Plant–soil interactions in the expansion and native range of a poleward shifting plant species

Climate warming causes range shifts of many species toward higher latitudes and altitudes. However, range shifts of host species do not necessarily proceed at the same rates as those of their enemies and symbionts. Here, we examined how a range shifting plant species performs in soil from its original range in comparison with soil from the expansion range. Tragopogon dubius is currently expanding from southern into north-western Europe and we examined how this plant species responds to soil communities from its original and expansion ranges. We compared the performance of T. dubius with that of the closely related Tragopogon pratensis , which has a natural occurrence along the entire latitudinal gradient. Inoculation with the rhizosphere soil from T. dubius populations of the original range had a more negative effect on plant biomass production than inoculation with rhizosphere soil from the expansion range. Interestingly, the nonrange expander T. pratensis experienced a net negative soil effect throughout this entire range. The effects observed in this species pair may be due to release from soil born enemies or accumulation of beneficial soil born organisms. If this phenomenon applies broadly to other species, then range expansion may enable plants species to show enhanced performance.     

Characterizing common and range expanding species

Climate change and biological invasions have increased the likelihood of the establishment, growth, spread and survival of some species. With the recent and strongly motivated call for balanced consideration of the ecology and conservation of common and rare species, species undergoing range expansion are of particular interest. The notion of essential biodiversity variables has also re‐focussed attention on the value of abundance and occupancy observations for assessing conservation values, threats and targets – species rare or common, contracting or expanding. Changes in the distribution and abundance of common and range expanding species deserve further attention, not only to better understand their dynamics, but also as the basis for monitoring range expansions and their consequences. With the aim of characterizing range expansion, here we combine perspectives from typological approaches to studying species conservation status with perspectives from spread ecology and macroecology. Closer examination of the properties and dynamics of common species is important not only because declines in their conservation status are underappreciated, but also because those species becoming more common have significant implications for biodiversity, ecosystems and society.     

Arctic and boreal plant species decline at their southern range limits in the Rocky Mountains

Climate change is predicted to cause a decline in warm‐margin plant populations, but this hypothesis has rarely been tested. Understanding which species and habitats are most likely to be affected is critical for adaptive management and conservation. We monitored the density of 46 populations representing 28 species of arctic‐alpine or boreal plants at the southern margin of their ranges in the Rocky Mountains of Montana, USA, between 1988 and 2014 and analysed population trends and relationships to phylogeny and habitat. Marginal populations declined overall during the past two decades; however, the mean trend for 18 dicot populations was ?5.8% per year, but only ?0.4% per year for the 28 populations of monocots and pteridophytes. Declines in the size of peripheral populations did not differ significantly among tundra, fen and forest habitats. Results of our study support predicted effects of climate change and suggest that vulnerability may depend on phylogeny or associated anatomical/physiological attributes.     

Beyond climate control on species range: The importance of soil data to predict distribution of Amazonian plant species

Aim

To evaluate the relative importance of climatic versus soil data when predicting species distributions for Amazonian plants and to gain understanding of potential range shifts under climate change.

Location

Amazon rain forest.

Methods

We produced species distribution models (SDM) at 5‐km spatial resolution for 42 plant species (trees, palms, lianas, monocot herbs and ferns) using species occurrence data from herbarium records and plot‐based inventories. We modelled species distribution with Bayesian logistic regression using either climate data only, soil data only or climate and soil data together to estimate their relative predictive powers. For areas defined as unsuitable to species occurrence, we mapped the difference between the suitability predictions obtained with climate‐only versus soil‐only models to identify regions where climate and soil might restrict species ranges independently or jointly.

Results

For 40 out of the 42 species, the best models included both climate and soil predictors. The models including only soil predictors performed better than the models including only climate predictors, but we still detected a drought‐sensitive response for most of the species. Edaphic conditions were predicted to restrict species occurrence in the centre, the north‐west and in the north‐east of Amazonia, while the climatic conditions were identified as the restricting factor in the eastern Amazonia, at the border of Roraima and Venezuela and in the Andean foothills.

