A framework for understanding human‐driven vegetation change

Despite a major research focus on human‐mediated reshuffling of plant communities, no coherent framework unites the numerous types of changes in abundances and distributions of native and non‐native species that are driven by human activities. Human driven vegetation change can occur through: non‐native species introductions; population outbreaks or collapses; range expansions or contractions; and range shifts of both native and non‐native species. Boundaries among these different types of floristic changes are not always distinct because of an overlap in the ecological, climatic, and anthropogenic processes that underpin them. We propose a new framework that connects various human‐mediated causes of vegetation change, highlights the spatial scales at which drivers act and the temporal scale at which plant assemblages respond, and provides critical insights for identifying and appropriately managing these changes.     

Native range climate is insufficient to predict anuran invasive potential

Niche conservatism explains biological invasions worldwide. However, a plethora of ecological processes may lead invasive species to occupy environments that are different from those found within native ranges. Here, we assess the potential invadable areas of  the world’s most pervasive invasive amphibians: the cane toad, Rhinella marina?+?R. horribilis, and the North American bullfrog, Lithobates catesbeianus. The uncontrolled spread of such voracious, large-bodied, and disease-tolerant anurans has been documented to impact native faunas worldwide. To disentangle their invasion-related niche dynamics, we compared the predictive ability and distributional forecasts of ecological niche models calibrated with information from native, invaded and pooled (native?+?invaded) ranges. We found that including occurrences from invaded ranges improved model accuracy for both studied species. Non-native occurrences also accounted for 54% and 61% increase in the total area of potential distribution of the cane toad and bullfrog, respectively. Besides, the latter species occupied locations with climatic conditions that are more extreme than those found within its native range. Our results indicate that the occupancy of environments different from those found in native ranges increases the overall potential distribution of the studied invasive anuran species. Therefore, climate information on native ranges alone is insufficient to explain and anticipate the distributional patterns of invasion of cane toads and bullfrogs, underestimating predictions of potential invadable distribution. Moreover, such an observed expansion of realized niches towards occupancy of climates not found within native ranges also has clear implications for invasion risk assessments based on climate modelling worldwide.

     

Patterns of niche filling and expansion across the invaded ranges of an Australian lizard

Studies of realized niche shifts in alien species typically ignore the potential effects of intraspecific niche variation and different invaded‐range environments on niche lability. We incorporate our detailed knowledge of the native‐range source populations and global introduction history of the delicate skink Lampropholis delicata to examine intraspecific variation in realized niche expansion and unfilling, and investigate how alternative niche modelling approaches are affected by that variation. We analyzed the realized niche dynamics of L. delicata using an ordination method, ecological niche models (ENMs), and occurrence records from 1) Australia (native range), 2) New Zealand, 3) Hawaii, 4) the two distinct native‐range clades that were the sources for the New Zealand and Hawaii introductions, and 5) the species’ global range (including Lord Howe Island, Australia). We found a gradient of realized niche change across the invaded ranges of L. delicata: niche stasis on Lord Howe Island, niche unfilling in New Zealand (16%), and niche unfilling (87%) and expansion (14%) in Hawaii. ENMs fitted to native‐range data generally identified suitable climatic conditions at sites where the species has established non‐native populations, whereas ENMs based on native‐range source clades and non‐native populations had lower spatial transferability. Our results suggest that the extent to which realized niches are maintained during invasion does not depend on species‐level traits. When realized niche shifts are predominately due to niche unfilling, fully capturing species’ responses along climatic gradients by basing ENMs on native distributions may be more important for accurate invasion forecasts than incorporating phylogenetic differentiation, or integrating niche changes in the invaded range.     

