Patterns of alien plant diversity in the urban landscapes of global biodiversity hotspots: a case study from the Himalayas

In an age of Anthropocene, the urban landscapes are recognised as the ‘hotspots’ of human-mediated alien species introductions. As the cities provide an ideal natural experimental system to investigate the patterns of alien plant diversity in urban landscapes, the present study aimed to unravel the taxonomic, biogeographic and ecological patterns of alien flora of Srinagar—one of the largest urban centres in the Himalayan biodiversity hotspot. The alien flora of Srinagar comprises 325 species, constituting ca.35% of total flora of the city. Out of the 325 alien species documented, 157 species (43%) were recorded to be under cultivation, while 168 species (57%) were growing in the wild (i.e., outside cultivation); those growing in the wild, in turn, comprised 110 cultivation escapes and 58 accidentally introduced plant species. Biogeographically, two-third of the alien plant diversity reported from Srinagar is native to Asia-Temperate. This indicates that climatic similarity between Asia-Temperate and Kashmir Himalayas facilitate in flourishing similar floristic diversity. The study highlights a relatively higher proportion of herbaceous growth form in the aliens growing in the wild (80%) than those under cultivation (43%). Similarly, 82% of the alien species under cultivation had a perennial life span, but those growing in the wild were dominated by annuals (44%). Currently, 45 species are growing as casuals and 124 species are naturalised (including 105 naturalised non-invasive and 19 naturalised invasive). Along the continuum of casual-naturalised-invasive categories, the contribution of cultivation escapes and accidently introduced aliens contrastingly shows decreasing and increasing trends respectively. Interestingly, the results revealed that the human practice of stopping cultivation of alien escapes increased rapidly as we move along the continuum. Thus, the present study has investigated the patterns of alien plant diversity in the urban landscape of Srinagar, and the results obtained offer scientific insights toward better scientific understanding and management of plant invasions in this Himalayan city, with wider policy implications for neighbouring urbanised landscapes in the Himalayas and other mountainous regions across the world.     

Changes in diversity along a successional gradient in a Costa Rican upper montane Quercus forest

Changes in terrestrial vascular plant diversity along a successional gradient were studied in a Costa Rican upper montane Quercus forest. In 1991 and 1992 species presence and cover were recorded in 12 successional 0.1 ha forest plots. A total of 176 species in 122 genera and 75 families were found. Asteraceae was the most speciose family. With the help of TWINSPAN three successional phases were classified: (i) Early Secondary Forest (ESF, 145 spp.), (ii) Late Secondary Forest (LSF, 130 spp.) and (iii) Primary Forest (PF, 96 spp.). Detrended Correspondence Analysis (DCA) species ordination using DECORANA illustrates that different ecological species groups can be distinguished along the time sequence. Alpha diversity (Shannon-Wiener index, among others) in ESF and LSF was significantly greater than in PF. This is probably explained by downslope migration of numerous sub(alpine) species to cleared and recently abandoned montane sites. Beta diversity applying Sørensen's similarity coefficients declined during succession. Using linear regression, the minimum time required for floristic recovery following disturbance and abandonment was calculated at 65.9 years. A comparison with other studies shows that secondary forests in upper montane Costa Rica can be as diverse as in neotropical lowlands.     

Observations on the floristic,life-form,leaf-size spectra and habitat diversity of vegetation in the Bhimber hills of Kashmir Himalayas

Vegetation analysis provides the prerequisites to understand the overall community structure and function of any ecosystem and is a fundamental requirement for the precise evaluation of biodiversity. Although many studies have assessed floristic attributes of specific areas, there are still unexplored regions, as is the case of the mountain region in the Kashmir Himalayas. Current research highlighted the recent findings of the scientific characterization of floristic and ecological aspects on the forest flora found in the Bhimber hills, Pakistan. Floristically, a total of 93 species belonging to 80 genera in 41 families were recorded. The species distribution patterns across the families were disproportionate with half of the species contributed by 8 families and 25 families were monotypic. Based on the floristic analysis, Asteraceae was the largest family with 12% of species followed by Poaceae with (11%) species. PAST software, a multivariate ecological community analysis was used to classify the species similarities and differences among the different habitat types. According to the habitat wise distribution, 21% of species were growing in the natural forest habitat, while 15% of species were dispersed in highly distributed habitats along roadsides and 8% on pedestrians. In terms of functional diversity, the herbaceous growth form was dominant (58%). The biological spectrum revealed therophytes as the dominant life form as it indicates the disturbed habitat vegetation. The phytogeographical analysis revealed that the maximum (69%) species were native, while the minimum (31%) species were exotic. Thus, the study of these functional and habitat diversity patterns can significantly improve our understanding of the ecological aspects of the flora in the geographical location. This information may additionally be useful in devising management plans to ensure sustainable utilization and better management of forest landscapes in this Himalayan region.     