Main conclusions

Our results revealed that soil data are a more important predictor than climate of plant species range in Amazonia. The strong control of species ranges by edaphic features might reduce species’ abilities to track suitable climate conditions under a drought‐increase scenario. Future challenges are to improve the quality of soil data and couple them with process‐based models to better predict species range dynamics under climate change.     

Interactive effects of global change drivers as determinants of the link between soil biodiversity and ecosystem functioning

Biodiversity, both aboveground and belowground, is negatively affected by global changes such as drought or warming. This loss of biodiversity impacts Earth's ecosystems, as there is a positive relationship between biodiversity and ecosystem functioning (BEF). Even though soils host a large fraction of biodiversity that underlies major ecosystem functions, studies exploring the relationship between soil biodiversity and ecosystem functioning (sBEF) as influenced by global change drivers (GCDs) remain scarce. Here we highlight the need to decipher sBEF relationships under the effect of interactive GCDs that are intimately connected in a changing world. We first state that sBEF relationships depend on the type of function (e.g., C cycling or decomposition) and biodiversity facet (e.g., abundance, species richness, or biomass) considered. Then, we shed light on the impact of single and interactive GCDs on soil biodiversity and sBEF and show that results from scarce studies studying interactive effects range from antagonistic to additive to synergistic when two individual GCDs cooccur. This indicates the need for studies quantitatively accounting for the impacts of interactive GCDs on sBEF relationships. Finally, we provide guidelines for optimized methodological and experimental approaches to study sBEF in a changing world that will provide more valuable information on the real impact of (interactive) GCDs on sBEF. Together, we highlight the need to decipher the sBEF relationship in soils to better understand soil functioning under ongoing global changes, as changes in sBEF are of immediate importance for ecosystem functioning.     

Endemic species and ecosystem sensitivity to climate change in Namibia

We present a first assessment of the potential impacts of anthropogenic climate change on the endemic flora of Namibia, and on its vegetation structure and function, for a projected climate in ～2050 and ～2080. We used both niche‐based models (NBM) to evaluate the sensitivity of 159 endemic species to climate change (of an original 1020 plant species modeled) and a dynamic global vegetation model (DGVM) to assess the impacts of climate change on vegetation structure and ecosystem functioning. Endemic species modeled by NBM are moderately sensitive to projected climate change. Fewer than 5% are predicted to experience complete range loss by 2080, although more than 47% of the species are expected to be vulnerable (range reduction >30%) by 2080 if they are assumed unable to migrate. Disaggregation of results by life‐form showed distinct patterns. Endemic species of perennial herb, geophyte and tree life‐formsare predicted to be negatively impacted in Namibia, whereas annual herb and succulent endemic species remain relatively stable by 2050 and 2080. Endemic annual herb species are even predicted to extend their range north‐eastward into the tree and shrub savanna with migration, and tolerance of novel substrates. The current protected area network is predicted to meet its mandate by protecting most of the current endemicity in Namibia into the future. Vegetation simulated by DGVM is projected to experience a reduction in cover, net primary productivity and leaf area index throughout much of the country by 2050, with important implications for the faunal component of Namibia's ecosystems, and the agricultural sector. The plant functional type (PFT) composition of the major biomes may be substantially affected by climate change and rising atmospheric CO₂– currently widespread deciduous broad leaved trees and C₄ PFTs decline, with the C₄ PFT particularly negatively affected by rising atmospheric CO₂ impacts by ～2080 and deciduous broad leaved trees more likely directly impacted by drying and warming. The C₃ PFT may increase in prominence in the northwestern quadrant of the country by ～2080 as CO₂ concentrations increase. These results suggest that substantial changes in species diversity, vegetation structure and ecosystem functioning can be expected in Namibia with anticipated climate change, although endemic plant richness may persist in the topographically diverse central escarpment region.     

