Continental rain forest fragments in Singapore resist invasion by exotic plants

Abstract Aim In general, the plant communities of oceanic islands suffer more from exotic plant invasions than their continental equivalents. At least part of this difference may be contributed by differences in non‐biological factors, such as the antiquity and intensity of human impacts and the absence of internal barriers to dispersal, rather than differences in inherent invasibility. We tested the resistance of species‐rich continental rain forests to plant invasion on a small, continental island that has been subject to prolonged and intensive human impact. Location Singapore is a 683‐km² equatorial island <1 km from the Asian mainland and with a population of 4 million people. It has a continental biota but has been subject to human impacts as intense as on any oceanic island. Methods We sampled twenty‐nine sites in seven vegetation types, ranging from urban wasteland to fragments of primary lowland rain forest. In each sample plot, all plant species were identified, exotic cover was estimated, and a range of environmental variables measured. Additional qualitative surveys for exotic invasion were made in other forest areas in Singapore. The data were analysed by Spearman's rank correlation coefficient. Results The number of exotic species recorded at a site was unrelated to the number of native species. Across all sites, percentage canopy opening had the highest correlation with the number of exotic species, while soil pH (which largely reflects the incorporation of calcareous construction wastes) had the highest correlation if the mangrove sites were excluded. There were no exotics in mangrove forest and only a tropical American, bird‐dispersed shrub, Clidemia hirta (L.) D. Don (Melastomataceae: Koster's Curse), in primary and tall secondary forest patches. The species‐poor early stages of woody plant succession on highly degraded soils were also very resistant to exotic plant invasion. Main conclusions Long‐isolated rain forest fragments in an exotic‐dominated continental island landscape resist invasion by exotic plants, suggesting that the problems on oceanic islands may reflect an inherently greater invasibility. This study also adds to the increasing evidence that the floras of tropical rain forest fragments in South‐east Asia are remarkably resilient on a time‐scale of decades to a century or more.     

Across island and continents, mammals are more successful invaders than birds (Reply)

Aim  The establishment success of exotic species is calculated as the fraction of introduced species that have become established, and invasion success is estimated as the fraction of established species that have spread significantly from their points of introduction. Records on species introductions are highly incomplete, so strong conclusions about the tens rule and invasibility of island and continents cannot be drawn.
Location  Global.
Methods  Using Jeschke (2008) as an example, we explain the issue of the inadequacy of data to draw conclusions about the tens rule and invasibility of island and continents.
Results  Lack of adequate data.
Main Conclusion  Jeschke (2008) probably overestimates the establishment and invasion success rates, so his conclusions about violation of the tens rule and that islands are not more susceptible to invasion than continents are misleading.     

Spatial Scaling of Non-Native Fish Richness across the United States

A major goal and challenge of invasion ecology is to describe and interpret spatial and temporal patterns of species invasions. Here, we examined fish invasion patterns at four spatially structured and hierarchically nested scales across the contiguous United States (i.e., from large to small: region, basin, watershed, and sub-watershed). All spatial relationships in both richness and fraction between species groups (e.g., natives vs. exotics) were positive at large scales. However, contrary to predictions using null/neutral models, the patterns at small scales were hump-shaped (unimodal), not simply negative. The fractions of both domestic (introduced among watersheds within the USA) and foreign (introduced from abroad) exotics increased with area across scales but decreased within each scale. The foreign exotics exhibited the highest dominance (lowest evenness) and spatial variation in distribution, followed by domestic exotics and natives, although on average natives still occupy larger areas than domestic and foreign exotics. The results provide new insight into patterns and mechanisms of fish species invasions at multiple spatial scales in the United States.     

Local and regional abundance of exotic plant species on Mediterranean islands: are species traits important?

Aim We assess the importance of three relevant and readily obtainable life‐history traits (dispersal syndrome, stem height and growth form) and biogeographical origin (European vs. non‐European) on the local and regional abundance of over 400 exotic plant species across eight Mediterranean islands. Location The Mediterranean islands of Lesbos, Rhodes, Crete, Malta, Corsica, Sardinia, Majorca and Minorca. Methods We adopt two abundance criteria for each exotic species: the proportion of islands in which the species occurs (regional abundance), and a qualitative estimate of species abundance within each of five islands (local abundance). Subsequently, we assess the relationship between local and regional abundance, as well as the role of key life‐history traits on both regional and local abundance. These analyses were undertaken separately for the European exotics and the non‐European exotics. Results Only 10.9% of the species occur on more than four islands, and only four species are present on all eight islands. Both local and regional abundances were higher for the non‐European than the European species. Local and regional abundances were positively correlated, particularly for exotics with non‐European origins. Wind‐dispersed species tended to have higher regional abundance than species dispersed by other means but this trend only occurred for local abundance on two islands — Corsica and Majorca. Neither a species’ growth form nor its stem height explained trends in regional or local abundance. Conclusions Although wind‐dispersed exotics are more widespread in the Mediterranean, plant life‐history traits appear to play a lesser role in invasion success than area of biogeographical origin. In general, exotic species of non‐European origin were more abundant at both local and regional scales. Invasion patterns should be interpreted at both local and regional scales, but the stochastic nature of biological invasions may limit deterministic interpretations of invasion patterns, especially if islands are studied in isolation.     

