Livestock reduces juvenile tree growth of alien invasive species with a minimal effect on natives: a field experiment using exclosures

Many alien invasive tree species were originally introduced to their non-native ranges for use in forestry and as urban trees. These alien species were selected for their fast growth and not necessarily for possessing mechanisms which deter browsing. Instead, many tree species native to semiarid areas of the world evolved mechanisms which deter browsing, presumably at the cost of slower growth. In a semiarid rangeland we observed that livestock exclusion greatly promoted the growth of juveniles of several alien species but not of native species, and we hypothesized that this increase in growth of aliens was due to livestock preference for alien and not native trees. With the objective of quantifying our observations and understanding the mechanism underlying the increased growth rates of alien juvenile trees under livestock exclusion, we assessed growth and browsing levels in juveniles of two alien invasive and four abundant native tree species within three parcels where livestock was excluded and three parcels with livestock at 0.20 cattle equivalents.ha^?1. Alien species grew around four-fold faster under livestock exclusion than with livestock and, as predicted, received five times more browsing than natives. Instead, native species did not significantly increase their growth rate with livestock exclusion. The results support our hypothesis and the implications for management would be that stocking paddocks with livestock to browse existing alien juveniles and re-growth of felled adults should be effective in delaying invasions of trees used for forestry without significantly affecting the growth of the most abundant native trees.     

外来树种的耗水特征及其对环境水资源的影响

外来树种常被作为"先锋树种"广泛引种,在荒山造林和水土保持方面发挥了积极作用;然而,不少外来树种具有高耗水和强抗旱的水分利用特性,耗水量大、水分消耗快,若不加甄别地引种会影响甚至危害本土森林生态系统的健康。目前,部分被引种的外来树种已造成入侵;同时,在当前水资源缺乏的形势下,外来树种的引种安全问题已成为全球关注的生态环境问题。本文综述了外来树种在生态系统、整树、叶片及枝条水平的耗水特性,分析了外来树种对引种地水资源的影响。前期的研究结果显示,被引种的外来树种在生态系统及整树水平的耗水量普遍高于乡土树种,但对水资源的影响却因受控于多个因素如环境、气候、植物个体等而不尽相同;同时还显示,外来树种在叶片和枝条水平的耗水特性更多地体现了其应对环境变化的水分利用策略。研究还就提高外来树种耗水特性研究的准确性和提升外来树种危害风险的预警能力方面,提出了若干研究方向,旨在为森林管理者和政府部门平衡外来树种的经济效益和生态效应提供借鉴和指导,以期降低外来树种引种风险,增进生态安全。     

Soil type, microsite, and herbivory influence growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna

Naturalization of Schinus molle (Anacardiaceae) has been observed in semi arid savanna of the Northern Cape Province of South Africa. However, with high dispersal ability, the species is expected to achieve greater densities and invade more widely. The study involved a field manipulation experiment over 14 months using a factorial block design to examine transplanted seedlings in different savanna environments. The experiments examine the effects of soil type (sandy and clay), microsite, and herbivores on seedling performance (establishment, growth and survival). Seedlings were grown in a greenhouse and individually transplanted into four treatment groups: in open grassland, under tree canopies, and with and without cages to exclude large herbivores (cattle and game). The same experiment was repeated in two different soil types: coarse sand and fine-textured clay soil. Results suggest that protection provided by canopies of large indigenous Acacia trees facilitates S. molle invasion into semi-arid savanna. In the field, S. molle seedlings performed considerably better beneath canopies of indigenous Acacia trees than in open areas regardless of soil type. Whether exposed or protected from large herbivores, no seedlings planted in open grassland survived the first winter. Although, seedlings grew better and had higher survival rates beneath tree canopies than in the open sites, exposure to large herbivores significantly decreased heights and canopy areas of seedlings compared with those protected from large herbivores. The effect was greater on clay soil than on sandy soil. The results suggest that low temperature (frost), and possibly inter-specific competition with grasses, may limit S. molle seedling establishment, survival and growth away from tree canopies in semi arid savannas. Low soil nutrient status and browsing may also delay growth and development of this species. The invasive potential of S. molle is thus greatest on fertile soils where sub-canopy microsites are present and browsing mammals are absent.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

More widespread alien tree species do not have larger impacts on regeneration of native tree species in a tropical forest reserve

