Casuarina: biogeography and ecology of an important tree genus in a changing world

Important insights on the invasion ecology of woody plants are emerging from the study of model groups, but it is important to test how widely such results can be generalised. We examined whether drivers of introduction and invasion in the genus Casuarina are similar to those identified for other groups. We reviewed the literature, mapped current global distributions, and modelled the potential global distribution of the genus. We assessed the rationale behind introductions, impacts of invasions, and the evolution of management approaches. Casuarinas have been introduced to about 150 countries. Ten of the 14 species in the genus have been introduced outside their native ranges, but only three species are recorded as naturalized or invasive (C. equisetifolia, C. cunninghamiana and C. glauca). Species with large native ranges are more likely to be introduced, and the three invasive species have the largest native ranges and the most records of introduction. There were no clear relationships between life-history traits (e.g. seed mass, plant height, or resprouting ability) and the extent of invasion. About 8 % of the Earth’s land surface is potentially suitable for casuarinas and large-scale plantings in some climatically suitable areas have not yet resulted in large-scale invasions; there is a substantial global Casuarina invasion debt. Experiences in Florida and the Mascarene Islands highlight that casuarinas have the potential to transform ecosystems with significant control costs. The challenge is to develop management approaches that minimise the impacts of invasions while preserving economic, environmental and cultural values of species in their introduced ranges.     

A standardized set of metrics to assess and monitor tree invasions

Scientists, managers, and policy-makers need functional and effective metrics to improve our understanding and management of biological invasions. Such metrics would help to assess progress towards management goals, increase compatibility across administrative borders, and facilitate comparisons between invasions. Here we outline key characteristics of tree invasions (status, abundance, spatial extent, and impact), discuss how each of these characteristics changes with time, and examine potential metrics to describe and monitor them. We recommend quantifying tree invasions using six metrics: (a) current status in the region; (b) potential status; (c) the number of foci requiring management; (d) area of occupancy (AOO) (i.e. compressed canopy area or net infestation); (e) extent of occurrence (EOO) (i.e. range size or gross infestation); and (f) observations of current and potential impact. We discuss how each metric can be parameterised (e.g. we include a practical method for classifying the current stage of invasion for trees following Blackburn’s unified framework for biological invasions); their potential management value (e.g. EOO provides an indication of the area over which management is needed); and how they can be used in concert (e.g. combining AOO and EOO can provide insights into invasion dynamics; and we use potential status and threat together to develop a simple risk analysis tool). Based on these metrics, we propose a standardized template for reporting tree invasions that we hope will facilitate cross-species and inter-regional comparisons. While we feel this represents a valuable step towards standardized reporting, there is an urgent need to develop more consistent metrics for impact and threat, and for many specific purposes additional metrics are still needed (e.g. detectability is required to assess the feasibility of eradication).     

Propagule pressure and native species richness effects drive invasibility in tropical dry forest seedling layers

Understanding the factors that encourage or inhibit plant invasions is vital to focusing limited invasive control efforts within areas where they are most practical and cost-effective. To extend the range of contexts in which invasibility is studied and aid the development of practical strategies to limit damaging plant invasions, we set out to test the relative importance of native species richness, native seedling density, and invasive propagule pressure, on the invasibility of artificial assemblages of naturally occurring tropical woody seedling communities. Our greenhouse mesocosms included a species pool of twelve trees and woody shrubs native to South Florida's tropical hardwood hammocks, and an increasingly prevalent noxious woody invader of this system, Ardisia elliptica. We found that invader propagule pressure was the single most important factor determining community invasibility. We also revealed a positive relationship between invasibility and native species richness in our polyculture mesocosms. Because A. elliptica biomass production significantly differed among different native monocultures and was not related to overyielding in native polycultures, we suggest that the effect of species richness on invasibility in this experiment was the result of sampling effects rather than a true effect of diversity.Three broad findings hold potential for application in preventing and controlling plant invasions, especially in the seedling layers of tropical dry forests: (1) effective invasive control efforts will likely benefit from measures to minimize propagule pressure; (2) managers might do well to prioritize invasive monitoring and removal efforts on the most diverse habitats within a management region; and (3) while more data are necessary to further understand our finding of a lack of association between productivity and invasibility, management regimes aimed at maximizing primary productivity might not increase invasibility, and in fact, strategies for controlling invasive plants via the management of ecosystem productivity may be ineffective.     

