From introduction to equilibrium: reconstructing the invasive pathways of the Argentine ant in a Mediterranean region

Determining the geographical range of invasive species is an important component of formulating effective management strategies. In the absence of detailed distributional data, species distribution models can provide estimates of an invasion range and increase our understanding of the ecological processes acting at various spatial scales. We used two complementary approaches to evaluate the influence of historical and environmental factors in shaping the distribution of the Argentine ant ( Linepithema humile ), a widespread, highly invasive species native to South America. Occurrence data were combined with environmental data at incremental spatial scales (extent and resolution) to predict the suitable range of the ant invasion using ecological niche models. In addition, we also used a spread model that simulated the jump dispersal of the species to identify the most plausible scenarios of arrival of L. humile in the NE Iberian Peninsula at local scales. Based on the results of both modelling practices, we suggest that L. humile might have reached its maximum geographic range at regional scales in the NE Iberian Peninsula. However, the species does not appear in equilibrium with the environment at small spatial scales, and further expansions are expected along coastal and inland localities of the Costa Brava. Long-distance jumps are ultimately responsible for the spread of the Argentine ant in the area. Overall, our study shows the utility of combining niche based models with spread models to understand the dynamics of species' invasions.     

Environmental data sets matter in ecological niche modelling: an example with Solenopsis invicta and Solenopsis richteri

Aim  In response to a recent paper suggesting the failure of ecological niche models to predict between native and introduced distributional areas of fire ants ( Solenopsis invicta ), we sought to assess methodological causes of this failure.
Location  Ecological niche models were developed on the species' native distributional area in South America, and projected globally.
Methods  We developed ecological niche models based on six different environmental data sets, and compared their respective abilities to anticipate the North American invasive distributional area of the species.
Results  We show that models based on the 'bioclimatic variables' of the WorldClim data set indeed fail to predict the full invasive potential of the species, but that models based on four other data sets could predict this potential correctly.
Main conclusions  The difference in predictive abilities appears to centre on the complexity of the environmental variables involved. These results emphasize important influences of environmental data sets on the generality and ability of ecological niche models to anticipate novel phenomena, and offer a simpler explanation for the lack of predictive ability among native and invaded distributional areas than that of niche shifts.     

Conclusions about niche expansion in introduced Impatiens walleriana populations depend on method of analysis

Determining the degree to which climate niches are conserved across plant species' native and introduced ranges is valuable to developing successful strategies to limit the introduction and spread of invasive plants, and also has important ecological and evolutionary implications. Here, we test whether climate niches differ between native and introduced populations of Impatiens walleriana, globally one of the most popular horticultural species. We use approaches based on both raw climate data associated with occurrence points and ecological niche models (ENMs) developed with Maxent. We include comparisons of climate niche breadth in both geographic and environmental spaces, taking into account differences in available habitats between the distributional areas. We find significant differences in climate envelopes between native and introduced populations when comparing raw climate variables, with introduced populations appearing to expand into wetter and cooler climates. However, analyses controlling for differences in available habitat in each region do not indicate expansion of climate niches. We therefore cannot reject the hypothesis that observed differences in climate envelopes reflect only the limited environments available within the species' native range in East Africa. Our results suggest that models built from only native range occurrence data will not provide an accurate prediction of the potential for invasiveness if applied to areas containing a greater range of environmental combinations, and that tests of niche expansion may overestimate shifts in climate niches if they do not control carefully for environmental differences between distributional areas.     

