Diversity and distribution of genetic variation in gammarids: Comparing patterns between invasive and non‐invasive species

Biological invasions are worldwide phenomena that have reached alarming levels among aquatic species. There are key challenges to understand the factors behind invasion propensity of non‐native populations in invasion biology. Interestingly, interpretations cannot be expanded to higher taxonomic levels due to the fact that in the same genus, there are species that are notorious invaders and those that never spread outside their native range. Such variation in invasion propensity offers the possibility to explore, at fine‐scale taxonomic level, the existence of specific characteristics that might predict the variability in invasion success. In this work, we explored this possibility from a molecular perspective. The objective was to provide a better understanding of the genetic diversity distribution in the native range of species that exhibit contrasting invasive propensities. For this purpose, we used a total of 784 sequences of the cytochrome c oxidase subunit I of mitochondrial DNA (mtDNA‐COI) collected from seven Gammaroidea, a superfamily of Amphipoda that includes species that are both successful invaders (Gammarus tigrinus, Pontogammarus maeoticus, and Obesogammarus crassus) and strictly restricted to their native regions (Gammarus locusta, Gammarus salinus, Gammarus zaddachi, and Gammarus oceanicus). Despite that genetic diversity did not differ between invasive and non‐invasive species, we observed that populations of non‐invasive species showed a higher degree of genetic differentiation. Furthermore, we found that both geographic and evolutionary distances might explain genetic differentiation in both non‐native and native ranges. This suggests that the lack of population genetic structure may facilitate the distribution of mutations that despite arising in the native range may be beneficial in invasive ranges. The fact that evolutionary distances explained genetic differentiation more often than geographic distances points toward that deep lineage divergence holds an important role in the distribution of neutral genetic diversity.     

Impact assessment revisited: improving the theoretical basis for management of invasive alien species

The theoretical underpinnings of the assessment of invasive alien species impacts need to be improved. At present most approaches are unreliable to quantify impact at regional scales and do not allow for comparison of different invasive species. There are four basic problems that need to be addressed: (1) Some impacted ecosystem traits are spatially not additive; (2) invader effects may increase non-linearly with abundance or there may be effect thresholds impairing estimates of linear impact models; (3) the abundance and impact of alien species will often co-vary with environmental variation; and (4) the total invaded range is an inappropriate measure for quantifying regional impact because the habitat area available for invasion can vary markedly among invasive species. Mathematical models and empirical data using an invasive alien plant species (Heracleum mantegazzianum) indicate that ignoring these issues leads to impact estimates almost an order of magnitude from the real values. Thus, we propose a habitat-sensitive formula for regional impact assessment that is unaffected by non-linearity. Furthermore, we make some statistical suggestions on how to assess invader effects properly and we discuss the quantification of the invaded range. These improvements are crucial for impact assessment with the overall aim of prioritizing management of invasive species.     

A Panax-centric view of invasive species

Plant-centric sampling provides a novel approach to quantifying the potential impact of invasive species on native plant species. The aim of this study was to determine the level of exposure of individuals and populations of Panax quinquefolius to invasive plant species using this approach in thirty natural ginseng populations. A high level of invasion was found with 63–70% of ginseng populations containing at least one invasive species. Approximately one-third of all individuals were found in close proximity to invasive plants. The most prevalent invasive species were Rosa multiflora and Berberis thunbergii. The exposure to invasives of plants in different size classes varied among populations. Invasive species presence increased with greater ginseng population sizes and presence of harvest. The abundance of invasives plants within forest interiors near this valuable medicinal herb suggests that the economic and ecological costs of competitive interactions with native species could be high.     

Invasion by Solidago species has limited impacts on soil seed bank communities

Increasing attention in invasion biology is being paid to measuring and understanding the impacts of invasive species. For plant invasions, however, the impact of invasion on soil seed bank communities has been under-studied. At six sites in southern Germany, we investigated whether areas invaded by Solidago gigantea and Solidago canadensis experienced a reduction in seed bank species richness, size and diversity, and a change in species composition compared to adjacent uninvaded areas. We found no overall effect of invasion on seed bank size, or on species richness and diversity. Seed bank size significantly decreased from 0–5 cm to 5–10 cm depth in both invaded and uninvaded areas. A significant amount of variation in species composition was explained by invasion, but it was only one-tenth of that explained solely by site effects. Our study suggests that invasion by Solidago species may not have the same impacts on the soil seed banks of native species as other invasive perennial forbs that have so far been studied.     