Biological invasions in rapidly urbanizing areas: a case study of Beijing, China

Urbanization is widely recognized as a major factor promoting biological invasions worldwide. In this article, we provide insights into the patterns of biological invasions in Beijing, one of the largest and quickly urbanizing cities of the world, by developing a comprehensive list of naturalized and invasive flora and their associate traits (e.g., distribution, life form, habitat, or geographic origin). One hundred and twelve naturalized (including 48 invasive) plants have been identified within the Beijing Municipality. Most of the naturalized and invasive plants belong to four families (Asteraceae, Poaceae, Amaranthaceae, and Euphorbiaceae) and are annual herbs that preferentially grow in disturbed sites. North and South America are the main contributors to the naturalized and invasive flora of Beijing. As expected, those Beijing districts that have recently experienced the highest human population growth, urban expansion, and the largest economic growth are also those with the highest number of naturalized and invasive species. Urban expansion is predicted to continue in the near term making additional invasions likely that will significantly increase the proportion of introduced species in Beijing’s flora. An integrated management strategy for the whole municipality is urgently needed that includes comprehensive scientific research that documents the extent of invasions and their effects on Beijing’s economy and environment.     

Pine invasions in the southern hemisphere: modelling interactions between organism,environment and disturbance

Current theories of plant invasion have been criticized for their limited heuristic and predictive value. We explore the heuristic and predictive potential of a model which explicitly simulates the mechanisms of plant invasion. The model, a spatially-explicit individual-based simulation, is applied to the invasion of pine trees (Pinus spp.; Pinaceae) in three vegetation types in the southern hemisphere. The model simulates factors which have been invoked as major determinants of invasive success: plant traits, environmental features and disturbance level. Results show that interactions between these determinants of invasive success are at least as important as the main effects. The complexity of invasions has promoted the belief that many factors must be invoked to explain invasions. This study shows that by incorporating interactions and mechanisms into our models we can potentially reduce the number of factors needed to predict plant invasions. The importance of interactions, however, means that predictions about invasions must be context-specific. The search for all-encompassing rules for invasions is therefore futile. The model presented here is of heuristic value since it improves our understanding of invasions, and of management value since it defines the data and models needed for predicting invasions.     

Genetic tools in the management of invasive mammals: recent trends and future perspectives

1. Invasive non-native species are now considered to be one of the greatest threats to biodiversity worldwide. Therefore, efficient and cost-effective management of species invasions requires robust knowledge of their demography, ecology and impacts, and genetic-based techniques are becoming more widely adopted in acquiring such knowledge.
2. We focus on the use of genetic tools in the applied management of mammalian invasions globally, as well as on their inherent advantages and disadvantages. We cover tools that are used in: 1) detecting and monitoring mammalian invaders; 2) identifying origins and invasive pathways; 3) assessing and quantifying the negative impacts of invaders; and 4) population management and potential eradication of invasive mammals.
3. We highlight changes in sequencing technologies, including how the use of techniques such as Sanger sequencing and microsatellite genotyping, for monitoring and tracing invasive pathways respectively, are now giving way to the use of high-throughput sequencing methods. These include the emergence of environmental DNA (eDNA) metabarcoding for the early detection of invasive mammals, and single nucleotide polymorphisms or whole genomes to trace the sources of invasive populations. We are now moving towards trials of genome-editing techniques and gene drives to control or eradicate invasive rodents.
4. Genetic tools can provide vital information that may not be accessible with non-genetic methods, for the implementation of conservation policies (e.g. early detection using systematic eDNA surveillance, the identification of novel pathogens). However, the lack of clear communication of novel genetic methods and results (including transparency and reproducibility) to relevant stakeholders can be prohibitive in translating these findings to appropriate management actions. Geneticists should engage early with stakeholders to co-design experiments in relation to management goals for invasive mammals.

