Commercially Important Trees as Invasive Aliens – Towards Spatially Explicit Risk Assessment at a National Scale

Alien species that are desirable and commercially important in parts of the landscape, but damaging invaders in other parts, present a special challenge for managers, planners, and policy-makers. Objective methods are needed for identifying areas where control measures should be focussed. We analysed the distribution of forestry plantations and invasive (self-sown) stands of Acacia mearnsii and Pinus spp. in South Africa; these two taxa account for 60% of the area under commercial plantations and 54% of the area invaded by alien trees and shrubs. The distribution of commercial forestry plantations and invasive stands of these taxa were mapped and the data was digitised and stored as Geographic Information System (GIS) (Arc/Info) layers. A series of environmental parameters were derived from GIS layers of climate, topography, geology, land use, and natural vegetation. The current distribution of the two taxa was subdivided into three groups according to the degree of invasion, planting history and the precision of the data collection. We used regression-tree analysis to relate, for each taxon, the distribution of invasive stands with environmental variables, and to derive habitat suitability maps for future invasion. The current distribution of invasive stands in South Africa was largely influenced by climatic factors. At a national scale, the distribution of large commercial plantations was a poor predictor of areas invaded by both taxa. Using environmental factors identified by the regression trees, we found that 6.6% and 9.8% of natural habitats currently not invaded and untransformed by urbanisation or agriculture are suitable for invasion by Pinus spp. and A. mearnsii, respectively. We then derived guidelines for policy on alien plant management based on vegetation type, degree of transformation, extent of invasion, and the risk of future alien spread. These factors were used to identify demarcated areas where these alien species can be grown with little risk of invasions, and areas where special measures are needed to manage spread from plantations.     

The ecology of Lagarostrobos franklinii (Hook.f.) Quinn (Podocarpaceae) in Tasmania. 2. Population structure and spatial pattern

Analysis of stand structure and spatial patterning indicates that Lagarostrobos franklinii (Huon pine) regenerates intermittently in response to canopy disturbance. Its ability to resprout rapidly from fallen trees is likely to be important in the continued dominance of gaps by this species in competition with faster-growing tree species. The largely riverine distribution pattern of this species is best explained by rapid vegetative spread down river systems and slow dispersal into canopy gaps away from river edges. The probability of recurrent disturbance within the lifespan of this species is high, implying that competitive exclusion of Huon pine is unlikely once it establishes at a site.     

Long‐distance dispersal of non‐native pine bark beetles from host resources

1. Dispersal and host detection are behaviours promoting the spread of invading populations in a landscape matrix. In fragmented landscapes, the spatial arrangement of habitat structure affects the dispersal success of organisms. 2. The aim of the present study was to determine the long distance dispersal capabilities of two non‐native pine bark beetles (Hylurgus ligniperda and Hylastes ater) in a modified and fragmented landscape with non‐native pine trees. The role of pine density in relation to the abundance of dispersing beetles was also investigated. 3. This study took place in the Southern Alps, New Zealand. A network of insect panel traps was installed in remote valleys at known distances from pine resources (plantations or windbreaks). Beetle abundance was compared with spatially weighted estimates of nearby pine plantations and pine windbreaks. 4. Both beetles were found ≥25 km from the nearest host patch, indicating strong dispersal and host detection capabilities. Small pine patches appear to serve as stepping stones, promoting spread through the landscape. Hylurgus ligniperda (F.) abundance had a strong inverse association with pine plantations and windbreaks, whereas H. ater abundance was not correlated with distance to pine plantations but positively correlated with distance to pine windbreaks, probably reflecting differences in biology and niche preferences. Host availability and dispersed beetle abundance are the proposed limiting factors impeding the spread of these beetles. 5. These mechanistic insights into the spread and persistence of H. ater and H. ligniperda in a fragmented landscape provide ecologists and land managers with a better understanding of factors leading to successful invasion events, particularly in relation to the importance of long‐distance dispersal ability and the distribution and size of host patches.     

