Biological invasions and scientific objectivity: Reply to Cassey et al. (2005)

Abstract We disagree with the assertion that recent human‐caused invasions differ substantially from historic natural invasions in their magnitudes and impacts on ecological processes. The position that exotic species are inherently ‘bad’ and should be eradicated is an ethical judgement, usually based on the naturalist fallacy or xenophobic prejudice; it is not a scientific judgement. The role of scientists in studying invasive species should be to gather, interpret and communicate information as accurately and objectively as possible.     

Molecular genetic approaches to elucidate the ecological and evolutionary issues associated with biological invasions

Biological invasions may cause serious damage to the native environments and threaten the native biodiversity. Molecular genetic approaches have been found to be powerful tools for investigating the ecological and evolutionary aspects of biological invasions because the genetic structure and level of genetic variation of an invasive species are changed following its invasion. The present article reviews the use of molecular markers in addressing various aspects of invasive species. The application of these techniques has shown that many invasive species are actually "cryptic" species – species whose uniqueness is only recognizable at the genetic level. An estimation of the actual number of invasive species is essential when evaluating its ecological and economic impacts. Molecular genetic approaches have also enabled the source populations of invasive species to be identified. Reconstructions of invasion histories are crucial to preventing future invasions and conserving the native biodiversity, while comparisons of genetic variations between the native and introduced populations provide valuable opportunities to elucidate the mechanisms of rapid adaptation demonstrated by many invasive species.     

Conservation in a cup of water: estimating biodiversity and population abundance from environmental DNA

Three mantras often guide species and ecosystem management: (i) for preventing invasions by harmful species, ‘early detection and rapid response’; (ii) for conserving imperilled native species, ‘protection of biodiversity hotspots’; and (iii) for assessing biosecurity risk, ‘an ounce of prevention equals a pound of cure.’ However, these and other management goals are elusive when traditional sampling tools (e.g. netting, traps, electrofishing, visual surveys) have poor detection limits, are too slow or are not feasible. One visionary solution is to use an organism’s DNA in the environment (eDNA), rather than the organism itself, as the target of detection. In this issue of Molecular Ecology, Thomsen et al. (2012) provide new evidence demonstrating the feasibility of this approach, showing that eDNA is an accurate indicator of the presence of an impressively diverse set of six aquatic or amphibious taxa including invertebrates, amphibians, a fish and a mammal in a wide range of freshwater habitats. They are also the first to demonstrate that the abundance of eDNA, as measured by qPCR, correlates positively with population abundance estimated with traditional tools. Finally, Thomsen et al. (2012) demonstrate that next‐generation sequencing of eDNA can quantify species richness. Overall, Thomsen et al. (2012) provide a revolutionary roadmap for using eDNA for detection of species, estimates of relative abundance and quantification of biodiversity.     

Distinct invasion strategies operating within a natural annual plant system

Alien plant species are known to have a wide range of impacts on recipient communities, from resident species’ exclusions to coexistence with resident species. It remains unclear; however, if this variety of impacts is due to different invader strategies, features of recipient communities or both. To test this, we examined multiple plant invasions of a single ecosystem in southwestern Australia. We used extensive community data to calculate pairwise segregation between target alien species and many co‐occurring species. We related segregation to species’ positions along community trait hierarchies and identified at least two distinct invasion strategies: ‘exploiters’ which occupy high positions along key trait hierarchies and reduce local native species diversity (particularly in nutrient‐enriched situations), and ‘coexisters’ who occupy intermediate trait positions and have no discernable impact on native diversity. We conclude that trait hierarchies, linked to measures of competition, can provide valuable insights about the processes driving different invasion outcomes.     