     

动物入侵的行为机制

行为特征可在外来动物建立种群和扩张过程中发挥重要作用,因此,要正确理解动物入侵,常常需要仔细研究其行为机制。20世纪80年代以来,随着动物入侵规模在世界各地的迅速加剧,有关其行为机制的研究也受到了广泛关注。最近一些研究表明,一些入侵动物种内攻击和觅食等行为具有可塑性,因此它们能够灵活应对多变的环境条件,这对于种群的建立和维持至关重要;入侵动物与土著物种发生行为互作时,往往占据优势,从而取代土著物种,并有助于其地域扩张;入侵动物长距离扩散可以提高其地域扩张速度,许多行为可与扩散行为结合进一步促进扩张。今后需要加强对入侵动物的行为分析,使之全面地融合到生物入侵的研究之中。这不仅可以提高对外来物种入侵的预警和治理能力,而且为探索动物行为的奥秘以及动物间行为互作在物种进化中的意义提供了独特的机会。     

Invasions by ladybugs,ladybirds, and other predatory beetles

Species of predatory Coleoptera have become abundant in new geographic regions recently, raising concerns for invaded ecosystems. We address this topic by focusing on invasive alien ladybird beetles (Coccinellidae; known also as ladybugs). Humans appear directly or indirectly responsible for all or most ladybird invasions. Factors hypothesized to have promoted ladybird invasions include genetic diversity (e.g., for polymorphism), phenotypic plasticity, adaptation and genetic shift, generalized diet and habitat preferences, flexible life history and reproduction, large body size, and release from enemies. Factors such as climate, habitat and prey availability, and biotic resistance may sometimes prevent or slow ladybird invasions. Indigenous species (e.g., herbivores) may suffer from invasions, and biological control programs may be affected. Species of indigenous ladybirds throughout the world are reported to have declined in abundance following ladybird invasions, with increased competition and/or intraguild predation most often hypothesized or inferred. Similar recent studies especially of ground beetles (Carabidae) also make clear the potential of invasive alien predatory Coleoptera to disrupt invaded natural and agricultural ecosystems.     

The risk of invasion by angiosperms peaks at intermediate levels of human influence

Biological invasions are a growing threat to biodiversity. The control and eradication of exotic species established in earnest are of limited success despite high financial investments. Anticipating biological invasions based on species’ suitabilities is a cost-effective strategy given it helps identifying areas where exotic species can prosper, which can then translate in improving management and conservation efforts. Based on information from 191 invasive angiosperm species worldwide, we used ecological niche models to identify areas at high risk of invasion (cumulative predicted distribution of invasive species) in Mexico. Further, we explored the importance of bioclimatic and human influence variables as drivers of the distribution of invasive species and analyzed the status of the currently recognized priority conservation sites in Mexico. We found that areas with intermediate human activity scores had a high risk of invasion. Additionally, we found that many of the current priority conservation sites in Mexico had a high risk of invasion. Our findings contribute to disentangling the factors that drive environment susceptibility to invasions and urge management strategies to minimize the impacts of biological invasions in priority conservation sites.     

海洋动物档案式标志及其定位方法研究进展

档案式标志是一种可存储数据档案的电子标志,应用于水生动物尤其是大洋高度洄游性鱼类的研究,是获取海洋动物长时间、大空间尺度的活动信息和环境数据的有力工具.自20世纪90年代以来,档案式标志被大量用于海洋动物研究,取得了一系列成果,其不足之处在于需要回收标志来获取采样数据.90年代末,分离式卫星档案标志的问世解决了数据回收的问题,不再依赖于渔业捕捞,扩展了海洋动物研究的广度和深度.基于光亮度定位是研究海洋动物洄游路线的关键,其重点在于匹配光亮度与特定天顶角(如日出、日落),有3种方法：固定参照法、可变参照法和映射法.在过去的20年内,光亮度定位方法有所发展,并取得了一定的成果,但纬度向的定位还不够精确,有继续发展的空间.本文还总结了档案式标志定位的现存问题,展望了档案式标志及其定位方法的研究方向,并认为存在档案式标志小型化和传感器集成化、定位方法由单一光亮度反演向多因子反演的趋势.     

