Exotic plant species invade diversity hot spots: the alien flora of northwestern Kenya

We analysed the distribution of native and alien plant species across 20 ecogeographic zones of northwestern Kenya. The source pool for the majority of aliens was Europe and America. Thus, the source pool has a biogeographic bias which explains the low proportion of aliens in the tropics: most species in the European or American source pool are not well adapted to tropical conditions. As expected, native and alien plant species showed an area effect. Correcting for this area effect. species rich zones showed a higher proportion of alien plant species in their flora. At the analysed scale, species richness of native plant communities does not increase the resistance to invasions and alien plant species invade diversity hotspots. Compared to the other ecogeographic zone, the urban area around Nairobi showed an increased richness in alien and native plant species. This is very similar to findings in Europe, although the history of urbanisation is much shorter in Kenya. The species turnover between zones (β-diversity) shows a similar pattern in native and alien plant species. Within a very short time scale the alien plant species mapped the biogeographic patterns of natives, although the geography of human activities influences the propagule pressure.     

Applying the dark diversity concept to plants at the European scale

Large‐scale biodiversity maps are essential to macroecology. However, between‐region comparisons can be more useful if patterns of observed species richness are supplemented by variations in dark diversity – the absent portion of the species pool. We aim to quantify and map plant diversity across Europe by using a measure that accounts for both observed and dark diversity. To do this we need to delimit suitable species pools, and evaluate the potential and limitation of a large‐scale dataset. We used Atlas Florae Europaeae (ca 20% of European plant species mapped within 50 × 50 km grid cells) and defined for each grid cell several species pools by applying various geographical and environmental filters: geographic species pool (number of species within 500 km radius), biogeographic species pool (additionally incorporating species distribution patterns, i.e. dispersion fields), site‐specific species pool (additionally integrating environmental preferences of species based on species co‐occurrence). We integrated dark diversity and observed diversity at a relative scale to calculate the completeness of site diversity: logistic expression of observed and dark diversity. We tested whether our results are robust against regional variation in data availability. We used independent regional databases to test if Atlas Florae Europaeae is a representative subset of total species richness. Environmental filtering was the most influential determinant of species pool size with more species filtered out in southern Europe. Both observed and dark diversity adhered to the well‐known latitudinal gradient, but completeness of site diversity varied throughout Europe with no latitudinal trend. Dark diversity patterns were fairly insensitive to variations in regional sampling intensity. Atlas Florae Europaeae represented well the total variation in plant diversity. In summary, dark diversity and completeness of site diversity add valuable information to broad‐scale diversity patterns since observed diversity is expressed at a relative scale.     

Species pools,community completeness and invasion: disentangling diversity effects on the establishment of native and alien species

Invasion should decline with species richness, yet the relationship is inconsistent. Species richness, however, is a product of species pool size and biotic filtering. Invasion may increase with richness if large species pools represent weaker environmental filters. Measuring species pool size and the proportion realised locally (completeness) may clarify diversity‐invasion relationships by separating environmental and biotic effects, especially if species’ life‐history stage and origin are accounted for. To test these relationships, we added seeds and transplants of 15 native and alien species into 29 grasslands. Species pool size and completeness explained more variation in invasion than richness alone. Although results varied between native and alien species, seed establishment and biotic resistance to transplants increased with species pool size, whereas transplant growth and biotic resistance to seeds increased with completeness. Consequently, species pools and completeness represent multiple independent processes affecting invasion; accounting for these processes improves our understanding of invasion.     

Community Completeness: Linking Local and Dark Diversity within the Species Pool Concept

Biodiversity of ecological communities has been examined widely. However, comparisons of observed species richness are limited because they fail to reveal what part of the differences are caused by natural variation in species pool size and what part is due to dark diversity – the absence of suitable species from a species pool. In other words, conventional biodiversity inventories do not convey information about how complete local plant communities are. We therefore propose the community completeness concept – a new perspective on the species pool framework. In order to ascertain community completeness, we need to estimate the extent of dark diversity, for which several methods are under development. We recommend the Community Completeness Index based on a log-ratio (or logistic) expression: ln(observed richness/dark diversity). This metric offers statistical advantages over other methods (e.g. the proportion of observed richness from the species pool). We discuss how community completeness can be related to long-term and successional community stability, landscape properties and disturbance patterns as well as to a variety of biotic interactions within and among trophic levels. The community completeness concept is related to but distinctive from the alpha-beta-gamma diversity approach and the community saturation phenomenon. The Community Completeness Index is a valuable metric for comparing biodiversity of different ecosystems for nature conservation. It can be used to measure the success of ecological restoration and vulnerability to invasion by alien species. In summary, community completeness is an interface between observed local observed species richness and dark diversity, which can be useful both in theoretical and applied biodiversity research.     