Can Invasive Species Facilitate Native Species? Evidence of How, When, and Why These Impacts Occur

Although the predatory and competitive impacts of biological invasions are well documented, facilitation of native species by non-indigenous species is frequently overlooked. A search through recent ecological literature found that facilitative interactions between invasive and native species occur in a wide range of habitats, can have cascading effects across trophic levels, can re-structure communities, and can lead to evolutionary changes. These are critical findings that, until now, have been mostly absent from analyses of ecological impacts of biological invasions. Here I present evidence for several mechanisms that exemplify how exotic species can facilitate native species. These mechanisms include habitat modification, trophic subsidy, pollination, competitive release, and predatory release. Habitat modification is the most frequently documented mechanism, reflecting the drastic changes generated by the invasion of functionally novel habitat engineers. Further, I predict that facilitative impacts on native species will be most likely when invasive species provide a limiting resource, increase habitat complexity, functionally replace a native species, or ameliorate predation or competition. Finally, three types of facilitation (novel, substitutive, and indirect) define why exotic species can lead to facilitation of native species. It is evident that understanding biological invasions at the community and ecosystem levels will be more accurate if we integrate facilitative interactions into future ecological research. Since facilitative impacts of biological invasions can occur with native endangered species, and can have wide-ranging impacts, these results also have important implications for management, eradication, and restoration.Contribution Number 2293, Bodega Marine Laboratory, University of California at Davis.     

洲际入侵植物生态位稳定性研究进展

人类活动引起的大规模洲际物种交换与生物入侵, 改变了当地生态系统结构与功能, 使生物多样性受到日益严重的威胁。本文通过综合分析主要国家和地区入侵植物的地理起源, 发现洲际入侵主要包括东亚—北美、东亚—南美、欧洲—南非、欧洲—北美、欧洲—东亚、北美—大洋洲等, 这些洲际入侵造成的后果往往比陆内入侵更为严重。利用物种分布模型(SDMs)预测入侵物种潜在分布范围是有效管理和提早预防生物入侵的重要依据, 但这些模型的一个关键假定是: 入侵物种的生态位在空间和时间上是保守的、稳定的。然而, 对于远离原产地种群并能快速适应新生境的洲际入侵植物来说, 生态位可能发生显著的变化。入侵种能否在入侵地保持原有的生态位, 取决于制约其生态分布的限制因素和生态过程在不同地区间是否发生变化。本文中作者总结了洲际入侵与陆内入侵的生态与进化过程的异同点, 认为这些限制物种原产地分布的因素如扩散限制、种间互作、适应性进化、生态可塑性和种群遗传特性等均可能导致入侵物种生态位的改变。建议下一步的研究应该重视: (1)对生态位属性进行多尺度的研究, 包括时间、空间、环境或系统发育等几个方面; (2)对比生态位稳定与发生偏移的物种特性, 确定什么样的入侵物种更容易改变原有的生态位; (3)进行生态位时间动态格局研究, 探讨生态位变化的倾向、历史速率和偏移程度, 以便判定生态位变化趋势。这些研究结果将会进一步提高物种分布模型的预测能力, 有助于更为准确地揭示气候变化和物种入侵对生物多样性的影响。     

外来种入侵的过程、机理和预测

生物入侵是指某种生物从原来的分布区域扩展到一个新的（通常也是遥远的）地区,在新的区域里,其后代可以繁殖、扩散并持续维持下去,生物入侵成功的原因,即与入侵者本身的生物学,生态学特征有关,也与群落的脆弱性有关,入侵者可能较本地种的竞争能力强,更适应当地的环境,有的入侵者还可以改变环境,使之对已有利,而不利于本地种。缺乏天敌制约。群落的稳定性低和异常的环境扰动往往导致生物入侵,生物入侵的预测包括哪一种外来种会变成入侵种？哪些生态系统区域会被入侵？影响程度如何？入侵种的扩散态势如何等内容,对有关的理论和模型作了评介。     

Is invasion success explained by the enemy release hypothesis?