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.     

Exotic Vertebrate Fauna in the Remote and Pristine Sub-Antarctic Cape Horn Archipelago, Chile

Pristine wilderness is a scarce global resource, but exotic species are so common that they constitute a principal component of worldwide ecological change. The relationship between these two topics, invasion and remoteness, was the impetus behind five years of fieldwork aimed at identifying the assemblage and range of exotic vertebrates in Cape Horn, Chile, identified as one of the world's most pristine wilderness areas. While the archipelago has extremely low human population density and vast tracts of undisturbed land, we discovered that several terrestrial vertebrate groups were dominated by exotic species. Native birds were diverse (approx. 154 spp), and only 1.3% (or two spp.) were introduced. In contrast, exotic terrestrial mammals (12 spp.) and freshwater fish (three spp.) outnumbered their native counterparts, constituting 55% and 75% of the assemblages. Using GIS, we found that not all areas were impacted equally, largely due to intensity of human occupation. Human settled islands (Navarino and Tierra del Fuego) hosted the greatest number of exotics, but humans alone did not explain observed patterns. Remote islands also had introduced species. North American beavers (Castor canadensis), American minks (Mustela vison) and feral domestic dogs and cats were particularly widespread, and their range in isolated parts of the study area raised important ecological and management questions. In conclusion, the Cape Horn Archipelago retained areas free of exotic vertebrates, particularly parts of the Cape Horn and Alberto D’Agostini National Parks, but at many sites introduced species were overwhelming native biota and altering these previously remote natural ecosystems.     

Exotic species richness and native species endemism increase the impact of exotic species on islands

Aim Exotic species pose one of the most significant threats to biodiversity, especially on islands. The impacts of exotic species vary in severity among islands, yet little is known about what makes some islands more susceptible than others. Here we determine which characteristics of an island influence how severely exotic species affect its native biota. Location We studied 65 islands and archipelagos from around the world, ranging from latitude 65° N to 54° S. Methods We compiled a global database of 10 island characteristics for 65 islands and determined the relative importance of each characteristic in predicting the impact of exotic species using multivariate modelling and hierarchical partitioning. We defined the impact of exotic species as the number of bird, amphibian and mammal (BAM) species listed by the International Union for Conservation of Nature (IUCN) as threatened by exotics, relative to the total number of BAM species on that island. Results We found that the impact of exotic species is more severe on islands with more exotic species and a greater proportion of native species that are endemic. Unexpectedly, the level of anthropogenic disturbance did not influence an island's susceptibility to the impacts of exotic species. Main conclusions By coupling our results with studies on the introduction and establishment of exotic species, we conclude that colonization pressure, or invasion opportunities, influences all stages of the invasion process. However, species endemism, the other important factor determining the impact of exotic species, is not known to contribute to introduction and establishment success on islands. This demonstrates that different factors correlate with the initial stages of the invasion process and the subsequent impacts of those invaders, highlighting the importance of studying the impacts of exotic species directly. Our study helps identify islands that are at risk of impact by exotics and where investment should focus on preventing further invasions.     

A multi-scale assessment of the occurrence of exotic plants on the Olympic Peninsula,Washington

Abstract. Exotic plants were surveyed in 208 plots within the Dungeness and Hoh river watersheds on the Olympic Peninsula, Washington, USA. Landscape patch types included uplands (clearcuts, young and mature forests) and riparian zones (cobble bars, shrub patches, riparian forests, and alder flats). Patterns of exotic plant invasions were assessed between watersheds, between riparian and upland areas, among patch types, and within clearcuts. 52 exotic plant species were encountered, accounting for 23% of the flora in each watershed. In both watersheds, exotic species richness was approximately 33% greater in riparian zones than in uplands, and mean number and cover of exotic species were > 50% greater in riparian zones than in uplands. Among landscape patch types, richness and mean number and cover of exotics was highest in young riparian patches, intermediate in clearcuts and riparian forests, and lowest in young and mature forests. The exception to this was Hoh alder flats, which had the highest mean cover of exotic plants. Cover of exotic plants peaked in uplands 3 to 7 yr after clearcutting, then decreased with increased canopy closure. Disturbance type and time since disturbance were major factors influencing invasibility. Landscape patch size, position within watershed (distance from patch to human population centers, major highway, or river mouth), and environmental variables (slope, aspect, and elevation) were not important indicators of landscape patch invasibility. Riparian zones facilitated movement of exotic plants through landscapes, but did not appear to act as sources of exotic plants for undisturbed upland areas.     

