Economics of invasive species policy and management

This article examines the use of economic analysis to inform bioinvasion management, with particular focus on forest resources. Economics is key for understanding invasion processes, impacts, and decision-making. Biological invasions are driven by and affect economic activities at multiple scales and stages of an invasion. Bioeconomic modeling seeks to inform how resources can be optimally allocated across invasion management activities—including prevention, surveillance programs for early detection and management, and controlling invasion populations and spread—to minimize the long-term costs and damages. Economic analysis facilitates understanding of decisions by public and private decision-makers, gaps between these, and the design of policies to achieve socially desirable outcomes. Private decision-makers may undercontrol invasions relative to socially optimal levels, because they generally account for their own costs and benefits of control but less often for broader ecosystem impacts or future spread across the landscape. Economic analysis considers approaches for increasing private invasion management and evaluates feedbacks between ecological and economic systems that can affect policy outcomes. Future research should continue evaluation and design of control strategies across the biosecurity continuum and across species to enhance cost-effectiveness, better incorporate uncertainty into policy design, increase focus on incentives and behavioral tools to influence private behaviors that affect invasion spread, and incorporate invasive species consideration within broader systems-focused science. In addition, challenges in valuing biodiversity and ecosystem service impacts and the costs and effectiveness of control measures are key data gaps. Greater collaboration between decision-makers and researchers will facilitate development and communication of usable economic research.     

Defining the twig fungal communities of Fraxinus species and Fraxinus excelsior genotypes with differences in susceptibility to ash dieback

Ash dieback disease (caused by Hymenoscyphus fraxineus) has affected European ash species (Fraxinus spp.) in recent decades. However, some Asian and American species of Fraxinus and certain genotypes of Fraxinus excelsior are less affected by the disease. We used ITS1-metabacoding to explore the drivers influencing diversity and composition of the twig fungal communities of Fraxinus species and F. excelsior genotypes. Our results revealed that fungi in the classes Eurotiomycetes and Dothideomycetes were among the most prevalent taxa in both Fraxinus species and F. excelsior genotypes. The diversity of the fungal communities differed significantly among Fraxinus species and could be explained by seed origin. Neither host genotype nor season had a significant effect on the community diversity of F. excelsior genotypes. On the other hand, the composition of twig fungal communities differed significantly among host species and among F. excelsior genotypes, and in F. excelsior there was also a significant effect of season on the composition of the fungal community. We did not find a clear effect of ash dieback susceptibility on either diversity or composition of fungal communities in twigs of Fraxinus species, although the effect was significant on the composition of fungal communities among F. excelsior genotypes. Our results demonstrated differences in fungal communities among species of Fraxinus and of F. excelsior genotypes, suggesting specific relationship between individual host genotypes and endophytic fungi.     

The invasive ash dieback pathogen Hymenoscyphus pseudoalbidus exerts maximal infection pressure prior to the onset of host leaf senescence

Shoot dieback disease of European ash caused by the ascomycete Hymenoscyphus pseudoalbidus threatens ash on a continental scale. A spore sampler placed in a diseased ash forest in Southern Norway, coupled with microscopy and DNA-based fungal species-specific real-time PCR assays, was employed to profile diurnal and within-season variation in infection pressure by ascospores of H. pseudoalbidus and the potentially co-existing non-pathogenic Hymenoscyphus albidus. Hymenoscyphus pseudoalbidus was found to be predominant in the stand. Massive simultaneous liberation, by active discharge of pathogen ascospores in the morning, peaked in mid-Jul. to mid-Aug. Accumulation of pathogen DNA on leaflets of current-year leaves reached a high level plateau phase before appearance of autumn coloration, suggesting that pathogen establishment in leaves is terminated before the onset of leaf senescence.     

Dynamic economic analysis on invasive species management: Some policy implications of catchability

The problem of controlling invasive species has emerged as a global issue. In response to invasive species threats, governments often propose eradication. This article challenges the eradication view by studying optimal strategies for controlling invasive species in a simple dynamic model. The analysis mainly focuses on deriving policy implications of catchability in a situation where a series of controlling actions incurs operational costs that derive from the fact that catchability depends on the current stock size of invasive species. We analytically demonstrate that the optimal policy changes drastically, depending on the sensitivity of catchability in response to a change in the stock size, as well as on the initial stock. If the sensitivity of catchability is sufficiently high, the constant escapement policy with some interior target level is optimal. In contrast, if the sensitivity of catchability is sufficiently low, there could exist a threshold of the initial stock which differentiates the optimal action between immediate eradication and giving-up without any control. In the intermediate range, immediate eradication, giving-up without any control, or more complex policies may be optimal. Numerical analysis is employed to present economic intuitions and insights in both analytically tractable and intractable cases.     