There is insufficient information regarding the factors affecting the environmental impacts of alien species. In particular, little is known about whether there is any relationship between the invasiveness (establishment and spread) of an introduced species and its per capita impact. We experimentally assessed the relationship between the extent of spread of up to 29 alien plant species and their impact on recruitment of native tree species in Amani Botanical Garden, Tanzania. We also studied the effects of allelochemicals of selected alien on native plant species to assess potential mechanisms of impact. We found no relationship between the extent of spread of an alien tree species and their impact on seed germination, seedling survival, and seedling communities of native trees in their understory, and no indication that allelochemicals consistently explain their effects on recruitment of the studied species. These results suggest that extent of spread cannot be used as a proxy for impact. Hence, managers should continue assessing both the spread and the impact of alien species when prioritizing alien species for management.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

Strategic light manipulation as a restoration strategy to reduce alien grasses and encourage native regeneration in Hawaiian mesic forests

Question: Is there a light level at which alien grass biomass is reduced while still supporting growth and survival of native woody species, allowing for native species regeneration in abandoned pastures? Location: Island of Hawaii, USA. Methods: In a two‐part study we examined the effect of light availability on common native woody and alien grass species found in secondary forests in Hawaii. A field survey was conducted to examine the relationship between light availability and canopy type (open pasture, planted canopy and secondary forest) on understory grass biomass and litter accumulation. We then experimentally manipulated light levels to determine the effect of light availability on growth and survival of six native woody species and three alien grasses. Low‐light (5%), medium‐light (10%) and high‐light (20‐30%) treatments were created using shade structures erected beneath the existing secondary koa canopy. Results: In the field survey, alien grass biomass was greatest under the open pasture and lowest in the secondary forest. There was a positive correlation between understory light availability and alien grass biomass. In the experimental study, large reductions in relative growth rates were documented for all of the grass species and four of the six woody species under the lowest light level. Although growth at 5% light is substantially reduced, survival is still high (84‐100%), indicating that these species may persist under closed canopy. Conclusion: Low‐light conditions result in the greatest reduction in alien grass biomass while creating an environment in which native woody species can grow and survive.     

Temporal shifts in interactions between alien trees and the alien Argentine ant on native ants

Synergistic effects between various stressors on native biodiversity are poorly understood, especially adverse synergisms between different invasive organisms. While it is known that alien trees and the invasive Argentine ant Linepithema humile individually impact indigenous ant assemblages, little is known about how the impacts of these two types of aliens have a joint effect, and even less about the temporal dynamics of this interaction. We found that the Argentine ant benefited from invasion by alien trees through creation of less extreme environmental conditions at the hottest and coolest times of the year. We also found that the impacts of the two types of aliens were synergistic on the native ant assemblage. However, this synergistic impact also varied in intensity throughout the year, especially so in the more open natural sites. The alien tree canopy, when dense, created more constant environmental conditions throughout the year, leading to more constant Argentine ant levels and consequently more even impacts on the native ants. The various native ants varied considerably in their response to the impacts of both types of aliens, with many formicine ants being particularly sensitive. Our results show that the synergistic impacts, both negative and positive, of the two alien types are particular to each native ant species. Furthermore, the intensity of the adverse synergism is highly variable across the year.     

Post-fire spread of alien plant species in a mixed broad-leaved forest of the Insubric region

How do tree species regenerate and which ecological conditions are required after forest fire in the Insubric region of the Alps? Are indigenous stand-forming tree species resistant over the invasion of alien plant species after such a disturbance? We addressed these questions in a case study in the Swiss canton of Ticino. In April 2006, a surface fire with severe intensity burnt a forest area of 55 ha on a south-facing slope (400-800 m.a.s.l.). The dominant trees in the investigated area were chestnut (Castanea sativa Mill.), beech (Fagus sylvatica L.) and deciduous oaks (Quercus spp.) mixed with tree species of intermediate height. Vegetation data were collected in May and August 2009 by systematic sampling. Resprouting of the survived trees and generative regeneration were analysed by counting vegetative shoots from sprouting stools, of seedling age, height and damage rate, respectively. Different vegetation structures related to low or high fire intensity were clearly visible three years after the forest fire, creating various habitats for both new invaders and seedlings of the stand-forming trees. The dominant chestnut was the only tree species that regenerated effectively by sprouting from stools. Seedlings of the stand-forming trees grew in high abundance under shadow conditions close to their mother trees which provided the seed source. In contrast, pioneer trees invaded patches where full light was available. Under such conditions the two main woody alien plant species, Ailanthus altissima and Robinia pseudoacacia, grew in high abundance. Due to the different ecological requirements of indigenous and alien tree seedlings, not any interaction between the two groups was detected.     

