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.     

Segregation,nestedness and homogenisation in plant communities dominated by native and alien species

ABSTRACT

Background: Highly modified landscapes offer the opportunity to assess how environmental factors influence the integration of alien plant species into native vegetation communities and determine the vulnerability of different communities to invasion.

Aims: To examine the importance of biotic and abiotic drivers in determining whether alien plant species segregate spatially from native plant communities or become integrated and lead to biotic homogenisation.

Methods: Ordination and classification of a floristic survey of over 1200 systematically located 6 m × 6 m plots were used to examine how plant community segregation, nestedness and homogenisation varied in relation to climate, environmental and human-related factors across Banks Peninsula, New Zealand.

Results: The analyses of community structure indicated that native and alien plant communities were spatially and ecologically segregated due to different responses primarily to an anthropogenic impact gradient and secondly to environmental factors along an elevation gradient. Human-land use appeared most strongly linked to the distribution of alien species and was associated with increased vegetation homogenisation. However, despite spatial segregation of alien and native plant communities, biotic homogenisation not only occurred in highly managed grasslands but also in relatively less managed shrublands and forest.

Conclusions: The role played by anthropogenic factors in shaping alien and native plant species community structure should not be ignored and, even along a marked environmental gradient, if the recipient sites have a long history of human-related disturbance, biotic homogenisation is often strong.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

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.     

Herbivory is related to taxonomic isolation, but not to invasiveness of tropical alien plants

Aim  The enemy release hypothesis is often invoked to explain why some alien plant species become invasive. Here, we investigated relationships between invasiveness, taxonomic isolation and leaf herbivory for tropical alien plant species introduced to a botanical garden in East Africa.
Location  Amani Botanical Garden, East Usambara mountains, northeast Tanzania.
Methods  We measured the proportion of leaves damaged, and the percentage leaf area damaged on individuals of 28 alien plant species. We extracted data on the presence/absence of native congeners and the number of native confamilial species from an inventory of the East Usambara flora. We also obtained data on planting effort for 26 species, from historical records. Linear and generalized linear models were used to analyse the relationships between invasiveness, herbivory and taxonomic isolation.
Results  Mean proportion of leaves damaged per species was significantly explained by taxonomic isolation; proportion of leaves damaged increased with the number of native confamilial species and was greater, on average, for species with native congeners than those without native congeners. The mean percentage of leaf area damaged per species could not be explained by any variables considered in this study. There was no relationship between the degree of herbivory or taxonomic isolation and alien plant species invasiveness, but more-invasive species did have a significantly greater planting effort than less-invasive species.
Conclusions  The role herbivores play in controlling alien plant invasions has been investigated relatively little in the tropics. In this study, although the amount of herbivory suffered by alien plants was related to taxonomic isolation, we found no evidence for leaf-feeding invertebrates having a significant role in invasion, suggesting that other factors may be responsible for differences in species success.     

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.     

Bird species richness is associated with phylogenetic relatedness,plant species richness,and altitudinal range in Inner Mongolia

Bird species richness is mediated by local, regional, and historical factors, for example, competition, environmental heterogeneity, contemporary, and historical climate. Here, we related bird species richness with phylogenetic relatedness of bird assemblages, plant species richness, topography, contemporary climate, and glacial‐interglacial climate change to investigate the relative importance of these factors. This study was conducted in Inner Mongolia, an arid and semiarid region with diverse vegetation types and strong species richness gradients. The following associated variables were included as follows: phylogenetic relatedness of bird assemblages (Net Relatedness Index, NRI), plant species richness, altitudinal range, contemporary climate (mean annual temperature and precipitation, MAT and MAP), and contemporary‐Last Glacial Maximum (LGM) change in climate (change in MAT and change in MAP). Ordinary least squares linear, simultaneous autoregressive linear, and Random Forest models were used to assess the associations between these variables and bird species richness across this region. We found that bird species richness was correlated negatively with NRI and positively with plant species richness and altitudinal range, with no significant correlations with contemporary climate and glacial–interglacial climate change. The six best combinations of variables ranked by Random Forest models consistently included NRI, plant species richness, and contemporary‐LGM change in MAT. Our results suggest important roles of local ecological factors in shaping the distribution of bird species richness across this semiarid region. Our findings highlight the potential importance of these local ecological factors, for example, environmental heterogeneity, habitat filtering, and biotic interactions, in biodiversity maintenance.     