Altered stream-flow regimes and invasive plant species: the Tamarix case

Aim  To test the hypothesis that anthropogenic alteration of stream-flow regimes is a key driver of compositional shifts from native to introduced riparian plant species.
Location  The arid south-western United States; 24 river reaches in the Gila and Lower Colorado drainage basins of Arizona.
Methods  We compared the abundance of three dominant woody riparian taxa (native Populus fremontii and Salix gooddingii , and introduced Tamarix ) between river reaches that varied in stream-flow permanence (perennial vs. intermittent), presence or absence of an upstream flow-regulating dam, and presence or absence of municipal effluent as a stream water source.
Results  Populus and Salix were the dominant pioneer trees along the reaches with perennial flow and a natural flood regime. In contrast, Tamarix had high abundance (patch area and basal area) along reaches with intermittent stream flows (caused by natural and cultural factors), as well as those with dam-regulated flows.
Main conclusions  Stream-flow regimes are strong determinants of riparian vegetation structure, and hydrological alterations can drive dominance shifts to introduced species that have an adaptive suite of traits. Deep alluvial groundwater on intermittent rivers favours the deep-rooted, stress-adapted Tamarix over the shallower-rooted and more competitive Populus and Salix . On flow-regulated rivers, shifts in flood timing favour the reproductively opportunistic Tamarix over Populus and Salix , both of which have narrow germination windows . The prevailing hydrological conditions thus favour a new dominant pioneer species in the riparian corridors of the American Southwest. These results reaffirm the importance of reinstating stream-flow regimes (inclusive of groundwater flows) for re-establishing the native pioneer trees as the dominant forest type.     

Reproductive mode and life cycle of the ash dieback pathogen Hymenoscyphus pseudoalbidus

Ash dieback caused by the fungal pathogen Hymenoscyphus pseudoalbidus is currently ravaging in Europe, killing Fraxinus excelsior and Fraxinus angustifolia trees of all age classes. The aim of this work was to elucidate aspects of the reproduction biology of this fungal pathogen and its cryptic, non-pathogenic sister species Hymenoscyphus albidus. The mating type (MAT) locus of both species was identified, partly sequenced and characterized. Whereas a heterothallic MAT organization was detected in H. pseudoalbidus, H. albidus was shown to be structurally homothallic. The molecular MAT determination of H. pseudoalbidus was confirmed by crossing experiments on sterile ash petioles. Crossings of strains exhibiting alternate MAT idiomorphs produced fertile apothecia whereas crosses of strains with identical MAT idiomorphs were never successful. Offspring genotyping with microsatellites (MSs) and the MAT marker confirmed that both parental strains were involved in apothecia formation. In addition, polymorphic MS were shown to follow Mendelian inheritance. However, for yet unknown reasons the MAT ratio of progenies of one successful cross revealed a significant segregation distortion. Based on the MAT sequences of H. pseudoalbidus a multiplex PCR was developed, allowing for a quick and reliable MAT determination. The PCR was applied to screen the MAT ratio of two H. pseudoalbidus populations derived from the country of the disease outbreak in Poland and two populations from the disease periphery in Switzerland. None of the screened populations showed a significant deviation from the 1:1 ratio, expected under random mating. Therefore, an initial clonal distribution through asexually produced conidiospores as observed for other fungal pathogens holds not true for H. pseudoalbidus. Instead, our data is highly supportive for a distribution through ascospores. Leaf petioles collected in the field were thoroughly analyzed for the number of different colonizing strains and their mating behavior. Up to eight different H. pseudoalbidus genotypes were found on a single petiole. Cross-fertilizations of strains on the same petiole and fertilizations of unknown strains from outside were found, indicating that fertilization is mediated by spermatia. The presented study complements our understanding of the life cycle of this highly destructive pathogen. The possibility to perform sexual crosses in the lab provides ample opportunities for further genetic studies of H. pseudoalbidus and related species in the future.     

