (SDMs) most often relate species occurrence data to abiotic variables, but few SDM studies include biotic predictors to help explain species distributions. Therefore, most predictions of species distributions under future climates assume implicitly that biotic interactions remain constant or exert only minor influence on large‐scale spatial distributions, which is also largely expected for species with high competitive ability. We examined the extent to which variance explained by SDMs can be attributed to abiotic or biotic predictors and how this depends on species traits. We fit generalized linear models for 11 common tree species in Switzerland using three different sets of predictor variables: biotic, abiotic, and the combination of both sets. We used variance partitioning to estimate the proportion of the variance explained by biotic and abiotic predictors, jointly and independently. Inclusion of biotic predictors improved the SDMs substantially. The joint contribution of biotic and abiotic predictors to explained deviance was relatively small (~9%) compared to the contribution of each predictor set individually (~20% each), indicating that the additional information on the realized niche brought by adding other species as predictors was largely independent of the abiotic (topo‐climatic) predictors. The influence of biotic predictors was relatively high for species preferably growing under low disturbance and low abiotic stress, species with long seed dispersal distances, species with high shade tolerance as juveniles and adults, and species that occur frequently and are dominant across the landscape. The influence of biotic variables on SDM performance indicates that community composition and other local biotic factors or abiotic processes not included in the abiotic predictors strongly influence prediction of species distributions. Improved prediction of species' potential distributions in future climates and communities may assist strategies for sustainable forest management.     

The flowering pattern of the perennial herb Lobularia maritima: an unusual case in the Mediterranean basin

In plant communities of the Mediterranean Basin most plant species reach their blooming peak in spring and have characteristically short flowering periods of two-three months. The perennial herb Lobularia maritima represents an exception to these characteristics, because it flowers for almost 10 months, and has its flowering peak in autumn. In this five-year study, we describe the flowering pattern of L. maritima at the population and community levels. Despite the unusually extended flowering period of L. maritima, the species showed characteristic low among-year variability in the length of the flowering period but large interannual variation in the distribution of flowers throughout the flowering period. The flowering pattern (unimodal or bimodal) of L. maritima individuals differed among the five years, suggesting that L. maritima plants are plastic enough to tailor their flowering to variable environmental conditions. We conclude that flowering phenology of L. maritima represents a very particular case in the plant community studied, and the influence of abiotic and biotic factors on the phenology of this species is discussed.     

Species distribution models for Peruvian plantcutter improve with consideration of biotic interactions

Biotic interactions have been controversial in distributional ecology, mainly in regards to whether they have effects over broad extents, with the negative view known as the Eltonian noise hypothesis (ENH). In this study, we evaluated the ENH for Phytotoma raimondii, a restricted‐range Peruvian endemic bird species: we developed models based on 1) only abiotic conditions, 2) only host plant distributions, and 3) both abiotic conditions and host plant distributions; models were evaluated with partial receiver operating characteristic test and Akaike information criteria metrics. We rejected the ENH for this case: biotic interactions improved the model. The frequency with which exceptions to the ENH are detected has important implications for distributional ecology and methods for estimating distributions of species.