Native range climate is insufficient to predict anuran invasive potential

Niche conservatism explains biological invasions worldwide. However, a plethora of ecological processes may lead invasive species to occupy environments that are different from those found within native ranges. Here, we assess the potential invadable areas of  the world’s most pervasive invasive amphibians: the cane toad, Rhinella marina?+?R. horribilis, and the North American bullfrog, Lithobates catesbeianus. The uncontrolled spread of such voracious, large-bodied, and disease-tolerant anurans has been documented to impact native faunas worldwide. To disentangle their invasion-related niche dynamics, we compared the predictive ability and distributional forecasts of ecological niche models calibrated with information from native, invaded and pooled (native?+?invaded) ranges. We found that including occurrences from invaded ranges improved model accuracy for both studied species. Non-native occurrences also accounted for 54% and 61% increase in the total area of potential distribution of the cane toad and bullfrog, respectively. Besides, the latter species occupied locations with climatic conditions that are more extreme than those found within its native range. Our results indicate that the occupancy of environments different from those found in native ranges increases the overall potential distribution of the studied invasive anuran species. Therefore, climate information on native ranges alone is insufficient to explain and anticipate the distributional patterns of invasion of cane toads and bullfrogs, underestimating predictions of potential invadable distribution. Moreover, such an observed expansion of realized niches towards occupancy of climates not found within native ranges also has clear implications for invasion risk assessments based on climate modelling worldwide.

     

Shifting global invasive potential of European plants with climate change

Global climate change and invasions by nonnative species rank among the top concerns for agents of biological loss in coming decades. Although each of these themes has seen considerable attention in the modeling and forecasting communities, their joint effects remain little explored and poorly understood. We developed ecological niche models for 1804 species from the European flora, which we projected globally to identify areas of potential distribution, both at present and across 4 scenarios of future (2055) climates. As expected from previous studies, projections based on the CGCM1 climate model were more extreme than those based on the HadCM3 model, and projections based on the a2 emissions scenario were more extreme than those based on the b2 emissions scenario. However, less expected were the highly nonlinear and contrasting projected changes in distributional areas among continents: increases in distributional potential in Europe often corresponded with decreases on other continents, and species seeing expanding potential on one continent often saw contracting potential on others. In conclusion, global climate change will have complex effects on invasive potential of plant species. The shifts and changes identified in this study suggest strongly that biological communities will see dramatic reorganizations in coming decades owing to shifting invasive potential by nonnative species.     

海南岛霸王岭热带天然林景观中主要木本植物关键种的潜在分布

在较大的空间尺度上生态位模型是预测物种潜在分布的有效途径之一。为了探讨在热带天然林景观中木本植物(限于乔木和灌木)主要关键种的潜在分布,在对海南岛霸王岭的热带天然林进行按公里网格样方调查的基础上,采用演替地位和最大潜在高度两个功能性指标对物种进行了功能群划分,并在功能群框架下运用优势度指数法进行了关键种的确定;采用基于地理信息系统(Geographic information system, GIS)的基于规则集合预测的遗传算法(Algorithm for rule-set prediction, GARP)生态位模型对主要关键种的地理分布进行了预测,并应用受试者工作特征分析进行了模型精度验证;应用多元线性回归分析对影响各关键种潜在分布的关键因子进行了确定。结果表明:除了顶极次林层乔木功能群和顶极主林层乔木功能群外,在先锋种功能群、顶极灌木种功能群和顶极超冠层乔木功能群中采用优势度指数法划分出的关键种较为理想;一般来讲,在进行预测的8个关键种中,除了先锋主林层乔木种海南杨桐(Adinandra hainanensis),其它3个先锋种毛稔(Melastoma sanquiueum)、银柴(Aporosa chinensis)和枫香(Liquidambar formosana) 在研究区北部、西部以及西南部均具有较高的发生概率,而顶极种除了顶极超冠层乔木种南亚松(Pinus merkusii)外,九节(Psychotria rubra)、高脚罗伞(Ardisia quinquegona)和海南椎(Castanopsis hainanensis)具有相似的潜在分布格局,在研究区中部、东南部和南部地区具有较高的发生概率;相关分析表明极端最低温、年均温、极端最高温、年均降水量、海拔和坡向6大因子是影响研究区关键种潜在分布的关键因子;精度检验表明,GARP模型对8个关键种的潜在分布预测效果均较好,而其中又以银柴和海南椎的预测精度最高。     

Detrimental effects of highly efficient interference competition: invasive Argentine ants outcompete native ants at toxic baits

