Deconstructing the native–exotic richness relationship in plants

Aim Classic theory suggests that species‐rich communities should be more resistant to the establishment of exotic species than species‐poor communities. Although this theory predicts that exotic species should be less diverse in regions that contain more native species, macroecological analyses often find that the correlation between exotic and native species richness is positive rather than negative. To reconcile results with theory, we explore to what extent climatic conditions, landscape heterogeneity and anthropogenic disturbance may explain the positive relationship between native and exotic plant richness. Location Catalonia (western Mediterranean region). Methods We integrated floristic records and GIS‐based environmental measures to make spatially explicit 10‐km grid cells. We asked whether the observed positive relationship between native and exotic plant richness (R²= 0.11) resulted from the addition of several negative correlations corresponding to different environmental conditions identified with cluster analysis. Moreover, we directly quantified the importance of common causal effects with a structural equation modelling framework. Results We found no evidence that the relationship between native and exotic plant richness was negative when the comparison was made within environmentally homogeneous groups. Although there were common factors explaining both native and exotic richness, mainly associated with landscape heterogeneity and human pressure, these factors only explained 17.2% of the total correlation. Nevertheless, when the comparison was restricted to native plants associated with human‐disturbed (i.e. ruderal) ecosystems, the relationship was stronger (R²= 0.52) and the fraction explained by common factors increased substantially (58.3%). Main conclusions While our results confirm that the positive correlation between exotic and native plant richness is in part explained by common extrinsic factors, they also highlight the great importance of anthropic factors that – by reducing biotic resistance – facilitate the establishment and spread of both exotic and native plants that tolerate disturbed environments.     

Proportion of exotics and relatedness of host species mediate the positive effect of plant richness on the species richness of fruit flies

1. All else being equal, the greater the local species richness of plants, the greater the number of associated herbivore species. Because most herbivore insects feed on a subset of closely related plant species, plant phylogenetic diversity is expected to play a key role in determining the number of herbivore species. What is not well known, however, is how an increase in the species richness of exotic plants affects the species richness of herbivores. 2. In this study, we used plant–fruit fly interactions to investigate the influence of the proportion and species richness of exotic host plants on the species richness of herbivorous insects. We also tested whether the phylogenetic diversity of host plants increases when the number of exotic plant species increases. 3. We found that the species richness of fruit flies is more accurately predicted by the richness of native host plants than by total plant species richness (including both native and exotic species). The proportion of exotic host species and the phylogenetic diversity of host plants had negative and positive effects, respectively, on the species richness of fruit flies. 4. Our findings suggest that a positive effect of plant richness on herbivore richness occurs only when an increase in plant diversity involves plant species with which native herbivores share some evolutionary history.     

An Essay on Some Topics Concerning Invasive Species

Abstract There is a tendency for both scientists and lay people to regard invading alien species as inherently ‘bad’ and native species as inherently ‘good.’ Past invasions occurred commonly without human assistance. They rarely caused large, lasting decreases in species richness or ecological damage. Current invasions provide opportunities for scientific study. They are unintentional, uncontrolled experiments, which can provide insights into attributes of successful colonists, relationships with native species, and impacts on the structure and function of ecological systems.     

Controls on pathogen species richness in plants' introduced and native ranges: roles of residence time, range size and host traits

Introduced species escape many pathogens and other enemies, raising three questions. How quickly do introduced hosts accumulate pathogen species? What factors control pathogen species richness? Are these factors the same in the hosts' native and introduced ranges? We analysed fungal and viral pathogen species richness on 124 plant species in both their native European range and introduced North American range. Hosts introduced 400 years ago supported six times more pathogens than those introduced 40 years ago. In hosts' native range, pathogen richness was greater on hosts occurring in more habitat types, with a history of agricultural use and adapted to greater resource supplies. In hosts' introduced range, pathogen richness was correlated with host geographic range size, agricultural use and time since introduction, but not any measured biological traits. Introduced species have accumulated pathogens at rates that are slow relative to most ecological processes, and contingent on geographic and historic circumstance.     

