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.     

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.     

Distance decay of similarity among European urban floras: the impact of anthropogenic activities on β diversity

Aim We examine how two categories of non‐native species (archaeophyte and neophyte, introduced before and after ad 1500, respectively) have had different impacts on β diversity across European urban floras. Our goal is to use the unique biological perspective provided by urban areas, and the contrasting historical and geographical perspectives provided by archaeophytes and neophytes, to infer how non‐native species will impact upon β diversity in the future. Location Twenty‐two urban areas located in seven European countries. Methods We used the β‐sim dissimilarity index to estimate the level of β diversity for 231 unique pair‐wise combinations of 22 urban floras. We examined bivariate plots of dissimilarity by geographical separation of city centres to evaluate distance decay of similarity for native species, archaeophytes and neophytes. Results Based on average percentages, 52.8% (SD = 8.2%) of species in the urban floras were identified as non‐native with 28.3% (SD = 6.9%) classified as neophytes and 24.5% (SD = 4.9%) as archaeophytes. Relative to native species, across urban floras, archaeophytes were associated with higher compositional similarity and weaker distance decay patterns, whereas neophytes were associated with lower compositional similarity and stronger distance decay patterns. Main conclusions Across European urban floras, archaeophytes and neophytes occurred in similar numbers but archaeophytes were consistently associated with lower β diversity and neophytes with higher β diversity. Thus, the impact of non‐native species on β diversity can be determined, at least in part, through their historical and geographical associations with anthropogenic activities. If archaeophytes represent the long‐term biogeographical outcome for human commensal species, neophytes could develop similar patterns. The consequences, however, are likely to be more substantial ecologically and geographically due to the increasing numbers of neophytes and their global anthropogenic associations. Nevertheless, at present, our findings suggest that, based on occurrence information, neophytes have not achieved this state with European urban floras retaining regionally distinct assemblages of neophytes.     

Alien and native species in Central European urban floras: a quantitative comparison

The paper provides quantitative information on the occurrence of alien species in Central European cities and analyses factors determining the richness of alien and native floras in this habitat type. Data for 54 cities (25 Polish, 24 German, 4 Czech and 1 Austrian) were gathered, and the representation of archaeophytes (i.e. aliens introduced before 1500 ad ), neophytes (introduced after that date) and native species was expressed. In an average city there were 87.4 archaeophytes (15.2% of the city flora) and 172.4 neophytes (25.2%) giving a total of 259.7 for alien species (40.3%). The number of native species averaged 386.5. The numbers of species in each category of immigration status increased significantly with city size. For neophytes, the species-area relationship showed a higher slope (0.49) on log–log axes than for both archaeophytes (0.16) and native species (0.30). Not only the number, but also the relative contribution of neophytes to the total flora, increased with city size, indicating that neophytes are the group which are most closely associated with human activity. On the other hand, archaeophytes were better represented in smaller cities, as they were confined to rural environment. A step-wise multiple regression was used to test for environmental variables acting as significant predictors, and explained between 40 and 65% of variation in the species numbers for particular categories of immigration status, providing the best fit for neophytes. City size was the best predictor for each characteristic, except of the proportion of total aliens, where the percentage of explained variability was low (8.2%), with latitude being the only significant predictor. Temperature was another highly significant predictor for the number of archaeophytes and total aliens, reflecting the origin of aliens in warmer areas. There was an effect of region on some flora characteristics. Polish cities had significantly higher proportion of archaeophytes and of total aliens than German cities. It is concluded that the occurrence of native and alien species in urban floras follows rather different pattern.     