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. Previous studies that examined the mechanisms by which Argentine ants attain ecological dominance showed that superior interference and exploitation competition are key to the successful displacement of native ant species. The objective of this research was to test the hypothesis that effective interference competition by Argentine ants may also be detrimental to the survival of Argentine ant colonies where Argentine ants and native ants compete at toxic baits used to slow the spread of Argentine ants. To study this hypothesis, we examined the competitive interactions between Argentine ants and native odorous house ants, Tapinoma sessile, in the presence and absence of toxic baits. Results showed that Argentine ants aggressively outcompete T. sessile from toxic baits through efficient interference competition and monopolize bait resources. This has severe negative consequences for the survival of Argentine ants as colonies succumb to the toxic effects of the bait. In turn, T. sessile avoid areas occupied by Argentine ants, give up baits, and consequently suffer minimal mortality. Our results provide experimental evidence that highly efficient interference competition may have negative consequences for Argentine ants in areas where toxic baits are used and may provide a basis for designing innovative management programs for Argentine ants. Such programs would have the double benefit of selectively eliminating the invasive species while simultaneously protecting native ants from the toxic effects of baits.     

Effect of Climate Change on Invasion Risk of Giant African Snail (Achatina fulica Férussac, 1821: Achatinidae) in India

The Giant African Snail (Achatina fulica) is considered to be one the world’s 100 worst invasive alien species. The snail has an impact on native biodiversity, and on agricultural and horticultural crops. In India, it is known to feed on more than fifty species of native plants and agricultural crops and also outcompetes the native snails. It was introduced into India in 1847 and since then it has spread all across the country. In this paper, we use ecological niche modeling (ENM) to assess the distribution pattern of Giant African Snail (GAS) under different climate change scenarios. The niche modeling results indicate that under the current climate scenario, Eastern India, peninsular India and the Andaman and Nicobar Islands are at high risk of invasion. The three different future climate scenarios show that there is no significant change in the geographical distribution of invasion prone areas. However, certain currently invaded areas will be more prone to invasion in the future. These regions include parts of Bihar, Southern Karnataka, parts of Gujarat and Assam. The Andaman and Nicobar and Lakshadweep Islands are highly vulnerable to invasion under changed climate. The Central Indian region is at low risk due to high temperature and low rainfall. An understanding of the invasion pattern can help in better management of this invasive species and also in formulating policies for its control.     

紫茎泽兰Eupatorium adenophorum Spreng.在中国入侵分布预测

原产于墨西哥的紫茎泽兰Eupatorium adenophorum Spreng.作为一个有害的外来物种在印度、新西兰和澳大利亚生长已有很长时间。在中国,尤其是在南方和西南地区其蔓延速度之快,带来了不可忽视的经济和社会后果。我们采用了生态位模拟新方法来预测紫茎泽兰可能入侵的范围。据此,预测的潜在分布区包括该植物在中国境内已分布的省份及未来华中、华东易受入侵的区域。     

Climate change and the potential distribution of an invasive shrub, Lantana camara L

The threat posed by invasive species, in particular weeds, to biodiversity may be exacerbated by climate change. Lantana camara L. (lantana) is a woody shrub that is highly invasive in many countries of the world. It has a profound economic and environmental impact worldwide, including Australia. Knowledge of the likely potential distribution of this invasive species under current and future climate will be useful in planning better strategies to manage the invasion. A process-oriented niche model of L. camara was developed using CLIMEX to estimate its potential distribution under current and future climate scenarios. The model was calibrated using data from several knowledge domains, including phenological observations and geographic distribution records. The potential distribution of lantana under historical climate exceeded the current distribution in some areas of the world, notably Africa and Asia. Under future scenarios, the climatically suitable areas for L. camara globally were projected to contract. However, some areas were identified in North Africa, Europe and Australia that may become climatically suitable under future climates. In South Africa and China, its potential distribution could expand further inland. These results can inform strategic planning by biosecurity agencies, identifying areas to target for eradication or containment. Distribution maps of risk of potential invasion can be useful tools in public awareness campaigns, especially in countries that have been identified as becoming climatically suitable for L. camara under the future climate scenarios.     