Upward elevation and northwest range shifts for alpine Meconopsis species in the Himalaya–Hengduan Mountains region

Climate change may impact the distribution of species by shifting their ranges to higher elevations or higher latitudes. The impacts on alpine plant species may be particularly profound due to a potential lack of availability of future suitable habitat. To identify how alpine species have responded to climate change during the past century as well as to predict how they may react to possible global climate change scenarios in the future, we investigate the climatic responses of seven species of Meconopsis, a representative genus endemic in the alpine meadow and subnival region of the Himalaya–Hengduan Mountains. We analyzed past elevational shifts, as well as projected shifts in longitude, latitude, elevation, and range size using historical specimen records and species distribution modeling under optimistic (RCP 4.5) and pessimistic (RCP 8.5) scenarios across three general circulation models for 2070. Our results indicate that across all seven species, there has been an upward shift in mean elevation of 302.3 m between the pre‐1970s (1922–1969) and the post‐1970s (1970–2016). The model predictions suggest that the future suitable climate space will continue to shift upwards in elevation (as well as northwards and westwards) by 2070. While for most of the analyzed species, the area of suitable climate space is predicted to expand under the optimistic emission scenario, the area contracts, or, at best, shows little change under the pessimistic scenario. Species such as M. punicea, which already occupy high latitudes, are consistently predicted to experience a contraction of suitable climate space across all the models by 2070 and may consequently deserve particular attention by conservation strategies. Collectively, our results suggest that the alpine high‐latitude species analyzed here have already been significantly impacted by climate change and that these trends may continue over the coming decades.     

Early signs of range disjunction of submountainous plant species: an unexplored consequence of future and contemporary climate changes

Poleward and upward species range shifts are the most commonly anticipated and studied consequences of climate warming. However, these global responses to climate change obscure more complex distribution change patterns. We hypothesize that the spatial arrangement of mountain ranges and, consequently, climatic gradients in Europe, will result in range disjunctions. This hypothesis was investigated for submountainous forest plant species at two temporal and spatial scales: (i) under future climate change (between 1950–2000 and 2061–2080 periods) at the European scale and (ii) under contemporary climate change (between 1914–1987 and 1997–2013 periods) at the French scale. We selected 97 submountainous forest plant species occurring in France, among which distribution data across Europe are available for 25 species. By projecting future distribution changes for the 25 submountainous plant species across Europe, we demonstrated that range disjunction is a likely consequence of future climate change. To assess whether it is already taking place, we used a large forest vegetation‐plot database covering the entire French territory over 100 years (1914–2013) and found an average decrease in frequency (?0.01 ± 0.004) in lowland areas for the 97 submountainous species – corresponding to a loss of 6% of their historical frequency – along with southward and upward range shifts, suggesting early signs of range disjunctions. Climate‐induced range disjunctions should be considered more carefully since they could have dramatic consequences on population genetics and the ability of species to face future climate changes.     

Small mammal controls on the climate‐driven range shift of woody plant species

Climate change is resulting in shifts in species’ ranges as species inhabit new climatically suitable areas. A key factor affecting range‐shifts is the interaction with predators. Small mammals, being primary seed predators and dispersers in forest ecosystems, may play a major role in determining which plant species will successfully expand and the rate at which range‐shifts will occur. Plants dispersing seeds beyond the species’ current range limits will encounter seed predators to which these seeds are novel; however, empirical studies of seed predator–novel seed interactions are lacking. The aims of our study were to: 1) quantify seed selection by small mammals presented with ‘novel’ seeds; 2) quantify the post‐selection fate of ‘novel’ seeds; and 3) identify seed traits that affect seed selection and post‐selection seed fate. We designed a field experiment exposing small mammal communities to novel seeds produced by plants expected to shift their ranges in response to climate change. We matched novel seeds with reference ‘familiar’ seeds and studied key steps defining interactions between small mammals and novel seeds. We found that the probability of selection of a novel seed varied among species and was, at times, higher than the selection probability of familiar seeds. Key traits that affected seed selection and the distance a seed was dispersed for caching were shell hardness and seed mass. We also found that 33% of dispersed seeds were cached in optimal germination sites (e.g. within fallen logs and buried under the leaf litter mat). Through seed emergence trials we found that emergence was higher for larger seeds, suggesting that the role of small mammals may be modulated by emergence rates. Our results suggest that the interaction between small mammals and novel seeds may have cascading effects on climate‐induced plant range shifts and community composition.     