Comparing naturalized alien plants and recipient habitats across an east–west gradient in the Mediterranean Basin

Aim To investigate alien plant species invasion levels in different habitats and alien species traits by comparing the naturalized flora in different areas of the same biogeographical region. Location Spain, Italy, Greece and Cyprus. Methods Comparison of floristic composition, species traits and recipient habitats of naturalized alien neophytes across an east–west gradient comprising four countries in the European Mediterranean basin. Results A total of 782 naturalized neophytes were recorded; only 30 species were present in all four countries. Although floristic similarity is low, the four alien floras share the same patterns of growth form (mostly herbs), life cycle (mostly perennials) and life form (mostly therophytes, hemicryptophytes and phanerophytes). The majority of the recipient habitats were artificial. Wetlands were the natural habitats, with the highest numbers of naturalized species. Floristic similarity analyses revealed: (1) the highest floristic similarity between Italy and Spain, both of which were more similar to Greece than to Cyprus; (2) two groups of floristic similarity between habitat categories in each country (Greece–Cyprus and Italy–Spain); (3) a higher degree of homogenization in the plant assemblages in different habitats in Greece and Cyprus and a lower degree of homogenization in those in Italy and Spain; and (4) a higher degree of homogenization in artificial and natural fresh‐water habitats than in the other natural habitats. Main conclusions The floristic similarity of naturalized neophytes between the four countries is low, although the overall analysis indicates that the western group (Italy–Spain) is separated from the eastern group (Greece–Cyprus). Similar patterns emerged regarding the life‐history traits and recipient habitats. The artificial habitats and the natural wet habitats are those that are invaded most and display the greatest homogenization in all four countries. Coastal habitats display a lower degree of homogenization but a high frequency of aliens. Dry shrubs and rocky habitats display a lower degree of homogenization and a low frequency of aliens.     

Ecological succession in areas degraded by bauxite mining indicates successful use of topsoil

Brazilian ironstone outcrops (cangas) are nutrient‐poor stressful habitat dominated by slow‐growing woody species with high biodiversity and unique evolutionary history. Mining has produced great impacts on this ecosystem. Spontaneous regeneration of abandoned canga mined areas has not been observed. One of the active methods most widely used for ecological restoration in environments where soil has been lost or severely degraded is topsoil transposition due to the physical, chemical, and microbiological improvement of the substrate, in addition to the seed bank. Thus, plant succession was monitored for 40 months after topsoil transposition in a canga area degraded by aluminum mining, without any other type of management. A completely randomized design with 70 permanent plots (1 × 1 m) was used. Annual phytosociological surveys were carried out and floristic and vegetational spectra were constructed with the life‐forms proposed by Raunkiaer. Floristic composition was compared with a reference site. Overall, 105 species were identified. Both flora and vegetation changed over time, increasing resemblance to the reference areas. The floristic and vegetational spectra after 4 years of topsoil deposition are similar to pristine ones. The vegetation spectrum showed an increase in the dominance of phanerophytes and hemicryptophytes, while therophytes reduced their proportion. The early successional stage is dominated by weeds, like in other canga restoration studies, but did not impede the native species regeneration. Cangas's species recruited well from transposed topsoil. Unlike other studies with fertilized topsoil, our findings show the efficiency of topsoil transposition to provide initial conditions for the ecological restoration of this ecosystem.     