Distorted perception of the spatial distribution of plant diversity through uneven collecting efforts: the example of Asteraceae in Australia

Aim Our aims were (1) to compare observed, estimated and predicted patterns of species richness using the Australian native Asteraceae as an example, (2) to identify candidates for hotspots of diversity for the study group, and (3) to examine the distortion of our perception of the spatial distribution of species richness through uneven or misdirected sampling efforts. Location Australia. Methods Based on data from Australia’s Virtual Herbarium, we calculated and visualized observed species richness, the Chao1 estimate of richness, the C index of collecting completeness, and an estimate of richness derived from environmental niche modelling for grid cells at a resolution of 1°. The 20 cells with the highest diversity values were used to define hotspots of diversity. Results Uneven collecting activity results in misleading diversity patterns for the family Asteraceae. While observed species richness is much higher in central Australia than in other parts of the arid interior, this is an artefact resulting from the area being a hotspot of collecting activity. The mountain ranges of south‐eastern Australia and Tasmania are candidates for unbiased hotspots of species richness. Main conclusions Vast areas of the Australian interior are insufficiently sampled on a local scale, although most of them can be expected to be relatively species poor. Some areas in the south‐east and south‐west of the continent remain undersampled relative to their high species richness. Observed species numbers, estimators and environmental niche‐modelling all have their unique advantages and disadvantages for the inference of patterns of diversity.     

Contrasting response of native and alien plant species richness to environmental energy and human impact along alpine elevation gradients

Aim We tested whether the species–energy and species–human relationships vary between native and both naturalized and casual alien species richness when other environmental variables had been taken into account. Location Trento Province, a region (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided into 156 contiguous (c. 37.5 km²) cells and ranging in elevation from 66 to 3769 m. Methods Data were separated into three subsets, representing richness of natives, naturalized aliens and casual aliens and separately related to temperature, human population and various environmental correlates of plant species diversity. We applied ordinary least squares and simultaneous autoregressive regressions to identify potential contrasting responses of the three plant status subsets and hierarchical partitioning to evaluate the relative importance of the predictor variables. Results Variation in alien plant species richness along the region was almost entirely explained by temperature and human population density. The relationships were positive but strongly curvilinear. Native species richness was less strongly related to either factor but was positively related to the presence of calcareous bedrock. Native species richness had a decelerating positive relationship with temperature (R²= 55%), whereas naturalized and casual aliens had a positive accelerating relationship explaining 86% and 62% of the variation in richness, respectively. Native species richness had a positive decelerating relationship with population density (R²= 42%), whilst both alien subsets had a positive accelerating relationship. Main conclusions Alien species richness was higher in areas with the most rich and diverse assemblages of native species. Areas at high altitudes are not especially prone to alien invasion due to energy constraints, low propagule pressure and disturbance, even considering a potential increased in temperature. Thus, if we consider future environmental change, we should expect a stronger response of aliens than natives in the currently warm, urbanized, low‐altitude areas than in cold, high‐altitude areas where human population density is low.     

Assessing the level of plant invasion: A multi-scale approach based on vegetation plots

Recent studies highlight the importance of selecting the appropriate scale and indices of invasion level for evaluating the abundance and impact of alien plants. Our survey considers the use of vegetation plot databases compared with floristic checklists to address invasion patterns regarding alien–native relationships across vegetation types by means of a multi-scale approach. We analysed the alien–native richness relationship in 1077 vegetation plots from the Basque Country (N. Spain) at ecosystem level and phytosociological class and alliance levels. According to our results, the alien species richness (Alo)–native species richness (Nat) relationship is variable and depends not only on the scale but also on the vegetation type. In contrast with other multi-scale approaches, no negative correlation has been detected at any studied level. The strong correlation existing between plot number and cumulative Alo and cumulative Nat highlights the constraints of using checklists to generalize invasion patterns. Our results demonstrate that the combined use of both relative alien species richness and relative alien species cover facilitates the understanding of invasion patterns across plant communities at different scales. In addition to climate, disturbance and propagule pressure, habitat type proved to be an important filter for alien species, capable of explaining such patterns.     