Species richness and interacting factors control invasibility of a marine community

Anthropogenic vectors have moved marine species around the world leading to increased invasions and expanded species'' ranges. The biotic resistance hypothesis of Elton (in The ecology of invasions by animals and plants, 1958) predicts that more diverse communities should have greater resistance to invasions, but experiments have been equivocal. We hypothesized that species richness interacts with other factors to determine experimental outcomes. We manipulated species richness, species composition (native and introduced) and availability of bare space in invertebrate assemblages in a marina in Monterey, CA. Increased species richness significantly interacted with both initial cover of native species and of all organisms to collectively decrease recruitment. Although native species decreased recruitment, introduced species had a similar effect, and we concluded that biotic resistance is conferred by total species richness. We suggest that contradictory conclusions in previous studies about the role of diversity in regulating invasions reflect uncontrolled variables in those experiments that modified the effect of species richness. Our results suggest that patches of low diversity and abundance may facilitate invasions, and that such patches, once colonized by non-indigenous species, can resist both native and non-indigenous species recruitment.     

Feeding and breeding across host plants within a locality by the widespread thrips Frankliniella schultzei, and the invasive potential of polyphagous herbivores

Abstract. Polyphagous insect herbivores could be expected to perform relatively well in new areas because of their ability to exploit alternative resources. We investigated relative abundance patterns of the polyphagous thrips species Frankliniella schultzei , which is characteristically found on plants from many different families, to establish the role of different host plant species in a single locality where the species is not indigenous (Brisbane, south-eastern Queensland, Australia). F. schultzei females and larvae were always present in flowers (where oviposition takes place) and never on leaves of the eight plant species that we surveyed regularly over one year. They were present in flowers of Malvaviscus arboreus in much higher densities than for any other host. F. schultzei females were more fecund and larvae developed faster on floral tissue diets of M . arboreus than on those of other hosts. M. arboreus is therefore regarded as the 'primary' host plant of F. schultzei in the locality that we investigated. The other species are regarded as 'minor' hosts. Available evidence indicates a common geographical origin of F. schultzei and M. arboreus . F. schultzei may therefore be primarily adapted to M. arboreus . The flowers of the minor species on which F. schultzei is also found may coincidentally share some features of the primary host. Adult thrips may therefore accumulate on minor hosts and breed there, but to a lesser extent than on the primary host. The general implications for investigating polyphagous host relationships and interpreting the ecology of these species as generalist invaders are spelt out.     

Minimum residence time, biogeographical origin, and life cycle as determinants of the geographical extent of naturalized plants in continental Chile

The geographical extent of exotic plant species is a major component of invasiveness, which has been explained by intrinsic attributes of the plants, such as growth rate, reproductive type, life form, and biogeographical origin. We assessed quantitatively life cycle and biogeographical origin as determinants of the geographical distribution of naturalized plants in continental Chile, using minimum residence time (MRT) as an estimator of introduction date. We assembled a database with information on 428 plants (principally herbs) in continental Chile, corresponding to 61% of the exotic naturalized flora. For each species we recorded: (1) minimum residence time (introduction date or first recorded date in the country); (2) biogeographical origin (American, Eurasian, others); (3) life cycle (annual, perennial, others); (4) number of Chilean regions occupied by the plant. We found that 82 species (19%) have been recorded in only one region of Chile, while only three species have been found in all 13 regions of the country. About 89% of the species (381) have been found only in central Chile (Regions IV to VIII), while the remaining 11% (47) are found only in the northern (Regions I to III) or southern parts of the country (Regions IX to XII). We detected significant differences in regional spread of naturalized plants according to minimum residence time: those species with shorter MRT had more limited spread ranges than those with longer MRT. Biogeographical origin and life cycle did not explain geographical extent in Chile. This study shows that historical factors are more important than biological ones in determining the geographical extent of naturalized plants in continental Chile. Thus, caution should be exercised when assigning value to biological attributes that may confer invasiveness to naturalized plants.     

BIODIVERSITY RESEARCH: Native‐exotic species richness relationships across spatial scales and biotic homogenization in wetland plant communities of Illinois,USA

Aim To examine native‐exotic species richness relationships across spatial scales and corresponding biotic homogenization in wetland plant communities. Location Illinois, USA. Methods We analysed the native‐exotic species richness relationship for vascular plants at three spatial scales (small, 0.25 m² of sample area; medium, 1 m² of sample area; large, 5 m² of sample area) in 103 wetlands across Illinois. At each scale, Spearman’s correlation coefficient between native and exotic richness was calculated. We also investigated the potential for biotic homogenization by comparing all species surveyed in a wetland community (from the large sample area) with the species composition in all other wetlands using paired comparisons of their Jaccard’s and Simpson’s similarity indices. Results At large and medium scales, native richness was positively correlated with exotic richness, with the strength of the correlation decreasing from the large to the medium scale; at the smallest scale, the native‐exotic richness correlation was negative. The average value for homogenization indices was 0.096 and 0.168, using Jaccard’s and Simpson’s indices, respectively, indicating that these wetland plant communities have been homogenized because of invasion by exotic species. Main Conclusions Our study demonstrated a clear shift from a positive to a negative native‐exotic species richness relationship from larger to smaller spatial scales. The negative native‐exotic richness relationship that we found is suggested to result from direct biotic interactions (competitive exclusion) between native and exotic species, whereas positive correlations likely reflect the more prominent influence of habitat heterogeneity on richness at larger scales. Our finding of homogenization at the community level extends conclusions from previous studies having found this pattern at much larger spatial scales. Furthermore, these results suggest that even while exhibiting a positive native‐exotic richness relationship, community level biotas can/are still being homogenized because of exotic species invasion.     

