Predicting changes in the distribution and abundance of species under environmental change

Environmental changes are expected to alter both the distribution and the abundance of organisms. A disproportionate amount of past work has focused on distribution only, either documenting historical range shifts or predicting future occurrence patterns. However, simultaneous predictions of abundance and distribution across landscapes would be far more useful. To critically assess which approaches represent advances towards the goal of joint predictions of abundance and distribution, we review recent work on changing distributions and on effects of environmental drivers on single populations. Several methods have been used to predict changing distributions. Some of these can be easily modified to also predict abundance, but others cannot. In parallel, demographers have developed a much better understanding of how changing abiotic and biotic drivers will influence growth rate and abundance in single populations. However, this demographic work has rarely taken a landscape perspective and has largely ignored the effects of intraspecific density. We advocate a synthetic approach in which population models accounting for both density dependence and effects of environmental drivers are used to make integrated predictions of equilibrium abundance and distribution across entire landscapes. Such predictions would constitute an important step forward in assessing the ecological consequences of environmental changes.     

Relative abundance distributions in plant communities: effects of species richness and of spatial scale

Abstract. Relative abundance distributions (RADs) are an important feature of community structure, but little is known of the factors affecting which type of RAD is observed in a particular community. We examined the influences of species richness and of spatial scale on the RAD of plant communities. The effect of species richness was examined by analysing simulated communities generated under the Broken stick model, the Sequential breakage model, and a randomized version of Niche pre-emption model. In all cases, when there were few species in the community the data only occasionally gave the best fit to the model that was used to generate it. With 40–65 species, a best fit was obtained for the correct model in about 75 % of cases, almost irrespective of the model. Effects of spatial scale were examined in data from four dune slacks and from two semi-arid grasslands, by analysing biomass values at a range of sample sizes. The model that best fitted the whole sample differed between the four slacks and between the slacks and the semi-arid grasslands. The change in which model of RAD fitted best, as sample size was reduced, varied between sites and between habitat types. At the smallest sample sizes, the Zipf-Mandelbrot model often fitted, and in the slack sites the Broken stick also, though neither fitted (in the vegetation examined) at larger spatial scales. It is concluded the RAD is affected by species richness and by spatial scale, in ways that currently do not enable simple prediction. RADs can theoretically give information on the processes such as resource partitioning, immigration and competition that have structured the community, but they are a blunt tool for this purpose.     

Body size patterns in Drosophila inhabiting a mesocosm: interactive effects of spatial variation in temperature and abundance

Body size is a major component of fitness. However, the relative contributions of different factors to optimal size, and the determinants of spatial and temporal variation in size, have not been fully established empirically. Here, we use a mesocosm of a Drosophilidae assemblage inhabiting decaying nectarines to investigate the influence of spatial variation in temperature on adult body size in Drosophila simulans Sturtevant. Two treatments were established; one in the sun where developing larvae were exposed to high temperatures and the other in the shade where temperature conditions were milder. The simple developmental effects of temperature differences (i.e. larger flies are likely to emerge from cooler environments), or the simple effects of stressful temperatures (i.e. high temperatures yield wing abnormalities and smaller flies), were overridden by interactive effects between temperature and larval density. Emergences were lower in the sun than shade, probably as a result of temperature-induced mortality. However, flies attained the same final sizes in the shade and sun. In addition, abnormally winged flies were clustered in the shaded treatments. In the shade treatments, where emergences were higher than in the sun, stressful conditions as a result of high larval density likely resulted in wing abnormalities and small size. Consequently, there was little spatial variation in size across the mesocosm, but substantial spatial variation in abundance. Under natural conditions both mortality and non-lethal effects of temperature and/or crowding are likely to play a role in the evolution of body size.     