Ecological Niche Transferability Using Invasive Species as a Case Study

Species distribution modeling is widely applied to predict invasive species distributions and species range shifts under climate change. Accurate predictions depend upon meeting the assumption that ecological niches are conserved, i.e., spatially or temporally transferable. Here we present a multi-taxon comparative analysis of niche conservatism using biological invasion events well documented in natural history museum collections. Our goal is to assess spatial transferability of the climatic niche of a range of noxious terrestrial invasive species using two complementary approaches. First we compare species’ native versus invasive ranges in environmental space using two distinct methods, Principal Components Analysis and Mahalanobis distance. Second we compare species’ native versus invaded ranges in geographic space as estimated using the species distribution modeling technique Maxent and the comparative index Hellinger’s I. We find that species exhibit a range of responses, from almost complete transferability, in which the invaded niches completely overlap with the native niches, to a complete dissociation between native and invaded ranges. Intermediate responses included expansion of dimension attributable to either temperature or precipitation derived variables, as well as niche expansion in multiple dimensions. We conclude that the ecological niche in the native range is generally a poor predictor of invaded range and, by analogy, the ecological niche may be a poor predictor of range shifts under climate change. We suggest that assessing dimensions of niche transferability prior to standard species distribution modeling may improve the understanding of species’ dynamics in the invaded range.     

Occupancy versus colonization–extinction models for projecting population trends at different spatial scales

Understanding spatiotemporal population trends and their drivers is a key aim in population ecology. We further need to be able to predict how the dynamics and sizes of populations are affected in the long term by changing landscapes and climate. However, predictions of future population trends are sensitive to a range of modeling assumptions. Deadwood‐dependent fungi are an excellent system for testing the performance of different predictive models of sessile species as these species have different rarity and spatial population dynamics, the populations are structured at different spatial scales, and they utilize distinct substrates. We tested how the projected large‐scale occupancies of species with differing landscape‐scale occupancies are affected over the coming century by different modeling assumptions. We compared projections based on occupancy models against colonization–extinction models, conducting the modeling at alternative spatial scales and using fine‐ or coarse‐resolution deadwood data. We also tested effects of key explanatory variables on species occurrence and colonization–extinction dynamics. The hierarchical Bayesian models applied were fitted to an extensive repeated survey of deadwood and fungi at 174 patches. We projected higher occurrence probabilities and more positive trends using the occupancy models compared to the colonization–extinction models, with greater difference for the species with lower occupancy, colonization rate, and colonization:extinction ratio than for the species with higher estimates of these statistics. The magnitude of future increase in occupancy depended strongly on the spatial modeling scale and resource resolution. We encourage using colonization–extinction models over occupancy models, modeling the process at the finest resource‐unit resolution that is utilizable by the species, and conducting projections for the same spatial scale and resource resolution at which the model fitting is conducted. Further, the models applied should include key variables driving the metapopulation dynamics, such as the availability of suitable resource units, habitat quality, and spatial connectivity.     

From richer to poorer: successful invasion by freshwater fishes depends on species richness of donor and recipient basins

Evidence for the theory of biotic resistance is equivocal, with experiments often finding a negative relationship between invasion success and native species richness, and large‐scale comparative studies finding a positive relationship. Biotic resistance derives from local species interactions, yet global and regional studies often analyze data at coarse spatial grains. In addition, differences in competitive environments across regions may confound tests of biotic resistance based solely on native species richness of the invaded community. Using global and regional data sets for fishes in river and stream reaches, we ask two questions: (1) does a negative relationship exist between native and non‐native species richness and (2) do non‐native species originate from higher diversity systems. A negative relationship between native and non‐native species richness in local assemblages was found at the global scale, while regional patterns revealed the opposite trend. At both spatial scales, however, nearly all non‐native species originated from river basins with higher native species richness than the basin of the invaded community. Together, these findings imply that coevolved ecological interactions in species‐rich systems inhibit establishment of generalist non‐native species from less diverse communities. Consideration of both the ecological and evolutionary aspects of community assembly is critical to understanding invasion patterns. Distinct evolutionary histories in different regions strongly influence invasion of intact communities that are relatively unimpacted by human actions, and may explain the conflicting relationship between native and non‐native species richness found at different spatial scales.     

Local range boundaries vs. large‐scale trade‐offs: climatic and competitive constraints on tree growth

Species often respond to human‐caused climate change by shifting where they occur on the landscape. To anticipate these shifts, we need to understand the forces that determine where species currently occur. We tested whether a long‐hypothesised trade‐off between climate and competitive constraints explains where tree species grow on mountain slopes. Using tree rings, we reconstructed growth sensitivity to climate and competition in range centre and range margin tree populations in three climatically distinct regions. We found that climate often constrains growth at environmentally harsh elevational range boundaries, and that climatic and competitive constraints trade‐off at large spatial scales. However, there was less evidence that competition consistently constrained growth at benign elevational range boundaries; thus, local‐scale climate‐competition trade‐offs were infrequent. Our work underscores the difficulty of predicting local‐scale range dynamics, but suggests that the constraints on tree performance at a large‐scale (e.g. latitudinal) may be predicted from ecological theory.     

