Evidence of biotic resistance to exotic plant invasion in degraded Bornean forests

Intact tropical forests are generally considered to be resistant to invasions by exotic species, although the shrub Clidemia hirta (Melastomataceae) is highly invasive in tropical forests outside its native range. Release from natural enemies (e.g., herbivores and pathogens) contributes to C. hirta invasion success where native melastomes are absent, and here we examine the role of enemies when C. hirta co-occurs with native Melastomataceae species and associated herbivores and pathogens. We study 21 forest sites within agricultural landscapes in Sabah, Malaysian Borneo, recording herbivory rates in C. hirta and related native Melastoma spp. plants along two 100-m transects per site that varied in canopy cover. Overall, we found evidence of enemy release; C. hirta had significantly lower herbivory (median occurrence of herbivory per plant = 79% of leaves per plant; median intensity of herbivory per leaf = 6% of leaf area) than native melastomes (93% and 20%, respectively). Herbivory on C. hirta increased when closer to native Melastoma plants with high herbivory damage, and in more shaded locations, and was associated with fewer reproductive organs on C. hirta. This suggests host-sharing by specialist Melastomataceae herbivores is occurring and may explain why invasion success of C. hirta is lower on Borneo than at locations without related native species present. Thus, natural enemy populations may provide a “biological control service” to suppress invasions of exotic species (i.e., biotic resistance). However, lower herbivory pressures in more open canopy locations may make highly degraded forests within these landscapes more susceptible to invasion.     

Passive restoration in Araucaria Forest: useful ecological indicators in monitoring successional advancement in exotic tree plantation landscapes

Monitoring successional advancement is a complex field involving a constant search for applied ecological indicators which facilitate monitoring of secondary forests for both active and passive restoration. In this study, the authors investigate the successional advancement of floristics and tree structure within Araucaria Forest (AF) fragments under passive restoration in a context where exotic tree plantations (mainly Pinus L. genus) dominate the landscape. The ecological indicators used were floristic dissimilarity (β‐diversity inferences), indicator species, ecological groups of species, basal area, and species abundance distribution (SAD) models (α‐diversity inferences). A total of 182 tree species belonging to 91 genera and 43 botanical families were identified. A high β diversity was verified for which each site has indicator species (for the locations CD—Dicksonia sellowiana; CO—Cryptocarya aschersoniana; and PG—Pinus taeda), where pioneer species contributed to much of the abundance. Different SAD models are useful for describing passive restoration sites in exotic tree plantation landscapes, namely Lognormal, Mandelbrot, and Zipf. SAD models together with basal area, taxonomic group (e.g. Myrtaceae assemblage), and tree abundance in ecological groups are strategic ecological indicators for monitoring successional advancement in AF.     

Restoration of Themeda australis swards suppresses soil nitrate and enhances ecological resistance to invasion by exotic annuals

Understanding processes that underlie ecological resistance to weed invasion is critical for sustainable restoration of invaded plant communities. Experimental studies have demonstrated that invasive nitrophilic annuals can be controlled by addition of carbon to reduce soil nitrate concentrations, sometimes leading to enhanced establishment of native plants. However, effects of carbon supplements on soil nitrate are temporary, and the longer-term value of carbon supplementation as a restoration tool is dependent on the resistance of the re-established ecosystem to repeat invasion. We investigated whether re-established swards of the tussock grass Themeda australis (R.Br.) Stapf (a natural understorey dominant in mesic grassy woodlands of SE Australia) could suppress soil nitrate concentrations, and through this or other means, could impart ongoing resistance to exotic invasion in restored woodlands. In a remnant invaded by exotic annuals, we applied three plot treatments (carbon supplements, annual spring burns and untreated control) and two seed treatments (± Themeda seed) in a replicated, factorial design. Within 3 years, successful establishment of Themeda swards on burnt and carbon-supplemented plots was associated with a reduction in soil nitrate to levels comparable with non-invaded, Themeda-dominated reference sites in the region (<3 mg/kg), and significantly reduced exotic cover compared with unseeded plots. By contrast, on plots not seeded with Themeda, soil nitrate increased after cessation of carbon addition and exotic cover returned to levels comparable with untreated control plots, despite a high cover of other native perennial grasses. Few persistent effects of carbon supplements or spring burning on soil nutrients were evident 9–19 months after cessation of these treatments. Results suggest that Themeda is a keystone species that regulates nitrate cycling, thereby imparting ecological resistance to invasion by nitrophilic annuals.     

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.     