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。     

Long-term impact of exotic ants on the native ants of Madeira

Abstract.  1. The earliest exotic records for two notorious invasive ants, the big-headed ant ( Pheidole megacephala ) and the Argentine ant ( Linepithema humile ), both come from the Atlantic islands of Madeira, where the two species underwent population explosions in the 1850s and 1890s respectively. Researchers have long assumed that these invaders spread across all of Madeira and exterminated most or all native ants, despite no research actually documenting such impact.
2. Re-examination of first-hand nineteenth century accounts suggest that P. megacephala and L. humile may never have spread beyond coastal lowland areas, representing < 10% of Madeira's land area. In 2002, native ants dominated most of Madeira; P. megacephala and L. humile were restricted to ≈ 0.3% and ≈ 6% of Madeira's land area respectively.
3. Of the 10 native ant species known from Madeira, only one ( Temnothorax wollastoni ) was not present in 1999–2002 surveys. Although exotic ants may have exterminated T. wollastoni , it seems likely that this species still survives.
4. Thus, even after 150 or more years of residence, P. megacephala and L. humile have come to occupy only a small part of Madeira, and appear to have had little impact.
5. Most of Madeira may be too cool for P. megacephala and perhaps too moist for L. humile to dominate. Also, Madeira's vast natural areas may generally lack weedy vegetation that can support high densities of plant-feeding Hemiptera critical for the ecological dominance of invasive ants. Finally, a dominant native ant, Lasius grandis , inhabiting ≈ 84% of Madeira, may actively exclude P. megacephala and L. humile .     

Equilibrium or not? Modelling potential distribution of invasive species in different stages of invasion

Aim The assumption of equilibrium between organisms and their environment is a standard working postulate in species distribution models (SDMs). However, this assumption is typically violated in models of biological invasions where range expansions are highly constrained by dispersal and colonization processes. Here, we examined how stage of invasion affects the extent to which occurrence data represent the ecological niche of organisms and, in turn, influences spatial prediction of species’ potential distributions. Location Six ecoregions in western Oregon, USA. Methods We compiled occurrence data from 697 field plots collected over a 9‐year period (2001–09) of monitoring the spread of invasive forest pathogen Phytophthora ramorum. Using these data, we applied ecological‐niche factor analysis to calibrate models of potential distribution across different years of colonization. We accounted for natural variation and uncertainties in model evaluation by further investigating three hypothetical scenarios of varying equilibrium in a simulated virtual species, for which the ‘true’ potential distribution was known. Results We confirm our hypothesis that SDMs calibrated in early stages of invasion are less accurate than models calibrated under scenarios closer to equilibrium. SDMs that are developed in early stages of invasion tend to underpredict the potential range compared to models that are built in later stages of invasion. Main conclusions A full environmental niche of invasive species cannot be effectively captured with data from a realized distribution that is restricted by processes preventing full occupancy of suitable habitats. If SDMs are to be used effectively in conservation and management, stage of invasion needs to be considered to avoid underestimation of habitats at risk of invasion.     

提高生态位模型转移能力来模拟入侵物种 的潜在分布

生态位模型利用物种分布点所关联的环境变量去推算物种的生态需求, 模拟物种的分布。在模拟入侵物种分布时, 经典生态位模型包括模型构建于物种本土分布地, 然后将其转移并投射至另一地理区域, 来模拟入侵物种的潜在分布。然而在模型运用时, 出现了模型的转移能力较低、模拟的结果与物种的实际分布不相符的情况, 由此得出了生态位漂移等不恰当的结论。提高生态位模型的转移能力, 可以准确地模拟入侵物种的潜在分布, 为入侵种的风险评估提供参考。作者以入侵种茶翅蝽(Halyomorpha halys)和互花米草(Spartina alterniflora)为例, 从模型的构建材料(即物种分布点和环境变量)入手, 全面阐述提高模型转移能力的策略。在构建模型之前, 需要充分了解入侵物种的生物学特性、种群平衡状态、本土地理分布范围及物种的生物历史地理等方面的知识。在模型构建环节上, 物种分布点不仅要充分覆盖物种的地理分布和生态空间的范围, 同时要降低物种采样点偏差; 环境变量的选择要充分考虑其对物种分布的限制作用、各环境变量之间的空间相关性, 以及不同地理种群间生态空间是否一致, 同时要降低环境变量的空间维度; 模型构建区域要真实地反映物种的地理分布范围, 并考虑种群的平衡状态。作者认为, 在生态位保守的前提下, 如果模型是构建在一个合理方案的基础上, 生态位模型的转移能力是可以保证的, 在以模型转移能力较低的现象来阐述生态位分化时需要引起注意。     