How robust is the Australian Weed Risk Assessment protocol? A test using pine invasions in the Northern and Southern hemispheres

The Australian Weed Risk Assessment protocol (WRA) is often considered the standard approach for pre-border screening of new plant introductions. Here we assess its robustness against three key criteria: ability to discriminate success or failure of species at three stages of the invasion process (introduction, naturalisation and spread); sensitivity to taxonomic range and target region; and dependence on knowledge of invasive behaviour elsewhere. We address these issues by retrospectively testing the WRA using pine (Pinus) introductions to New Zealand and Great Britain. For both regions we calculated WRA scores for 115 species, and classified all species according to whether they had been introduced, which of these had naturalised, and the extent of their naturalised distribution (spread). Using regression models, we assessed whether WRA scores could predict success at each stage. We repeated this procedure using WRA scores calculated without information on species naturalisation behaviour elsewhere. In both regions, the WRA could discriminate among species in the same genus at the introduction and naturalisation stages, but not at the spread stage. The outcome at the naturalisation stage depended on prior knowledge of naturalisation behaviour elsewhere. Without this information the WRA may be unable to distinguish among closely related species, and should be used cautiously where data on invasive behaviour elsewhere is lacking. Human selection played a strong role in the invasion process both through introducing pine species likely to naturalise in New Zealand and Great Britain in the first instance, and subsequent use of many of these species for forestry in the target regions.     

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

Summary The spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.     

Gringos En El Bosque: Introduced Tree Invasion in a Native Nothofagus/Austrocedrus Forest

We studied invasion into native Nothofagus/Austrocedrus forest by many introduced tree species planted between 1910 and 1940 in plantations near the center of Isla Victoria, in northern Patagonia. We located virtually all individuals of these species in 30 ha of forest in two series of transects at increasing distances from the plantations. Although these species included many reported as highly invasive elsewhere, we found little evidence for invasion on Isla Victoria, with many invasive species utterly failing to invade native forest. There was a notable decline with distance in number of introduced individuals, but wind direction appeared to be unimportant. Pseudotsuga menziesii and Juniperus communis were the only 2 species represented by many individuals, while 4 pine species plus Araucaria araucana were far less numerous and 6 other species were found fewer than 10 times each. Even those species found repeatedly were represented overwhelmingly by small individuals, and the great majority of introduced individuals were found not in native forest proper, but in somewhat open areas such as road verges, small remnant pastures, and deer trails. Invasion may be occurring, but too slowly to be clearly evidenced yet because of the longevity of the dominant native trees. Factors that may be stopping or slowing invasion include competition in gaps with native species, browsing by introduced deer, unfavorable soil, allelopathy, and natural enemies or other idiosyncratic factors for particular species. The absence of substantial invasion so far is no guarantee against future invasion, particularly if some major natural or anthropogenous disturbance were to occur.     

<Emphasis Type="Italic">Prunus serotina</Emphasis> unleashed: invader dominance after 70 years of forest development

Propagule pressure and disturbance have both been found to facilitate invasion. Therefore, knowledge on the history of introduction and disturbance is vital for understanding an invasion process, and research should focus on areas in which the invasive species has not been deliberately introduced or managed to study unconfounded colonization patterns. Comparing the outcome of such spontaneous colonization processes for different ecosystems might provide a useful framework for setting management priorities for invasive species that enter new, uninvaded areas. We focused on the 70-year spontaneous spread of the invasive tree species Prunus serotina in a pine forest in the Netherlands. To reconstruct the invasion pattern, we combined historical maps, tree ring analysis, spatially explicit tree inventory data, seed density data, and regeneration data for both native and non-native species. Prunus serotina was the only species that showed successful regeneration: the species was present throughout the forest in the tree, shrub, and herb layer. Native species were not able to outgrow the seedling stage. Our data demonstrate that P. serotina is a gap-dependent species with high seed production that builds up a seedling bank. We also compared the results of this study with a similar study on P. serotina colonization in a deciduous forest in Belgium, where P. serotina invasion was not successful. The sharp contrast between the outcomes of the two invasion processes shows the importance of studying an invasive species and the recipient ecosystem jointly and made us raise the hypothesis that herbivore pressure may facilitate P. serotina invasion.     