Vegetative incompatibility in the ash dieback pathogen Hymenoscyphus pseudoalbidus and its ecological implications

Pairings were carried out between isolates of the ash dieback pathogen Hymenoscyphus pseudoalbidus (Chalara fraxinea) to determine whether vegetative incompatibility (mycelial self–nonself recognition) reactions could be discriminated. On malt agar (MA) and ash sapwood agar (ASA) distinct compatible and incompatible reactions were observed. Compatible (C- or c-) reactions were characterised by full or partial colony intermingling along the junction line. Incompatible (G- or g-) reactions were characterised by a gap ca. 1–3 mm wide along the junction line. On MA a distinct narrow brown line or L-reaction was observed with some gap reactions, often comprising reticulate mycelium and conidia-producing phialides. When eleven isolates from a dieback site at Lower Wood, Norfolk were paired on ASA 53 of 54 reactions between different isolates gave incompatible reactions. A similar result was obtained with smaller samples from two further sites in Norfolk and Kent. This indicates that for local populations in the UK most genetic individuals of H. pseudoalbidus are likely to be vegetatively incompatible. The implications for the ecology and genetics of H. pseudoalbidus are discussed.     

Climate niche shift in invasive species: the case of the brown anole

Anolis sagrei, a Cuba and Bahama native lizard, is a successful invader in Florida and adjacent areas. Herein, we focus on conservatism in its climate niche axes and possible congruencies with its natural history properties. The not mutually exclusive hypotheses of the present study explaining its northern range limit are: (1) climatic conditions within species' native and invasive ranges are identical; (2) the species is pre‐adapted to novel conditions as a result of historical climate variations; and (3) only some niche axes limit the species' invasive distribution and the observed pattern is explained by an interplay between the potential niche within its native range and life‐history. Species distribution models for native and invasive distributions were built on ten bioclimatic variables. Using Schoener's niche overlap index, the degree of niche conservatism among variables was identified. Significances of hypothesis (1) were tested using null‐model approaches. Possible climatic pre‐adaptations were evaluated by comparing its actual tolerance within its invasive range with that of the Last Glacial Maximum (LGM) within its native range (hypothesis 2). Results of (1) and (2) are discussed in relation to natural history, approaching hypothesis 3. We detect varying overlaps in niche axes, indicating that natural history properties are associated with conservative niche axes. Climatic comparisons with LGM of native and current conditions of invasive range suggest that pre‐adaptations are unlikely. Possible shifts in the fundamental niche of the species may have been facilitated by enhanced genetic diversity in northern invasive populations. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 943–954.     

Progression of ash canopy thinning and dieback outward from the initial infestation of emerald ash borer (Coleoptera: Buprestidae) in southeastern Michigan

Our objective was to characterize the rate at which ash (Fraxinus spp.) trees decline in areas adjacent to the leading edge of visible ash canopy thinning due to emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). Trees in southeastern Michigan were surveyed from 2003 to 2006 for canopy thinning and dieback by comparing survey trees with a set of 11 standard photographs. Freeways stemming from Detroit in all directions were used as survey transects. Between 750 and 1,100 trees were surveyed each year. A rapid method of sampling populations of emerald ash borer was developed by counting emerald ash borer emergence holes with binoculars and then felling trees to validate binocular counts. Approximately 25% of the trees surveyed for canopy thinning in 2005 and 2006 also were sampled for emerald ash borer emergence holes using binoculars. Regression analysis indicates that 41-53% of the variation in ash canopy thinning can be explained by the number of emerald ash borer emergence holes per tree. Emerald ash borer emergence holes were found at every site where ash canopy thinning averaged > 40%. In 2003, ash canopy thinning averaged 40% at a distance of 19.3 km from the epicenter of the emerald ash borer infestation in Canton. By 2006, the point at which ash trees averaged 40% canopy thinning had increased to a distance of 51.2 km away from Canton. Therefore, the point at which ash trees averaged 40% canopy thinning, a state of decline clearly visible to the average person, moved outward at a rate of 10.6 km/yr during this period.     