Species pool size and invasibility of island communities: a null model of sampling effects

The success of alien species on oceanic islands is considered to be one of the classic observed patterns in ecology. Explanations for this pattern are based on lower species richness on islands and the lower resistance of species‐poor communities to invaders, but this argument needs re‐examination. The important difference between islands and mainland is in the size of species pools, not in local species richness; invasibility of islands should therefore be addressed in terms of differences in species pools. Here I examine whether differences in species pools can affect invasibility in a lottery model with pools of identical native and exotic species. While in a neutral model with all species identical, invasibility does not depend on the species pool, a model with non‐zero variation in population growth rates predicts higher invasibility of communities of smaller pools. This is because of species sampling; drawing species from larger pools increases the probability that an assemblage will include fast growing species. Such assemblages are more likely to exclude random invaders. This constitutes a mechanism through which smaller species pools (such as those of isolated islands) can directly underlie differences in invasibility.     

Effects of native species diversity and resource additions on invader impact

Theory and empirical work have demonstrated that diverse communities can inhibit invasion. Yet, it is unclear how diversity influences invader impact, how impact varies among exotics, and what the relative importance of diversity is versus extrinsic factors that themselves can influence invasion. To address these issues, we established plant assemblages that varied in native species and functional richness and crossed this gradient in diversity with resource (water) addition. Identical assemblages were either uninvaded or invaded with one of three exotic forbs: spotted knapweed (Centaurea maculosa), dalmatian toadflax (Linaria dalmatica), or sulfur cinquefoil (Potentilla recta). To determine impacts, we measured the effects of exotics on native biomass and, for spotted knapweed, on soil moisture and nitrogen levels. Assemblages with high species richness were less invaded and less impacted than less diverse assemblages. Impact scaled with exotic biomass; spotted knapweed had the largest impact on native biomass compared with the other exotics. Although invasion depressed native biomass, the net result was to increase total community yield. Water addition increased invasibility (for knapweed only) but had no effect on invader impact. Together, these results suggest that diversity inhibits invasion and reduces impact more than resource additions facilitate invasion or impact.     

Landscape-scale patterns of biological invasions in shoreline plant communities

Little is known about the patterns and dynamics of exotic species invasions at landscape to regional spatial scales. We quantified the presence (identity, abundance, and richness) and characteristics of native and exotic species in estuarine strandline plant communities at 24 sites in Narragansett Bay, Rhode Island, USA. Our results do not support several fundamental predictions of invasion biology. Established exotics (79 of 147 recorded plant species) were nearly indistinguishable from the native plant species (i.e. in terms of growth form, taxonomic grouping, and patterns of spatial distribution and abundance) and essentially represent a random sub-set of the current regional species pool. The cover and richness of exotic species varied substantially among quadrats and sites but were not strongly related to any site-level physical characteristics thought to affect invasibility (i.e. the physical disturbance regime, legal status, neighboring habitat type, and substrate characteristics). Native and exotic cover or richness were not negatively related within most sites. Across sites, native and exotic richness were positively correlated and exotic cover was unrelated to native richness. The colonization and spread of exotics does not appear to have been substantially reduced at sites with high native diversity. Furthermore, despite the fact that the Rhode Island strandline system is one of the most highly-invaded natural plant communities described to date, exotic species, both individually and as a group, currently appear to pose little threat to native plant diversity. Our findings are concordant with most recent, large-scale investigations that do not support the theoretical foundation of invasion biology and generally contradict small-scale experimental work.     