Predicting spread of invasive macrophytes in New Zealand lakes using indirect measures of human accessibility

1. Models predicting invasive macrophyte spread between lakes provide an important tool for focusing proactive management efforts to lakes deemed susceptible to invasion. However, challenges to forecasting macrophyte spread include wide physiological tolerances of invasive macrophytes and a lack of information on the relative importance of the various human vectors (e.g. boating traffic). In New Zealand, three invasive species that reproduce vegetatively, Ceratophyllum demersum, Lagarosiphon major, Egeria densa, and a single species that reproduces sexually, Utricularia gibba, are currently spreading across the lake landscape at a great cost to the local ecology and economy. 2. In this study, we first examined whether variables that indirectly describe weed spread via human access and use, as well as a lake’s position in the landscape, could describe the distribution of these four species using a boosted regression trees (BRT) modelling approach. Then, as these invasive species have not reached their full invasion potential, we examined how giving more influence to infected lakes at the edge of the invasion front, and including all lakes across New Zealand as background samples, simulating ‘absences beyond the invasion front’, influenced our ability to forecast the potential for new lakes to be invaded. 3. The BRT models identified that variables characterising human access and use, as well as lake position, were associated with the occurrence of the three vegetatively reproducing macrophytes. Weed occurrence was more likely when there was a highway in the vicinity, human population density was high and if the lake was large (c. 55 km²). But in the single case of U. gibba, temperature was the variable that best explained occurrence. This is consistent with the suggestion that U. gibba is predominantly dispersed by waterbirds, rather than human activity. 4. But for all four species, the BRT models based on the recorded observations alone predicted observed invasions with low prediction probabilities and did not forecast further spread. By contrast, when observations at the edge of the invasion front were upweighted, and additional background lakes implemented into the model, recorded observations were predicted and additional lakes were forecast to be at risk, suggesting that these models better captured the current and potential distribution of these macrophyte species. 5. The use of variables that characterise weed spread could provide similar insights into other systems where survey information on the nature, strength and direction of invasion vectors is lacking. Furthermore, when weighting the data, many lakes across New Zealand were forecasted to be at risk of invasion. The advantage of weighing the presence data was that insights into the potential for a species to spread were obtained. The probabilistic estimates of risk, as derived from the models, together with other information for prioritising lakes, can be used to focus surveillance and protection efforts.     

Impact of avian frugivores on dispersal and recruitment of the invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> in an agricultural landscape

Although seed dispersal is considered to be a key process determining the spatial structure and spread of non-native plant populations, few studies have explicitly addressed the link between dispersal vector behaviour, seed distribution and seedling recruitment to gain insight into the process of exotic species invasion within a fragmented landscape context. The present study analyses the relationship between avian frugivory and spatial patterns of seed deposition and seedling recruitment for an expanding population of the invasive Prunus serotina in a hedgerow network landscape in Flanders, Belgium. We quantified fruit production, observed frugivores, and determined the spatial distribution of bird droppings and P. serotina seedlings. A relatively diverse assemblage of frugivores visited P. serotina seed trees, with Columba palumbus and Turdus merula being by far the most important dispersers. Landscape structure strongly affected dispersal vector behaviour and the spatial distribution of perching birds, droppings and seedlings. Frugivorous birds non-randomly dispersed seeds to perching sites and an association between perching birds, seed deposition and seedling recruitment was found. Results indicate that landscape structure contributes to non-random seed deposition of P. serotina by common local frugivores. Cutting the larger seed trees is proposed as the most feasible measure to slow down the invasion rate.     