A recent trend in invasion ecology relates the success of non‐indigenous species (NIS) to reduced control by enemies such as pathogens, parasites and predators (i.e. the enemy release hypothesis, ERH). Despite the demonstrated importance of enemies to host population dynamics, studies of the ERH are split – biogeographical analyses primarily show a reduction in the diversity of enemies in the introduced range compared with the native range, while community studies imply that NIS are no less affected by enemies than native species in the invaded community. A broad review of the invasion literature implies at least eight non‐exclusive explanations for this enigma. In addition, we argue that the ERH has often been accepted uncritically wherever (i) NIS often appear larger, more fecund, or somehow ‘better’ than either congeners in the introduced region, or conspecifics in the native range; and (ii) known enemies are conspicuously absent from the introduced range. However, all NIS, regardless of their abundance or impact, will lose natural enemies at a biogeographical scale. Given the complexity of processes that underlie biological invasions, we argue against a simple relationship between enemy ‘release’ and the vigour, abundance or impact of NIS.     

Contrasting functional trait syndromes underlay woody alien success in the same ecosystem

We performed a comprehensive comparative study of functional traits in coexisting alien and native woody species in order to examine the strategies related to resource use and dispersion underlying alien success in mountain Chaco woodlands of central Argentina. Our approach integrated seemingly contrasting pieces of evidence in the region. We specifically assessed whether (i) the ‘functional acquisitive trend’ previously observed along a broad environmental gradient accounts for woody alien naturalization when considering a single mesic ecosystem; or (ii) more than one trait syndrome is important among alien species, which would be more in line with the context‐dependent nature of biological invasions at a local scale. Fifteen vegetative and regenerative traits were measured on the most common 14 native and 11 alien woody species. We compared the attributes of (i) native and alien species and (ii) between native species and two contrasting groups of alien species identified in the previous analysis. The overall trait comparison (i) showed that, in terms of vegetative attributes, woody alien species tend to be on average more acquisitive than native species. However, (ii) two contrasting syndromes were revealed among alien species: a group of seven deciduous species with acquisitive attributes; and a group of four evergreen species showing markedly more conservative attributes than the first group. The functional attributes of ‘conservative aliens’ completely overlapped with the range observed for native species, except for an exclusive dispersal phenology and a stronger tendency to clonal spread. Acquisitive aliens, in turn, proved to be beyond the range of attributes of native species, at the acquisitive extreme, as they did in previous comparisons. Despite their importance, general trends in plant functional attributes across regions and ecosystems can sometimes obscure trends at more local scales that are nevertheless important for the understanding and management of particular systems. Our study concurs with previous general trends when looking at the overall comparison between native and alien species, but unveils contrasting functional strategies among alien species when examining their attributes more closely, even within the same ecosystem.     

新建川藏铁路(雅安-昌都段)沿线外来入侵植物种类及分布特征

横断山区为全球生物多样性热点地区之一, 也是全国生态屏障的重要组成部分。新建川藏铁路雅安至昌都段横跨横断山核心地区, 铁路建设形成的交通网络将沿线生物多样性热点区域与外界相连, 导致生物入侵风险陡增。为获得区域内外来入侵植物的种类及分布特征信息, 为即将开始的铁路工程建设、生态保护及生态修复等工作提供参考, 我们在雅安-昌都段内选择43个位点各进行长度1 km、宽度20 m的样线调查。研究结果显示: 雅安-昌都段共发现外来入侵植物58种, 隶属于18科42属, 其中出现频度最高的种类依次是牛膝菊(Galinsoga parviflora)、秋英(Cosmos bipinnatus)和鬼针草(Bidens pilosa)。从危害等级来看, 其中10种为恶性入侵种, 16种为严重入侵种, 8种为局部入侵种, 15种为一般入侵种, 9种为有待观察种, 超过半数种类具有明显入侵性。原产地分析结果显示美洲是该区域外来入侵植物的主要原产地。基于海拔及主要河流区段的比较研究发现: 入侵植物的种类数量呈现出明显的由东向西、由低海拔向高海拔逐渐递减的趋势, 该分布格局是环境因子和人类活动共同作用的结果。结合铁路沿线入侵现状和生境特征, 本文分析了铁路建设可能造成的外来植物入侵风险, 并针对入侵的防范提出了相应的建议。     