Relationship type affects the reliability of dispersal distance estimated using pedigree inferences in partially sampled populations: A case study involving invasive American mink in Scotland

Estimating dispersal—a key parameter for population ecology and management—is notoriously difficult. The use of pedigree assignments, aided by likelihood‐based software, has become popular to estimate dispersal rate and distance. However, the partial sampling of populations may produce false assignments. Further, it is unknown how the accuracy of assignment is affected by the genealogical relationships of individuals and is reflected by software‐derived assignment probabilities. Inspired by a project managing invasive American mink (Neovison vison), we estimated individual dispersal distances using inferred pairwise relationships of culled individuals. Additionally, we simulated scenarios to investigate the accuracy of pairwise inferences. Estimates of dispersal distance varied greatly when derived from different inferred pairwise relationships, with mother–offspring relationship being the shortest (average = 21 km) and the most accurate. Pairs assigned as maternal half‐siblings were inaccurate, with 64%–97% falsely assigned, implying that estimates for these relationships in the wild population were unreliable. The false assignment rate was unrelated to the software‐derived assignment probabilities at high dispersal rates. Assignments were more accurate when the inferred parents were older and immigrants and when dispersal rates between subpopulations were low (1% and 2%). Using 30 instead of 15 loci increased pairwise reliability, but half‐sibling assignments were still inaccurate (>59% falsely assigned). The most reliable approach when using inferred pairwise relationships in polygamous species would be not to use half‐sibling relationship types. Our simulation approach provides guidance for the application of pedigree inferences under partial sampling and is applicable to other systems where pedigree assignments are used for ecological inference.     

Geographical variation in the spatial synchrony of a forest-defoliating insect: isolation of environmental and spatial drivers

Despite the pervasiveness of spatial synchrony of population fluctuations in virtually every taxon, it remains difficult to disentangle its underlying mechanisms, such as environmental perturbations and dispersal. We used multiple regression of distance matrices (MRMs) to statistically partition the importance of several factors potentially synchronizing the dynamics of the gypsy moth, an invasive species in North America, exhibiting outbreaks that are partially synchronized over long distances (approx. 900 km). The factors considered in the MRM were synchrony in weather conditions, spatial proximity and forest-type similarity. We found that the most likely driver of outbreak synchrony is synchronous precipitation. Proximity played no apparent role in influencing outbreak synchrony after accounting for precipitation, suggesting dispersal does not drive outbreak synchrony. Because a previous modelling study indicated weather might indirectly synchronize outbreaks through synchronization of oak masting and generalist predators that feed upon acorns, we also examined the influence of weather and proximity on synchrony of acorn production. As we found for outbreak synchrony, synchrony in oak masting increased with synchrony in precipitation, though it also increased with proximity. We conclude that precipitation could synchronize gypsy moth populations directly, as in a Moran effect, or indirectly, through effects on oak masting, generalist predators or diseases.     

Type and spatial structure of distribution data and the perceived determinants of geographical gradients in ecology: the species richness of African birds

Aim Studies exploring the determinants of geographical gradients in the occurrence of species or their traits obtain data by: (1) overlaying species range maps; (2) mapping survey‐based species counts; or (3) superimposing models of individual species’ distributions. These data types have different spatial characteristics. We investigated whether these differences influence conclusions regarding postulated determinants of species richness patterns. Location Our study examined terrestrial bird diversity patterns in 13 nations of southern and eastern Africa, spanning temperate to tropical climates. Methods Four species richness maps were compiled based on range maps, field‐derived bird atlas data, logistic and autologistic distribution models. Ordinary and spatial regression models served to examine how well each of five hypotheses predicted patterns in each map. These hypotheses propose productivity, temperature, the heat–water balance, habitat heterogeneity and climatic stability as the predominant determinants of species richness. Results The four richness maps portrayed broadly similar geographical patterns but, due to the nature of underlying data types, exhibited marked differences in spatial autocorrelation structure. These differences in spatial structure emerged as important in determining which hypothesis appeared most capable of explaining each map's patterns. This was true even when regressions accounted for spurious effects of spatial autocorrelation. Each richness map, therefore, identified a different hypothesis as the most likely cause of broad‐scale gradients in species diversity. Main conclusions Because the ‘true’ spatial structure of species richness patterns remains elusive, firm conclusions regarding their underlying environmental drivers remain difficult. More broadly, our findings suggest that care should be taken to interpret putative determinants of large‐scale ecological gradients in light of the type and spatial characteristics of the underlying data. Indeed, closer scrutiny of these underlying data — here the distributions of individual species — and their environmental associations may offer important insights into the ultimate causes of observed broad‐scale patterns.     

