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.     

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

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

Species richness and phylogenetic diversity of native and non‐native species respond differently to area and environmental factors

Aim

To test whether native and non‐native species have similar diversity–area relationships (species–area relationships [SARs] and phylogenetic diversity–area relationships [PDARs]) and whether they respond similarly to environmental variables.

Location

United States.

Methods

Using lists of native and non‐native species as well as environmental variables for >250 US national parks, we compared SARs and PDARs of native and non‐native species to test whether they respond similarly to environmental conditions. We then used multiple regressions involving climate, land cover and anthropogenic variables to further explore underlying predictors of diversity for plants and birds in US national parks.

Results

Native and non‐native species had different slopes for SARs and PDARs, with significantly higher slopes for native species. Corroborating this pattern, multiple regressions showed that native and non‐native diversity of plants and birds responded differently to a greater number of environmental variables than expected by chance. For native species richness, park area and longitude were the most important variables while the number of park visitors, temperature and the percentage of natural area were among the most important ones for non‐native species richness. Interestingly, the most important predictor of native and non‐native plant phylogenetic diversity, temperature, had positive effects on non‐native plants but negative effects on natives.

Main conclusions

SARs, PDARs and multiple regressions all suggest that native and non‐native plants and birds responded differently to environmental factors that influence their diversity. The agreement between diversity–area relationships and multiple regressions with environmental variables suggests that SARs and PDARs can be both used as quick proxies of overall responses of species to environmental conditions. However, more importantly, our results suggest that global change will have different effects on native and non‐native species, making it inappropriate to apply the large body of knowledge on native species to understand patterns of community assembly of non‐native species.

     

The impact of an invading alien grass (Agropyron cristatum) on species turnover in native prairie

Abstract. Alien invasions typically reduce species richness of habitats, but few studies have examined their effects on species turnover, the difference in species composition between localities. Agropyron cristatum (L.) Gaertn. (crested wheat grass) has been planted on 6–10 million ha of North American prairie, and is invading native prairie. We studied the invasion of A. cristatum into native prairie by measuring species composition along a gradient from maximum to minimum A. cristatum abundance. As A. cristatum increased, the abundance of most common native species decreased, but one appeared to be unaffected (Bouteloua gracilis (H.B.K.) Lag.), and another (Poa sandbergii Vasey) increased. The effect of A. cristatum on species turnover was investigated by examining species–area curves for areas from 0.5 m² to 8.0 m². Species diversity was reduced by 35% at high A. cristatum abundances at all areas. A. cristatum reduced the intercept of the species–area curve, but not the slope, suggesting that A. cristatum affected species turnover proportionally in all areas and habitats. This unusual result may indicate a homogeneous environment where species are distributed randomly. A. cristatum produced almost twice as many seeds as all native grasses combined. The number of seeds collected of native grasses and A. cristatum was highly correlated with the number of seed heads immediately nearby, but not with transect position. This suggests most seeds were dispersed over distances less than 5 m. In sum, the invasion of native prairie by A. cristatum might be related to high rates of seed production, and has the effect of decreasing species turnover by reducing the intercept of the species–area curve.     

Influence of settlement time, human population, park shape and age, visitation and roads on the number of alien plant species in protected areas in the USA

Abstract. I examined a data set of 77 protected areas in the USA (including national and state parks) to determine which of the following variables most strongly influence alien plant species richness: park area, climate (temperature and precipitation), native species richness, visitation rate, local human population size, total road length, park shape and duration of European settlement. Many of these predictor variables are intercorrelated, so I used multiple regression to help separate their effects. In support of previous studies, native species richness was the best single predictor of alien species richness, probably because it was a good estimator of both park area and habitat diversity available for establishment of alien species. Other significant predictors of alien species richness were years of occupation of the area by European settlers and the human population size of adjacent counties. Climate, visitation rate, road length and park shape did not influence alien species richness. The proportion of alien species (alien richness/native richness) is inversely related to park area, in agreement with a previous study. By identifying which variables are most important in determining alien species richness, such findings suggest ways to reduce alien species establishment.     

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.     

A century of the ornamental plant trade and its impact on invasion success

We identify a significant relationship between domestic market-based propagule pressure, as measured both in presence in the British horticultural market and in seed prices of ornamental plant species, with success in invasion. We employ a multispecies temporal approach and use a Generalized Estimation Equation model comparing ornamental non-native species introduced into Britain which started to invade with species introduced but not known outside cultivation. Historical nursery catalogues gave information on the availability and prices of seeds of 506 ornamental species in the British horticultural market every 20 years from 1885 to 1985. Higher market frequency and cheap prices of seeds were more significant and had a greater impact on the invading probability 20 years later than at the date of listing in a nursery catalogue. Our results suggest that national economic factors are an important part of the explanation for the invasiveness of ornamental plant species, and hence for the development of potential solutions.     

Invasions by alien plant species of the agro-pastoral ecotone in northern China: Species-specific and environmental determinants

The establishment, reproduction, dispersal, and distribution of alien plants are affected by various factors during the transition from being newly introduced in a habitat to being invasive. In the agro-pastoral ecotone of northern China, comprising farmlands and natural grasslands, the biological characteristics of alien plant species were the key intrinsic factors (propagation characteristics and competitive ability), followed by such extrinsic factors as human interference and environmental heterogeneity. Among biological characteristics, the life form may be an important and useful indicator of the invasive ability of a species, and the risk of invasion is greater from alien species that are poisonous, inedible, and have traits that facilitate wide dispersal. Farmlands may serve as initial shelters for alien species, from which they spread into neighbouring habitats, whereas natural grassland may act as a barrier to plant invasions. Management practices detrimental to grasslands, including overgrazing, reclamation, and road construction, often facilitate the invasions; therefore, counter measures such as reseeding and a ban on grazing need special attention. Environmental factors including precipitation, nutrients, prevailing winds, fires, and topography may be other factors that promote or block the process of invasion. In studying ways of preventing or controlling such invasions, alien plants with short life cycle, prolific seed production, and strong competitiveness, deserve particular attention and so do human activities that may damage the environment and fragile habitats.     