Floristic composition and biological spectrum of Keran - a remote valley of northwestern Himalaya

The Himalayas are one of the youngest and richest ecosystems on earth with a variety of species and forest types due to the varying altitude, topographic, and climatic conditions. The present study was aimed to investigate the floristic composition and biological spectrum of Keran valley, a region nestled in the northwestern frontiers of Himalayan biodiversity hotspot. Floristically, a total of 183 species were recorded, out of which 55 were aliens. Herbaceous growth form was dominant (67%) and therophytes were the dominant (37%) life form. Likewise, mesophylls (34%) followed by nanophylls (29%) and microphylls (27%) were major leaf size categories; and the simple leaf lamina type was recorded in majority of the species (65%). Almost 1/3rd of the total species pool was growing in the forest habitats, while as the 2/3rd occur within the rest 9 habitat types which were largely human-modified. The vegetation phenology observed during different seasons revealed that most of the species were dominant in summer season (37%), followed by spring (29%), autumn (21%) and winter (13%). The present study provides baseline information on the plant diversity in this hitherto unexplored region of northwestern Himalaya.     

Qualitative variation in roadside weed vegetation along an urban–rural road gradient

Information on mechanisms of differentiation or homogenization of urban floras is deficient, despite their importance for urban nature conservation and urban land management. Roads, as a major human promoter of urbanization, can be an initial habitat for plants dispersed by transportations. We assumed that variation in weed vegetation along urban–rural roadside gradients is small, particularly in curbside cracks. We classified vegetation occurring in curbside cracks along the National Route 3 in southern Japan and compared the characteristics of the vegetation types recognized. Species in curbside cracks were recorded on 40 plots. Three vegetation types were classified, in part related to surrounding land-use types. Although the Shannon–Wiener diversity index and the number of native species clearly differed among the vegetation types in the curbside cracks, no significant differences between the three floristic vegetation types were found in the number of non-natives and invasive alien species. This may result from the small specificity and complexity of landscape structures, due to the continuity and connectivity of paved-road networks. Of the 122 species, 44 were non-natives or invasive alien species. The vegetation types generally involved the same plant families, with large numbers of species from the Asteraceae and Poaceae, regardless of vegetation types, but frequency of occurrence of the two families clearly differed between natives and non-natives and invasive alien species. Ephemeral non-natives and invasive alien species, particularly Asteraceae and Poaceae, seem to have advantageous dispersal strategies or low habitat dependency facilitating their occurrence in curbside cracks, regardless of adjacent land uses and the urban–rural landscape gradient. Expansion of these species may cause a homogenization of regional floras along roads.     

浙江天台山七子花群落特征的初步研究

天台山的七子花林,常见的种子植物有１２５种,隶属５１科,１０２属。其中热带类型３６属,温带类型５２属。按Ｒａｕｎｋｉａｅｒ生活型统计,高位芽植物占７０．８％,地面芽植物占２２．３％,地下芽植物占４．６％,一年生植物占２．３％,高位芽植物的比例与本省常绿落叶阔叶混交林的生活型谱近似。叶级以小型叶为主,占５３．１％。叶型以单叶比例最大,占７８．５％。群落结构可分为乔木层、灌木层、草本层和地被层。     

Floristic surveys of hydrocarbon-polluted sites in some Cameroonian cities (Central Africa)

Soils pollution with hydrocarbons is on the increase, especially in developing countries like Cameroon. Identifying plant species capable of growing on hydrocarbon-polluted soils remains an essential step in phytoremediation, a low cost, solar energy driven clean up technique in which living green plants are used. Here, we report floristic surveys conducted in 4 cities of Cameroon to identify plants with phytoremediation capacities. A total of 13 hydrocarbon-polluted sites were surveyed in each city using the quadrat method. Unpolluted sites served as the control. 106 species belonging to 76 genera and 30 families were identified on hydrocarbon-polluted sites. Plant diversity was shown to be higher (166 species contained in 125 genera and 50 families) in the control sites. Poaceae, Cyperaceae, Asteraceae and Amaranthaceae families have a higher taxonomic richness on polluted sites (16, 15, 10 and 8 taxa respectively). Shannon diversity index of the hydrocarbon-polluted sites (1.6 to 2.7 bits/ind.) were significantly lower than the control sites (2.7 to 3.2 bit/ind.). Based on a relative frequency > 10% and abundance > 7%, our results show that more than 15 plant species could be tested in preliminary trials to determine their phytoremediation/cleaning-up capacities on hydrocarbon-polluted soils in the tropics.     