Properties of native plant communities do not determine exotic success during early forest succession

Considerable research has been devoted to understanding how plant invasions are influenced by properties of the native community and to the traits of exotic species that contribute to successful invasion. Studies of invasibility are common in successionally stable grasslands, but rare in recently disturbed or seral forests. We used 16 yr of species richness and abundance data from 1 m² plots in a clearcut and burned forest in the Cascade Range of western Oregon to address the following questions: 1) is invasion success correlated with properties of the native community? Are correlations stronger among pools of functionally similar taxa (i.e. exotic and native annuals)? Do these relationships change over successional time? 2) Does exotic abundance increase with removal of potentially dominant native species? 3) Do the population dynamics of exotic and native species differ, suggesting that exotics are more successful colonists? Exotics were primarily annual and biennial species. Regardless of the measure of success (richness, cover, biomass, or density) or successional stage, most correlations between exotics and natives were non‐significant. Exotic and native annuals showed positive correlations during mid‐succession, but these were attributed to shared associations with bare ground rather than to direct biotic interactions. At peak abundance, neither cover nor density of exotics differed between controls and plots from which native, mid‐successional dominants were removed. Tests comparing nine measures of population performance (representing the pace, magnitude, and duration of population growth) revealed no significant differences between native and exotic species. In this early successional system, local richness and abundance of exotics are not explained by properties of the native community, by the presence of dominant native species, or by superior colonizing ability among exotics species. Instead natives and exotics exhibit individualistic patterns of increase and decline suggesting similar sets of life‐history traits leading to similar successional roles.     

Seed predation as a barrier to alien conifer invasions

Interactions between exotic plants and animals can play a major role in determining success or failure of plant introductions. Seed predation has been seen as important in explaining biotic resistance to plant invasion, but this hypothesis has rarely been tested. We studied seed predation on exotic forest plants on an island in Patagonia, Argentina where 43 pine species, including 60% of the world’s known invasive Pinaceae, were introduced ca. 80 years ago, but where exotics attain relatively high densities only near the original plantings. To test if seed predation limits exotic conifer establishment in this area, we compared seed predation in areas close to plantations (colonized by exotics) and far from them (not invaded). Seeds of exotics were preferred over seeds of native species, possibly because exotic seeds are bigger. Predation was more intense in areas far from plantations than in areas close to them, substantially reducing the chances of exotic seed establishment. Using automatic cameras, we found that both rodents and birds preyed on exotic seeds. This study suggests that native seed predators can be an important component of biological resistance to plant invasion.     

群落可侵入性及其影响因素

 可侵入性用于评价群落易遭受生物入侵的程度,受外来种死亡率、区域气候、干扰水平、生态系统抵抗入侵的能力、本地种竞争和抗干扰能力等因素的影响。当前对群落或区域间可侵入性的比较常以外来种数量或丰度为据,然而这两者仅代表了群落内单一的动态过程,不足以作为衡量群落可侵入性的广泛标准。借助一个描述外来种数量的简单模型阐明,由于影响可侵入性因素的复杂性,各地之间的可侵入性几乎不可比较。并从入侵过程、入侵种特性及本地种、本地群落对入侵的抵抗性几方面对群落可侵入性进行了阐述分析,其中着重介绍入侵生态中几个重要的概念,如可侵     

Characterizing the microhabitats of exotic species in Illinois shale barrens

Like native species, exotics form part of the mosaic structure of plant communities. However, their role in these communities is unclear. Thus, we ask, are invading exotic species present in recognizable microhabitats within plant communities? We compared the microhabitat of exotic species and random locations to determine if exotic plants occupied a recognizable subset of habitats within three barren communities. Fourteen exotic species were identified. The factors that characterized their habitats varied between barrens and among species. Considered as a group, exotic microhabitats comprised a subset of available habitats within the barrens, specifically, edge habitats rather than the open conditions of the barrens interior. Soil properties such as low temperature or neutral pH, and high litter or woody species cover, were common factors associated with the occurrence of exotics. Individual exotic species occupied different habitats within those identified for exotics as a group, but the range of variation between the different species was not generally significant. Within each barren, exotics occupied a distinct subset of microhabitats. This distribution suggests that exotics are not widely dispersed within these areas. Site conditions may be restricting the establishment of exotics to specific microhabitats, and in addition, some exotics may be altering the microhabitats that they colonize. Therefore, management efforts toward maintaining open conditions in the barrens may also discourage further encroachment by exotics into these areas.     

What do human economies,large islands and forest fragments reveal about the factors limiting ecosystem evolution?

What factors limit ecosystem evolution? Like human economies, ecosystems are arenas where agents compete for locally limiting resources. Like economies, but unlike genes, ecosystems are not units of selection. In both economies and ecosystems, productivity, diversity of occupations or species and intensity of competition presuppose interdependence among many different agents. In both, competitive dominants need abundant, varied resources, and many agents’ products or services, to support the activity and responsiveness needed to maintain dominance. Comparing different‐sized land masses suggests that productivity is lower on islands whose area is too small to maintain some of the interdependences that maintain diversity, productivity and competitiveness in mainland ecosystems. Islands lacking the rare, metabolically active dominants that make competition so intense in mainland ecosystems are more easily invaded by introduced exotics. Studies of islets in reservoirs identify mechanisms generating these phenomena. These phenomena suggest how continued fragmentation will affect future ‘natural’ ecosystems.     