Invasiveness of alien plants in Brussels is related to their phylogenetic similarity to native species

Aim Understanding the processes that drive invasion success of alien species has received considerable attention in current ecological research. From an evolutionary point of view, many studies have shown that the phylogenetic similarity between the invader species and the members of the native community may be an important aspect of invasiveness. In this study, using a coarse‐scale systematic sampling grid of 1 km², we explore whether the occupancy frequency of two groups of alien species, archaeophytes and neophytes, in the urban angiosperm flora of Brussels is influenced by their phylogenetic relatedness to native species. Location The city of Brussels (Belgium). Methods We used ordinary least‐squares regressions and quantile regressions for analysing the relationship between the occupancy frequency of alien species in the sampled grid and their phylogenetic distance to the native species pool. Results Alien species with high occupancy frequency in the sampled grid are, on average, more phylogenetically related to native species than are less frequent aliens, although this relationship is significant only for archaeophytes. In addition, as shown by the quantile regressions, the relationship between phylogenetic relatedness to the native flora and occupancy frequency is much stronger for the most frequent aliens than for rare aliens. Main conclusions Our data suggest that it is unlikely that species with very low phylogenetic relatedness to natives will become successful invaders with very high distribution in the area studied. To the contrary, under future climate warming scenarios, present‐day urban aliens of high occupancy frequency are likely to become successful invaders even outside urban areas.     

Compositional similarity among urban floras within and across continents: biogeographical consequences of human-mediated biotic interchange

Anthropogenic activities have weakened biogeographical barriers to dispersal resulting in the global spread and establishment of an increasing number of non‐native species. We examine the broad‐scale consequences of this phenomenon based on an analysis of compositional similarity across urban floras in the northeastern United States and Europe. We test the prediction that homogenization of species composition is uniquely defined within vs. between continents based on the time and place of origin of non‐native species. In this case, for archaeophytes and neophytes in Europe (introduced before and after ad 1500, respectively) and non‐native species originating from within and outside the United States. More species in urban floras were shared within than between continents. Within Europe, archaeophytes shared more species across urban floras compared with neophytes; strong associations were not observed for non‐native species across US urban floras. Between the two continents, non‐native species in the United States that originated from outside the United States shared species primarily with archaeophytes but also with European natives and neophytes. These results suggest that the direction of biotic interchange was unidirectional with species moving primarily from Europe to the United States with archaeophytes playing a primary and non‐native species originating from outside the two continents a secondary role as a homogenizing source. Archaeophytes, based on combination of biogeographical, evolutionary, and ecological factors in association with a long history of anthropogenic influence, appear to have played a prominent role in the continental and intercontinental homogenization of species composition. This suggests that the uniform homogenization of the Earth's biota is not imminent and is presently directed by a combination of biogeographically defined anthropogenic and historical factors.     

Alien plant species and factors of invasiveness of anthropogenic vegetation in the Northwestern Balkans — a phytosociological approach

We studied the anthropogenic vegetation of the Northwest Balkans in order to determine its susceptibility to invasion by alien plant species. We compiled a dataset of 3089 vegetation plots sampled between 1939 and 2009, recording a set of variables for each sample plot in order to determine which factors have the most effect on a habitat’s vulnerability to invaders. We calculated the proportion of native species, archaeophytes and neophytes for each plot. We used regression tree models to determine the site conditions of the most invaded anthropogenic habitats. The sample plots contained an average of 12.7% alien plant species, with a low proportion of archaeophytes (4.3%) and 8.4% neophytes. Local habitat conditions proved to have the largest effect, rather than climatic variables or propagule pressure. The proportion of archaeophytes follows a different pattern than that seen in central and northern Europe, indicating that macroecological factors are more important. Neophytes show a similar distribution to other European locations.     