Overstorey tree species regulate colonization by native and exotic plants: a source of positive relationships between understorey diversity and invasibility

The North American woody species, Prunus serotina Ehrh., is an aggressive invader of forest understories in Europe. To better understand the plant invasion process, we assessed understorey plants and Prunus serotina seedlings that have colonized a 35-year-old replicated common-garden experiment of 14 tree species in south-western Poland. The density and size of established (> 1 year old) P. serotina seedlings varied among overstorey species and were related to variation in light availability and attributes of the understorey layer. In a multiple regression analysis, the density of established P. serotina seedlings was positively correlated with light availability and understorey species richness and negatively correlated with understorey species cover. These results suggest that woody invader success is adversely affected by overstorey shading and understorey competition for resources. Simultaneously, however, invader success may generally be positively associated with understorey species richness because both native and invasive plant colonization respond similarly to environmental conditions, including those influenced by overstorey tree species. Identification of characteristics of forests that increase their susceptibility to invasion may allow managers to target efforts to detect invasives and to restore forests to states that may be less invasible.     

Invasibility of mature and 15-year-old deciduous forests by exotic plants

High species richness, resource availability and disturbance are community characteristics associated with forest invasibility. We categorized commonly measured community variables, including species composition, topography, and landscape features, within both mature and 15-year-old clearcuts in West Virginia, USA. We evaluated the importance of each variable for predicting the degree of forest invasion by early-establishing exotic invasive plants. Biotic variables, including overall richness (excluding exotic invasive species) and mutually exclusive native and exotic non-invasive species richness, were the strongest indicators of invasibility. Sites that were located on northeast-facing slopes, more mesic conditions, or in clearcuts were more likely to be invaded by exotic invasive plants. Invasion of clearcut sites was more dependent on available microsites (e.g., lower solar radiation, northeast-facing slopes, and lower elevations) within each site than on the condition of the surrounding landscape, whereas invasion into the mature forests was dependent more on the surrounding landscape (e.g., proximity to paved roads). Our results indicate that exotic invasive plant species in our study area respond similarly as other plant species to resource availability and that competitive interactions are relatively unimportant. Current invasion into this landscape is more likely to be a passive reaction to site conditions instead of a driver of change.     

The impact of an exotic social wasp (Vespula germanica) on the native arthropod community of north-west Patagonia, Argentina: an experimental study

Abstract 1. Biological invasions are usually thought to have a negative impact on native communities. However, data supporting this idea are often based on comparative studies between invaded and non‐invaded areas, and are spatially and temporally limited. 2. The present study experimentally assessed the impact of an exotic wasp, Vespula germanica, on the native arthropod community of north‐west Patagonia during 3 years in an area of 80 ha. Vespula germanica is an exotic social vespid that invaded north‐west Patagonia 20 years ago. It has been suggested that its populations affect native arthropods because of its broad diet and also because Patagonia lacks natural enemies and potential competitors for these wasps. 3. Using wasp‐specific toxic baits, V. germanica abundance was reduced in five sites of native woodlands during 3 consecutive years. The abundance, species richness, and composition of arthropods between non‐poisoned (control) and poisoned sites was then compared, both before and after the wasps were poisoned. 4. Wasp abundance represented 6% of the total arthropod catches in non‐poisoned sites and was reduced, on average, by 50% in the treated areas. The abundance, species richness, and composition of the arthropod community (305 species, 24 600 individuals) did not differ between control areas and areas where the abundance of V. germanica was reduced. Significant differences in response variables were found only before wasp poisoning had begun and were related to variations among sites. 5. These results suggest that V. germanica is not affecting the local arthropod assemblages, contradicting past work in other regions. The low relative abundance of wasps in Patagonia, when compared with other invaded regions, might explain the findings. 6. The present study provides further evidence for the importance of large‐scale experimental work with before/after comparisons to fully understand the impact of invaders on natural communities.     