Segmenting boaters based on level of transience: outreach and policy implications for the prevention of aquatic invasive species

Preventing the spread of aquatic invasive species (AIS) is a priority for many ecologists and natural resource managers. Because some boaters pose greater risk to spreading AIS, targeted outreach to them may be strategic. This study used data from a Wisconsin boater survey and examined whether boaters who differ in transience level, a proxy for risk, differed in terms of compliance with AIS prevention behaviors and other factors that could inform policy and targeted outreach. Segmenting based on transience yielded differences in terms of behavior, knowledge and communication habits. The results indicate that outreach may need to clarify behaviors related to live bait. Partnerships with lake associations and interpersonal communication may help improve compliance and clarify bait disposal behaviors, as it was found that highly transient boaters discuss AIS more with family and friends, and they hear more about AIS from lake associations. Differences in the three groups confirm their distinctness, suggesting that representatives from each category should be included in public management discussions when inclusivity is desired.     

Quantifying the recent expansion of native invasive rush species in a UK upland environment

Rushes, such as soft rush (Juncus effusus L.), hard rush (Juncus inflexus L.), and compact rush (Juncus conglomeratus L.) have become problem species within upland grasslands across the United Kingdom and the coastal grasslands of western Norway. Indeed, being largely unpalatable to livestock and having a vigorous reproductive ecology means that they can rapidly come to dominate swards. However, rush dominance results in a reduction in grassland biodiversity and farm productivity. Anecdotal evidence from the United Kingdom suggests that rush cover within marginal upland grasslands has increased considerably in recent decades. Yet, there is currently no published evidence to support this observation. Here, we use recent and historical Google Earth imagery to measure changes in rush frequency over a 13-year period within four survey years: 2005, 2009, 2015, and 2018. During each survey year, we quantified rush presence or absence using a series of quadrats located within 300 upland grassland plots in the West Pennine Moors, United Kingdom. Data were analysed in two stages, first, by calculating mean rush frequencies per sample year using all the available plot-year combinations (the full dataset), and second by examining differences in rush frequency using only the plots for which rush frequency data were available in every sample year (the continuous dataset). The full dataset indicated that rush frequency has increased by 82% between 2005 and 2018. Similarly, the continuous dataset suggested that rush frequency has increased by 174% over the same period, with the increases in frequency being statistically significant (p < .05) between 2005–2018 and 2009–2018. We discuss the potential drivers of rush expansion in the West Pennine Moors, the ecological and agronomic implications of grassland rush infestations, and priorities for future research.     

Fungal communities associated with species of Fraxinus tolerant to ash dieback,and their potential for biological control

Ash dieback, caused by the fungus Hymenoscyphus fraxineus, has threatened ash trees in Europe for more than two decades. However, little is known of how endophytic communities affect the pathogen, and no effective disease management tools are available. While European ash (Fraxinus excelsior) is severely affected by the disease, other more distantly related ash species do not seem to be affected. We hypothesise that fungal endophytic communities of tolerant ash species can protect the species against ash dieback, and that selected endophytes have potential as biocontrol agents. These hypotheses were tested by isolating members of the fungal communities of five tolerant ash species, and identifying them using ITS regions. Candidate endophytes were tested by an in vitro antagonistic assay with H.fraxineus. From a total of 196 isolates we identified 9 fungal orders, 15 families, and 40 species. Fungi in orders Pleosporales, such as Boeremia exigua and Diaporthe spp., and Hypocreales (e.g., Fusarium sp.), were recovered in most communities, suggesting they are common taxa. The in vitro antagonistic assay revealed five species with high antagonistic activity against H. fraxineus. These endophytes were identified based on ITS region as Sclerostagonospora sp., Setomelanomma holmii, Epicoccum nigrum, B. exigua and Fusarium sp. Three of these taxa have been described previously as antagonists of plant pathogenic microbes, and are of interest for future studies of their potential as biological control agents against ash dieback, especially for valuable ash trees in parks and urban areas.     

Density-dependent effects on the dieback of exotic species Spartina anglica in coastal China

The exotic species Spartina anglica C.E. Hubbard has been experiencing dieback in coastal China during the past decade, while it is invading other places in the world. Thus, an explanation for the dieback of the populations in coastal China would be helpful for controlling invasions of S. anglica in other places of the world. Both field and greenhouse experiments were conducted to determine the possible role of density in the dieback of the populations in China. The density effects were tested by maintaining six density levels in field experiments and three density levels in greenhouse experiments over 7 months. The results showed that growth, asexual reproduction and biomass accumulation in medium density were significantly higher than those in other density treatments for field experiments. The growth and asexual reproduction in high-density patches were lowest among the treatments in the field. Data showed the same pattern for the greenhouse experiments. The results implied that density regulation had a great effect on most of the studied parameters. Because the species is surviving at high density along the Chinese coast, and has a much lower growth rate due to strong intraspecific competition, and also because high-density populations may decline into low density populations due to resource exhaustion, high plant density might be one of the important factors in the dieback of S. anglica.     