Diversity of native and alien vascular plant species of dry grasslands in central Europe

Question: Which factors determine diversity of native and alien vascular plant species in semi‐natural dry grasslands? Location: Northern limestone Alps to the southern rim of the Bohemian massif in northern Austria. Methods: In 70 randomly chosen dry grassland patches (0.008 ha ‐ 7 ha) we sampled a complete inventory of vascular plant species at each site. We analysed the correlation between species diversity of natives, archaeophytes (pre‐1500 aliens) and neophytes (post‐1500 aliens). We used GLM to study the relationship of species number (natives, neophytes, archaeophytes) to five explanatory variables (altitude, within habitat diversity, habitat diversity of adjacent areas, within land‐use diversity and land‐use in adjacent areas). Orthogonal components of these variables were derived with a PCA and used in the models. We also tested the influence of minimum residence time (MRT) and the covariables origin, mode of introduction and life form on the number of grassland sites with neophytes with analogous GLMs. Results: Native species diversity species was positively correlated with the species diversity of new, but not old invaders. GLM explains 70% of the variance in the number of native species. Patch size explained the largest part of the variation in the number of native species. PCA axes 1 and 3 were significantly related to the number of native species. Axis 1was related to on‐site habitat and land‐use diversity. The GLM of the archaeophyte diversity explains 18% of the variance. Altitude and presence of fields and grassland in the neighbourhood mainly explained archaeophyte species diversity. The GLM of neophyte diversity explains 12% of the variance. The number of neophytes was positively related to that of archaeophytes. Only PCA axis 3, which is mainly influenced by adjacent land‐use types, showed a relationship with neophytes. MRT, mode of introduction and region of origin (but not life form) were significantly related to the number of grassland sites invaded by neophytes, explaining 35% of the variance. Conclusion: Most factors governing native species diversity are not significantly related to alien species diversity. Additional determinants of the local scale diversity of alien species exist such as region of origin and historical factors (MRT, mode of introduction).     

Determinants of native and alien species richness in the urban flora of Rome

Until now, analytical studies on European urban floras have mostly concentrated on the central and north‐western parts of the continent. In this paper, factors determining species richness of urban flora were studied for the city of Rome, Italy, based on a comprehensive floristic survey carried out between 1985 and 1994, and updated in 2005. All species were recorded in grid cells of 1.6 km² and classified into native and alien (the latter divided into archaeophytes and neophytes). The grids were classified with respect to the prevailing habitat type, area available to vegetation, level of disturbance and geographical position within the city. Data were analysed using minimal adequate models. Total species number was determined by habitat and its interaction with position on the north‐west gradient; other variables explained much less variance. Holding other variables constant, the average species number per grid cell was highest in archaeological sites and parks, followed by woodlands and rivers, and grasslands and recent developments. Residential areas and the historical centre were poorest in species number. Towards the north of the city, species richness in corresponding habitats increases because of higher landscape heterogeneity and closer association with diaspore pools in the surroundings. Native species make up on average 84% of the total species numbers, and trends opposite to those for the total number of species were found for the proportional representation of aliens. The occurrence of alien and native species in the flora of Rome is driven by similar factors, but factors that increase representation of aliens decrease that of natives and vice versa. The representation of neophytes and native species in grid cells was easier to explain (74% of variation accounted for) than that of archaeophytes (27%); this result reflects that in terms of ecology and response to factors, archaeophytes take an intermediate position between native plants and neophytes. Proportional representation of neophytes decreased with increasing area available to vegetation, reflecting that semi‐natural vegetation is better developed where less fragmented.     

Archaeophytes in Britain

The concept of 'archaeophytes' (alien taxa which became established in a study area before AD  1500) is widely used in floristic analyses in central and northern Europe, but few authors have applied it to the British flora. Six criteria for the recognition of archaeophytes are outlined, drawing upon evidence of fossil and recent history, current habitat and European and extra-European distribution. These are used to identify 157 probable archaeophytes in Britain. Only five of these are known from fossil records from the Neolithic; most are first recorded in archaeological contexts in the Late Bronze Age, Iron Age, Roman or Medieval periods. As a group, archaeophytes (unlike neophytes) have declined in Britain in the 20th century. Comparison of the accepted status of these species in the Czech Republic, Germany, Poland and Finland demonstrates that over 50% are treated as archaeophytes in central Europe, but in Finland many are absent or only present as casual introductions. Species regarded as archaeophytes in these countries but as natives in Britain are also reviewed. The indirect nature of the evidence used to identify archaeophytes means that it is usually impossible to be certain about the history of a species; in particular, archaeophytes which have successfully invaded semi-natural habitats are likely to be overlooked as natives. The suggestion that a species is an archaeophyte is best regarded as a hypothesis to be tested by further studies. There is considerable scope for archaeological investigations aimed at addressing these botanical problems.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 257−294.     