Vulnerability of riparian zones to invasion by exotic vascular plants

We compared the invasibility of riparian plant communities high on river banks with those on floodplain floors for four South African rivers. Analyses of abundant and significant riparian species showed that the floors have 3.1 times more exotic plants than the banks. The percent exotics ranges from 5% to 11% of total species richness for the banks, and from 20% to 30% for the floors. Species richness and percent exotics are negatively correlated for the banks, but not correlated for the floors.Despite great differences in climate, species richness, and landuse history, the percentages of exotic plants in three rivers in the Pacific Northwest of the USA and one river in southwestern France are similar to those in South Africa (24-30% vs. 20-30%, respectively). Furthermore, the high proportions of exotic species in these riparian plant communities are comparable to those reported for vascular plant communities on islands. We conclude that the macro-channel floor regions of the riparian zones of South African rivers are highly vulnerable to invasion by exotic vascular plants.     

Species richness of sawfly–host plant associations at higher taxonomic levels

Insect–plant interactions are important to understanding the evolutionary mechanisms responsible for most of the diverse plant‐feeding insects. We aggregated data on sawfly–host plant associations and other resource associations from existing sources to address the following questions: (i) Is there a general correlation between host diversity and sawfly species richness? (ii) Is the pattern of host plant use consistent across sawfly lineages? and (iii) Is there a phylogenetically significant shift in species richness among sawflies? Our analysis comprised 8567 sawfly species, including 2087 species with host plant and other records. In total, there were 2126 records of host usage for sawflies, the overwhelming majority of which were sawflies using angiosperms as resources. Rosales are used by most of the species in sawfly families or subfamilies. We found that there was a strong correlation between the number of host plant orders and the species richness of sawfly families and subfamilies. To examine the points at which sawflies have experienced significant shifts in species richness, we compared sister taxon species richness. Several positive and negative shifts in species richness among sawflies were related to their range of host plant usage and specialized niche, respectively. In general, we found that most of the sawfly families and subfamilies used several orders as host plants, but mainly core eudicots, although some families or subfamilies were specialized on pteridophytes or gymnosperms.     

Sympatric host races of the European corn borer: adaptation to host plants and hybrid performance

The European corn borer (ECB), Ostrinia nubilalis, is a major pest of maize crops. In Europe, two sympatric host races are found: one feeds on maize (Zea mays) and the other mainly on mugwort (Artemisia vulgaris). The two host races are genetically differentiated, seldom crossing in the laboratory or in the field, and females preferentially lay eggs on their native host species. We conducted two independent experiments, in field and greenhouse conditions, to determine whether the two host races are locally adapted to their host species. The effect of larval density and the performance of hybrids were also investigated. Despite some differences in overall larval feeding performance, both experiments revealed consistent patterns of local adaptation for survival and for larval weight in males. In females the same trend was observed but with weaker statistical support. F1 hybrids did not seem to be disadvantaged compared with the two parental races. Overall, our results showed that both host races are physiologically adapted to their native host. The fitness trade-off between the two host plants provides a potential driving force for ecological speciation in this species.     

Species richness and spatial distribution of blackflies (Diptera: Simuliidae) in streams of Central Amazonia, Brazil

1. The spatial distribution and species richness of blackflies were evaluated at 58 stream sites in Central Amazonia, Brazil. Samples were taken along a north–south axis of approximately 130 km and a east–west axis of approximately 220 km.
2. Based on stream-site characteristics, the occurrence of larvae of the six most frequently collected species was highly predictable (79.3–91.5% accuracy in prediction of occurrence). The predictive value of stream size and the presence of impoundments agrees with results of similar work in the Holarctic Region, suggesting a general responses of blackflies to environmental parameters.
3. Although only 19.0% of interstream variation in species richness was explained by a regression model, results suggested that species richness was greater in larger, cooler, faster, covered streams with rocky beds than in smaller, warmer, slower, open streams with sandy bottoms. Overall, the species richness of blackflies (11 species in total) was lower than in the temperate zone suggesting, for some taxa at least, that aquatic communities do not follow the terrestrial pattern of greater species richness in the tropics.     

植物外来种对土壤理化特性的影响

植物外来种对土壤特性可产生深刻的影响。植物外来种可影响土壤的温度、湿度、土壤侵蚀等土壤物理特性 ;同样会影响土壤的含氮量、有机质、金属元素等化学特性。植物外来种对土壤特性的影响是十分复杂的 ,在不同的环境里同一植物外来种可产生不同的影响。总的来说 ,植物外来种对土壤特性的影响有正面、中性和负面 3种效应