Advancing impact prediction and hypothesis testing in invasion ecology using a comparative functional response approach

Invasion ecology urgently requires predictive methodologies that can forecast the ecological impacts of existing, emerging and potential invasive species. We argue that many ecologically damaging invaders are characterised by their more efficient use of resources. Consequently, comparison of the classical ‘functional response’ (relationship between resource use and availability) between invasive and trophically analogous native species may allow prediction of invader ecological impact. We review the utility of species trait comparisons and the history and context of the use of functional responses in invasion ecology, then present our framework for the use of comparative functional responses. We show that functional response analyses, by describing the resource use of species over a range of resource availabilities, avoids many pitfalls of ‘snapshot’ assessments of resource use. Our framework demonstrates how comparisons of invader and native functional responses, within and between Type II and III functional responses, allow testing of the likely population-level outcomes of invasions for affected species. Furthermore, we describe how recent studies support the predictive capacity of this method; for example, the invasive ‘bloody red shrimp’ Hemimysis anomala shows higher Type II functional responses than native mysids and this corroborates, and could have predicted, actual invader impacts in the field. The comparative functional response method can also be used to examine differences in the impact of two or more invaders, two or more populations of the same invader, and the abiotic (e.g. temperature) and biotic (e.g. parasitism) context-dependencies of invader impacts. Our framework may also address the previous lack of rigour in testing major hypotheses in invasion ecology, such as the ‘enemy release’ and ‘biotic resistance’ hypotheses, as our approach explicitly considers demographic consequences for impacted resources, such as native and invasive prey species. We also identify potential challenges in the application of comparative functional responses in invasion ecology. These include incorporation of numerical responses, multiple predator effects and trait-mediated indirect interactions, replacement versus non-replacement study designs and the inclusion of functional responses in risk assessment frameworks. In future, the generation of sufficient case studies for a meta-analysis could test the overall hypothesis that comparative functional responses can indeed predict invasive species impacts.     

Assessing the risk of freshwater fish introductions into the Iberian Peninsula

1. Preventing the introduction of species likely to become invaders is the best management option to deal with biological invasions. A data set consisting of native, introduced and species not currently present in Iberian Peninsula (n = 167 species) was used to identify freshwater fish species that are likely to be introduced and become successful invaders in the near future. 2. Principal component analysis (PCA) of species traits was used to determine species likely to be introduced, assuming that the traits of species introduced in the future will resemble those of previously introduced species. The likelihood of introduction was calculated as the proportion of neighbour species (in the space defined by the PCA) that have been introduced to the Iberian Peninsula and, together with metrics related to different stages of invasion, was used to construct a region‐specific risk index (Iberian risk index). 3. Introduced species had higher index values compared with native species or species currently absent from the region. The Iberian risk index was positively related to the results of an independent risk analysis for freshwater fish as well as to the geographical spread of species previously introduced to the Iberian Peninsula. 4. Iberian risk index values were used to establish a cut‐off value for estimating the probability of a successful invasion. This threshold value was used to construct a list of 20 species to be included in a ‘watch list’ to prevent freshwater fish invasions in the Iberian Peninsula.     

Conservation Implications of Invasion by Plant Hybridization

The increasing number of invasive exotic plant species in many regions and the continuing alteration of natural ecosystems by humans promote hybridization between previously allopatric species; among both native as well as between native and introduced species. We review the ecological factors and mechanisms that promote such hybridization events and their negative consequences on biological diversity. Plant invasions through hybridization may occur in four different ways: hybridization between native species, hybridization between an exotic species and a native congener, hybridization between two exotics and by the introduction and subsequent spread of hybrids. The main harmful genetic effect of such hybrids on native species is the loss of both genetic diversity and of locally adapted populations, such as rare and threatened species. The spread of aggressive hybrid taxa can reduce the growth of, or replace, native species. The main factor promoting the formation of hybrids is species dispersal promoted by humans. However, the success and spread of hybrids is increased by disturbance and fragmentation of habitats, thus overcoming natural crossing barriers, and range expansions due to human activity. There are differences in flowering, pollination and seed dispersal patterns between parental species and hybrids. Hybrid resistance to pathogens and herbivores may also enhance the success of hybrids. To predict the mechanisms and consequences of invasions mediated by hybridization, extensive data on hybrid ecology and biology are needed, as well as carefully designed field experiments focused on the comparative ecology of parental populations and hybrids.     