Biogeographical contrasts to assess local and regional patterns of invasion: a case study with two reciprocally introduced exotic maple trees

Quantitative comparisons of distribution and abundance of exotic species in their native and non‐native ranges represent a first step when studying invaders. However, this approach is rarely applied 2 particularly to tree species. Using biogeographical contrasts coupled with regional dispersal surveys, we assessed whether two exotic maple tree species, Acer negundo and Acer platanoides, can be classified as invasive in the non‐native regions surveyed. We also examined the importance of biogeography in determining the degree of invasion by exotic species using this reciprocal approach. Local‐scale surveys were conducted in a total of 34 forests to compare density, relative abundance, age structure of native and introduced populations, and whether the two introduced maple species negatively affected native tree species density. Regional‐scale surveys of a total of 136 forests were then conducted to assess distribution in the introduced regions. Introduced populations of A. negundo were denser than populations measured in their native range and negatively related to native tree species density. Age structure did not differ between regions for this species. At the regional scale, this species has invaded most of the riparian corridors sampled in France. Conversely, the density of A. platanoides introduced populations was similar to that of native populations and was not related to native tree species density. Although seedling recruitment was higher away than at home, this species has invaded only 9% of the forests sampled in southern Ontario, Canada. Although reported invasive, these two exotic maple species differed in their relative demographic parameters and regional spread. Acer negundo is currently invasive in southern France while A. platanoides is not aggressively invasive in southern Ontario. Importantly, this study effectively demonstrates that biogeography through structured contrasts provide a direct means to infer invasion of exotic species.     

Prefire grazing by cattle increases postfire resistance to exotic annual grass (Bromus tectorum) invasion and dominance for decades

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Conspecific tolerance and heterospecific competition as mechanisms for overcoming resistance to invasion by an intertidal crab

The success of the invasive Asian shore crab, Hemigrapsus sanguineus, stems partly from its ability to exclude established crab species from preferred rocky and cobble intertidal habitat. Here, we assessed preference and competition for habitat types (cobble vs. sand) for H. sanguineus and two competitor species; the previous invasive green crab, Carcinus maenas, and the native rock crab, Cancer irroratus, in New England. In simple laboratory experiments, we paired similarly sized heterospecifics and conspecifics from each species, and also grouped combinations of C. maenas and H. sanguineus in a series of four-individual sets in order to dissect the outcome of intra- and inter-specific competition at different densities. Individually, all three species preferred cobble substrate. With paired conspecifics, H. sanguineus individuals would cohabitate in cobble, whereas C. maenas and C. irroratus individuals each excluded conspecifics from cobble. In heterospecific pairs, H. sanguineus excluded both C. maenas and C. irroratus from cobble. C. maenas and C. irroratus, were equally likely to exclude the other species, but rarely excluded H. sanguineus. In larger assemblages, H. sanguineus preferentially grouped under cobble, whereas C. maenas were more evenly distributed among habitat types. These observations demonstrate that conspecific tolerance and heterospecific competition can be effective, complementary mechanisms for overcoming invasion resistance. Such mechanisms help explain the well-studied success of H. sanguineus following its introduction into New England coastal habitats, and the resulting exclusion of preexisting crab species.     

Native tree regeneration in native tree plantations: understanding the contribution of Araucaria angustifolia to biodiversity conservation in the threatened Atlantic Forest in Argentina

Deforestation is a global process that has strongly affected the Atlantic Forest in South America, which has been recognised as a threatened biodiversity hotspot. An important proportion of deforested areas were converted to forest plantations. Araucaria angustifolia is a native tree to the Atlantic Forest, which has been largely exploited for wood production and is currently cultivated in commercial plantations. An important question is to what extent such native tree plantations can be managed to reduce biodiversity loss in a highly diverse and vulnerable forest region . We evaluated the effect of stand age, stand basal area, as a measure of stand density, and time since last logging on the density and richness of native tree regeneration in planted araucaria stands that were successively logged over 60 years, as well as the differences between successional groups in the response of plant density to stand variables. We also compared native tree species richness in planted araucaria stands to neighbouring native forest. Species richness was 71 in the planted stands (27 ha sampled) and 82 in native forest (18 ha sampled) which approximate the range of variation in species richness found in the native forests of the study area. The total abundance and species richness of native trees increased with stand age and time since last logging, but ecological groups differed in their response to such variables. Early secondary trees increased in abundance with stand age 3–8 times faster than climax or late secondary trees. Thus, the change in species composition is expected to continue for a long term. The difference in species richness between native forest and planted stands might be mainly explained by the difference in plant density. Therefore, species richness in plantations can contribute to local native tree diversity if practices that increase native tree density are implemented.     