Investigating the genetic load of an emblematic invasive species: the case of the invasive harlequin ladybird Harmonia axyridis

Introduction events can lead to admixture between genetically differentiated populations and bottlenecks in population size. These processes can alter the adaptive potential of invasive species by shaping genetic variation, but more importantly, they can also directly affect mean population fitness either increasing it or decreasing it. Which outcome is observed depends on the structure of the genetic load of the species. The ladybird Harmonia axyridis is a good example of invasive species where introduced populations have gone through admixture and bottleneck events. We used laboratory experiments to manipulate the relatedness among H. axyridis parental individuals to assess the possibility for heterosis or outbreeding depression in F₁ generation offspring for two traits related to fitness (lifetime performance and generation time). We found that inter‐populations crosses had no major impact on the lifetime performance of the offspring produced by individuals from either native or invasive populations. Significant outbreeding depression was observed only for crosses between native populations for generation time. The absence of observed heterosis is indicative of a low occurrence of fixed deleterious mutations within both the native and invasive populations of H. axyridis. The observed deterioration of fitness in native inter‐population crosses most likely results from genetic incompatibilities between native genomic backgrounds. We discuss the implications of these results for the structure of genetic load in H. axyridis in the light of the available information regarding the introduction history of this species.     

Physiological and genomic characterisation of Luteimonas fraxinea sp. nov., a bacterial species associated with trees tolerant to ash dieback

A group of isolates of the genus Luteimonas was characterised, which represented a specific component of the healthy core microbiome of Fraxinus excelsior in forest districts with a high infection rate of H. fraxineus, the causal agent of ash dieback. Based on phylogenomic and phenotypic analyses, a clear differentiation from related Luteimonas species was shown. Comparisons of the overall genome relatedness indices with the closest phylogenetic neighbours resulted in values below the recommended species cut-off levels. In addition, differences in several physiological and chemotaxonomic traits allowed a clear demarcation from the type strains of closely related species. Conclusively, the strain group was considered to represent a novel species in the genus Luteimonas, for which the name Luteimonas fraxinea sp. nov. is proposed, with strain D4P002^T (=DSM 113273^T = LMG 32455^T) as the type strain. A functional analysis of the genome revealed features particularly associated with attachment, biofilm production and motility, indicating the ability of D4P002^T to effectively colonise ash leaves. In nursery trials, ash seedlings inoculated with this strain showed suppression of the pathogen over a period of three years. This effect was accompanied by a significant shift in the bacterial microbiome of the plants. Altogether, the exclusive occurrence in the microbiome of tolerant ash trees, the genetic background and the results of the inoculation experiment suggest that strain D4P002^T may suppress the penetration and spreading of H. fraxineus in or on ash leaves via colonisation resistance or trigger a priming effect of plant defences against the pathogen.     

Influences of size and sex on invasive species aggression and native species vulnerability: a case for modern regression techniques

Animal behaviour is of fundamental importance but is often overlooked in biological invasion research. A problem with such studies is that they may add pressure to already threatened species and subject vulnerable individuals to increased risk. One solution is to obtain the maximum possible information from the generated data using a variety of statistical techniques, instead of solely using simple versions of linear regression or generalized linear models as is customary. Here, we exemplify and compare the use of modern regression techniques which have very different conceptual backgrounds and aims (negative binomial models, zero-inflated regression, and expectile regression), and which have rarely been applied to behavioural data in biological invasion studies. We show that our data display overdispersion, which is frequent in ecological and behavioural data, and that conventional statistical methods such as Poisson generalized linear models are inadequate in this case. Expectile regression is similar to quantile regression and allows the estimation of functional relationships between variables for all portions of a probability distribution and is thus well suited for modelling boundaries in polygonal relationships or cases with heterogeneous variances which are frequent in behavioural data. We applied various statistical techniques to aggression in invasive mosquitofish, Gambusia holbrooki, and the concomitant vulnerability of native toothcarp, Aphanius iberus, in relation to individual size and sex. We found that medium sized male G. holbrooki carry out the majority of aggressive acts and that smaller and medium size A. iberus are most vulnerable. Of the regression techniques used, only negative binomial models and zero-inflated and expectile Poisson regressions revealed these relationships.