Alien and native plant life‐forms respond differently to human and climate pressures

Aim To investigate whether differences in the elevational trend in native and alien species richness were dependent on climate or human pressures. Specifically we tested whether life‐form and/or alien/native status modifies the response of plant species richness to human population and temperature along: (1) a complete elevational gradient, and (2) within separate elevational bands that, by keeping temperature within a narrow range, elucidate the effects of human pressures more clearly. Location Two provinces (c. 7507 km²) on the southern border of the European Alps (Italy), subdivided into 240 contiguous sampling cells (c. 35.7 km²). Methods We used an extensive dataset on alien and native species richness across an elevation gradient (20–2900 m a.s.l.). Richness of natives and naturalized aliens were separately related to temperature, human population and Raunkiaer life‐form using general linear mixed models. Life‐form describes different plant strategies for survival during seasons with adverse cold/arid conditions. Results The relationship between species richness and temperature for natives was strongly dependent on life‐form, while aliens showed a consistent positive trend. Similar trends across alien and native life‐forms were found for the relationship between species richness and human population along the whole gradient and within separate elevational bands. Main conclusions The absence of life‐form‐dependent responses amongst aliens supports the hypothesis that the distribution of alien plant species richness was more related to propagule pressure and availability of novel niches created by human activities than to climatic filtering. While climate change will potentially contribute to relaxing species thermal constraints, the response of alien species to future warming will also be contingent on changes in anthropogenic pressures.     

Native and alien floras in urban habitats: a comparison across 32 cities of central Europe

Aim To determine relative effects of habitat type, climate and spatial pattern on species richness and composition of native and alien plant assemblages in central European cities. Location Central Europe, Belgium and the Netherlands. Methods The diversity of native and alien flora was analysed in 32 cities. In each city, plant species were recorded in seven 1‐ha plots that represented seven urban habitat types with specific disturbance regimes. Plants were classified into native species, archaeophytes (introduced before ad 1500) and neophytes (introduced later). Two sets of explanatory variables were obtained for each city: climatic data and all‐scale spatial variables generated by analysis of principal coordinates of neighbour matrices. For each group of species, the effect of habitat type, climate and spatial variables on variation in species composition was determined by variation partitioning. Responses of individual plant species to climatic variables were tested using a set of binomial regression models. Effects of climatic variables on the proportion of alien species were determined by linear regression. Results In all cities, 562 native plant species, 188 archaeophytes and 386 neophytes were recorded. Proportions of alien species varied among urban habitats. The proportion of native species decreased with increasing range and mean annual temperature, and increased with increasing precipitation. In contrast, proportions of archaeophytes and neophytes increased with mean annual temperature. However, spatial pattern explained a larger proportion of variation in species composition of the urban flora than climate. Archaeophytes were more uniformly distributed across the studied cities than the native species and neophytes. Urban habitats rich in native species also tended to be rich in archaeophytes and neophytes. Main conclusions Species richness and composition of central European urban floras are significantly affected by urban habitat types, climate and spatial pattern. Native species, archaeophytes and neophytes differ in their response to these factors.     

Observed and dark diversity of alien plant species in Europe: estimating future invasion risk

Invasion by alien species is a growing concern for nature conservation. We estimated the level of invasion by alien plant species and future invasion risks at the European scale. We used a pan-European atlas and eight regional plant atlases to determine the distribution of alien and native plant richness. In addition, we estimated alien and native dark diversity (species currently absent from a site but present in the surrounding region and able to colonize the site). We used relative diversity metrics to indicate current and future risks by alien species: relative alien richness (compared to native species), alien and native completeness (log-ratio of observed to dark diversity) and completeness difference between alien and native species. Observed and relative richness of alien species were greatest in NW Europe; this suggests that sites in NW Europe could be more disturbed. Observed alien and native species richness show clear regional hotspots; the distribution of completeness values is dispersed, indicating local hotspots. Northern Europe has relatively lower alien completeness, likely because potential invaders inhabit the region but have not yet reached many localities, thereby suggesting a risk of future invasion. A greater number of potential alien species in the region increases the probability that some alien species could have detrimental impacts. Both alien richness and completeness are positively correlated with native richness and completeness, respectively, indicating that both groups share similar distribution patterns. Alien species diversity metrics in Europe are related positively to human population density and agricultural land-use. We suggest that the dark diversity concept can broaden our understanding of alien species diversity and future invasion risks.     