Potential selection in native grass populations by exotic invasion

Ecological impacts of invasive plant species are well documented, but the genetic response of native species to invasive dominance has been often overlooked. Invasive plants can drastically alter site conditions where they reach dominance, potentially exerting novel selective pressures on persistent native plant populations. Do native plant populations in old exotic invasions show evidence of selection when compared to conspecific populations in adjacent, noninvaded areas? We employ amplified fragment length polymorphism (AFLP) analysis to screen a large number of loci from two native grass species (Hesperostipa comata (Trin. & Rupr.) Barkworth and Sporobolus airoides Torr.) that occur in old infestations of the invasive forb Acroptilon repens. We then compare observed locus by locus F_ST values with distributions of F_ST estimated from simulation models under expectation of neutrality. We also compare the proportion of loci possibly linked to selection and those not linked to selection which exhibit parallel trends in divergence between two community types (invaded, noninvaded). Few loci (H. comata, 2.6%; S. airoides, 8.7%) in the two native grasses may be linked to genes under the influence of selection. Also, loci linked to selection showed a greater portion of parallel trends in divergence than neutral loci. Genetic similarities between community types were less than genetic similarity within community types suggesting differentiation in response to community alteration. These results indicate that a small portion of scored AFLP loci may be linked to genes undergoing selection tied to community dominance by an invasive species. We propose that native plants in communities dominated by exotic invasives may be undergoing natural selection.     

Dwarf bamboos affect the regeneration of zoochorous trees by providing habitats to acorn-feeding rodents

The effects of dwarf bamboos (Sasa spp.) on the regeneration of trees in a natural hardwood forest were studied by analysing the spatial dispersion of seedlings and saplings of anemochores (Acer palmatum var. matsumurae, Fraxinus lanuginosa, and Carpinus laxiflora) and zoochores (Quercus mongolica var. grosseserrata and Q. serrata). Relative photosynthetic photon flux density at 10 cm above ground was significantly correlated with the coverage of dwarf bamboos (r=0.661, P<0.001). Seedlings were abundant and were randomly distributed in the anemochores, other than the shade-intolerant species C. laxiflora which was significantly more sparse in sites with dense Sasa than in sites where Sasa was rare. Distribution of saplings was also random in the shadetolerant anemochores A. palmatum var. matsumurae and F. lanuginosa but aggregated in sites with sparse Sasa in the shade-intolerant anemochore C. laxiflora. In contrast to the anemochores, seedlings of zoochores were very few and were distributed in sites with sparse Sasa. Saplings were also aggregated and negatively correlated with Sasa cover in the shade-intolerant species Q. serrata and the tolerant species Q. mongolica var. grosseserrata. The acorns put on the forest floor in a site with dense Sasa were quickly removed by small rodents such as Apodemus speciosus and A. argenteus. Trap census of rodents revealed that those mammals prefer the dense Sasa habitat to the sparse Sasa habitat. This suggests that the dwarf bamboos strongly affect the regeneration of zoochorous trees not only by shading the seedlings but also by providing habitats to acorn-feeding small mammals.     

Vulnerability of Mediterranean Basin ecosystems to climate change and invasion by exotic plant species

Aim To assess at a broad scale the vulnerability of Mediterranean vegetation to alien plant invasion under different climatic and disturbance scenarios. Location We simulated the vegetation biogeography and dynamics on five of the main islands of the Mediterranean Basin: Mallorca, Corsica, Sardinia, Crete and Lesvos. Methods We used LPJ‐GUESS, a generalized ecosystem model based on dynamic processes describing establishment, competition, mortality and ecosystem biogeochemistry. We simulated the vegetation distribution and dynamics using a set of plant functional types (PFTs) based on bioclimatic and physiological parameters, which included tree and shrub PFTs defined especially for the Mediterranean. Additionally, two invasive PFTs, an invasive tree type and an invasive herb type, were defined and used to estimate the vulnerability to invasion of a range of different ecosystems. The model was used to simulate climate changes and associated changes in atmospheric [CO₂] to 2050 according to two SpecialReport on Emissions Scenarios climate scenarios (A1Fi and B1) combined with mean disturbance intervals of 3 and 40 years. Results The simulations and scenarios showed that the effect of climate change alone is likely to be negligible in many of the simulated ecosystems, although not all. The simulated progression of an invasion was highly dependent on the initial ecosystem composition and local environmental conditions, with a particular contrast between drier and wetter parts of the Mediterranean, and between mountain and coastal areas. The rate of ecosystem disturbance was the main factor controlling susceptibility to invasion, strongly influencing vegetation development on the shorter time scale. Main conclusions Further invasion into Mediterranean island ecosystems is likely to be an increasing problem: our simulations predict that, in the longer term, almost all the ecosystems will be dominated by exotic plants irrespective of disturbance rates.     

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.     