Scale dependence of temperature as an abiotic driver of species' distributions

Aim Scale dependence of patterns and processes remains one of the major unresolved problems in ecology. The responses of ecosystems to environmental stressors are reported to be strongly scale dependent, but projections of the effects of climate change on species' distributions are still restricted to particular scales and knowledge about scale dependence is lacking. Here we propose that the scale dependence of those species' niche dimensions related to climate change is strongly related to the strength of climatic cross‐scale links. More specifically, we hypothesize that the strong cross‐scale links between micro‐ and macroclimatic conditions are related to high cross‐scale similarity (low scale dependence) of species' realized temperature niches and, thus, species' spatial distributions. Location This study covers seven orders of magnitude of spatial scale, ranging from local‐scale (below a metre) and regional‐scale (kilometre) investigations in central European wetland ecosystems to continental‐scale (thousands of kilometres) studies of species' distributions. Methods We combined data on the spatial occurrence of species (vegetation records at local and regional scales, digitized distribution maps at the continental scale) with information about the corresponding temperature regime of vascular plant species occurring in environmentally stable wetland ecosystems characterized by strong cross‐scale links between micro‐ and macroclimatic conditions. Results We observed high cross‐scale similarity of the characteristics of species temperature niches across seven orders of magnitude of spatial scale. However, the importance of temperature as an abiotic driver decreased nonlinearly with decreasing scale, suggesting greater importance of additional (biotic) drivers of species' occurrence at small spatial scales. Main conclusions We report high cross‐scale similarity of realized temperature niches for species inhabiting ecosystems where small‐scale environmental noise is low and cross‐scale links between micro‐ and macroclimatic conditions are strong. By highlighting a strong relationship between abiotic and biotic cross‐scale similarity, our results will help to improve niche‐based species distribution modelling, one of the major assessment tools for determining the ecological effects of climate change.     

Congener diversity,topographic heterogeneity and human‐assisted dispersal predict spread rates of alien herpetofauna at a global scale

Understanding the factors that determine rates of range expansion is not only crucial for developing risk assessment schemes and management strategies for invasive species, but also provides important insight into the ability of species to disperse in response to climate change. However, there is little knowledge on why some invasions spread faster than others at large spatiotemporal scales. Here, we examine the effects of human activities, species traits and characteristics of the invaded range on spread rates using a global sample of alien reptile and amphibian introductions. We show that spread rates vary remarkably among invaded locations within a species, and differ across biogeographical realms. Spread rates are positively related to the richness of native congeneric species and human‐assisted dispersal in the invaded range but are negatively correlated with topographic heterogeneity. Our findings highlight the importance of environmental characteristics and human‐assisted dispersal in developing robust frameworks for predicting species' range shifts.     

Reduced vertebrate diversity independent of spatial scale following feral swine invasions

Biological invasions often have contrasting consequences with reports of invasions decreasing diversity at small scales and facilitating diversity at large scales. Thus, previous literature has concluded that invasions have a fundamental spatial scale‐dependent relationship with diversity. Whether the scale‐dependent effects apply to vertebrate invaders is questionable because studies consistently report that vertebrate invasions produce different outcomes than plant or invertebrate invasions. Namely, vertebrate invasions generally have a larger effect size on species richness and vertebrate invaders commonly cause extinction, whereas extinctions are rare following invertebrate or plant invasions. In an agroecosystem invaded by a non‐native ungulate (i.e., feral swine, Sus scrofa), we monitored species richness of native vertebrates in forest fragments ranging across four orders of magnitude in area. We tested three predictions of the scale‐dependence hypothesis: (a) Vertebrate species richness would positively increase with area, (b) the species richness y‐intercept would be lower when invaded, and (c) the rate of native species accumulation with area would be steeper when invaded. Indeed, native vertebrate richness increased with area and the species richness was 26% lower than should be expected when the invasive ungulate was present. However, there was no evidence that the relationship was scale dependent. Our data indicate the scale‐dependent effect of biological invasions may not apply to vertebrate invasions.     