Niche shifts during the global invasion of the Asian tiger mosquito, Aedes albopictus Skuse (Culicidae), revealed by reciprocal distribution models

Aim Niche‐based distribution models are often used to predict the spread of invasive species. These models assume niche conservation during invasion, but invasive species can have different requirements from populations in their native range for many reasons, including niche evolution. I used distribution modelling to investigate niche conservatism for the Asian tiger mosquito (Aedes albopictus Skuse) during its invasion of three continents. I also used this approach to predict areas at risk of invasion from propagules originating from invasive populations. Location Models were created for Southeast Asia, North and South America, and Europe. Methods I used maximum entropy (Maxent ) to create distribution models using occurrence data and 18 environmental datasets. One native model was created for Southeast Asia; this model was projected onto North America, South America and Europe. Three models were created independently for the non‐native ranges and projected onto the native range. Niche overlap between native and non‐native predictions was evaluated by comparing probability surfaces between models using real data and random models generated using a permutation approach. Results The native model failed to predict an entire region of occurrences in South America, approximately 20% of occurrences in North America and nearly all Italian occurrences of A. albopictus. Non‐native models poorly predict the native range, but predict additional areas at risk for invasion globally. Niche overlap metrics indicate that non‐native distributions are more similar to the native niche than a random prediction, but they are not equivalent. Multivariate analyses support modelled differences in niche characteristics among continents, and reveal important variables explaining these differences. Main conclusions The niche of A. albopictus has shifted on invaded continents relative to its native range (Southeast Asia). Statistical comparisons reveal that the niche for introduced distributions is not equivalent to the native niche. Furthermore, reciprocal models highlight the importance of controlling bi‐directional dispersal between native and non‐native distributions.     

Habitat modification inhibits conspecific seedling recruitment in populations of an invasive ecosystem engineer

Invasive species that strongly modify their physical habitat are a particular management concern. Theoretical models predict that habitat modification could speed spread rates or allow invasion of sites that would otherwise resist invasion. There are few empirical tests of this hypothesis, however. We tested whether habitat modification by invading Spartina alterniflora populations facilitates conspecific seedling recruitment and spatial spread in Willapa Bay, WA, USA. Established S. alterniflora individuals strongly modified their local physical environment. Hydrologic flow, porewater salinity, and light availability were decreased while sediment NH₄ ⁺ increased with increasing S. alterniflora stem density. The S. alterniflora seed bank was greater and spring seedlings were denser within meadows of S. alterniflora than on unvegetated tideflats. However, almost all seedling recruitment after 1 year occurred on tideflats or on meadow edge plots where the above ground S. alterniflora biomass had been removed. Instead of facilitating invasive spread, ecosystem engineering in this system appears to create conditions that inhibit local seedling recruitment. These results suggest that the influence of ecosystem engineering on invasive spread is highly contingent on the relative spatial scales of habitat modification, environmental heterogeneity, and propagule availability. Control activities could change these spatial relationships, however, inadvertently promoting invasive recruitment.     

入侵害虫蔗扁蛾在我国的潜在分布区

【目的】蔗扁蛾是危害巴西木、甘蔗等园林植物和经济作物的重要入侵害虫。该虫于20世纪90年代初在我国被发现,现已分布在海南、广东和上海等19个省市,并有迅速扩散蔓延的趋势。对入侵害虫的潜在分布区进行预测,可为实施害虫监测和管理提供参考。【方法】根据蔗扁蛾已有分布点的记录,分别在4种地理区域构建Maxent生态位模型,并采用加权平均值法对其进行整合,进而分析蔗扁蛾在我国的潜在分布区。【结果】基于4种地理区域构建的Maxent模型对我国南部地区的预测结果基本一致,4种模型的预测差异主要在新疆北部和西南部、黑龙江东部和西部、吉林西部、山西中部等地区。整合模型显示,华东和华南地区以及东部沿海地区具有较大的分布可能性。【结论】蔗扁蛾在我国尤其是南方具有较大的潜在分布空间。这些地区应警惕蔗扁蛾的入侵,同时采取应对措施防止其进一步扩散。     

Pine pest aphids of the genus Eulachnus (Hemiptera: Aphididae: Lachninae): how far can their range extend?