哈尔滨市主要城市绿地特征和耦合关系

采用样方法调查了哈尔滨市17块绿地的植物多样性,研究城市绿地生态系统中本土植物多样性对外来植物的影响规律。结果如下:哈尔滨市绿地有75种外来植物,外来栽培植物42种,外来非栽培植物33种。在哈尔滨市绿地植物群落中,外来栽培植物的平均盖度、平均重要值和平均出现频度均较高;外来非栽培种的平均盖度、平均重要值较低,而平均出现频度较高。绿地本土植物物种数与外来栽培种、外来非栽培种的物种数极显著正相关(P<0.01),与外来非栽培种的平均出现频度显著负相关(P<0.05)。绿地乔木层本土植物的香农威尔指数、物种丰富度与外来栽培种、外来非栽培种的物种数均显著负相关(P<0.05),乔木层本土植物的均匀度与外来非栽培种的平均出现频度极显著负相关(P<0.01);灌木层本土植物的物种丰富度与外来栽培种的平均重要值显著负相关(P<0.05);草本层本土植物的均匀度与外来非栽培种的平均相对盖度显著正相关(P<0.05)。随绿地总面积的增加,外来栽培种和外来非栽培种的物种数增加,平均出现频度均呈现显著降低的趋势。随建成时间增加,外来栽培种的平均出现频度呈非线形变化,外来非栽培种的平均出现频度呈现降低的趋势。应提高哈尔滨市绿地本土植物的多样性,严格控制新建成的、总面积较小的绿地中外来植物的引种栽培管理,以防止外来植物发展成为危害本地生态系统的入侵植物。     

Landscape context modulates alien plant invasion in Mediterranean forest edges

Natural habitats in human-altered landscapes are especially vulnerable to biological invasions, especially in their edges. We aim to understand the influence of landscape and local characteristics on biological invasions by exploring the level of plant invasion and alien species traits in forest edges in highly urbanized landscapes. We identified all plant species in 73 paired plots in the edge and 50 m towards the interior of the forest. We explored the association between alien species richness and similarity in species composition between edge and interior plots with landscape and local variables, using generalized linear models and variance partitioning techniques. Then, we performed Fourth-corner analyses to explore the association between alien plant traits and local and landscape variables. In contrast to native species richness, alien species richness was more affected by the surrounding landscape than by the local characteristics of the edge. Road proximity was positively associated with alien species richness and proportion and was its most important correlate, whereas disturbance was negatively associated with native species richness and was its most influential factor. Alien plant traits were also primarily associated with landscape characteristics. For instance, species of Mediterranean origin and introduced for agriculture were associated with higher agriculture use in the landscape. Our findings suggest that risk analyses of habitat vulnerability to invasion must consider the landscape context in order to successfully predict highly invaded areas and identify potentially successful invaders.     

Widespread plant species: natives versus aliens in our changing world

Estimates of the level of invasion for a region are traditionally based on relative numbers of native and alien species. However, alien species differ dramatically in the size of their invasive ranges. Here we present the first study to quantify the level of invasion for several regions of the world in terms of the most widely distributed plant species (natives vs. aliens). Aliens accounted for 51.3% of the 120 most widely distributed plant species in North America, 43.3% in New South Wales (Australia), 34.2% in Chile, 29.7% in Argentina, and 22.5% in the Republic of South Africa. However, Europe had only 1% of alien species among the most widespread species of the flora. Across regions, alien species relative to native species were either as well-distributed (10 comparisons) or more widely distributed (5 comparisons). These striking patterns highlight the profound contribution that widespread invasive alien plants make to floristic dominance patterns across different regions. Many of the most widespread species are alien plants, and, in particular, Europe and Asia appear as major contributors to the homogenization of the floras in the Americas. We recommend that spatial extent of invasion should be explicitly incorporated in assessments of invasibility, globalization, and risk assessments.     