A multi-scale approach to identify invasion drivers and invaders’ future dynamics

Climate, land use and disturbances are well known drivers of invasion. However, their relative influence may change across spatial scales, where climate is expected to be the main filter at broad scales; land use is expected to have more influence at intermediate scales, and disturbance, at fine ones. Understanding the underlying processes that drive invasion patterns at different spatial scales is thus crucial to be able to anticipate the future spread of invaders. Here, we quantified the relative importance of climate, land use, and disturbance on the distribution of the invasive trees Ailanthus altissima and Robinia pseudoacacia, across three nested spatial scales, namely global, country (Spain) and riverbank (three riparian riverbanks). To do so, for each species and scale, we built ensemble species distribution models. We also identified their range filling and inferred the most suitable areas in Spain for them to spread. In general, our study confirms that climate acts as an initial coarse filter of species distribution, whilst both climate and land use were important at the country scale; at the riverbank scale human-mediated disturbances gained importance. However, R. pseudoacacia and A. altissima showed differences in their degree of range filling, where A. altissima has a higher potential for range expansion in the near future. Overall, the integration of different scales into invasion studies shows a great potential to enrich our understanding of species-habitat relationships, and to help anticipate their future dynamics.     

Predicted effects of climate factors on mountain species are not uniform over different spatial scales

The selection of relevant factors and appropriate spatial scale(s) is fundamental when modelling species response to climate change. We evaluated whether the effects of climate factors on species distribution/occurrence are consistently modelled over different spatial scales in birds, and used a two‐scale approach to identify species–climate correlations unlikely to represent causal effects. We used passerine birds inhabiting mountain grassland in the Apennines (Italy) as a model. We surveyed four grassland species at 400 sampling points, and built habitat selection models (territory scale) and distribution models (seven algorithms, landscape scale). We compared the effect of climatic predictors on occurrence/distribution highlighted by models over the two spatial scales, and with the effects supposed a priori based on the climatic niche of each species. Models at the territory level included at least one climatic predictor for three species; the observed effect of climatic predictors was seldom consistent with supposed effects. At the broadest scale, distribution models for all species included climatic predictors, with varying consistence with supposed effects and findings at the finer scale. Despite the importance of climate for species distribution, occurrence could be more directly related to other factors, with important implications for understanding/predicting the impacts of climate/environmental changes. Our approach revealed key variables for grassland birds, and highlighted the scale‐dependent perceived importance of climate. At the local scale, climate effects were weak or hard to interpret. We found a general lack of consistence between supposed and observed effects at the territory level, and between landscape and territory models. Our results show the importance of predicting the potential effect of climatic factors prior to the analyses, carefully selecting ecologically meaningful variables and scales, and evaluating the nature and scale of climate–species links. We call for caution when predicting under future climates, especially when mechanistic effects and consistency across scales are lacking.     

Predicting current and future biological invasions: both native and invaded ranges matter

The classical approach to predicting the geographical extent of species invasions consists of training models in the native range and projecting them in distinct, potentially invasible areas. However, recent studies have demonstrated that this approach could be hampered by a change of the realized climatic niche, allowing invasive species to spread into habitats in the invaded ranges that are climatically distinct from those occupied in the native range. We propose an alternative approach that involves fitting models with pooled data from all ranges. We show that this pooled approach improves prediction of the extent of invasion of spotted knapweed (Centaurea maculosa) in North America on models based solely on the European native range. Furthermore, it performs equally well on models based on the invaded range, while ensuring the inclusion of areas with similar climate to the European niche, where the species is likely to spread further. We then compare projections from these models for 2080 under a severe climate warming scenario. Projections from the pooled models show fewer areas of intermediate climatic suitability than projections from the native or invaded range models, suggesting a better consensus among modelling techniques and reduced uncertainty.     