Temporal variability in the spatial and environmental determinants of functional metacommunity organization – stream fish in a human‐modified landscape

1. Quantifying the relative importance of environmental filtering versus regional spatial structuring has become an intensively studied area in the context of metacommunity ecology. However, most studies have evaluated the role of environmental and spatial processes using taxonomic data sets of single snapshot surveys. 2. Here, we examined temporal changes in patterns and possible processes behind the functional metacommunity organization of stream fishes in a human‐modified landscape. Specifically, we (i) studied general changes in the functional composition of fish assemblages among 40 wadeable stream sites during a 3‐year study period in the catchment area of Lake Balaton, Hungary, (ii) quantified the relative importance of spatial and environmental factors as determinants of metacommunity structure and (iii) examined temporal variability in the relative role of spatial and environmental processes for this metacommunity. 3. Partial triadic analysis showed that assemblages could be effectively ordered along a functional gradient from invertebrate consuming species dominated by the opportunistic life‐history strategy, to assemblages with a diverse array of functional attributes. The analysis also revealed that functional fish assemblage structure was moderately stable among the sites between the sampling periods. 4. Despite moderate stability, variance partitioning using redundancy analyses (RDA) showed considerable temporal variability in the contribution of environmental and spatial factors to this pattern. The analyses also showed that environmental variables were, in general, more important than spatial ones in determining metacommunity structure. Of these, natural environmental variables (e.g. altitude, velocity) proved to be more influential than human‐related effects (e.g. pond area, % inhabited area above the site, nutrient enrichment), even in this landscape with relatively low variation in altitude and stream size. 5. Pond area was, however, the most important human stressor variable that was positively associated with the abundance of non‐native species with diverse functional attributes. The temporal variability in the relative importance of environmental and spatial factors was probably shaped by the release of non‐native fish from fish ponds to the stream system during flood events. 6. To conclude, both spatial processes and environmental control shape the functional metacommunity organization of stream fish assemblages in human‐modified landscapes, but their importance can vary in time. We argue, therefore, that metacommunity studies should better consider temporal variability in the ecological mechanisms (e.g. dispersal limitation, species sorting) that determine the dynamics of landscape‐level community organization.     

Monitoring and modeling the invasion of the fast spreading alien Senecio inaequidens DC. in an alpine region

We modeled the distribution of the South African alien Senecio inaequidens DC. in the Aosta Valley, Western Italian Alps, using data extracted from the Regional floristic database and from an intensive field survey carried out in years 2009–2010. The aims of the work were (1) to evaluate whether the species is in the introduction, colonization, or establishment stage of invasion, (2) to detect the environmental factors that drive the invasion process, and (3) to highlight the potential range of distribution of the alien species. The modeling framework was a stepwise generalized linear model (GLM), using gridded presence/absence data and environmental predictors such as topography, climate, land use, and anthropogenic and natural disturbances. GLM were fit both with and without an additional independent variable to take into account current dispersal limitations. S. inaequidens displayed a very fast spread in the Aosta Valley in the years 1990–2010. The species was positively associated with roads and rivers, southern slopes, and negatively with elevation. However, it was found at an elevation of 1600 m, showing the ability to reach higher elevations than those observed for other invasive alien species, and confirming to be pre-adapted to mountain conditions. The difference between the species distribution models, with and without dispersal constraints, suggested that the availability of seed sources still limits the potential distribution of the species, rather than the environmental variables, and that the realized regional niche differs to a great extent from the equilibrium niche. When limitations to the seed source cease (i.e., in the establishment stage), the species will likely invade large areas that are currently characterized by pastures and grasslands with native species of high agricultural importance. The invasion of S. inaequidens should therefore be considered a serious threat, due to its potential to invade mountain regions, and in particular to colonize habitats used for grazing and forage, thus leading to a high risk for cattle and human health. We discuss the relevance of the results both concerning communication with the public and to support local eradication and control activities. The inclusion of S. inaequidens in the “black list” of the regional law for the conservation of alpine flora (L.R. 45/2009) will help to transfer the information and support invasion control, in particular at medium elevations.     