Local vs regional factors as determinants of the invasibility of indigenous forest fragments by alien plant species

Both local and regional filters can determine the invasion of alien species into native plant communities. However, their relative importance is essentially unknown. We used plot data from fragments of indigenous forests in southeastern New Zealand to infer which factors are important in explaining invasibility, measured as alien species richness. Twenty-eight predictor variables comprising both local factors (stand structure and soil) and regional ones (climate and land cover) were assessed. Reduction or increase in deviance in linear models was assessed, both individually and with a forward and backward stepwise variable selection procedure using the Akaike information criterion (AIC).
We found that higher alien species richness was mainly associated with forest fragments of small area in warm and dry climates and where there were only small areas of surrounding indigenous forest. Local soil and stand structure variables had considerably smaller effects on alien species richness than the regional land cover and climate variables. Alien species richness showed no relationship with native species richness. We conclude that in the forest fragments investigated here, of the variables included in the analyses, regional land cover and climate variables are potentially important drivers for alien species richness at plot level. This has implications for projections of alien species spread in the future under different climate change and land use scenarios.     

Large‐scale dark diversity estimates: new perspectives with combined methods

Large‐scale biodiversity studies can be more informative if observed diversity in a study site is accompanied by dark diversity, the set of absent although ecologically suitable species. Dark diversity methodology is still being developed and a comparison of different approaches is needed. We used plant data at two different scales (European and seven large regions) and compared dark diversity estimates from two mathematical methods: species co‐occurrence (SCO) and species distribution modeling (SDM). We used plant distribution data from the Atlas Florae Europaeae (50 × 50 km grid cells) and seven different European regions (10 × 10 km grid cells). Dark diversity was estimated by SCO and SDM for both datasets. We examined the relationship between the dark diversity sizes (type II regression) and the overlap in species composition (overlap coefficient). We tested the overlap probability according to the hypergeometric distribution. We combined the estimates of the two methods to determine consensus dark diversity and composite dark diversity. We tested whether dark diversity and completeness of site diversity (log ratio of observed and dark diversity) are related to various natural and anthropogenic factors differently than simple observed diversity. Both methods provided similar dark diversity sizes and distribution patterns; dark diversity is greater in southern Europe. The regression line, however, deviated from a 1:1 relationship. The species composition overlap of two methods was about 75%, which is much greater than expected by chance. Both consensus and composite dark diversity estimates showed similar distribution patterns. Both dark diversity and completeness measures exhibit relationships to natural and anthropogenic factors different than those exhibited by observed richness. In summary, dark diversity revealed new biodiversity patterns which were not evident when only observed diversity was examined. A new perspective in dark diversity studies can incorporate a combination of methods.     

Areas of high conservation value at risk by plant invaders in Georgia under climate change

Invasive alien plants (IAP) are a threat to biodiversity worldwide. Understanding and anticipating invasions allow for more efficient management. In this regard, predicting potential invasion risks by IAPs is essential to support conservation planning into areas of high conservation value (AHCV) such as sites exhibiting exceptional botanical richness, assemblage of rare, and threatened and/or endemic plant species. Here, we identified AHCV in Georgia, a country showing high plant richness, and assessed the susceptibility of these areas to colonization by IAPs under present and future climatic conditions. We used actual protected areas and areas of high plant endemism (identified using occurrences of 114 Georgian endemic plant species) as proxies for AHCV. Then, we assessed present and future potential distribution of 27 IAPs using species distribution models under four climate change scenarios and stacked single‐species potential distribution into a consensus map representing IAPs richness. We evaluated present and future invasion risks in AHCV using IAPs richness as a metric of susceptibility. We show that the actual protected areas cover only 9.4% of the areas of high plant endemism in Georgia. IAPs are presently located at lower elevations around the large urban centers and in western Georgia. We predict a shift of IAPs toward eastern Georgia and higher altitudes and an increased susceptibility of AHCV to IAPs under future climate change. Our study provides a good baseline for decision makers and stakeholders on where and how resources should be invested in the most efficient way to protect Georgia's high plant richness from IAPs.     