Factors relating to regional and local success of exotic plant species in their new range

Aim To estimate invasiveness of exotic plant species, many studies have used the frequency of occurrence within a defined region. This measure is informative on how widespread exotics are, however, it does not inform on their local dominance, which is crucial for conservation of biodiversity and ecosystem functioning. The aim of the present study is to determine if regional frequency of occurrence of exotic plant species indeed is indicative of their local dominance. We also determined which plant traits and other factors predict regional and local frequencies best. Location The Netherlands. Methods We used information on exotic plant species established in The Netherlands and compared traits relating to their frequency of occurrence regionally (the entire country) and their frequency of dominance locally (in 1–100 m² quadrats). We created minimal adequate models with factors explaining regional frequency and frequency of local dominance of 111 exotic plant species in The Netherlands. Results The model that used plant traits to explain regional frequency of exotic plant species differed from the models that best explained their frequency of local dominance. Regionally, the factors that correlated with frequency were: life form, height, polyploidy, length of flowering season, residence time, human use and origin. The factors that correlated to frequency of local dominance were lateral vegetative spread and residence time. Main conclusions We conclude that plant traits relating to the regional frequency of exotic plant species differ from those that relate to their frequency of local dominance. The implication of our results is that predictive studies on plant invasiveness based on regional frequencies may not be indicative of the local impacts. Since the prediction of local impacts is crucial for conservation and risk assessment, our study emphasized the need for better information on the local abundance of exotic invaders.     

Relatedness with plant species in native community influences ecological consequences of range expansions

Global warming is enabling many plant species to expand their range to higher latitudes and altitudes, where they may suffer less from natural aboveground and belowground enemies. Reduced control by natural enemies can enable climate warming‐induced range expanders to gain an advantage in competition with natives and become disproportionally abundant in their new range. However, so far studies have only examined individual growth of range expanders, which have common congeneric plant species in their new range. Thus it is not known how general is this reduced effect of above‐ and belowground enemies and how it operates in communities, where multiple plant species also interact with each other. Here we show that range‐expanding plant species with and without congenerics in the invaded habitats differ in their ecological interactions in the new range. In a community‐level experiment, range‐expanding plant species, both with and without congenerics, suppressed the growth of a herbivore. However, only range expanders without congenerics reduced biomass production of the native plant species. In the present study, range expanders without congenerics allocated more biomass aboveground compared to native plant species, which can explain their competitive advantage. Competitive interaction and also biomass allocation of native plants and their congeneric range expanders were similar. Our results highlight that information about species phylogenetic relatedness with native flora can be crucial for improving predictions about the consequences of climate warming‐induced range expansions.     

Annual temperature variation as a time machine to understand the effects of long‐term climate change on a poleward range shift

Range shifts due to annual variation in temperature are more tractable than range shifts linked to decadal to century long temperature changes due to climate change, providing natural experiments to determine the mechanisms responsible for driving long‐term distributional shifts. In this study we couple physiologically grounded mechanistic models with biogeographic surveys in 2 years with high levels of annual temperature variation to disentangle the drivers of a historical range shift driven by climate change. The distribution of the barnacle Semibalanus balanoides has shifted 350 km poleward in the past half century along the east coast of the United States. Recruits were present throughout the historical range following the 2015 reproductive season, when temperatures were similar to those in the past century, and absent following the 2016 reproductive season when temperatures were warmer than they have been since 1870, the earliest date for temperature records. Our dispersal dependent mechanistic models of reproductive success were highly accurate and predicted patterns of reproduction success documented in field surveys throughout the historical range in 2015 and 2016. Our mechanistic models of reproductive success not only predicted recruitment dynamics near the range edge but also predicted interior range fragmentation in a number of years between 1870 and 2016. All recruits monitored within the historical range following the 2015 colonization died before 2016 suggesting juvenile survival was likely the primary driver of the historical range retraction. However, if 2016 is indicative of future temperatures mechanisms of range limitation will shift and reproductive failure will lead to further range retraction in the future. Mechanistic models are necessary for accurately predicting the effects of climate change on ranges of species.     