Local population genetic structure of the montane caddisfly Drusus discolor is driven by overland dispersal and spatial scaling

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Modelling the spatial dynamics of a recovering species: the Grey Heron Ardea cinerea in France

In many countries, the implementation of measures to protect declining species has led to the recolonization of some areas by bird species within their historic range. There is, however, a lack of quantitative tools for exploring and projecting such large-scale dynamics. Here, we present an approach that makes use of both census and habitat suitability data and integrates them into a spatially explicit model of colonization dynamics (cellular automaton). The Grey Heron Ardea cinerea , which has recently recolonized much of France following legal protection measures, was used as a focal species. Data analysis allowed us to quantify the positive effects of habitat quality on the probability of colonization, and to uncover both positive and negative density-dependent colony settlement processes. Projective simulations then allowed us to determine an inflection point in global colonization dynamics and predict a significant reduction in the colonization rate during the next 50 years. This makes it possible to project the spatial distribution of the species at different timescales, and to estimate the carrying capacity of France for the species (around 1500 colonies).     

Integrating species distribution models and interacting particle systems to predict the spread of an invasive alien plant

Aim We demonstrate how to integrate two widely used tools for modelling the spread of invasive plants, and compare the performance of the combined model with that of its individual components using the recent range dynamics of the invasive annual weed Ambrosia artemisiifolia L. Location Austria. Methods Species distribution models, which deliver habitat‐based information on potential distributions, and interacting particle systems, which simulate spatio‐temporal range dynamics as dependent on neighbourhood configurations, were combined into a common framework. We then used the combined model to simulate the invasion of A. artemisiifolia in Austria between 1990 and 2005. For comparison, simulations were also performed with models that accounted only for habitat suitability or neighbourhood configurations. The fit of the three models to the data was assessed by likelihood ratio tests, and simulated invasion patterns were evaluated against observed ones in terms of predictive discrimination ability (area under the receiver operating characteristic curve, AUC) and spatial autocorrelation (Moran’s I). Results The combined model fitted the data significantly better than the single‐component alternatives. Simulations relying solely on parameterized spread kernels performed worst in terms of both AUC and spatial pattern formation. Simulations based only on habitat information correctly predicted infestation of susceptible areas but reproduced the autocorrelated patterns of A. artemisiifolia expansion less adequately than did the integrated model. Main conclusions Our integrated modelling approach offers a flexible tool for forecasts of spatio‐temporal invasion patterns from landscape to regional scales. As a further advantage, scenarios of environmental change can be incorporated consistently by appropriately updating habitat suitability layers. Given the susceptibility of many alien plants, including A. artemisiifolia, to both land use and climate changes, taking such scenarios into account will increasingly become relevant for the design of proactive management strategies.     

Distribution of aquatic vascular plants in lowland rivers: separating the effects of local environmental conditions, longitudinal connectivity and river basin isolation