Native and alien plant species richness in relation to spatial heterogeneity on a regional scale in Germany

Aim The aim of our study was to reveal relationships between richness patterns of native vs. alien plant species and spatial heterogeneity across varying landscape patterns at a regional scale. Location The study was carried out in the administrative district of Dessau (Germany), covering around 4000 km². Methods Data on plant distribution of the German vascular flora available in grid cells covering 5′ longitude and 3′ latitude (c. 32 km²) were divided into three status groups: native plants, archaeophytes (pre 1500 AD aliens) and neophytes (post 1500 AD aliens). Land use and abiotic data layers were intersected with 125 grid cells comprising the selected area. Using novel landscape ecological methods, we calculated 38 indices of landscape composition and configuration for each grid cell. Principal components analysis (PCA) with a set of 29 selected, low correlated landscape indices was followed by multiple linear regression analysis. Results PCA reduced 29 indices to eight principal components (PCs) that explained 80% cumulative variance. Multiple linear regression analysis was highly significant and explained 41% to 60% variance in plant species distribution (adjusted R²) with three significant PCs (tested for spatial autocorrelation) expressing moderate to high disturbance levels and high spatial heterogeneity. Comparing the significance of the PCs for the species groups, native plant species richness is most strongly associated with riverine ecosystems, followed by urban ecosystems, and then small‐scale rural ecosystems. Archaeophyte and neophyte richness are most strongly associated with urban ecosystems, followed by small‐scale rural ecosystems and riverine ecosystems for archaeophytes, and riverine ecosystems and small‐scale rural ecosystems for neophytes. Main conclusions Our overall results suggest that species richness of native and alien plants increases with moderate levels of natural and/or anthropogenic disturbances, coupled with high levels of habitat and structural heterogeneity in urban, riverine, and small‐scale rural ecosystems. Despite differences in the order of relevance of PCs for the three plant groups, we conclude that at the regional scale species richness patterns of native plants as well as alien plants are promoted by similar factors.     

Alien and native plants show contrasting responses to climate and land use in Europe

Aim We tested whether the distribution and cover of alien plant species in Europe was related to human disturbance and microclimate. Location Surveys were conducted at 13 sites across Europe, each containing a pair of landscapes with different land‐use intensities. Methods Sampling locations were chosen based on land use and microclimate at two scales: land use was characterized at the patch and landscape scale; climate was expressed as regional and local temperature. The slope of each sample location was derived from a digital elevation model. Cover of plant species was measured using point counts and analysed using mixed effect models. Species were classified as native, archaeophytes and neophytes (pre‐ versus post‐ad 1500 immigrants). Due to the zero inflation observed in the alien groups, their cover was analysed conditional on their presence. Results Anthropogenic disturbance was a significant explanatory variable, increasing the presence and cover of alien species and decreasing the cover of native species. Alien presence was increased in sites under agricultural management, while their cover responded to land use at both local and landscape scales (and to their interaction), such that only natural habitats in semi‐natural landscapes had low alien cover. Microclimate was important for neophytes, with presence concentrated around mesic conditions. Slope was relevant for archaeophytes and native species, suppressing the former group and promoting the latter one. Main conclusions We found that, at the European scale, the distribution of alien plants is related to anthropogenic disturbance more than to microclimatic differences. The presence of neophytes, however, was influenced by climate at local and regional scales, with the highest incidence under mesic conditions. The different patterns observed for the presence and cover of alien species suggest different mechanisms acting during their establishment and spread. They also suggest that to counteract the expansion of alien species natural habitats may need to be maintained at landscape scales.     