Contrasting alien and native plant species–area relationships: the importance of spatial grain and extent

Aim The proportion of alien plant species in floras is increasingly being used to indicate the threat of invasions to native species and/or the homogenization of biodiversity. However, this indicator is only valuable if it is independent of the spatial extent and grain of observation. This study tested the equivalence of native and alien species–area relationships (SARs) in order to assess the support for scale invariance in the proportion of alien species in floras. Location England, UK. Methods Nested SARs were generated by assessing the richness of native and alien plant species drawn from the New atlas of the British and Irish flora for six areas comprising 100, 400, 900, 1600, 2500 and 3600 km² with each larger area containing all smaller areas. Five replicate sets of nested areas encompassing northern, southern, eastern, western and central regions were chosen. For each set of nested areas, the log‐transformed species richness was regressed on log‐transformed area to fit a power function to the SAR. Results Native and alien plant SARs reveal consistent differences in slope, highlighting that the proportion of alien species is a function of spatial grain. Aliens are more rare than natives and have higher spatial turnover leading to faster accumulation of species as area increases. However, equivalent samples drawn from a larger spatial extent reveal similar alien and native SARs. Main conclusions The significant differential scale dependence in native and alien species richness observed in this study reflects dissimilar influences of regional drivers such as habitat, but potentially also propagule pressure and introduction history, that leads to the relative rarity and high spatial turnover of alien species. Maps of invasion hotspots that identify areas where the proportion of the alien flora is particularly high should therefore be treated with considerable caution since patterns across most grains used for species monitoring will be scale dependent.     

Grazing, Environmental Heterogeneity, and Alien Plant Invasions in Temperate Pampa Grasslands

Temperate humid grasslands are known to be particularly vulnerable to invasion by alien plant species when grazed by domestic livestock. The Flooding Pampa grasslands in eastern Argentina represent a well-documented case of a regional flora that has been extensively modified by anthropogenic disturbances and massive invasions over recent centuries. Here, we synthesise evidence from region-wide vegetation surveys and long-term exclosure experiments in the Flooding Pampa to examine the response of exotic and native plant richness to environmental heterogeneity, and to evaluate grazing effects on species composition and diversity at landscape and local community scales. Total plant richness showed a unimodal distribution along a composite stress/fertility gradient ranging several plant community types. On average, more exotic species occurred in intermediate fertility habitats that also contained the highest richness of resident native plants. Exotic plant richness was thus positively correlated with native species richness across a broad range of flood-prone grasslands. The notion that native plant diversity decreases invasibility was supported only for a limited range of species-rich communities in habitats where soil salinity stress and flooding were unimportant. We found that grazing promoted exotic plant invasions and generally enhanced community richness, whereas it reduced the compositional and functional heterogeneity of vegetation at the landscape scale. Hence, grazing effects on plant heterogeneity were scale-dependent. In addition, our results show that environmental fluctuations and physical disturbances such as large floods in the pampas may constrain, rather than encourage, exotic species in grazed grasslands.     

Distributions of exotic plants in eastern Asia and North America

Although some plant traits have been linked to invasion success, the possible effects of regional factors, such as diversity, habitat suitability, and human activity are not well understood. Each of these mechanisms predicts a different pattern of distribution at the regional scale. Thus, where climate and soils are similar, predictions based on regional hypotheses for invasion success can be tested by comparisons of distributions in the source and receiving regions. Here, we analyse the native and alien geographic ranges of all 1567 plant species that have been introduced between eastern Asia and North America or have been introduced to both regions from elsewhere. The results reveal correlations between the spread of exotics and both the native species richness and transportation networks of recipient regions. This suggests that both species interactions and human-aided dispersal influence exotic distributions, although further work on the relative importance of these processes is needed.     

Differential processes underlying the roadside distributions of native and alien plant assemblages

Although biological invasion often alters ecosystem properties and community composition at different scales, considerable uncertainty still exists regarding the underlying mechanisms that regulate the spread of alien species into new habitats. An alien invasion is generally achieved through multiple processes from multiple sources; this type of invasion often prevents us from understanding of the dispersal mechanisms. Here, we aim to disentangle the processes of alien invasion by focusing on a single migration source. We surveyed the distribution of alien and native plant species in Shiretoko National Park, located in northern Japan. We measured the coverage of each species and the environmental and spatial factors in 362 quadrats established along roadsides. We found 101 native species and 35 alien species (γ-diversity) throughout the quadrats. The local species richness (α-diversity) was higher for the alien species (6.1 species) than for the native species (3.2 species). There was a significant negative correlation in α-diversity between native and alien species. Moreover, the α-diversity and distance from the nearest town (migration source) showed a negative relationship for alien assemblages while the native assemblages showed the opposite trend. These results suggest that the alien species are expanding their distribution outward from the town, resulting in a decrease in the α-diversity of native species in localities close to the migration source. Overall, our results emphasize that roadsides could unintentionally act as corridors for alien species, even in protected areas. Careful consideration is thus needed for utilizing these human-created habitats even though they were designed for conservation and management purposes.     