Naturalization of introduced plants: ecological drivers of biogeographical patterns

CONTENTS: Summary 383 I. Introduction 383 II. The introduction-naturalization-invasion continuum for conceptualizing biological invasions 384 III. The biogeographical background for studying naturalization: variation among populations and regions 385 IV. Factors determining naturalization in plants 388 Acknowledgements 392 References 392 SUMMARY: The literature on biological invasions is biased in favour of invasive species - those that spread and often reach high abundance following introduction by humans. It is, however, also important to understand previous stages in the introduction-naturalization-invasion continuum ('the continuum'), especially the factors that mediate naturalization. The emphasis on invasiveness is partly because most invasions are only recognized once species occupy large adventive ranges or start to spread. Also, many studies lump all alien species, and fail to separate introduced, naturalized and invasive populations and species. These biases impede our ability to elucidate the full suite of drivers of invasion and to predict invasion dynamics, because different factors mediate progression along different sections of the continuum. A better understanding of the determinants of naturalization is important because all naturalized species are potential invaders. Processes leading to naturalization act differently in different regions and global biogeographical patterns of plant invasions result from the interaction of population-biological, macroecological and human-induced factors. We explore what is known about how determinants of naturalization in plants interact at various scales, and how their importance varies along the continuum. Research that is explicitly linked to particular stages of the continuum can generate new information that is appropriate for improving the management of biological invasions if, for example, potentially invasive species are identified before they exert an impact.     

Invisible invaders: non-pathogenic invasive microbes in aquatic and terrestrial ecosystems

Although the number of studies on invasive plants and animals has risen exponentially, little is known about invasive microbes, especially non-pathogenic ones. Microbial invasions by viruses, bacteria, fungi and protists occur worldwide but are much harder to detect than invasions by macroorganisms. Invasive microbes have the potential to significantly alter community structure and ecosystem functioning in diverse terrestrial and aquatic ecosystems. Consequently, increased attention is needed on non-pathogenic invasive microbes, both free-living and symbiotic, and their impacts on communities and ecosystems. Major unknowns include the characteristics that make microbes invasive and properties of the resident communities and the environment that facilitate invasions. A comparison of microbial invasions with invasions of macroorganisms should provide valuable insights into general principles that apply to invasions across all domains of life and to taxon-specific invasion patterns. Invasive microbes appear to possess traits thought to be common in many invasive macroorganisms: high growth rate and resource utilization efficiency, and superior competitive abilities. Invading microorganisms are often similar to native species, but with enhanced performance traits, and tend to spread in lower diversity communities. Global change can exacerbate microbial invasions; therefore, they will likely increase in the future.     

Modeling the spread of invasive species using dynamic network models

Spread dynamics of biological invasions are influenced by both the availability and spatial arrangement of suitable habitat. As such, invasive spread can be considered to occur across a network of nodes, representing patches of suitable habitat, with linkages representing the potential for movement between habitat patches. While static network models can provide valuable insight into the potential framework of nodes and linkages across which spread could occur, they offer little information on the actual spatiotemporal dynamics of range expansion processes. Here, we explore the development and application of dynamic network models (DNMs) to model the spread of invasive species. DNMs accommodate temporal dynamics in the utilization of nodes and the connections between them and can flexibly perform simulations at the spatial scales of observational data. As case studies, we develop DNMs to simulate the spread of a generalist forest pathogen and the hemlock woolly adelgid (Adelges tsugae Annand). We highlight the utility of DNMs for identifying habitat patches that contribute most to spread across the landscape and for visualizing emergent spread dynamics. While currently underutilized in ecology as compared to static network models, DNMs are potentially applicable to numerous research and management questions relevant to biological invasions and the more general phenomena of range expansion.     