Climatic Niche Conservatism and Biogeographical Non-Equilibrium in Eschscholzia californica (Papaveraceae), an Invasive Plant in the Chilean Mediterranean Region

Species climate requirements are useful for predicting their geographic distribution. It is often assumed that the niche requirements for invasive plants are conserved during invasion, especially when the invaded regions share similar climate conditions. California and central Chile have a remarkable degree of convergence in their vegetation structure, and a similar Mediterranean climate. Such similarities make these geographic areas an interesting natural experiment for testing climatic niche dynamics and the equilibrium of invasive species in a new environment. We tested to see if the climatic niche of Eschscholzia californica is conserved in the invaded range (central Chile), and we assessed whether the invasion process has reached a biogeographical equilibrium, i.e., occupy all the suitable geographic locations that have suitable conditions under native niche requirements. We compared the climatic niche in the native and invaded ranges as well as the projected potential geographic distribution in the invaded range. In order to compare climatic niches, we conducted a Principal Component Analysis (PCA) and Species Distribution Models (SDMs), to estimate E. californica''s potential geographic distribution. We also used SDMs to predict altitudinal distribution limits in central Chile. Our results indicated that the climatic niche occupied by E. californica in the invaded range is firmly conserved, occupying a subset of the native climatic niche but leaving a substantial fraction of it unfilled. Comparisons of projected SDMs for central Chile indicate a similarity, yet the projection from native range predicted a larger geographic distribution in central Chile compared to the prediction of the model constructed for central Chile. The projected niche occupancy profile from California predicted a higher mean elevation than that projected from central Chile. We concluded that the invasion process of E. californica in central Chile is consistent with climatic niche conservatism but there is potential for further expansion in Chile.     

Accommodating scenarios of climate change and management in modelling the distribution of the invasive tree Schinus molle in South Africa

Determining the potential range of invasive alien species under current conditions is important. However, we also need to consider future distributions under scenarios of climate change and different management interventions when formulating effective long‐term intervention strategies. This paper combines niche modelling and fine‐scale process‐based modelling to define regions at high risk of invasion and simulate likely dynamics at the landscape scale. Our study species is Schinus molle (Peruvian pepper tree; Anacardiaceae), a native of central South America, introduced to South Africa in about 1850 where it was widely planted along roads. Localities of planted and naturalized trees were mapped along 5380 km of roads – a transect that effectively samples a large part of western South Africa. Correlative modelling was used to produce profiles of present and future environmental conditions characterizing its planted and naturalized ranges. A cellular‐automata simulation model was used to estimate the dynamics of S. molle under future climates and different management scenarios. The overall potential range of S. molle in the region is predicted to shrink progressively with predicted climate change. Some of the potential range of S. molle defined based on current conditions (including areas where it is currently highly invasive) is likely to become less favourable. The species could persist where it is well established long after conditions for recruitment have deteriorated. Some areas where the species is not widely naturalized now (notably the fynbos biome) are likely to become more favourable. Our modelling approach allows for the delineation of areas likely to be invaded in future by considering a range of factors at different scales that mediate the interplay of climatic variables and other drivers that define the dimensions of human intervention such as distance from planted trees and the density of planted plants, both of which affect propagule pressure.     

Anomalous outbreaks of an invasive defoliator and native bark beetle facilitated by warm temperatures,changes in precipitation and interspecific interactions

Biotic disturbance agents such as insects can be highly responsive to climatic change and have widespread ecological and economic impacts on forests. Quantifying the responses of introduced and native insects to climate, including how dynamics of one agent may mediate those of another, is important for forecasting disturbance and associated impacts on forest structure and function. We investigated drivers of outbreaks by larch casebearer Coleophora laricella, an invasive defoliator, and eastern larch beetle Dendroctonus simplex, a native, tree‐killing bark beetle, on tamarack Larix laricina from 2000 to in Minnesota, USA. We evaluated the utility of temporal, spatial and climatic variables in predicting the presence/absence of outbreaks of each insect in cells of rasterized aerial survey data. The role of defoliation by larch casebearer in outbreaks of eastern larch beetle was also investigated. For both species, the most important predictors of outbreak occurrence were proximity of conspecific outbreaks in space and time. For larch casebearer, outbreak occurrence was positively associated with spring precipitation and warmer growing seasons. Outbreak occurrence of eastern larch beetle was positively associated with warmer and dryer years and was more likely in cells with prior defoliation by larch casebearer. Our results demonstrate that climate can drive large scale outbreaks of introduced and non‐native disturbance agents on a single host species, and that interactions at the tree level between such agents may scale up to manifest across large temporal and spatial scales.     