Resolving the invasion paradox: pervasive scale and study dependence in the native‐alien species richness relationship

The degree to which plant communities are vulnerable to invasion by alien species has often been assessed using the relationship between native and alien plant species richness (NAR). Variation in the direction and strength of the NAR tends to be negative for small plot sizes and study extents, but positive for large plots and extents. This invasion paradox has been attributed to different processes driving species richness at different spatial scales. However, the focus on plot size has drawn attention away from other factors influencing the NAR, in part because the influence of other factors may be obscured by or interact with plot size. Here, we test whether variation in the NAR can be explained by covariates linked to community susceptibility to invasion and whether these interact with plot size using a quantitative meta‐analysis drawn from 87 field studies that examined 161 NARs. While plot size explained most variation, the NAR was less positive in grassland habitats and in the Australasian region. Other covariates did not show strong relationships with the NAR even after accounting for interactions with plot size. Instead, much of the unexplained variation is associated with article or author specific differences, suggesting the NAR depends strongly on how different authors choose their study system or study design.     

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 vegetation structure,landscape features and climate shape non-native plant richness and cover in New Zealand native shrublands

Aim

Studies investigating the determinants of plant invasions rarely examine multiple factors and often only focus on the role played by native plant species richness. By contrast, we explored how vegetation structure, landscape features and climate shape non-native plant invasions across New Zealand in mānuka and kānuka shrublands.

Location

New Zealand.

Method

We based our analysis on 247 permanent 20 × 20-m plots distributed across New Zealand surveyed between 2009 and 2014. We calculated native plant species richness and cumulative cover at ground, understorey and canopy tiers. We examined non-native species richness and mean species ground cover in relation to vegetation structure (native richness and cumulative cover), landscape features (proportion of adjacent anthropogenic land cover, distance to nearest road or river) and climate. We used generalized additive models (GAM) to assess which variables had greatest importance in determining non-native richness and mean ground cover and whether these variables had a similar effect on native species in the ground tier.

Results

A positive relationship between native and non-native plant species richness was not due to their similar responses to the variables examined in this study. Higher native canopy richness resulted in lower non-native richness and mean ground cover, whereas higher native ground richness was associated with higher native canopy richness. Non-native richness and mean ground cover increased with the proportion of adjacent anthropogenic land cover, whereas for native richness and mean ground cover, this relationship was negative. Non-native richness increased in drier areas, while native richness was more influenced by temperature.

Main Conclusions

Adjacent anthropogenic land cover seems to not only facilitate non-native species arrival by being a source of propagules but also aids their establishment as a result of fragmentation. Our results highlight the importance of examining both cover and richness in different vegetation tiers to better understand non-native plant invasions.     

Neophyte species richness at the landscape scale under urban sprawl and climate warming

Aim Land use and climate are two major components of global environmental change but our understanding of their simultaneous and interactive effects upon biodiversity is still limited. Here, we investigated the relationship between the species richness of neophytes, i.e. non‐native vascular plants introduced after 1500 AD, and environmental covariates to draw implications for future dynamics under land‐use and climate change. Location Switzerland, Central Europe. Methods The distribution of vascular plants was derived from a systematic national grid of 1 km² quadrates (n = 456; Swiss Biodiversity Monitoring programme) including 1761 species, 122 of which were neophytes. Generalized linear models (GLMs) were used to correlate neophyte species richness with environmental covariates. The impact of land‐use and climate change was thereafter evaluated by projections for the years 2020 and 2050 using scenarios of moderate and strong changes for climate warming (IPCC) and urban sprawl (NRP 54). Results Mean annual temperature and the amount of urban areas explained neophyte species richness best, with a high predictive power of the corresponding model (cross‐validated D² = 0.816). Climate warming had a stronger impact on the potential increase in the mean neophyte species richness (up to 191% increase by 2050) than ongoing urban sprawl (up to 10% increase) independently from variable interactions and model extrapolations to non‐analogue environments. Main conclusions In contrast to other vascular plants, the prediction of neophyte species richness at the landscape scale in Switzerland requires few variables only, and regions of highest species richness of the two groups do not coincide. The neophyte species richness is basically driven by climatic (temperature) conditions, and urban areas additionally modulate small‐scale differences upon this coarse‐scale pattern. According to the projections climate warming will contribute to the future increase in neophyte species richness much more than ongoing urbanization, but the gain in new neophyte species will be highest in urban regions.     