Evidence of climatic niche shift during biological invasion

Niche-based models calibrated in the native range by relating species observations to climatic variables are commonly used to predict the potential spatial extent of species' invasion. This climate matching approach relies on the assumption that invasive species conserve their climatic niche in the invaded ranges. We test this assumption by analysing the climatic niche spaces of Spotted Knapweed in western North America and Europe. We show with robust cross-continental data that a shift of the observed climatic niche occurred between native and non-native ranges, providing the first empirical evidence that an invasive species can occupy climatically distinct niche spaces following its introduction into a new area. The models fail to predict the current invaded distribution, but correctly predict areas of introduction. Climate matching is thus a useful approach to identify areas at risk of introduction and establishment of newly or not-yet-introduced neophytes, but may not predict the full extent of invasions.     

Niche tracking and rapid establishment of distributional equilibrium in the house sparrow show potential responsiveness of species to climate change

The ability of species to respond to novel future climates is determined in part by their physiological capacity to tolerate climate change and the degree to which they have reached and continue to maintain distributional equilibrium with the environment. While broad-scale correlative climatic measurements of a species' niche are often described as estimating the fundamental niche, it is unclear how well these occupied portions actually approximate the fundamental niche per se, versus the fundamental niche that exists in environmental space, and what fitness values bounding the niche are necessary to maintain distributional equilibrium. Here, we investigate these questions by comparing physiological and correlative estimates of the thermal niche in the introduced North American house sparrow (Passer domesticus). Our results indicate that occupied portions of the fundamental niche derived from temperature correlations closely approximate the centroid of the existing fundamental niche calculated on a fitness threshold of 50% population mortality. Using these niche measures, a 75-year time series analysis (1930-2004) further shows that: (i) existing fundamental and occupied niche centroids did not undergo directional change, (ii) interannual changes in the two niche centroids were correlated, (iii) temperatures in North America moved through niche space in a net centripetal fashion, and consequently, (iv) most areas throughout the range of the house sparrow tracked the existing fundamental niche centroid with respect to at least one temperature gradient. Following introduction to a new continent, the house sparrow rapidly tracked its thermal niche and established continent-wide distributional equilibrium with respect to major temperature gradients. These dynamics were mediated in large part by the species' broad thermal physiological tolerances, high dispersal potential, competitive advantage in human-dominated landscapes, and climatically induced changes to the realized environmental space. Such insights may be used to conceptualize mechanistic climatic niche models in birds and other taxa.     

Invasive termites in a changing climate: A global perspective

Termites are ubiquitous insects in tropical, subtropical, and warm temperate regions and play an important role in ecosystems. Several termite species are also significant economic pests, mainly in urban areas where they attack human‐made structures, but also in natural forest habitats. Worldwide, approximately 28 termite species are considered invasive and have spread beyond their native ranges, often with significant economic consequences. We used predictive climate modeling to provide the first global risk assessment for 13 of the world's most invasive termites. We modeled the future distribution of 13 of the most serious invasive termite species, using two different Representative Concentration Pathways (RCPs), RCP 4.5 and RCP 8.5, and two projection years (2050 and 2070). Our results show that all but one termite species are expected to significantly increase in their global distribution, irrespective of the climatic scenario and year. The range shifts by species (shift vectors) revealed a complex pattern of distributional changes across latitudes rather than simple poleward expansion. Mapping of potential invasion hotspots in 2050 under the RCP 4.5 scenario revealed that the most suitable areas are located in the tropics. Substantial parts of all continents had suitable environmental conditions for more than four species simultaneously. Mapping of changes in the number of species revealed that areas that lose many species (e.g., parts of South America) are those that were previously very species‐rich, contrary to regions such as Europe that were overall not among the most important invasion hotspots, but that showed a great increase in the number of potential invaders. The substantial economic and ecological damage caused by invasive termites is likely to increase in response to climate change, increased urbanization, and accelerating economic globalization, acting singly or interactively.     