Native–Exotic Species Richness Relationships Across Spatial Scales in a Prairie Restoration Matrix

In highly invaded ecosystems, restoration of native plant communities is dependent upon reducing exotic species relative to native species. Even so, in monitoring, the native–exotic species richness ratio has been shown to be scale‐dependent. Measurement at small spatial scales (<1 m²) can reveal a negative native–exotic richness relationship, where niche occupation may prevent invasion. Conversely, at larger scales, a positive correlation may exist, where environmental heterogeneity and equally favorable conditions may drive native–exotic relationships. Here, we compare slopes of native–exotic relationships across spatial scales in a prairie undergoing active restoration. The observed native–exotic richness ratios varied considerably over scales ranging from 1 to 1,000 m², emphasizing the importance of choosing a measurement scale that is most pertinent to the treatment and ecological mechanism used to evaluate restoration success. Our native–exotic richness slopes were positive over all scales, but lower than would be expected in a random community assembly, suggesting the influence of niche‐based competition. Correspondingly, our native–exotic cover slope was more negative than a null model; however, areas of frequent fire treatments showed a significant deviation from null only for richness, indicating that burning may enhance native–exotic competitive dynamics for number of species but not cover. The negative native–exotic cover relationships appear to be driven in this system mainly by exotic graminoids, across burn treatments and native functional groups, supporting the concept that frequent burning can alter the dominant competitive mechanism from coverage of these exotic grasses to an improved environment for germination and dispersal of more native species.     

Scale‐specific processes shape plant community patterns in subtropical coastal grasslands

Processes responsible for shaping community patterns act at specific spatial scales. In this study, we aimed at disentangling the effects of climate, soil and space as drivers of variation in a coastal grassland plant community. We were specifically interested in evaluating the relative influence of those processes at broad and fine spatial scales as well as when considering species groups with good and poor long‐distance dispersal capacity. We sampled grassland vegetation at 16 sites distributed along a latitudinal gradient of more than 500 km in subtropical southern Brazil and used variation partitioning procedures to ascertain the relative influence of climatic, edaphic and spatial processes on variation in species composition at different spatial scales, considering the entire community and subsets with only species from the Asteraceae family (good long‐distance dispersal) and Poaceae (poor long‐distance dispersal). Climatic filters were the most responsible for shaping grassland community composition at the broad scale, while edaphic filters showed higher importance at the fine scale. When not considering the influence of spatial scale, we observed higher influence of climate structured in space. Composition patterns of species with poor long‐distance dispersal (Poaceae) were more closely related to spatial variables than those of species with effective dispersal (Asteraceae). Our results stressed the importance of addressing different spatial scales to rightly ascertain the magnitude that different drivers exert on plant community assembly. Dividing the community into groups with different dispersal abilities proved useful for a more detailed understanding of the community assembly processes.     

Using multi-scale species distribution data to infer drivers of biological invasion in riparian wetlands

Aim Biological invasion is a major conservation problem that is of interest to ecological science. Understanding mechanisms of invasion is a high priority, heightened by the management imperative of acting quickly after species introduction. While information about invading species’ ecology is often unavailable, species distribution data can be collected near the onset of invasion. By examining distribution patterns of exotic and native plant species at multiple spatial scales, we aim to identify the scale (of those studied) that accounts for most variability in exotic species abundance, and infer likely drivers of invasion. Location River Murray wetlands, south‐eastern Australia. Methods A nested, crossed survey design was used to determine the extent of variation in wetland plant abundance, grazing intensity and water depth at four spatial scales (reaches, wetland clumps, wetlands, wetland sections), and among three Depth‐strata. We examined responses of exotic and native species groups (grouped into terrestrial and amphibious taxa), native weeds and 10 individual species using hierarchical ANOVA. Results As a group dominated by terrestrial taxa, exotic species cover varied at reach‐, wetland‐ and section‐scales. This likely reflects differences in abiotic characteristics and propagule pressure at these scales. Groups based on native species did not vary at any scale examined. Cover of 10 species mostly varied among and within wetlands (patterns unrelated to species’ origin or functional group), but species’ responses differed, despite individual plants being similar in size. While flora mostly varied among wetlands, exotic cover varied most among reaches (26%), which was attributed to hydrological modification and human activities. Main conclusions Multi‐scale surveys can rapidly identify factors likely to affect species’ distributions and can indicate where future research should be directed. By highlighting disproportionate variation in exotic cover among reaches, this study suggests that flow regulation and human‐mediated dispersal facilitate exotic plant invasion in River Murray wetlands.