Impacts of Myriophyllum aquaticum invasion in a Mediterranean wetland on plant and macro-arthropod communities

The invasion by alien macrophytes in aquatic ecosystems may produce a strong alteration of the native aquatic vegetation leading to heavy impacts for both plant and faunal native diversity. Myriophyllum aquaticum is an aquatic plant native of Southern America, invasive in several part of the world. We studied the effects of M. aquaticum invasion on plant and macro-arthropod communities in the canals around a protected wetland in the Mediterranean basin. We sampled plant and macro-arthropod communities in 10 transects in invaded and non-invaded tracts of the canals. We assessed the differences in plant and macro-arthropod species richness, diversity, taxonomic diversity and species composition between invaded and non-invaded habitats by means of univariate and multivariate analyses. Our study shows a significant loss of plant diversity between non-invaded to invaded sites, leading to communities numerically and taxonomically impoverished and highly divergent in the species composition. We also detected significant differences in arthropod species composition between invaded and non-invaded transects. Some taxa such as mosquitoes and malacostraca were more frequent in the M. aquaticum-dominated stands. Furthermore, the study shows a positive relation between invaded habitats and juvenile individuals of the invasive alien crayfish Procambarus clarkii.     

南宁老虎岭林区多种外来植物入侵程度与影响因子模型解释

了解森林环境中多种外来植物对多种环境因子的互作效应,可以更有针对性地应对外来入侵威胁。在南宁老虎岭林区分6个区进行样方调查,以样方所有外来植物的相对百分比作为外来植物入侵程度,利用以分区为随机截距的混合效应模型和一般线性模型来分析有关因素对入侵的影响,用R-effects包的互作效应图形化和数据提取来解释互作效应的复杂变化。多因素混合效应模型分析表明,路边对入侵的主效应为正且极显著(P=0.000),林冠郁蔽度和优势最大株高对入侵的主效应为负(P=0.000),土著物种丰富度对入侵的主效应不显著,但土著物种丰富度与路边的互作对入侵的效应极显著(P=0.007);路边的土著物种丰富度明显提升入侵抵抗性,但林内的土著物种丰富度只能微弱增加入侵抵抗性;林冠郁蔽度和优势最大株高的互作对入侵的效应极显著(P=0.004),但两个因素对入侵的限制作用非可加。一般线性模型分析表明,林龄和抚育时间对外来植物入侵的影响趋势不明显;未发现引进树种造林与乡土树种造林的林下外来植物入侵程度有差异;相对于林道的样方位置高低影响入侵程度,林道下方的样方较易被入侵。在监测或防控林业外来植物时,重点应放在低于林道的森林。     

Altitudinal distribution of native and alien plant species in roadside communities from central Argentina

Roadsides may homogenize the distribution of native species and act as corridors for the spread of alien taxa. We examined the variation in native and alien plant species richness and composition at two spatial scales defined by altitude and habitat type (edges and fill slopes), as well as the relationship between native and exotic species richness in roadside plant communities in mountains from central Argentina. Following a gradient from 1100 to 2200 m a.s.l. along a mountain road, plant species cover was recorded within sample plots of 30 m × 10 m systematically located at 100‐m altitude intervals on both roadside habitats. Although native species richness decreased with altitude and composition changed accordingly, the number of alien species peaked at both extremes of the elevation gradient and did not reflect an altitudinal replacement of chorological groups. The number of both native and alien species was higher in roadside edges, but a negative association between the richness of native and alien species occurred only on fill slopes, suggesting that roadside habitats differ in their susceptibility to plant species colonization and in the mechanisms driving native and alien species richness. Our results highlight the importance of altitude and roadside habitat as factors controlling plant species richness and composition along roadside communities in central Argentina. Although altitude acts as a filter for native plants, it apparently did not constrain the establishment of alien species along the studied roadsides, indicating that the influence of this road as a plant species corridor may increase with time, promoting the opportunities for aliens to expand their current distribution.