Global Invasion of Lantana camara: Has the Climatic Niche Been Conserved across Continents?

Lantana camara, a native plant from tropical America, is considered one of the most harmful invasive species worldwide. Several studies have identified potentially invasible areas under scenarios of global change, on the assumption that niche is conserved during the invasion process. Recent studies, however, suggest that many invasive plants do not conserve their niches. Using Principal Components Analyses (PCA), we tested the hypothesis of niche conservatism for L. camara by comparing its native niche in South America with its expressed niche in Africa, Australia and India. Using MaxEnt, the estimated niche for the native region was projected onto each invaded region to generate potential distributions there. Our results demonstrate that while L. camara occupied subsets of its original native niche in Africa and Australia, in India its niche shifted significantly. There, 34% of the occurrences were detected in warmer habitats nonexistent in its native range. The estimated niche for India was also projected onto Africa and Australia to identify other vulnerable areas predicted from the observed niche shift detected in India. As a result, new potentially invasible areas were identified in central Africa and southern Australia. Our findings do not support the hypothesis of niche conservatism for the invasion of L. camara. The mechanisms that allow this species to expand its niche need to be investigated in order to improve our capacity to predict long-term geographic changes in the face of global climatic changes.     

应用生态位模型研究外来入侵物种生态位漂移

由于基础生态位和实际生态位的改变,外来入侵物种在入侵地成功定殖、扩散后常会发生生态位漂移,而物种生态位漂移往往很难直接证明。生态位模型在假设入侵物种的生态位需求保守的前提下,以物种在其原产地的生态位需求为基础,预测其在入侵地的潜在分布,通过比较预测分布与实际分布的差异可以从一定程度上得到外来入侵物种的生态位是否发生漂移的间接证据。以我国入侵杂草胜红蓟在原产地的生态位需求为基础,应用生态位模型预测其在其他地区的潜在分布。研究结果表明,生态位模型可以很好地预测胜红蓟在亚太平洋地区和非洲地区的分布,但在我国,其预测分布与实际分布存在较大差别。胜红蓟在我国预测分布主要为云南、海南、台湾部分地区,而胜红蓟入侵我国后现已广泛分布于长江以南地区,其实际分布比预测分布广泛得多,由此推测胜红蓟在入侵我国后其生态位已经产生了漂移。     

Is There Any Evidence for Rapid,Genetically-Based,Climatic Niche Expansion in the Invasive Common Ragweed?

Climatic niche shifts have been documented in a number of invasive species by comparing the native and adventive climatic ranges in which they occur. However, these shifts likely represent changes in the realized climatic niches of invasive species, and may not necessarily be driven by genetic changes in climatic affinities. Until now the role of rapid niche evolution in the spread of invasive species remains a challenging issue with conflicting results. Here, we document a likely genetically-based climatic niche expansion of an annual plant invader, the common ragweed (Ambrosia artemisiifolia L.), a highly allergenic invasive species causing substantial public health issues. To do so, we looked for recent evolutionary change at the upward migration front of its adventive range in the French Alps. Based on species climatic niche models estimated at both global and regional scales we stratified our sampling design to adequately capture the species niche, and localized populations suspected of niche expansion. Using a combination of species niche modeling, landscape genetics models and common garden measurements, we then related the species genetic structure and its phenotypic architecture across the climatic niche. Our results strongly suggest that the common ragweed is rapidly adapting to local climatic conditions at its invasion front and that it currently expands its niche toward colder and formerly unsuitable climates in the French Alps (i.e. in sites where niche models would not predict its occurrence). Such results, showing that species climatic niches can evolve on very short time scales, have important implications for predictive models of biological invasions that do not account for evolutionary processes.