Predicting and explaining plant invasions through analysis of source area floras: some critical considerations

Comparing species that become invasive with others from the same regional species pool that do not invade raises several issues about the accuracy of analyses attempting to define the determinants of invasiveness. The delimitation of the source area and deciding which species group(s) to include are especially relevant in analyses focusing on species originating in Europe. Historical patterns of immigration of alien species into Europe must be considered since European floras comprise a complex mix of native species, historical introductions (archaeophytes) and relative newcomers (neophytes). We make three main points: (1) Archaeophytes (species introduced to Europe before the discovery of America) differ from natives in a number of traits and in historical association with people; it is misleading to lump archaeophytes with native taxa. (2) Taxa from climatically and geographically different regions, representing distinct floristic geoelements, need to be treated separately, and not as a homogenous pool of potential invaders. Restricting the source species pool to native taxa with comparable phytogeographical characteristics reduces the variation associated with chance of dispersal by humans from the source area. (3) For prediction, a clear distinction should be made between accuracy (the proportion of those found to be alien that were also predicted to be there) and reliability (or predictive value, the proportion of those predicted to become aliens that do so). Information accumulated over centuries by botanists in Central Europe provides an excellent opportunity to deal with these issues and avoid spurious results. To illustrate these issues, we revisit a recently published study of Central‐European plant species as invaders in two Argentinean provinces ( Prinzing et al., 2002 ) to explore and demonstrate the implications of the above points. We hope that future studies will build on these points to achieve more reliable predictions.     

Urban alien plants in temperate oceanic regions of Europe originate from warmer native ranges

When colonizing new areas, alien plant species success can depend strongly on local environmental conditions. Microclimatic barriers might be the reason why some alien plant species thrive in urban areas, while others prefer rural environments. We tested the hypothesis that the climate in the native range is a good predictor of the urbanity of alien species in the invaded range. The relationship between climate in the native range and the percentage of artificially sealed surfaces (urbanity) at the occurrences of 24 emerging alien plant species, in European areas with a temperate climate (termed oceanic Europe) was evaluated. We found that alien species growing in more urban environments originated from warmer or drier native ranges than found in oceanic Europe. These results have strong conservation implications as climate-warming will likely lift climatic barriers that currently constrain numerous alien plant species to cities, boosting the role of cities as points of entry for invasive plants.

     

Interaction of local and landscape features in the conservation of Hungarian arable weed diversity

Question: How do local and landscape management contribute to weed diversity in Hungarian winter cereal fields? Location: Central Hungary. Methods: Vascular plants were sampled in 18 winter cereal fields along an intensification gradient according to nitrogen fertilization, in the first cereal rows (edge) and in the interior part of the fields. Weed species were divided into groups according to their residence time in Central Europe (native species, archaeophytes, neophytes) and nitrogen preference (low to medium, LMNP, and high, HNP species). The percentage of semi‐natural habitats was calculated in the 500 m radius circle. Effects of fertilizer use, transect position and semi‐natural habitats were estimated by general linear mixed models. Results: We recorded 149 weed species. Fertilizer had a negative impact on the species richness of archaeophytes and LMNP species, and on the cover of native weeds. There was greater species richness and weed cover at the edge of the fields than in the centre. A higher percentage of seminatural habitats around the arable fields resulted in greater total species richness, especially of archaeophytes and LMNP species. We found an interaction between the percentage of semi‐natural habitats and transect position for species richness of archaeophytes and LMNP species. Conclusions: Reduced use of fertilizers and a high percentage of semi‐natural habitats would support native and archaeophyte weed diversity even in winter cereal fields, while large amounts of fertilizer may promote invasion of neophytes. However, the beneficial effect of the semi‐natural habitats and greater species pool on the arable flora may prevail only in the crop edges.     

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.     

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.     