Biogeographical variation in community response to root allelochemistry: novel weapons and exotic invasion

Centaurea diffusa is one of the most destructive invasive weeds in the western USA and allelopathy appears to contribute to its invasiveness ( Callaway & Aschehoug 2000 ). Here we identify a chemical from the root exudates of C. diffusa, 8‐hydroxyquinoline, not previously reported as a natural product, and find that it varies biogeographically in its natural concentration and its effect as an allelochemical. 8‐Hydroxyquinoline is at least three times more concentrated in C. diffusa‐invaded North American soils than in this weed's native Eurasian soils and has stronger phytotoxic effects on grass species from North America than on grass species from Eurasia. Furthermore, experimental communities built from North American plant species are far more susceptible to invasion by C. diffusa than communities built from Eurasian species, regardless of the biogeographical origin of the soil biota. Sterilization of North American soils suppressed C. diffusa more than sterilization of Eurasian soils, indicating that North American soil biota may also promote invasion by C. diffusa. Eurasian plants and soil microbes may have evolved natural resistance to 8‐hydroxyquinoline while North American plants have not, suggesting a remarkable potential for evolutionary compatibility and homeostasis among plants within natural communities and a mechanism by which exotic weeds destroy these communities.     

Introduction history and invasion success in exotic vines introduced to Australia

The ecological damage caused by invasive vines poses a considerable threat to many natural ecosystems. However, very little data are available for this potentially environmentally destructive functional group in Australia. In order to address this paucity of information, we assembled the first inventory of exotic vines that have become established in natural ecosystems across Australia. The influence that introduction history attributes, variables that relate to the introduction of a species to a new area, may have on the occurrence and distribution of exotic vines was also determined. We asked whether the continent of origin, reason for introduction, and residence time related to the prevalence and distribution of exotic vines across Australia. A total of 179 exotic climbing plant species from 40 different families were found to have become established across continental Australia. However, five families accounted for over 50% of these species. Most exotic vines originated from South America, and were introduced for ornamental purposes. The length of time in which an exotic vine had been present in its new range was significantly related to its distribution, with a positive relationship found between residence time and area of occupancy across the continent. No other introduction history attribute was significantly related to the area of occupancy, or distribution, of a species. This suggests that while the trends found among introduction history attributes are important in explaining the prevalence of exotic vines in Australia, only residence time is currently a useful predictor of their future success.     

Phytophagous insect fauna tracks host plant responses to exotic grass invasion

The high dependence of herbivorous insects on their host plants implies that plant invaders can affect these insects directly, by not providing a suitable habitat, or indirectly, by altering host plant availability. In this study, we sampled Asteraceae flower heads in cerrado remnants with varying levels of exotic grass invasion to evaluate whether invasive grasses have a direct effect on herbivore richness independent of the current disturbance level and host plant richness. By classifying herbivores according to the degree of host plant specialization, we also investigated whether invasive grasses reduce the uniqueness of the herbivorous assemblages. Herbivorous insect richness showed a unimodal relationship with invasive grass cover that was significantly explained only by way of the variation in host plant richness. The same result was found for polyphagous and oligophagous insects, but monophages showed a significant negative response to the intensity of the grass invasion that was independent of host plant richness. Our findings lend support to the hypothesis that the aggregate effect of invasive plants on herbivores tends to mirror the effects of invasive plants on host plants. In addition, exotic plants affect specialist insects differently from generalist insects; thus exotic plants affect not only the size but also the structural profile of herbivorous insect assemblages.     

Applying competition theory to invasion: resource impacts indicate invasion mechanisms in California shrublands

Despite widespread work documenting invasion, it remains a challenge to determine invasion mechanisms and incorporate them into invasive species management. Competition theory presents a strong model for evaluating the role of resource reduction and requirements in invasion. Additionally, alternative models suggest fluctuations in resources, niche differences, or non-resource priority effects are key factors determining invasion success. We propose a comparative framework that incorporates resource impacts of native and invasive species, performance in controlled invasion trials, and long-term natural invasion patterns to elucidate relative importance of these invasion mechanisms. To demonstrate this framework, we established monocultures of two representative native and two invasive plant species in Southern California’s coastal sage scrub (CSS), measured resource impacts (i.e., R*), and conducted invasion trials to test whether resource impacts predicted invasion success. We then related experimental results to field invasion patterns. Compared to exotic herbaceous species, native shrubs were associated with greater resource depletion of key resources: light, soil water (at multiple depths), and soil inorganic nitrogen (particularly at depth). In invasion trials, natives resisted invasion by the exotics, as resource depletion measures would predict. However, these results did not follow long-term natural invasion patterns indicating that these exotic species invade areas once dominated by native shrubland. Applying our results to the invasion framework, we conclude that disturbance, or a similar mechanism causing resources to fluctuate, is needed for exotics to invade CSS habitats. This resource-based comparative analysis of invasion mechanisms can point out important processes and help suggest effective management actions.