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The spread of the Argentine ant: environmental determinants and impacts on native ant communities

The increasing numbers of invasive species have stimulated the study of the underlying causes promoting the establishment and spread of exotic species. We tracked the spread of the highly invasive Argentine ant (Linepithema humile) along an environmental and habitat gradient on the northeastern Iberian Peninsula to determine the role of climatic, habitat and biotic variables on the rate of spread, and examine impact on native ant communities. We found the species well-established within natural environments. The mean annual rate of spread of the invasion (7.94 ± 2.99 m/year) was relatively low compared to other studies, suggesting that resistance posed by native ants in natural environments with no or low human disturbance might delay (although not prevent) the spread of the invasion irrespectively of the land-use type. Factors related to the distance to urban areas and characteristics of native and introduced populations explained the rate of spread of the invasion, while habitat-related variables determined the distribution of native ants and the impact of the Argentine ant on them. Native ant communities became more homogeneous following the invasion due to the decline of species richness and abundance. Only few species (Plagiolepis pygmaea and Temnothorax spp.) were able to cope with the spread of the invasion, and were possibly favored by the local extinction of other ant species. Taken together, our results indicate that land uses per se do not directly affect the spread of L. humile, but influence its invasive success by molding the configuration of native ant communities and the abiotic suitability of the site.     

Predicted range expansion of Chinese tallow tree (Triadica sebifera) in forestlands of the southern United States

Aim We present an integrated approach for predicting future range expansion of an invasive species (Chinese tallow tree) that incorporates statistical forecasting and analytical techniques within a spatially explicit, agent‐based, simulation framework. Location East Texas and Louisiana, USA. Methods We drew upon extensive field data from the US Forest Service and the US Geological Survey to calculate spread rate from 2003 to 2008 and to parameterize logistic regression models estimating habitat quality for Chinese tallow within individual habitat cells. We applied the regression analyses to represent population spread rate as a function of habitat quality, integrated this function into a logistic model representing local spread, and coupled this model with a dispersal model based on a lognormal kernel within the simulation framework. We simulated invasions beginning in 2003 based on several different dispersal velocities and compared the resulting spatial patterns to those observed in 2008 using cross Mantel’s tests. We then used the best dispersal velocity to predict range expansion to the year 2023. Results Chinese tallow invasion is more likely in low and flat areas adjacent to water bodies and roads, and less likely in mature forest stands and in pine plantations where artificial regeneration by planting seedlings is used. Forecasted invasions resulted in a distribution that extended from the Gulf Coast of Texas and Louisiana northward and westward as much as 300 km, representing approximately 1.58 million ha. Main conclusions Our new approach of calculating time series projections of annual range expansion should assist land managers and restoration practitioners plan proactive management strategies and treatments. Also, as field sampling continues on the national array of FIA plots, these new data can be incorporated easily into the present model, as well as being used to develop and/or improve models of other invasive plant species.     

Pisonia grandis monocultures limit the spread of an invasive ant—a case of carbohydrate quality?

The mechanisms by which invasive species are able to spread into and dominate natural communities are poorly understood and remain a focus of invasion research. In this quest, studying invasions that are limited by a controlling factor will be more informative than will studies documenting unabated spread and impacts. Some ant species are very successful invaders, and research demonstrating abiotic and biotic factors limiting their success has aided the understanding of invasion ecology. We report here a study showing the highly invasive African big headed ant Pheidole megacephala having a novel distribution on coral cays within Australia’s Great Barrier Reef. These patterns displayed a clear limitation of its distribution with monocultures of the tree Pisonia grandis. This distribution was contrary to the known environmental limitations of the ant, and the limitation could not be associated with an underlying abiotic determinant of the vegetation type. We present these distributional patterns, and following consideration of all known biotic and abiotic limitations of ant invasions we discuss the potential that the peculiar ecophysiology of P. grandis is the causal factor. Specifically, we suggest that the quality of carbohydrate supply to ants is a limitation to invasive spread in much the same way that carbohydrate quantity is known to affect ant population densities in other ecosystems.     

Niche shift can impair the ability to predict invasion risk in the marine realm: an illustration using Mediterranean fish invaders

Climatic niche conservatism, the tendency of species‐climate associations to remain unchanged across space and time, is pivotal for forecasting the spread of invasive species and biodiversity changes. Indeed, it represents one of the key assumptions underlying species distribution models (SDMs), the main tool currently available for predicting range shifts of species. However, to date, no comprehensive assessment of niche conservatism is available for the marine realm. We use the invasion by Indo‐Pacific tropical fishes into the Mediterranean Sea, the world's most invaded marine basin, to examine the conservatism of the climatic niche. We show that tropical invaders may spread far beyond their native niches and that SDMs do not predict their new distributions better than null models. Our results suggest that SDMs may underestimate the potential spread of invasive species and call for prudence in employing these models in order to forecast species invasion and their response to environmental change.     