A phylogenetic approach towards understanding the drivers of plant invasiveness on Robben Island,South Africa

Invasive plant species are a considerable threat to ecosystems globally and on islands in particular where species diversity can be relatively low. In this study, we examined the phylogenetic basis of invasion success on Robben Island in South Africa. The flora of the island was sampled extensively and the phylogeny of the local community was reconstructed using the two core DNA barcode regions, rbcLa and matK. By analysing the phylogenetic patterns of native and invasive floras at two different scales, we found that invasive alien species are more distantly related to native species, a confirmation of Darwin's naturalization hypothesis. However, this pattern also holds even for randomly generated communities, therefore discounting the explanatory power of Darwin's naturalization hypothesis as the unique driver of invasion success on the island. These findings suggest that the drivers of invasion success on the island may be linked to species traits rather than their evolutionary history alone, or to the combination thereof. This result also has implications for the invasion management programmes currently being implemented to rehabilitate the native diversity on Robben Island. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 142–152.     

Weed vegetation of arable land in Central Europe: Gradients of diversity and species composition

Question: What are the main broad‐scale spatial and temporal gradients in species composition of arable weed communities and what are their underlying environmental variables? Location: Czech Republic and Slovakia. Methods: A selection of 2653 geographically stratified relevés sampled between 1954–2003 was analysed with direct and indirect ordination, regression analysis and analysis of beta diversity. Results: Major changes in weed species composition were associated with a complex gradient of increasing altitude and precipitation and decreasing temperature and base status of the soils. The proportion of hemicryptophytes increased, therophytes and alien species decreased, species richness increased and beta diversity decreased with increasing altitude. The second most important gradient of weed species composition was associated with seasonal changes, resulting in striking differences between weed communities developed in spring and summer. In summer, weed communities tended to have more neophytes, higher species richness and higher beta diversity. The third gradient reflected long‐term changes in weed vegetation over past decades. The proportion of hemicryptophytes and neophytes increased, while therophytes and archaeophytes decreased, as did species richness over time. The fourth gradient was due to crop plants. Cultures whose management involves less disturbances, such as cereals, harboured less geophytes and neophytes, and had higher species richness but lower beta diversity than frequently disturbed cultures, such as root crops. Conclusions: Species composition of Central European weed vegetation is mainly influenced by broad‐scale climatic and edaphic factors, but its variations due to seasonal dynamics and long‐term changes in agricultural management are also striking. Crop plants and crop‐specific management affect it to a lesser, but still significant extent.     

Effects of different cultivation types on native and alien weed species richness and diversity in Moravia (Czech Republic)

Changes in weed species richness and beta-diversity are partly attributable to different types and intensity of disturbance and partly to broad-scale variation in environmental conditions. We compiled a data set of 434 vegetation plots of weed vegetation in root crop and cereal fields in Moravia (eastern Czech Republic) to compare the effects of environmental conditions and different disturbance regimes on species richness and beta-diversity. To detect changes in species richness, we related the variation in species richness to individual environmental conditions. To assess differences in beta-diversity between the vegetation of cereal and root crop fields, we used Whittaker's measure of beta-diversity. The relative importance of each environmental variable for the variation in species composition was evaluated using canonical correspondence analysis. All analyses were done for all vascular plant species and separately for native species, archaeophytes and neophytes. A comparison of weed vegetation of root crops and cereals showed a distinct dichotomy between these two types of weed vegetation. There was no significant difference in total species richness and native species richness; however, cereal fields were richer in archaeophytes and root crop fields were richer in neophytes. The beta-diversity of weed vegetation was higher in root crops. Environmental factors explained a significant part of the variability in richness of both natives and aliens. The richness of native species increased and beta-diversity decreased with increasing precipitation. The opposite relationship was found for archaeophytes, in both cereals and root crops. These results confirmed the importance of climatic factors and management practices for changes in weed species composition. They also showed a distinct pattern of species richness and beta-diversity of native and alien weed species.