Ecological significance of floristic composition and life forms of Riyadh region,Central Saudi Arabia

Riyadh region is one of the hardest habitats in Saudi Arabia with a hyper-arid climate. This study was conducted to investigate floristic composition and plant life forms of Riyadh region and their ecological significance. Work aimed to determine the prevailing plant families and biological spectrum of their components that reflects the phytoclimate and adaptation to hyper-arid conditions of the region. Work involved field surveys of different locations in Riyadh region where plant specimens were collected and identified. Collected plant species were then listed according to their families and data were used to assess the contribution of different plant families to the flora of the region. Study of life forms was conducted to classify recorded floristic elements into categories that reflect environmental conditions prevailing in the region. Prevailing plant families were Asteraceae (17.4%), Poaceae (11%), Brassicaceae (9.9%), and Fabaceae (7%). These percentages reflect wide ecological ranges especially for Asteraceae and Poaceae. High presence of species belonging to Astreaceae and Poaceae can be attributed to their adaptation to harsh conditions as well as to effective wind dispersal strategies of their diaspores. While the most frequent life form classes were therophytes with 52%, and chamaephytes with 30%, phanerophytes and hemicryptophytes represented 9% and 8%, respectively. Predominance of therophytes and chamaephytes over other life forms is a response to hyper arid climate with insufficient rainfall and the nature of region of few available microhabitats that can support high percentage of perennials.     

Phylogenetic beta diversity of native and alien species in European urban floras

Aim Human activities have weakened biogeographical barriers to dispersal, increasing the rate of introduction of alien plants. However, their impact on beta diversity and floristic homogenization is poorly understood. Our goal is to compare the phylogenetic beta diversity of native species with that of two groups of alien species, archaeophytes and neophytes (introduced before and after ad 1500, respectively), across European urban floras to explore how biological invasions affect phylogenetic turnover at a continental scale. Location Twenty European cities located in six countries between 49 and 53° N latitude in continental Europe and the British Isles. Methods To compare the phylogenetic beta diversity of native and alien species we use the average phylogenetic dissimilarity of individual floras from their group centroid in multivariate space. Differences in phylogenetic beta diversity among different species groups are then assessed using a randomization test for homogeneity of multivariate dispersions. Results Across European urban floras, and when contrasted with natives, archaeophytes are usually associated with lower levels of phylogenetic beta diversity while neophytes tend to increase phylogenetic differentiation. Main conclusions While archaeophytes tend to promote limited homogenization in phylogenetic beta diversity, because of their diverse geographical origin together with short residence times in the invaded regions, neophytes are not promoting biotic homogenization of urban floras across Europe. Therefore, in spite of the increasing rate of alien invasion, an intense phylogenetic homogenization of urban cities is not to be expected soon.     

Indigenous and Ammophila arenaria-dominated dune vegetation on the South African Cape coast

Abstract. Communities formed by the potentially invasive European Ammophila arenaria (marram grass) are compared with those dominated by indigenous dune plant species in coastal dune systems. Sampling of communities was carried out along the Cape coast for species richness, species diversity, importance values and species associations. The influence of soil and other environmental factors on vegetation were also compared. While species richness values in A. arenaria communities appear similar to those of indigenous dune plant communities, diversity indices are significantly lower. However, on the basis of importance values of individual species, A. arenaria does not show extreme dominance to the exclusion of other species, as it does on the North American Pacific coast, where it has also been introduced. Because of its growth in dense tufts, A. arenaria is accompanied mostly by small chamaephytes and therophytes, while indigenous stands support more phanerophytes. Moreover, A. arenaria forms weaker species associations than dominant indigenous dune plant species. The alien status of A. arenaria in South Africa is confirmed by applying classification and ordination analyses which failed to differentiate A. arenaria communities according to their geographical origin as achieved with indigenous communities. This may be attributed to the lack of vigorous indigenous plants in A. arenaria communities, which accounts for the low variety in species composition of A. arenaria communities along the coastline. With regard to environmental factors, A. arenaria communities were observed to be less sensitive to extrinsic factors, such as climate, than indigenous dune plant communities. Results confirm that A. arenaria is an alien plant species in South Africa, but do not imply its invasiveness in the present or near future.     