1. Changes in species distributions along rivers have rarely been observed independently of changes in environmental conditions and meaningful comparison between different catchments is made difficult by the limited geographical distribution of species. This study presents a new approach to quantify the effect of the spatial structure of lowland river networks on aquatic plant distribution and to explore the potential underlying processes using species life‐history characteristics. 2. Twenty‐five species of aquatic vascular plants recorded in 62 sites across five calcareous river basins were used to investigate (i) the temporal turnover of plant species, (ii) the habitat utilisation of species, (iii) the trade‐offs between different plant life‐history characteristics and (iv) the relationship between species life‐history characteristics and habitat utilisation. 3. The annual plant turnover within a 3‐year period was, although significant, extremely low. It suggests that results from spatial surveys conducted over 3 years should not be undermined by temporal changes. 4. Spatial connectivity along and between rivers was more important than in‐channel physical characteristics in shaping species assemblages. Neither chemical factors (ammonium, phosphate) nor extrinsic biotic competitors (filamentous green algae) significantly influenced plant distribution. 5. The most common combinations of life‐history characteristics were neither related to environmental conditions nor to spatial isolation. Instead, they could reflect natural selection processes associated with larger scales than those considered in this study. 6. Plant distribution was most strongly related to the dispersal and regeneration abilities of the plants, supporting the hypotheses relating to longitudinal connectivity. The hypothesis that different growth forms would be associated with different in‐channel physical features was not verified. As expected, there were no substantial differences in plant life‐history characteristics between river basins.     

Patterns of woody plant species richness in the Iberian Peninsula: environmental range and spatial scale

Aim Climate‐based models often explain most of the variation in species richness along broad‐scale geographical gradients. We aim to: (1) test predictions of woody plant species richness on a regional spatial extent deduced from macro‐scale models based on water–energy dynamics; (2) test if the length of the climate gradients will determine whether the relationship with woody species richness is monotonic or unimodal; and (3) evaluate the explanatory power of a previously proposed ‘water–energy’ model and regional models at two grain sizes. Location The Iberian Peninsula. Methods We estimated woody plant species richness on grid maps with c. 2500 and 22,500 km² cell size, using geocoded data for the individual species. Generalized additive models were used to explore the relationships between richness and climatic, topographical and substrate variables. Ordinary least squares regression was used to compare regional and more general water–energy models in relation to grain size. Variation partitioning by partial regression was applied to find how much of the variation in richness was related to spatial variables, explanatory variables and the overlap between these two. Results Water–energy dynamics generate important underlying gradients that determine the woody species richness even over a short spatial extent. The relationships between richness and the energy variables were linear to curvilinear, whereas those with precipitation were nonlinear and non‐monotonic. Only a small fraction of the spatially structured variation in woody species richness cannot be accounted for by the fitted variables related to climate, substrate and topography. The regional models accounted for higher variation in species richness than the water–energy models, although the water–energy model including topography performed well at the larger grain size. Elevation range was the most important predictor at all scales, probably because it corrects for ‘climatic error’ due to the unrealistic assumption that mean climate values are evenly distributed in the large grid cells. Minimum monthly potential evapotranspiration was the best climatic predictor at the larger grain size, but actual evapotranspiration was best at the smaller grain size. Energy variables were more important than precipitation individually. Precipitation was not a significant variable at the larger grain size when examined on its own, but was highly significant when an interaction term between itself and substrate was included in the model. Main conclusions The significance of range in elevation is probably because it corresponds to several aspects that may influence species diversity, such as climatic variability within grid cells, enhanced surface area, and location for refugia. The relative explanatory power of energy and water variables was high, and was influenced by the length of the climate gradient, substrate and grain size of the analysis. Energy appeared to have more influence than precipitation, but water availability is also determined by energy, substrate and topographic relief.     

Influence of edaphic factors on the spatial structure of inland halophytic communities: a case study in China