The interplay of disturbance,vegetation structure,and propagule pressure contributes to Pinus kesiya invasion in Tapia woodland,Madagascar

Pinus are among the highly invasive species that have spread outside their plantation area after their introduction in the Southern Hemisphere. The case of Pinus kesiya invasion is observed in the high plateau of Madagascar, inside the sclerophyll Tapia woodland which is dominated by the endemic Uapaca bojeri tree species. The analysis of this invasion was carried out using 375 plots of 100 m² each in Tapia woodland. Data on the vegetation structure, the plot characteristics and the propagule pressure were collected. We recorded a total of 740 pines distributed in 29.8% of the plots. The generalized linear model built on P. kesiya at the three different life stages allowed us to highlight a different explicative variable on the species’ presence and abundance separately. The factors explaining pine occurrence varied according to the pine life stage. In the seedling stage, the distance of the plot from the propagule source combined with the longitudinal position of the plot explained 18% of the pine presence. In the sapling and adult stages, the vegetation structure was the main important factor (22% and 11% of variation explained regarding presence and abundance). The frequency of U. bojeri and the degree of disturbance were the most important factors characterizing this vegetation structure. Based on these results, a strategy to control pine invasion in the Tapia woodland may focus on enrichment with U. bojeri and limitation of the plantation of P. kesiya in proximity.

     

Recovery of Native Grasslands after Removing Invasive Pines

The advance of exotic tree and shrub species is one of the main threats to conservation of the last relicts of natural grassland in South America; however, control actions in the region are still scarce and there are almost no evaluations of the recovery of natural ecosystems after removing invasive plants. Monitoring of the vegetation during the years after removal of invasive trees is critical in order to decide whether an active restoration strategy is necessary. The recovery of montane grassland four years after the control of a dense invasion of Aleppo pine (Pinus halepensis) is described in this study. Experimental clearing areas were followed during four years and compared to grassland controls. Variation was seen in the levels of recovery in function of the proximity of sectors of grassland that are free of invasive species and/or the density of invasive trees before control. Native species slowly replaced many exotic herbs that had appeared as pioneers, there was low recruitment of pine seedlings in spite of the quantity of seeds from trees that surrounded the clearings, and species richness and diversity were restored, including cover of the typical grasses in the controls. Recovery of grassland after felling was shown to be successful and does not seem to be seed limited if tree removal occurs early in the invasion process.     

Invasive plant species in the West Indies: geographical,ecological, and floristic insights

The level of invasion (number or proportion of invasive species) in a given area depends on features of the invaded community, propagule pressure, and climate. In this study, we assess the invasive flora of nine islands in the West Indies to identify invasion patterns and evaluate whether invasive species diversity is related to geographical, ecological, and socioeconomic factors. We compiled a database of invasive plant species including information on their taxonomy, origin, pathways of introduction, habitats, and life history. This database was used to evaluate the similarity of invasive floras between islands and to identify invasion patterns at regional (West Indies) and local (island) scales. We found a total of 516 alien plant species that are invasive on at least one of the nine islands studied, with between 24 to 306 invasive species per island. The invasive flora on these islands includes a wide range of taxonomic groups, life forms, and habitats. We detected low similarity in invasive species diversity between islands, with most invasive species (>60%) occurring on a single island and 6% occurring on at least five islands. To assess the importance of different models in predicting patterns of invasive species diversity among islands, we used generalized linear models. Our analyses revealed that invasive species diversity was well predicted by a combination of island area and economic development (gross domestic product per capita and kilometers of paved roadways). Our results provide strong evidence for the roles of geographical, ecological, and socioeconomic factors in determining the distribution and spread of invasive species on these islands. Anthropogenic disturbance and economic development seem to be the major drivers facilitating the spread and predominance of invasive species over native species.