A plant‐traits approach to assessing the success of alien weed species in irrigated Mediterranean orchards

In addition to their impact on natural habitats, invasive alien plants can have a significant negative effect on agricultural systems and cause economic losses. Flood‐irrigated orchards in the Mediterranean Basin are vulnerable to the invasion of alien weeds, primarily because of the traditional management practices used in the orchards, which are characterized by high soil moisture during the dry summer period, nutrient availability and high levels of disturbance. This study sought to determine whether their biological traits can explain the success of alien weed species. To answer this question, 408 floristic relevés were conducted in 136 flood‐irrigated orchards on the Plains of Lleida (Catalonia, NE of Spain). Richness and cover of native and alien weeds were compared. Furthermore, a set of biological traits were compared between successful and non‐successful weeds for the whole data and separately between native and alien weeds using logistic regression and classification trees. In flood‐irrigated orchards, alien species covered most of their area, even though the richness of alien species was lower than that of the native species. The most important species were C4 species with seeds dispersed by water, and on the other hand, rosulate and caespitose‐reptant hemicryptophytes with long flowering period. Most of these traits fitted with those of the invasive alien weeds, which were mostly C4 species with seeds dispersed by water. Perennial life form characterized successful native weeds. In this study, we discuss how the traditional management of flood irrigation in fruit‐tree orchards favours invasive alien weeds that have specific traits, acting as a reservoir for the spread of alien weeds into other crops and surrounding riparian habitats. We also propose changing management practices in order to avoid the selection of alien weeds and to promote native species.     

外来种隐蔽入侵： 类型及影响

隐蔽入侵（cryptic invasion）是指在形态上与土著种（或早期建群种）不能或难以区分的外来种在人们未觉察的状态下成功入侵的过程。人们对这类入侵方式往往视而不觉。本文综述了外来种隐蔽入侵的类型以及生态影响。隐蔽入侵的类型主要包括外来姊妹种形式的隐蔽入侵、 不同遗传支系的隐蔽入侵、 不同地理种群的隐蔽入侵以及“返传入”。其中, “返传入”目前还是一种假说。由于这类入侵外来种更容易与土著种（或早期建群种）杂交或基因渗透, 因而可对入侵种自身或土著种产生深远的生态影响。鉴于隐蔽入侵现象的广泛性, 建议进一步加强该方面的研究。     

Effects of invasive alien kahili ginger (Hedychium gardnerianum) on native plant species regeneration in a Hawaiian rainforest

Questions: Does the invasive alien Hedychium gardnerianum (1) replace native understory species, (2) suppress natural regeneration of native plant species, (3) increase the invasiveness of other non‐native plants and (4) are native forests are able to recover after removal of H. gardnerianum. Location: A mature rainforest in Hawai'i Volcanoes National Park on the island of Hawai'i (about 1200 m a.s.l.; precipitation approximately 2770 mm yr^?1). Study sites included natural plots without effects of alien plants, ginger plots with a H. gardnerianum‐dominated herb layer and cleared plots treated with herbicide to remove alien plants. Methods: Counting mature trees, saplings and seedlings of native and alien plant species. Using non‐parametric H‐tests to compare impact of H. gardnerianum on the structure of different sites. Results: Results confirmed the hypothesis that H. gardnerianum has negative effects on natural forest dynamics. Lower numbers of native tree seedlings and saplings were found on ginger‐dominated plots. Furthermore, H. gardnerianum did not show negative effects on the invasive alien tree species Psidium cattleianum. Conclusions: This study reveals that where dominance of H. gardnerianum persists, regeneration of the forest by native species will be inhibited. Furthermore, these areas might experience invasion by P. cattleianum, resulting in displacement of native canopy species in the future, leading to a change in forest structure and loss of other species dependent on natural rainforest, such as endemic birds. However, if H. gardnerianum is removed the native Hawaiian forest is likely to regenerate and regain its natural structure.     

Testing a facilitation model for ecosystem restoration: Does tree planting restore ground layer species in a grassy woodland?