中国农业生态系统外来种入侵及其管理现状

农业生态系统极易遭受外来生物入侵。作者根据文献资料和多年工作观察统计出入侵我国农业生态系统的外来生物共计92科175属239种, 其中植物155种, 动物55种, 微生物29种, 植物多为有意引入后逸生, 而动物和微生物则主要是无意引入。外来入侵种发生数量呈现从南到北、从东到西逐渐减少的趋势。这些入侵种中, 来源于美洲的最多(占45.04%), 其次是欧洲(22.90%); 菜地(包括温室大棚)和果园入侵种最多, 分别达64.85%和66.53%, 而半年期的秋熟旱地和夏熟旱地分别占34.31%和23.85%。其中17种外来杂草、10种害虫、7种病原菌为恶性有害生物, 应作为防除的重点目标。目前农业生态系统外来入侵物种的控制以化学防治为主, 但由于长期施用化学农药, 在侵入我国农田的入侵种中, 已有51种在世界不同地区演化出抗药性生物型, 因而需重视生物防治、农业和生态防治以及检疫等的综合应用。今后外来种对农业生态系统的入侵格局、机制和趋势, 入侵途径以及生物入侵和抗药性生物型对农业生态系统中有害生物群落演替的影响、转基因作物导致的生物入侵等问题值得关注。     

Cross-scale management strategies for optimal control of trees invading from source plantations

Biological invasion by non-native tree species can transform landscapes, and as a consequence, has received growing attention from researchers and managers alike. This problem is driven primarily by the naturalisation and invasion of tree species escaping from cultivation or forestry plantations. Furthermore, these invasions can be strongly influenced by the land-use matrix of the surrounding region, specific management of the source populations, and environmental conditions that influence seed dispersal or habitat quality for the invader. A major unresolved challenge for managing tree invasions in landscapes is how management should be deployed to contain or slow the spread of invading populations from one or more sources (e.g. plantations). We develop a spatial simulation model to test: (1) how to best prioritise the control of invasive tree populations spatially to slow or contain the biological invader when habitat quality varies in the landscape, and (2) how to allocate control effort among different management units when trees spread from many source populations. We first show that to slow down spread effectively, management strategy is less important than management effort. We then identify the conditions affecting the relative performance of different management strategies. At the landscape scale, targeting peripheral stands consistently yielded the best results whereas at the regional scale, management strategies needed to account for both habitat quality and tree life-history. Overall, our findings demonstrate that knowledge of how habitat affects tree life-history stages can improve management to contain or slow tree invasions by improving the spatial match between management effort and efficacy.     

Trophic cascades in an invaded ecosystem: native keystone predators facilitate a dominant invader in an estuarine community

Determining how various factors contribute to the invasibility of systems is essential for both understanding community formation and informing management of natural areas. Research demonstrating that predators can provide biotic resistance to invasions by consuming invasive species has led to the presence of healthy predator populations being associated with reduced invasion potential of ecosystems. However, predators structure communities in many ways and their presence could also potentially facilitate invasions if they decrease populations of native species that compete with or consume an invader. We considered these two impacts of predators on invasion by analyzing the effects of two keystone predators (Pisaster spp. and Enhydra lutris nereis) on two foundation species (a native mussel Mytilus californianus and the invasive exotic bryozoan Watersipora subtorquata, a putative competitor for space with Mytilus californianus). Both native predators were found to facilitate the invasion of the exotic bryozoan, and the rate of invasion was highest when both predators were present. Facilitation of W. subtorquata occurred via indirect mechanisms that partly involved the removal of a competitor (mussels) via predation. These results illustrate that although predators can provide biotic resistance to invasion, healthy predator populations do not always confer this advantage and in fact may facilitate invasions. Therefore, implementation of management actions to enhance populations of top predators could also potentially increase the invasibility of some ecosystems.     

Plant invasions in treeless vegetation of the Australian Alps

A total of 128 invasive plant species have been recorded in treeless vegetation in the Australian Alps. Most of these are forbs and most are uncommon. Cover of invasive species is generally minimal unless there has been gross disturbance to natural vegetation and soils. Although there is a significantly positive correlation between invasive and native species diversity, suggesting that conditions that allow numerous native species to co-exist also permit more plant invasions, altitude is the most important determinant of invasive species diversity. Only 22 of the 128 species have been recorded above 1800 m. Some plant communities (e.g. those with high pH or relatively nutrient-rich soils), however, seem to be vulnerable to invasions regardless of altitude. Most invasive species appear to have been introduced unintentionally (e.g. as seed attached to vehicles, animals and humans) but a few were introduced to assist with revegetation of disturbed soils and for amenity plantings in ski resorts, and have subsequently established in native vegetation. Treeless communities in the Australian Alps are likely to face increasing pressure from invasive species as a result of global warming and continued introduction of non-native plants to ski resort gardens. Whilst it may be difficult to prevent invasive species of low elevations migrating to higher elevations as temperatures rise, the risk of invasion from garden plants could be minimised through regulation. Non-native plants in ski resort gardens pose a far greater risk than most invasive species currently present in the Alps because they have been chosen for their capacity to survive at high altitudes.     