Impact: Toward a Framework for Understanding the Ecological Effects of Invaders

Although ecologists commonly talk about the impacts of nonindigenous species, little formal attention has been given to defining what we mean by impact, or connecting ecological theory with particular measures of impact. The resulting lack of generalizations regarding invasion impacts is more than an academic problem; we need to be able to distinguish invaders with minor effects from those with large effects in order to prioritize management efforts. This paper focuses on defining, evaluating, and comparing a variety of measures of impact drawn from empirical examples and theoretical reasoning. We begin by arguing that the total impact of an invader includes three fundamental dimensions: range, abundance, and the per-capita or per-biomass effect of the invader. Then we summarize previous approaches to measuring impact at different organizational levels, and suggest some new approaches. Reviewing mathematical models of impact, we argue that theoretical studies using community assembly models could act as a basis for better empirical studies and monitoring programs, as well as provide a clearer understanding of the relationship among different types of impact. We then discuss some of the particular challenges that come from the need to prioritize invasive species in a management or policy context. We end with recommendations about how the field of invasion biology might proceed in order to build a general framework for understanding and predicting impacts. In particular, we advocate studies designed to explore the correlations among different measures: Are the results of complex multivariate methods adequately captured by simple composite metrics such as species richness? How well are impacts on native populations correlated with impacts on ecosystem functions? Are there useful bioindicators for invasion impacts? To what extent does the impact of an invasive species depend on the system in which it is measured? Three approaches would provide new insights in this line of inquiry: (1) studies that measure impacts at multiple scales and multiple levels of organization, (2) studies that synthesize currently available data on different response variables, and (3) models designed to guide empirical work and explore generalities.     

Species Invasiveness in Biological Invasions: A Modelling Approach

The study of invasiveness, the traits that enable a species to invade a habitat, and invasibility, the habitat characteristics that determine its susceptibility to the establishment and spread of an invasive species, provide a useful conceptual framework to formulate the biological invasion problem in a modelling context. Another important aspect is the complex interaction emerging among the invader species, the noninvader species already present in the habitat, and the habitat itself. Following a modelling approach to the biological invasion problem, we present a spatially explicit cellular automaton model (Interacting Multiple Cellular Automata (IMCA)). We use field parameters from the invader Gleditsia triacanthos and the native Lithraea ternifolia in montane forests of central Argentina as a case study to compare outputs and performance of different models. We use field parameters from another invader, Ligustrum lucidum, and the native Fagara coco from the same system to run the cellular automaton model. We compare model predictions with invasion values from aerial photographs. We discuss in detail the importance of factors affecting species invasiveness, and give some insights into habitat invasibility and the role of interactions between them. Finally, we discuss the relevance of mathematical modelling for studying and predicting biological invasions. The IMCA model provided a suitable context for integrating invasiveness, invasibility, and the interactions. In the invasion system studied, the presence of an invader's juvenile bank not only accelerated the rate of invasion but was essential to ensure invasion. Using the IMCA model, we were able to determine that not only adult survival but particularly longevity of the native species influenced the spread velocity of the invader, at least when a juvenile bank is present. Other factors determining velocity of invasion detected by the IMCA model were seed dispersal distance and age of reproductive maturity. We derived relationships between species' adult survival, fecundity and longevity of both theoretical and applied relevance for biological invasions. Invasion velocities calculated from the aerial photographs agreed well with predictions of the IMCA model.     

Predicting the impacts of an introduced species from its invasion history: an empirical approach applied to zebra mussel invasions

1. Quantitative models of impact are lacking for the vast majority of known invasive species, particularly in aquatic ecosystems. Consequently, managers lack predictive tools to help them prioritise invasion threats and decide where they can most effectively allocate limited resources. Predictive tools would also enhance the accuracy of water quality assessments, so that impacts caused by an invader are not erroneously attributed to other anthropogenic stressors. 2. The invasion history of a species is a valuable guide for predicting the consequences of its introduction into a new environment. Regression analysis of data from multiple invaded sites can generate empirical models of impact, as is shown here for the zebra mussel Dreissena polymorpha. Dreissena's impacts on benthic invertebrate abundance and diversity follow predictable patterns that are robust across a range of habitat types and geographic regions. Similar empirical models could be developed for other invaders with a documented invasion history. 3. Because an invader's impact is correlated with its abundance, a surrogate model may be generated (when impact data are unavailable) by relating the invader's abundance to environmental variables. Such a model could help anticipate which habitats will be most affected by invasion. Lack of precision should not be a deterrent to developing predictive models where none exist. Crude predictions can be refined as additional data become available. Empirical modelling is a highly informative and inexpensive, but underused, approach in the management of aquatic invasive species.     