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.     

Diversity and distribution of genetic variation in gammarids: Comparing patterns between invasive and non‐invasive species

Biological invasions are worldwide phenomena that have reached alarming levels among aquatic species. There are key challenges to understand the factors behind invasion propensity of non‐native populations in invasion biology. Interestingly, interpretations cannot be expanded to higher taxonomic levels due to the fact that in the same genus, there are species that are notorious invaders and those that never spread outside their native range. Such variation in invasion propensity offers the possibility to explore, at fine‐scale taxonomic level, the existence of specific characteristics that might predict the variability in invasion success. In this work, we explored this possibility from a molecular perspective. The objective was to provide a better understanding of the genetic diversity distribution in the native range of species that exhibit contrasting invasive propensities. For this purpose, we used a total of 784 sequences of the cytochrome c oxidase subunit I of mitochondrial DNA (mtDNA‐COI) collected from seven Gammaroidea, a superfamily of Amphipoda that includes species that are both successful invaders (Gammarus tigrinus, Pontogammarus maeoticus, and Obesogammarus crassus) and strictly restricted to their native regions (Gammarus locusta, Gammarus salinus, Gammarus zaddachi, and Gammarus oceanicus). Despite that genetic diversity did not differ between invasive and non‐invasive species, we observed that populations of non‐invasive species showed a higher degree of genetic differentiation. Furthermore, we found that both geographic and evolutionary distances might explain genetic differentiation in both non‐native and native ranges. This suggests that the lack of population genetic structure may facilitate the distribution of mutations that despite arising in the native range may be beneficial in invasive ranges. The fact that evolutionary distances explained genetic differentiation more often than geographic distances points toward that deep lineage divergence holds an important role in the distribution of neutral genetic diversity.     

Ecological impacts of invasive alien plants: a meta-analysis of their effects on species, communities and ecosystems

Biological invasions cause ecological and economic impacts across the globe. However, it is unclear whether there are strong patterns in terms of their major effects, how the vulnerability of different ecosystems varies and which ecosystem services are at greatest risk. We present a global meta-analysis of 199 articles reporting 1041 field studies that in total describe the impacts of 135 alien plant taxa on resident species, communities and ecosystems. Across studies, alien plants had a significant effect in 11 of 24 different types of impact assessed. The magnitude and direction of the impact varied both within and between different types of impact. On average, abundance and diversity of the resident species decreased in invaded sites, whereas primary production and several ecosystem processes were enhanced. While alien N-fixing species had greater impacts on N-cycling variables, they did not consistently affect other impact types. The magnitude of the impacts was not significantly different between island and mainland ecosystems. Overall, alien species impacts are heterogeneous and not unidirectional even within particular impact types. Our analysis also reveals that by the time changes in nutrient cycling are detected, major impacts on plant species and communities are likely to have already occurred.     

Native and naturalized range size in Pinus: relative importance of biogeography,introduction effort and species traits

Aim Pine trees (genus Pinus) represent an ancient lineage, naturally occurring almost exclusively in the Northern Hemisphere, but introduced and widely naturalized in both hemispheres. As large trees of interest to forestry, they attract much attention and their distribution is well documented in both indigenous and naturalized ranges. This creates an opportunity to analyse the relationship between indigenous and naturalized range sizes in the context of different levels of human usage, biological traits and the characteristics of the environments of origin. Location Global. Methods We combined and expanded pre‐existing data sets for pine species distributions and pine species traits, and used a variety of regression techniques (including generalized additive models and zero‐inflated Poisson models) to assess which variables explained naturalized and indigenous range sizes. Results Indigenous and naturalized range sizes are positively correlated but there are many notable exceptions. Some species have large indigenous ranges but small or no naturalized ranges, whereas others have small indigenous ranges, but have naturalized in many regions. Indigenous range is correlated to factors such as seed size (?), age at first reproduction (?), and latitude (+, supporting Rapoport's rule), but also to the extent of coverage of species in the forestry literature (+). Naturalized range size is strongly influenced by the extent of coverage of species in the forestry literature (+), a proxy for propagule pressure. Naturalization was also influenced by average elevation in the indigenous range (?) and age at first reproduction (?). Main conclusions The macroecological and evolutionary pressures facing plant groups are not directly transferable between indigenous and naturalized ranges. In particular, there are strong biases in species naturalization and expansion in invasive ranges that are unrelated to factors determining indigenous range size. At least for Pinus, a new set of macroecological patterns are emerging which are profoundly influenced by humans.     

福建外来物种入侵现状及对经济社会和生态的影响

福建已成为中国遭受外来物种入侵最严重的省份之一。本文概述了福建主要入侵物种的数量、种类、地理起源、分布、引入途径和入侵特点,系统分析了福建外来物种入侵严重的原因和外来物种入侵对福建生态安全、经济及社会的影响,在此基础上提出了针对外来物种入侵的防控对策。