The negative ecological impacts of a globally introduced species decrease with time since introduction

While there is a long‐history of biological invasions and their ecological impacts have been widely demonstrated across taxa and ecosystems, our knowledge on the temporal dynamic of these impacts remains extremely limited. Using a meta‐analytic approach, we investigated how the ecological impacts of non‐native brown trout (Salmo trutta), a model species with a 170‐year‐long and well‐documented history of intentional introductions across the globe, vary with time since introduction. We first observed significant negative ecological impacts immediately after the species introduction. Second, we found that the negative ecological impacts decrease with time since introduction and that the average ecological impacts become nonsignificant more than one century after introduction. This pattern was consistent across other ecological contexts (i.e., geographical location, levels of biological organization, and methodological approach). However, overall negative ecological impacts were more pronounced at the individual and population levels and in experimental studies. While the mechanisms leading to this decrease remain to be determined, our results indicate that rapid response of native organisms (e.g. adaptation, but also local extinction) may play an important role in this dynamic. Changes in native species traits and local extinction can have important conservation implications. Therefore, we argue that the decline of the negative ecological impacts over time should not be used as an argument to neglect the negative impacts of biological invasions.     

Suitable areas for the invasion expansion of Xylocopa bees in South America

The introduction of exotic species into native ecosystems can be a cause for concern when those species are invasive. Invasive species cause ecological problems and have socio-cultural impacts on human health and the economy; for example, invasive bees may negatively impact their introduced ecosystem by spreading diseases or outcompeting native pollinators. Xylocopa spp. bees are diverse and distributed throughout the Neotropics. However, Xylocopa augusti (Lepeletier, 1841) and Xylocopa splendidula (Lepeletier, 1841) are not native to Mediterranean Chile. This study aimed to evaluate the invasive potential of these exotic species and predict the potential macroecological effects of their invasions. We also aimed to pinpoint possible distributions for these species throughout South America. We correlated biogeographic occurrence data with climatic variables for each species to model their potential distribution in both current and future scenarios. The models provide strong evidence that both species are changing their distributions: their ranges are expanding towards western South America, particularly Bolivia, Chile and Peru. We demonstrate an increase in niche overlap between these species and show there are new geographic areas vulnerable to the establishment of these invasive bees under current and future climate conditions. These data suggest that these bees may adapt their geographic distribution as the climate changes and pose a threat to native pollinators in new geographic areas.     

A neutral terminology to define 'invasive' species

The use of simple terms to articulate ecological concepts can confuse ideological debates and undermine management efforts. This problem is particularly acute in studies of nonindigenous species, which alternatively have been called ‘exotic’, ‘introduced’, ‘invasive’ and ‘naturalised’, among others. Attempts to redefine commonly used terminology have proven difficult because authors are often partial to particular definitions. In an attempt to form a consensus on invasion terminology, we synthesize an invasional framework based on current models that break the invasion process into a series of consecutive, obligatory stages. Unlike previous efforts, we propose a neutral terminology based on this framework. This ‘stage‐based’ terminology can be used to supplement terms with ambiguous meanings (e.g. invasive, introduced, naturalized, weedy, etc.), and thereby improve clarity of future studies. This approach is based on the concept of ‘propagule pressure’ and has the additional benefit of identifying factors affecting the success of species at each stage. Under this framework, invasions can be more objectively understood as biogeographical, rather than taxonomic, phenomena; and author preferences in the use of existing terminology can be addressed. An example of this recommended protocol might be: ‘We examined distribution data to contrast the characteristics of invasive species (stages IVa and V) and noninvasive species (stages III and IVb)’.     