Niche shift in invasive species: is it a case of “home away from home” or finding a “new home”?

In recent years, there has been a rather acrimonious debate on matters concerning the biology of invasive species, some as fundamental as the definition and what constitutes an invasive species. However, an abiding commonality of all invasive species is the fact that they have all moved away from their native ranges to newer and often non-native ranges. In plants, Lantana camara has shifted from its native South American range distribution to most other parts of the world. In animals, the African giant snail has dispersed from Africa to most parts of Asia. What do such niche shifts signify about the nature and quality of the habitats to which the invasive species have moved? In this paper, using the classical niche paradigm, we analyse if niche shifts of thirty-three of the world’s top invasive species constitute just moving from one habitat to another similar habitat somewhere on the earth (home away from home) or that they have moved to totally new habitats (different from their native home). Surprisingly, our results show that for 90% of the world’s top invasive species, movements have been largely restricted to homes away from home, rather than into alien homes. This clearly indicates the potential inertia that species might face in moving out of their fundamental niche. We discuss these results in the context of the overall debate on invasion biology and how niche conservatism may have played a role in dampening the rates of invasion.

     

玫瑰种群生物学研究进展

玫瑰(Rosa rugosa Thunb.)原产于我国东部沿海、日本、朝鲜半岛和俄罗斯远东地区,18世纪作为园艺种引入欧洲后逃逸并入侵至北海和波罗的海周边多个国家以及北美沙质海岸,而中国野生种群却在过去30年间持续萎缩,成为珍稀濒危物种。从玫瑰种群生物学角度,通过文献比较和综合,在阐明玫瑰生态学特性和野生分布变化的基础上,全面论述了玫瑰种群在我国的生境退化、种群动态、种子繁殖、遗传多样性、濒危机理、保育方面的成果和悬疑问题;并结合欧洲入侵种群分布范围和敏感生境、对本地群落和物种的影响、种子繁殖、遗传变异、种群扩张和模拟预测、管理和控制方面的研究进展,分析了濒危种群和入侵种群数量动态、群落组成、幼苗更新、遗传变异、管理策略方面的差异及其影响因素;进而提出未来的玫瑰研究可从濒危种群和入侵种群的比较研究、种群和灌丛的动态监测、适合度相关性状的变异及其遗传基础、基于种群生物学的保育或控制4个方面为切入点,集中探索玫瑰种群濒危和入侵动态的规律、遗传基础和主要驱动力,为玫瑰保育和管理提供理论依据,为相似物种的适应和进化机制研究提供例证。     

BIODIVERSITY RESEARCH: Genetic diversity in two introduced biofouling amphipods (Ampithoe valida & Jassa marmorata) along the Pacific North American coast: investigation into molecular identification and cryptic diversity

Aim We investigated patterns of genetic diversity among invasive populations of Ampithoe valida and Jassa marmorata from the Pacific North American coast to assess the accuracy of morphological identification and determine whether or not cryptic diversity and multiple introductions contribute to the contemporary distribution of these species in the region. Location Native range: Atlantic North American coast; Invaded range: Pacific North American coast. Methods We assessed indices of genetic diversity based on DNA sequence data from the mitochondrial cytochrome c oxidase subunit I (COI) gene, determined the distribution of COI haplotypes among populations in both the invasive and putative native ranges of A. valida and J. marmorata and reconstructed phylogenetic relationships among COI haplotypes using both maximum parsimony and Bayesian approaches. Results Phylogenetic inference indicates that inaccurate species‐level identifications by morphological criteria are common among Jassa specimens. In addition, our data reveal the presence of three well supported but previously unrecognized clades of A. valida among specimens in the north‐eastern Pacific. Different species of Jassa and different genetic lineages of Ampithoe exhibit striking disparity in geographic distribution across the region as well as substantial differences in genetic diversity indices. Main conclusions Molecular genetic methods greatly improve the accuracy and resolution of identifications for invasive benthic marine amphipods at the species level and below. Our data suggest that multiple cryptic introductions of Ampithoe have occurred in the north‐eastern Pacific and highlight uncertainty regarding the origin and invasion histories of both Jassa and Ampithoe species. Additional morphological and genetic analyses are necessary to clarify the taxonomy and native biogeography of both amphipod genera.     