Anthropogenic fires increase alien and native annual species in the Chilean coastal matorral

Aim We tested the hypothesis that anthropogenic fires favour the successful establishment of alien annual species to the detriment of natives in the Chilean coastal matorral. Location Valparaíso Region, central Chile. Methods We sampled seed rain, seedbank emergence and establishment of species in four paired burned and unburned areas and compared (using GLMM) fire resistance and propagule arrival of alien and native species. To assess the relative importance of seed dispersal and seedbank survival in explaining plant establishment after fire, we compared seed rain and seedbank structure with post‐fire vegetation using ordination analyses. Results Fire did not change the proportion of alien species in the coastal matorral. However, fire increased the number of annual species (natives and aliens) of which 87% were aliens. Fire reduced the alien seedbank and not the native seedbank, but alien species remained dominant in burned soil samples (66% of the total species richness). Seed rain was higher for alien annuals than for native annuals or perennials, thus contributing to their establishment after fire. Nevertheless, seed rain was less important than seedbank survival in explaining plant establishment in burned areas. Main conclusions Anthropogenic fires favoured alien and native annuals. Thus, fire did not increase the alien/native ratio but increased the richness of alien species. The successful establishment of alien annuals was attributable to their ability to maintain rich seedbanks in burned areas and to the greater propagule arrival compared to native species. The native seedbank also survived fire, indicating that the herbaceous community has become highly resilient after centuries of human disturbances. Our results demonstrate that fire is a relevant factor for the maintenance of alien‐dominated grasslands in the matorral and highlight the importance of considering the interactive effect of seed rain and seedbank survival to understand plant invasion patterns in fire‐prone ecosystems.     

怒江河谷入侵植物与乡土植物丰富度的分布格局与影响因子

外来物种入侵严重威胁着乡土植物多样性并削弱了生态系统服务功能。本文基于滇西北怒江河谷植被调查的样方数据, 从群落水平研究了乡土和入侵植物多样性的空间分布格局, 以及地形、气候、人类干扰等因子对两种格局的影响。本研究共记录到外来入侵植物26种, 隶属于13科21属; 乡土植物1,145种, 分属于158科628属。沿着怒江河谷, 入侵植物物种丰富度随纬度与海拔的增加而减少; 乡土物种丰富度则随纬度增加而增加, 并在海拔梯度上呈单峰格局。运用广义线性模型分析公路边缘效应(反映生境干扰)、气候、地形和土壤等环境因素对物种丰富度分布格局的影响。等级方差分离的结果显示, 公路两侧的生境干扰对入侵种和乡土种的丰富度格局均具有首要影响。在自然环境因子中, 降水量是入侵植物丰富度的主要限制因子, 而乡土物种丰富度则主要受到地形因子尤其是坡向的影响。结构方程模型的分析结果也表明, 乡土植物和入侵植物丰富度之间的负相关关系反映了二者对环境响应的差异。本文结果支持物种入侵的资源可利用性限制假说, 并强调了人类活动对生物多样性的负面影响; 乡土植物或已较好地适应了干旱河谷气候, 但并没有显示出对外来物种入侵的抵抗作用。     

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

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

Species richness of herbivores on exotic host plants increases with time since introduction of the host