Enemy release or invasional meltdown? Deer preference for exotic and native trees on Isla Victoria,Argentina

Abstract How interactions between exotic species affect invasion impact is a fundamental issue on both theoretical and applied grounds. Exotics can facilitate establishment and invasion of other exotics (invasional meltdown) or they can restrict them by re‐establishing natural population control (as predicted by the enemy‐release hypothesis). We studied forest invasion on an Argentinean island where 43 species of Pinaceae, including 60% of the world's recorded invasive Pinaceae, were introduced c. 1920 but where few species are colonizing pristine areas. In this area two species of Palearctic deer, natural enemies of most Pinaceae, were introduced 80 years ago. Expecting deer to help to control the exotics, we conducted a cafeteria experiment to assess deer preferences among the two dominant native species (a conifer, Austrocedrus chilensis, and a broadleaf, Nothofagus dombeyi) and two widely introduced exotic tree species (Pseudotsuga menziesii and Pinus ponderosa). Deer browsed much more intensively on native species than on exotic conifers, in terms of number of individuals attacked and degree of browsing. Deer preference for natives could potentially facilitate invasion by exotic pines. However, we hypothesize that the low rates of invasion currently observed can result at least partly from high densities of exotic deer, which, despite their preference for natives, can prevent establishment of both native and exotic trees. Other factors, not mutually exclusive, could produce the observed pattern. Our results underscore the difficulty of predicting how one introduced species will effect impact of another one.     

Darwin’s naturalization hypothesis up-close: Intermountain grassland invaders differ morphologically and phenologically from native community dominants

Darwin’s naturalization hypothesis predicts that successful invaders will tend to differ taxonomically from native species in recipient communities because less related species exhibit lower niche overlap and experience reduced biotic resistance. This hypothesis has garnered substantial support at coarse scales. However, at finer scales, the influence of traits and niche use on invasibility and invader impacts is poorly understood. Within grasslands of western Montana, USA, we compared morphological and phenological traits for five top exotic invasive forbs and five dominant native forbs using multivariate techniques to examine niche separation between exotics and natives. Exotic forbs differed from native forbs in multivariate space. Phenologically, native forbs synchronized vegetative growth with bolting and flowering early in spring. In contrast, exotics initiated vegetative growth concurrent with natives but bolted and flowered later. Morphologically, vegetative growth of exotics was three times shorter and narrower, but flowering stem growth was 35% taller and 65% wider than the natives. Collectively, these patterns suggest different strategies of resource uptake and allocation. Additionally, following wildfire, survival was four times higher for exotics compared to natives, and three times more of the surviving exotics flowered. The exotics we examined appeared to be exploiting an empty community-level niche. The resulting pattern of trait differences between exotics and natives suggests a predictable pattern of invasion and a predictable trajectory of community change. Our results illustrate how quantifying trait differences between invading exotics and natives at the within-community scale can improve understandings of community invasibility and invader impacts.     

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 biogeography of naturalization in alien plants

Aim  This paper reviews the main geographical determinants of naturalization in plants.
Location  Global.
Methods  Comparative studies of large data sets of alien floras are the main source of information on global patterns of naturalization.
Results  Temperate mainland regions are more invaded than tropical mainland regions but there seems to be no difference in invasibility of temperate and tropical islands. Islands are more invaded than the mainland. The number of naturalized species in temperate regions decreases with latitude and their geographical ranges increase with latitude. The number of naturalized species on islands increases with temperature. Naturalized species contribute to floristic homogenization, but the phenomenon is scale-dependent.
Main conclusions  Some robust patterns are evident from currently available data, but further research is needed on several aspects to advance our understanding of the biogeography of naturalization of alien plants. For example, measures of propagule pressure are needed to determine the invasibility of communities/ecosystems/regions. The patterns discussed in this paper are derived largely from numbers and proportions of naturalized species, and little is known about the proportion of introduced species that become naturalized. Further insights on naturalization rates, i.e. the proportion of aliens that successfully naturalize within regions, and on geographical and other determinants of its variation would provide us with better understanding of the invasion process. Comparative studies, and resulting generalizations, are almost exclusively based on numbers of species, but alien species differ in their impact on native biodiversity and ecosystem processes.