Elevational gradient and vegetation‐environmental relationships in the central Hyrcanian forests of northern Iran

Tertiary‐relict Hyrcanian (Caspian) forest along the shores of the southern Caspian Sea is a center of biodiversity. Still, there is little information on plant diversity patterns in this area. This study evaluated plant diversity, variation in life forms, and geographical distribution of the zonal vegetation types and their relationships with environmental variables, in the educational and experimental forest of Kheyrudkenar, an important protected area in the central Hyrcanian forest of northern Iran. For this purpose, 226 vegetation plots of 400 m² were laid out along two altitudinal transects from the lowlands (100 m a.s.l.) to the timberline (2000 m a.s.l.). Four vegetation types were identified using modified TWINSPAN, indirect and direct gradient analyses. Species‐related (species diversity indices, life form and phytogeographical elements) and environmental variables (climate, topographic and soil variables) were calculated and subjected to one‐way ANOVA among the vegetation types. Both constrained (CCA) and unconstrained (DCA) ordination analyses showed an almost identical variation of the floristic composition along their axes and demonstrated that there are two main gradients in the Hyrcanian forest. Elevation together with annual precipitation and mean annual temperature were the most important factors controlling the floristic composition in the area. Topographic features such as slope inclination and heat index were found to be important within an elevation zone/vegetation type. Soil physical and chemical properties were of secondary importance for the separation of the vegetation types. This knowledge will be useful for forest management and conservation practices in the Hyrcanian area with its distinct and unique flora and vegetation.     

Temporal floristic variations as indicator of environmental changes in the Tiber River in Rome

A comparison of the actual vascular flora of the Tiber in Rome with a similar floristic survey of the 1970s was carried out. Changes in species richness, floristic composition and structure occurred in the last 30 years were described and dynamics of abundance of each species was estimated. The total number of species decreased from 604 in 1975–1981 to 501 till date. The composition of the Tiber flora has partially changed. There has been a decrease in therophytes on the riverbed and an increase in hydrophytes especially eutrophic ones. Most of Mediterranean species and those associated with undisturbed land have been replaced by widely distributed, ruderal and alien species, as human influences have increased. The floristic variations were used as indicator to evaluate the effect of human impact on the development of site and to describe the main changes in river management and bank use occurred locally over the last decades.     

珠海市淇澳岛肉实树群落及其物种多样性特征

对珠海淇澳岛地区肉实树占优势的植物群落进行研究,结果表明：（1）肉实树群落中共有维管植物38科51属61种,种类组成具有极强的南亚热带性质;（2）年龄结构显示群落的主要优势种属于稳定型种群,群落总体处于稳定状态;群落的频度指数规律为A〉B〉C〉D〈E,与RaunKiaer频度规律相一致;（3）物种多样性指数为D=13．70,H=4．12,均匀度指数为E=0．39,E’=0．82,群落物种多样性指数和均匀度指数均较高,接近典型南亚热带常绿阔叶林顶极群落类型。     

Do urban areas act as foci for the spread of alien plant species? An assessment of temporal trends in the UK

Aim Given that urban landscapes often act as a point of entry for many non‐native species and urban development continues to increase as the human population rapidly expands, an understanding of the interaction between urbanization and non‐native plant species is important both in the control of potentially invasive species and in the conservation of native biodiversity. We investigated the spatial and temporal relationship between urban land cover and the distribution of non‐native species in Britain using two floristic data sets collected at two different time periods: 1987–88 and 2003–04. Location UK. Methods Using floristic data collected by the Botanical Society of the British Isles in 1987–88 (Monitoring Scheme) and 2003–04 (Local Change) in conjunction with habitat data obtained from the Land Cover Map of the UK, we conducted multiple regression analyses both within and between years on both groups of species (natives, neophytes and archaeophytes) and individual species. Results Neophytes (alien species introduced after 1500) were very strongly associated with urban land cover in both time periods and do not appear to be spreading out of urban habitats into the wider countryside. Archaeophytes (alien species introduced before 1500), however, showed a strong association with urban habitats in the earlier 1988 data set but no longer showed this association in the 2004 data set. Analysis at the individual species level showed that a large percentage of alien plant species, particularly archaeophytes, were not strongly associated with urban land cover or were negatively associated with such habitats. Main conclusions Our results suggest that there has been a reduction in the urban association of archaeophytes that is likely to have resulted from the recovery of archaeophytes associated with non‐urban (especially arable) habitats, following their decline in mid‐20th century, rather than from the movement of aliens into the wider countryside from urban habitats.