Abstract. In order to understand the influence of edaphic factors on the spatial structure of inland halophytic plant communities, a 2.6 km² study site, located on the lower fringe of the alluvial fan of the Hutubi River, in an arid region of China, was sampled and mapped. 105 patches were found to be homogeneous in species composition. Plant species and their coverage were recorded in each patch. 45 patches were randomly selected for the measurement of edaphic variables. A map with quadrat locations and boundaries of patches was digitized into a GIS and related to the vegetation and edaphic data matrices. CCA was used to evaluate the relative importance of edaphic factors in explaining the variation of the species assemblages and to identify the ecological preferences of species. The spatial structure of the communities and the main edaphic factors were analyzed using correlograms, Mantel correlograms and clustering under constraint of spatial contiguity. Gradient analysis showed that there are two distinct vegetation gradients in the study area, one of which is determined mainly by soil moisture (determined by depth to the water table), and the other by soil salinity (determined by electrical conductivity and hydrolytic alkalinity of the first soil layer). However, spatial analyses showed that at the sampling scale the halophytic communities in the study area are structured along one main spatial gradient determined by the water table level. Similar spatial autocorrelation structures between the factors related to the first soil layer and the communities, given our sampling scale, could not be detected. Our results suggest that the relative importance of the effects of different edaphic factors on the spatial structure of halophytic communities is scale-dependent. The partitioning of species variation indicates that in addition to edaphic factors, other factors, such as biotic interactions, may play an important role in structuring these communities.     

Disentangling the relative effects of environmental versus human factors on the abundance of native and alien plant species in Mediterranean sandy shores

Aim Mediterranean coastal sand dunes are characterized by both very stressful environmental conditions and intense human pressure. This work aims to separate the relative contributions of environmental and human factors in determining the presence/abundance of native and alien plant species in such an extreme environment at a regional scale. Location 250 km of the Italian Tyrrhenian coast (Region Lazio). Methods We analysed alien and native plant richness and fitted generalized additive models in a multimodel‐inference framework with comprehensive randomizations to evaluate the relative contribution of environmental and human correlates in explaining the observed patterns. Results Native and alien richness are positively correlated, but different variables influence their spatial patterns. For natives, human population density is the most important factor and is negatively related to richness. Numbers of natives are unexpectedly lower in areas with a high proportion of natural land cover (probably attributable to local farming practices) and, to a lesser degree, affected by the movement of the coastline. On the other hand, alien species richness is strongly related to climatic factors, and more aliens are found in sectors with high rainfall. Secondarily, alien introductions appear to be related to recent urban sprawl and associated gardening. Main conclusions Well‐adapted native species in a fragile equilibrium with their natural environment are extremely sensitive to human‐driven modifications. On the contrary, for more generalist alien species, the availability of limited resources plays a predominant role.     

天童常绿阔叶林中常绿与落叶物种的物种多度分布格局

物种多度分布是对群落内不同物种多度情况的数量描述, 作为理解群落性质的基石, 其形成机制受到广泛关注。常绿与落叶物种是两类有着不同物候性状与生长策略的物种集合, 它们普遍共存于常绿阔叶林中。在天童20 ha常绿阔叶林动态监测样地内, 虽然常绿物种在物种多度和胸高断面积等指标上占有绝对优势, 但其在物种丰富度上却不及落叶物种。分析两者在常绿阔叶林中的物种多度分布特征, 能够为理解常绿阔叶林内物种多样性的维持机制提供一个全新的视角。为此, 我们基于天童样地的植被调查数据, 一方面利用累积经验分布函数对两类生活型植物的物种多度分布进行描述, 使用Kolmogorov-Smirnov检验(K-S检验)判断其差异性; 另一方面, 采用纯统计模型、生态位模型和中性理论模型对二者的物种多度分布曲线进行拟合, 并基于K-S检验的结果以及AIC值进行最优模型的筛选。结果显示: (1)常绿与落叶物种的物种多度分布曲线间并无显著差异。(2)在选用的3类模型中, 中性理论模型对于两类物种多度分布曲线的拟合效果都最好, 而生态位模型的拟合效果则一般。从上述结果可以看出, 尽管常绿与落叶物种在物种数量和多度等方面均存在差异, 但它们却有着近似的物种多度分布格局以及相近的多样性维持机制。然而, 鉴于模型拟合的结果只能作为理解群落多样性构建机制的必要非充分条件, 故而只能初步判定中性过程对于常绿与落叶物种的物种多样性格局影响更大, 却不能排除或衡量诸如生态位分化等其他过程在两类生活型多样性格局形成中的贡献。