Climate change, anthropogenic disturbance and the northward range expansion of Lactuca serriola (Asteraceae)

Aim The distribution range of Lactuca serriola, a species native to the summer‐dry mediterranean climate, has expanded northwards during the last 250 years. This paper assesses the influence of climate on the range expansion of this species and highlights the importance of anthropogenic disturbance to its spread. Location Central and Northern Europe. Methods Data on the geographic distribution of L. serriola were assembled through a literature search as well as through floristic and herbarium surveys. Maps of the spread of L. serriola in Central and Northern Europe were prepared based on herbarium data. The spread was assessed more precisely in Germany, Austria and Great Britain by pooling herbarium and literature data. We modelled the bioclimatic niche of the species using occurrence and climatic data covering the last century to generate projections of suitable habitats under the climatic conditions of five time periods. We tested whether the observed distribution of L. serriola could be explained for each time period, assuming that the climatic niche of the species was conserved across time. Results The species has spread northwards since the beginning of the 19th century. We show that climate warming in Europe increased the number of sites suitable for the species at northern latitudes. Until the late 1970s, the distribution of the species corresponded to the climatically suitable sites available. For the last two decades, however, we could not show any significant relationship between the increase in suitable sites and the distributional range change of L. serriola. However, we highlight potential areas the species could spread to in the future (Great Britain, southern Scandinavia and the Swedish coast). It is predominantly non‐climatic influences of global change that have contributed to its rapid spread. Main conclusions The observation that colonizing species are not filling their climatically suitable range might imply that, potentially, other ruderal species could expand far beyond their current range. Our work highlights the importance of historical floristic and herbarium data for understanding the expansion of a species. Such historical distributional data can provide valuable information for those planning the management of contemporary environmental problems, such as species responses to environmental change.     

Detrital traits affect substitutability of a range‐expanding foundation species across latitude

Climate‐driven range shifts of foundation species could alter ecosystem processes and community composition by providing different resources than resident foundation species. Along the US Atlantic coast, the northward expanding foundation species, black mangrove Avicennia germinans, is replacing the dominant salt marsh foundation species, marsh cordgrass Spartina alterniflora. These species have distinct detrital attributes that ostensibly provide different resources to epifauna. We experimentally examined how detritus of these species affects decomposition and community composition in different habitat contexts at regional and local scales. First, we manipulated detritus identity (Avicennia, Spartina) at 13 sites across a 5° latitudinal gradient spanning mangrove, mixed marsh‐mangrove and salt marsh habitats. Across latitude, we found that Avicennia detritus decomposed 2–4 times faster than Spartina detritus, suggesting that detrital turnover will increase with mangrove expansion. Epifaunal abundance and richness increased 2–7 times from south to north (mangrove to salt marsh) and were equivalent between Avicennia and Spartina detritus except for crabs, a dominant taxonomic group that preferred Spartina detritus. Second, to examine the whether changing habitat context affected regional patterns, we manipulated detritus identity and surrounding habitat type (mangrove, salt marsh) at a single mixed site, also including inert mimics to separate structural and nutritional roles of detritus. Epifaunal richness was similar between the two detrital types, but crabs were 2–7 times more abundant in Spartina detritus due to its structural attributes. Surrounding habitat type did not influence decomposition rate or community patterns, which suggests that latitudinal influences, not surrounding habitat, drove the regional community patterns in the first experiment. Overall, mangrove expansion could alter epifaunal communities due to the lower structural value and faster turnover of mangrove detritus. As species shift with changing climate, understanding foundation species substitutability is critical to predict community change, but we must account for concomitant environmental changes that also modify communities.