Planning for the restoration of degraded ecosystems has a strong basis in facilitation successional theory, which, as applied in restoration practice, states that planting of structurally dominant tree species will assist the entry of other native species into a restored community. In Australia, tree planting has been widely applied in restoration of grassy woodland ecosystems. Trees have been postulated to reduce the cover and diversity of weed species, thus facilitating recolonization of native woodland species (indirect facilitation). The expected outcomes of this process include reduced species richness and abundance of exotic plant species and increased species richness and abundance/dominance of natives in areas beneath tree canopies, with these trends strengthening with time. To assess whether this was occurring, we carried out a comparative analysis of species assemblages found underneath and outside of planted tree canopies in sites replanted with juvenile canopy tree species 3–5 or 8–10 years previously. We sampled revegetated stands of Cumberland Plain Woodland, an endangered ecological community in Western Sydney, Australia. We found that neither the number nor abundance of native ground layer species beneath canopies increased as a result of trees being planted at sites of both ages. Where seed is limited, we predicted an increase in abundance of existing native species under planted tree canopies. On this point, the results were mixed and showed some natives with an increased abundance while others decreased. Exotic species richness showed the reverse of the expected pattern, being greater under tree canopies. These findings lend no support to the theory of indirect facilitation. We conclude that simple facilitation models may be inadequate to support planning of grassy woodland restoration and that those models incorporating successional time lags and restoration barriers are likely to be more informative about the development of communities initiated by tree planting.     

Biological invasion into the nested assemblage of tree–beetle associations on the oceanic Ogasawara Islands

Invasion by alien organisms is a common worldwide phenomenon, and many alien species invade native communities. Invasion by alien species is especially likely to occur on oceanic islands. To determine how alien species become integrated into island plant–insect associations, we analyzed the structure of tree–beetle associations using host plant records for larval feeding by wood-feeding beetles (Coleoptera: Cerambycidae) on the oceanic Ogasawara Islands in the northwestern Pacific Ocean. The host plant records comprised 109 associations among 28 tree (including 8 alien) and 26 cerambycid (including 5 alien) species. Of these associations, 41.3% involved at least one alien species. Most native cerambycid species feed on host trees that have recently died. Alien trees were used by as many native cerambycid species (but by significantly more alien cerambycid species) as were native trees. Native cerambycid species used as many alien tree species (but significantly more native tree species) as did alien cerambycids. Thus, we observed many types of interactions among native and alien species. A network analysis revealed a significant nested structure in tree–cerambycid associations regardless of whether alien species were excluded from the analysis. The original nested associations on the Ogasawara Islands may thus have accepted alien species.     

Observed and dark diversity of alien plant species in Europe: estimating future invasion risk

Invasion by alien species is a growing concern for nature conservation. We estimated the level of invasion by alien plant species and future invasion risks at the European scale. We used a pan-European atlas and eight regional plant atlases to determine the distribution of alien and native plant richness. In addition, we estimated alien and native dark diversity (species currently absent from a site but present in the surrounding region and able to colonize the site). We used relative diversity metrics to indicate current and future risks by alien species: relative alien richness (compared to native species), alien and native completeness (log-ratio of observed to dark diversity) and completeness difference between alien and native species. Observed and relative richness of alien species were greatest in NW Europe; this suggests that sites in NW Europe could be more disturbed. Observed alien and native species richness show clear regional hotspots; the distribution of completeness values is dispersed, indicating local hotspots. Northern Europe has relatively lower alien completeness, likely because potential invaders inhabit the region but have not yet reached many localities, thereby suggesting a risk of future invasion. A greater number of potential alien species in the region increases the probability that some alien species could have detrimental impacts. Both alien richness and completeness are positively correlated with native richness and completeness, respectively, indicating that both groups share similar distribution patterns. Alien species diversity metrics in Europe are related positively to human population density and agricultural land-use. We suggest that the dark diversity concept can broaden our understanding of alien species diversity and future invasion risks.     

Differential processes underlying the roadside distributions of native and alien plant assemblages

Although biological invasion often alters ecosystem properties and community composition at different scales, considerable uncertainty still exists regarding the underlying mechanisms that regulate the spread of alien species into new habitats. An alien invasion is generally achieved through multiple processes from multiple sources; this type of invasion often prevents us from understanding of the dispersal mechanisms. Here, we aim to disentangle the processes of alien invasion by focusing on a single migration source. We surveyed the distribution of alien and native plant species in Shiretoko National Park, located in northern Japan. We measured the coverage of each species and the environmental and spatial factors in 362 quadrats established along roadsides. We found 101 native species and 35 alien species (γ-diversity) throughout the quadrats. The local species richness (α-diversity) was higher for the alien species (6.1 species) than for the native species (3.2 species). There was a significant negative correlation in α-diversity between native and alien species. Moreover, the α-diversity and distance from the nearest town (migration source) showed a negative relationship for alien assemblages while the native assemblages showed the opposite trend. These results suggest that the alien species are expanding their distribution outward from the town, resulting in a decrease in the α-diversity of native species in localities close to the migration source. Overall, our results emphasize that roadsides could unintentionally act as corridors for alien species, even in protected areas. Careful consideration is thus needed for utilizing these human-created habitats even though they were designed for conservation and management purposes.     