Biological invasion risk assessment of Tuta absoluta: mechanistic versus correlative methods

The capacity to assess invasion risk from potential crop pests before invasion of new regions globally would be invaluable, but this requires the ability to predict accurately their potential geographic range and relative abundance in novel areas. This may be unachievable using de facto standard correlative methods as shown for the South American tomato pinworm Tuta absoluta, a serious insect pest of tomato native to South America. Its global invasive potential was not identified until after rapid invasion of Europe, followed by Africa and parts of Asia where it has become a major food security problem on solanaceous crops. Early prospective assessment of its potential range is possible using physiologically based demographic modeling that would have identified knowledge gaps in T. absoluta biology at low temperatures. Physiologically based demographic models (PBDMs) realistically capture the weather-driven biology in a mechanistic way allowing evaluation of invasive risk in novel areas and climes including climate change. PBDMs explain the biological bases for the geographic distribution, are generally applicable to species of any taxa, are not limited to terrestrial ecosystems, and hence can be extended to support ecological risk modeling in aquatic ecosystems. PBDMs address a lack of unified general methods for assessing and managing invasive species that has limited invasion biology from becoming a more predictive science.

     

The invasion and potential impact of the Asian House Gecko (Hemidactylus frenatus) in Australia

Hemidactylus frenatus is an Asian gecko that has spread pantropically to become one the world's most widespread reptiles. It has been established in Australia for approximately 50 years, but the last two decades have seen massive range expansion across settled areas of northern and eastern Australia; and this spread continues at pace. Disturbingly, H. frenatus is increasingly being detected in natural habitats in Australia, in some cases at high densities. Despite rampant spread, there has been little concern regarding the potential impact of this species on native geckos or natural systems more broadly. This is surprising given that Australia is a centre of gecko origin and diversity, and that H. frenatus has had well documented detrimental impacts on geckos in other parts of its introduced range. Here I review the biology and global distribution of H. frenatus, plot its spread in Australia over the five decades since establishment, and review the research on invasive populations of this species overseas and in Australia to assess potential impacts. I argue that Australia should be more concerned about H. Frenatus because: (i) it is spreading rapidly across northern, eastern and central Australia; (ii) it can invade natural habitats; (iii) it is a very strong competitor and may out‐compete Australian geckos in some situations; and (iv) it carries novel parasites that may impact native reptile species. Hemidactylus frenatus is here to stay and represents a potential threat to Australia's diversity and ecology. A key question is the degree to which it will invade natural habitats and what its impacts will be in these. Research is required to assess the current and potential impacts of H. frenatus in Australia so as to determine how these can be managed and the level of investment warranted.     

Niche conservatism and the potential for the crayfish Procambarus clarkii to invade South America

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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.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Invasion biology,ecology, and management of Frankliniella occidentalis in China

Frankliniella occidentalis is an economically important invasive pest worldwide, which can damage various horticultural crops and ornamental plants. F. occidentalis was first intercepted in Kunming, Yunnan province in 2000, and first reported to establish a population in Beijing, China in 2003. Since then, this pest is currently distributed across tens of provinces in mainland China and cause increasingly serious damage and loss. To control this pest, invasion biology, monitoring, and integrated pest management have been generally and intensively studied for 15 years in China. Furthermore, western flower thrips (WFT) as an important invasive insect pest, the research achievements on WFT has contributed to the promotion of technological innovation and development for invasive alien species management strategies and techniques in China. This review provides an overview for research on the biology, ecology, prevention, and management of this pest during 15 years in China. Meanwhile, China's “4E action” strategy on F. occidentalis is also discussed in this review.     