Aim Species richness of insect herbivores feeding on exotic plants increases with abundance as well as range size of the host in the area of introduction. The formation of these herbivore assemblages requires a certain amount of time, and the richness of insect faunas should also increase with the length of time an exotic plant has been present in the introduced range. Location Central Europe. Methods We analysed the variation in species richness of leaf‐chewing Lepidoptera larvae and sap‐sucking Auchenorrhyncha (Hemiptera) associated with 103 exotic woody plant species in Germany in relation to time since introduction, range size, growth form (trees versus shrubs), biogeographical origin (distance from Central Europe) and taxonomic isolation of the host plant (presence or absence of a native congener in the introduced area). Results Using simple correlation analyses we found for Lepidoptera and Auchenorrhyncha that species richness increased with time since introduction of the host plant. For the Lepidoptera the increase of species richness with time since introduction remained significant even after removing the effects of all other independent variables. Main conclusions Our results provide some evidence that assemblages of insects on exotic plants do not reach saturation within a time scale of few hundred years. This contrasts with previous findings for crop plants.     

Temperature-dependent performance of competitive native and alien invasive plant species

To assess the likely impacts of environmental change, the responses of two well-known invasive plant species, native Pueraria lobata and alien Humulus japonicus, to differences in growth temperature were studied in South Korea. Habitat preferences, physiological responses such as photosynthetic rates and chlorophyll contents, growth rates, and nutrient contents were quantified for each species. A competition experiment was conducted to evaluate the temperature preferences of the two species. All results indicated that the alien species H. japonicus can take advantage of elevated temperatures (35 °C) to enhance its competitive advantage against the native species P. lobata. While H. japonicus took advantage of elevated temperatures and preferred high-temperature areas, P. lobata showed reduced performance and dominance in high-temperature areas. Therefore, in future, due to global warming and urbanization, there are possibilities that H. japonicus takes advantage of elevated temperature against P. lobata that could lead to increased H. japonicus coverage over time. Therefore, consistent monitoring of both species especially where P. lobata is dominated are required because both species are found in every continents in the world. Controlling P. lobata requires thorough inspection of H. japonicus presence of the habitat in advance to prevent post P. lobata management invasion of H. japonicus.     

Comparing agglomerative clustering and three weed classification frameworks to assess the invasiveness of alien species across spatial scales

To prioritize weed management at the catchment scale, information is required on the species present, their relatively frequency, abundance, and likely spread and impact. The objective of this study was to classify the invasiveness of alien species that have invaded the Upper Burdekin Catchment in Queensland, Australia, at three spatial scales. A combination of three published weed classification frameworks and multivariate techniques were employed to classify species based on their frequency and cover at a range of spatial scales. We surveyed the Upper Burdekin Catchment for alien species, and for each species determined the following distribution indices — site frequency, total cover, transect frequency per site frequency and quadrat frequency per site frequency, cover per quadrat when present, cover per transect when present, and cover per site when present. These indices capture the effect of species abundance and frequency between sites (site frequency and total cover), within sites (transect frequency per site and cover per transect when present), and within transects (quadrat frequency per site frequency and cover per site). They were used to classify the species into seven groups using a hierarchical cluster analysis. The relationship between the indices was explored to determine how effective the small scale, site‐specific indices were at predicting the broader, landscape‐scale patterns. Strong correlations were observed between transect frequency per site and frequency (r² = 0.89) and cover per transect when present and total cover (r² = 0.62). This suggests that if a weed is abundant at the site level, it has the potential to occupy large areas of the catchment. The species groupings derived from the application of the three published weed classification frameworks were compared graphically to the groupings derived from the cluster analysis. One of the frameworks classified species into three groups. The other two frameworks classified species into four groups. There was a high degree of subjectivity in applying the frameworks to the survey data. Some of the data were of no relevance to the classification frameworks and were therefore ignored. We suggest that the weed classification frameworks should be used in conjunction with existing multivariate techniques to ensure that classifications capture important natural variations in observed data that may reflect invasion processes. The combined use of the frameworks and multivariate techniques enabled us to aggregate species into categories appropriate for management.