Invasion,alien control and restoration: Legacy effects linked to folivorous insects and phylopathogenic fungi

Invasive alien trees increase native tree stress and may increase attack by herbivores and pathogenic fungi. Alien tree removal should ameliorate such impacts. Here we compared the levels of damage by phylopathogenic fungi and folivorous insects on Brabejum stellatifolium and Metrosideros angustifolia (native trees) and Acacia mearnsii (invasive tree species) among near‐pristine, invaded and restored sites. Generally, foliar damage levels were higher at invaded than at near‐pristine sites. Damage levels at restored sites were similar, or even higher than those at invaded sites. Decreased native tree species richness did not explain these patterns, as restored sites had native tree species richness levels similar to those of near‐pristine sites. Increased host abundance and leaf nitrogen content did not significantly correlate with increased damage in most cases. Therefore, plant species richness recovers following restoration, but native trees still experience increased pressure from folivores and phylopathogenic fungi, which may even exceed levels experienced at invaded sites, thus impacting recovery trajectories.     

Do vascular plants and bryophytes respond differently to coniferous invasion of coastal heathlands?

Invasion by non-native conifers may pose a threat to local biodiversity, but knowledge about introduced conifer effects on Northern Hemisphere ecosystems is scarce. The coastal heathlands of north-west Europe are threatened by invasion of native and introduced tree species. We assess how spread of the introduced conifer Sitka spruce (Picea sitchensis (Bong.) Carr.) into European coastal heathlands affect two major functional groups; vascular plants and bryophytes, and how these effects relate to the environmental changes imposed by the developing tree canopies. We compared the impact of introduced Sitka spruce and native Scots pine (Pinus sylvestris L.) by analysing effects on species richness and turnover of vascular plants and bryophytes along fine-scale transects from individual tree stems into open heathland vegetation. Environmental impacts were assessed by measured environmental variables, and the responses of the two species groups were assessed by calculating changes in their respective mean Ellenberg indicator values. Species richness decreased beneath both conifers, related to decreased light and increased nitrogen and pH. Whereas vascular plants responded negatively to poor light conditions beneath dense and low Sitka spruce canopies, bryophytes were more negatively affected by the warmer and drier microclimates beneath Scots pine. Introduced Sitka spruce impacts the sub-canopy environment differently from the native Scots pine, and the two functional plant groups responded differently to these impacts. This suggests that future forests are likely to differ in species richness and composition, depending on whether succession is based on native or introduced coniferous trees.     

Frequent burning promotes invasions of alien plants into a mesic African savanna

Fire is both inevitable and necessary for maintaining the structure and functioning of mesic savannas. Without disturbances such as fire and herbivory, tree cover can increase at the expense of grass cover and over time dominate mesic savannas. Consequently, repeated burning is widely used to suppress tree recruitment and control bush encroachment. However, the effect of regular burning on invasion by alien plant species is little understood. Here, vegetation data from a long-term fire experiment, which began in 1953 in a mesic Zimbabwean savanna, were used to test whether the frequency of burning promoted alien plant invasion. The fire treatments consisted of late season fires, lit at 1-, 2-, 3-, and 4-year intervals, and these regularly burnt plots were compared with unburnt plots. Results show that over half a century of frequent burning promoted the invasion by alien plants relative to areas where fire was excluded. More alien plant species became established in plots that had a higher frequency of burning. The proportion of alien species in the species assemblage was highest in the annually burnt plots followed by plots burnt biennially. Alien plant invasion was lowest in plots protected from fire but did not differ significantly between plots burnt triennially and quadrennially. Further, the abundance of five alien forbs increased significantly as the interval (in years) between fires became shorter. On average, the density of these alien forbs in annually burnt plots was at least ten times as high as the density of unburnt plots. Plant diversity was also altered by long-term burning. Total plant species richness was significantly lower in the unburnt plots compared to regularly burnt plots. These findings suggest that frequent burning of mesic savannas enhances invasion by alien plants, with short intervals between fires favouring alien forbs. Therefore, reducing the frequency of burning may be a key to minimising the risk of alien plant spread into mesic savannas, which is important because invasive plants pose a threat to native biodiversity and may alter savanna functioning.