Ecological niche models of invasive seaweeds

Ecological niche models (ENMs) are commonly used to calculate habitat suitability from species’ occurrence and macroecological data. In invasive species biology, ENMs can be applied to anticipate whether invasive species are likely to establish in an area, to identify critical routes and arrival points, to build risk maps and to predict the extent of potential spread following an introduction. Most studies using ENMs focus on terrestrial organisms and applications in the marine realm are still relatively rare. Here, we review some common methods to build ENMs and their application in seaweed invasion biology. We summarize methods and concepts involved in the development of niche models, show examples of how they have been applied in studies on algae and discuss the application of ENMs in invasive algae research and to predict effects of climate change on seaweed distributions.     

Potential Distribution and Risk Assessment of an Invasive Plant Species: A Case Study of Hymenachne amplexicaulis in Australia

Given the limited resources available for weed management, a strategic approach is required to give the “best bang for your buck.” The current study incorporates: (1) a model ensemble approach to identify areas of uncertainty and commonality regarding a species invasive potential, (2) current distribution of the invaded species, and (3) connectivity of systems to identify target regions and focus efforts for more effective management. Uncertainty in the prediction of suitable habitat for H. amplexicaulis (study species) in Australia was addressed in an ensemble-forecasting approach to compare distributional scenarios from four models (CLIMATCH; CLIMEX; boosted regression trees [BRT]; maximum entropy [Maxent]). Models were built using subsets of occurrence and environmental data. Catchment risk was determined through incorporating habitat suitability, the current abundance and distribution of H. amplexicaulis, and catchment connectivity. Our results indicate geographic differences between predictions of different approaches. Despite these differences a number of catchments in northern, central, and southern Australia were identified as high risk of invasion or further spread by all models suggesting they should be given priority for the management of H. amplexicaulis. The study also highlighted the utility of ensemble approaches in indentifying areas of uncertainty and commonality regarding the species’ invasive potential.     

Identification and population genetic comparison of three ascidian species based on mtDNA sequences

Ascidians are sessile marine chordate invertebrates found along seashores worldwide and are typically regarded as invasive organisms. Knowledge concerning their global genetic structure and subsequent invasive potential is limited. Here, we identified three ascidians—Ciona robusta, Ciona savignyi, and Styela clava from the northeast region of China using morphological characteristics and mitochondrial cytochrome c oxidase subunit I (cox1) as genetic marker. We additionally used phylogenetics to aid in the identification of these three species. The results of a population genetic analysis showed that among the three species, the level of haplotype diversity was particularly high within C. savignyi, and nucleotide diversity varied moderately. We divided the three species separately into native and invasive populations using 170 cox1 sequences from global resources to explore population genetic structure and invasive potential. Although in the network analysis Ciona spp. formed haplogroups of native and invasive populations, some haplotypes were still shared. We found that the haplotypes did not cluster within the network of S. clava. Our AMOVA results also showed that Ciona spp. had a weak genetic structure, and less genetic differentiation was present in S. clava. These data suggest that there are extensive incursions of these three ascidians into different geographical regions. Global comparisons of ascidian populations will help in the understanding of their population genetic structure and invasive potential, hence providing important insights regarding conservation as well as management.     

Ecological consequences of invasion across the freshwater–marine transition in a warming world

The freshwater–marine transition that characterizes an estuarine system can provide multiple entry options for invading species, yet the relative importance of this gradient in determining the functional contribution of invading species has received little attention. The ecological consequences of species invasion are routinely evaluated within a freshwater versus marine context, even though many invasive species can inhabit a wide range of salinities. We investigate the functional consequences of different sizes of Corbicula fluminea—an invasive species able to adapt to a wide range of temperatures and salinity—across the freshwater–marine transition in the presence versus absence of warming. Specifically, we characterize how C. fluminea affect fluid and particle transport, important processes in mediating nutrient cycling (NH₄‐N, NO₃‐N, PO₄‐P). Results showed that sediment particle reworking (bioturbation) tends to be influenced by size and to a lesser extent, temperature and salinity; nutrient concentrations are influenced by different interactions between all variables (salinity, temperature, and size class). Our findings demonstrate the highly context‐dependent nature of the ecosystem consequences of invasion and highlight the potential for species to simultaneously occupy multiple components of an ecosystem. Recognizing of this aspect of invasibility is fundamental to management and conservation efforts, particularly as freshwater and marine systems tend to be compartmentalized rather than be treated as a contiguous unit. We conclude that more comprehensive appreciation of the distribution of invasive species across adjacent habitats and different seasons is urgently needed to allow the true extent of biological introductions, and their ecological consequences, to be fully realized.