Environment and dispersal paths override life strategies and residence time in determining regional patterns of invasion by alien plants

We describe a novel dissimilarity framework to analyze spatial patterns of species diversity and illustrate it with alien plant invasions in Northern Portugal. We used this framework to test the hypothesis that patterns of alien invasive plant species richness and composition are differently affected by differences in climate, land use and landscape connectivity (i.e. Geographic distance as a proxy and vectorial objects that facilitate dispersal such as roads and rivers) between pairs of localities at the regional scale. We further evaluated possible effects of plant life strategies (Grime's C-S-R) and residence time. Each locality consisted of a 1 km² landscape mosaic in which all alien invasive species were recorded by visiting all habitat types.Multi-model inference revealed that dissimilarity in species richness is more influenced by environmental distance (particularly climate), whereas geographic distance (proxies for dispersal limitations) is more important to explain dissimilarity in species composition, with a prevailing role for ecotones and roads. However, only minor differences were found in the responses of the three C-S-R strategies. Some effect of residence time was found, but only for dissimilarity in species richness. Our results also indicated that environmental conditions (e.g. climate conditions) limit the number of alien species invading a given site, but that the presence of dispersal corridors determines the paths of invasion and therefore the pool of species reaching each site. As geographic distances (e.g. ecotones and roads) tend to explain invasion at our regional scale highlights the need to consider the management of alien invasions in the context of integrated landscape planning. Alien species management should include (but not be limited to) the mitigation of dispersal pathways along linear infrastructures. Our results therefore highlight potentially useful applications of the novel multimodel framework to the anticipation and management of plant invasions.     

The elephant in the room: the role of failed invasions in understanding invasion biology

Most species introductions are not expected to result in invasion, and species that are invasive in one area are frequently not invasive in others. However, cases of introduced organisms that failed to invade are reported in many instances as anecdotes or are simply ignored. In this analysis, we aimed to find common characteristics between non‐invasive populations of known invasive species and evaluated how the study of failed invasions can contribute to research on biological invasions. We found intraspecific variation in invasion success and several recurring explanations for why non‐native species fail to invade; these included low propagule pressure, abiotic resistance, biotic resistance, genetic constraints and mutualist release. Furthermore, we identified key research topics where ignoring failed invasions could produce misleading results; these include studies on historical factors associated with invasions, distribution models of invasive species, the effect of species traits on invasiveness, genetic effects, biotic resistance and habitat invasibility. In conclusion, we found failed invasions can provide fundamental information on the relative importance of factors determining invasions and might be a key component of several research topics. Therefore, our analysis suggests that more specific and detailed studies on invasion failures are necessary.     

Granivory in California sage scrub: implications for common plant invaders and ecosystem conservation

Seed predation may influence community assembly and invasion dynamics when seed predators preferentially select some seed species over others. However, the role of different seed predators and their preferences for seeds in multiple ecological contexts, including the endangered California sage scrub ecosystem, remain unresolved making predictions about which processes limit or promote invasions difficult. In addition, selection criteria, specifically how seed size and biogeographic origin (native versus invasive) influences selection, requires further research across multiple granivore guilds. We examined seed predator preference for common native and invasive seeds across small and large seed classes using seed dish experiments with motion-sensor video observation. We also quantified the relative importance of ants, birds, and mammals as seed predators in sage scrub. Community-wide, granivores preferred the small-seeded invasive mustard Brassica nigra, avoided the large-seeded invasive grass Bromus rubens and the native large-seeded shrub Encelia californica, and did not show significant selection or avoidance for the small-seeded native shrub Salvia apiana. Birds, notably California towhees (Melozone crissalis), were the most frequent granivore while rodents and ants rarely removed seeds. This study reveals that birds are important seed predators in California sage scrub and have the potential to contribute to biotic resistance to mustard invasions (Brassica nigra and potentially others).