海洋外来物种入侵生态学研究

海洋外来物种入侵已成为最为严重的全球性环境问题之一。海洋生态系统类型多样、环境复杂,其生物入侵的监测、控制与管理难度相对较大。我国对陆地外来生物的入侵已开展了较为深入的研究,但对于海洋外来生物的入侵研究仍处于起步阶段,对其入侵监测、入侵机制、入侵危害的程度以及防治等问题缺乏基础数据。本文在分析国内外海洋外来生物入侵现状的基础上,概述其入侵生态学研究形势及相关成果,包括海洋外来物种的入侵途径、入侵过程、入侵生态效应以及全球变化对入侵的影响等。海洋外来生物的入侵可能对海洋生态系统造成直接或间接的影响,如种间竞争破坏生态环境、与土著种杂交造成遗传污染、病原生物及有毒藻类导致海洋生态灾害加剧等。此外,从政策和法规、入侵风险评估、监测和公共宣传教育、生物信息系统和有效管理机制等方面提出对我国海洋外来物种入侵的防治策略。本研究为我国海洋外来物种的进一步研究提供了参考。     

Protected areas offer refuge from invasive species spreading under climate change

Protected areas (PAs) are intended to provide native biodiversity and habitats with a refuge against the impacts of global change, particularly acting as natural filters against biological invasions. In practice, however, it is unknown how effective PAs will be in shielding native species from invasions under projected climate change. Here, we investigate the current and future potential distributions of 100 of the most invasive terrestrial, freshwater, and marine species in Europe. We use this information to evaluate the combined threat posed by climate change and invasions to existing PAs and the most susceptible species they shelter. We found that only a quarter of Europe's marine and terrestrial areas protected over the last 100 years have been colonized by any of the invaders investigated, despite offering climatically suitable conditions for invasion. In addition, hotspots of invasive species and the most susceptible native species to their establishment do not match at large continental scales. Furthermore, the predicted richness of invaders is 11%–18% significantly lower inside PAs than outside them. Invasive species are rare in long‐established national parks and nature reserves, which are actively protected and often located in remote and pristine regions with very low human density. In contrast, the richness of invasive species is high in the more recently designated Natura 2000 sites, which are subject to high human accessibility. This situation may change in the future, since our models anticipate important shifts in species ranges toward the north and east of Europe at unprecedented rates of 14–55 km/decade, depending on taxonomic group and scenario. This may seriously compromise the conservation of biodiversity and ecosystem services. This study is the first comprehensive assessment of the resistance that PAs provide against biological invasions and climate change on a continental scale and illustrates their strategic value in safeguarding native biodiversity.     

Constraints and release at different scales – The role of adaptation in biological invasions

Attempts to find a consensus on traits promoting the invasiveness of exotic species have agreed on the idiosyncrasy of successful invasions. Despite considerable efforts to integrate aspects of context-dependency into theories of invasions, none of them has provided an evolutionary perspective taking consistently into account the direction of environmental changes in terms of ‘constraint’ vs. ‘release’. Applying the filter theory of species sorting, I consider different filters at different scales explaining evolutionary changes during invasions. Within this hierarchical approach, the focus is on the factorial filters climate, abiotic environment and biotic environment, distinguishing trophic interactions and plant-plant interactions. This review summarizes the evidence of adaptive shifts from native to exotic ranges, thereby differentiating the direction of shifts with regard to either constrained or released situations. Following this systematic approach, the present paper identifies further trade-offs within hierarchical levels complementing already existing hypotheses such as those for biotic interactions. In particular, the role of climatic changes should more explicitly be linked with evolutionary responses during invasions. Studying exotic species successfully invading several regions with different environmental conditions will be a promising starting point to enlarge the understanding of context-dependency of invasions.