A simple approach to optimal control of invasive species

The problem of invasive species and their control is one of the most pressing applied issues in ecology today. We developed simple approaches based on linear programming for determining the optimal removal strategies of different stage or age classes for control of invasive species that are still in a density-independent phase of growth. We illustrate the application of this method to the specific example of invasive Spartina alterniflora in Willapa Bay, WA. For all such systems, linear programming shows in general that the optimal strategy in any time step is to prioritize removal of a single age or stage class. The optimal strategy adjusts which class is the focus of control through time and can be much more cost effective than prioritizing removal of the same stage class each year.     

Avoiding surprise effects on Surprise Island: alien species control in a multitrophic level perspective

Eradications of invasive alien species have generally benefited biodiversity. However, without sufficient planning, successful eradications can have unexpected and unwanted consequences for native species and ecosystems. In particular, the “surprise effect” is the rapid increase of hitherto unnoticed species following the sudden removal of an invasive alien that was exerting an ecological force on those species (predation, competition or herbivory, for example). The only way to prevent these undesired outcomes is to adapt the control programme following the characterization of the trophic relationships between the invasive alien species and the invaded communities, that is, to view the control with a holistic perspective. Here, we illustrate this point with the study of the role of the ship rat (Rattus rattus), which invaded a tropical pacific atoll, Surprise Island, New Caledonia. We assessed the risk of surprise effects during a pre-eradication phase of several years, and then adapted our eradication strategy accordingly.     

Minimizing invader impacts: striking the right balance between removal and restoration

Invasive species can cause severe damage in their introduced range; this damage often persists even after removal of the invader. In order to efficiently allocate a limited budget between invader removal and restoration of habitat from which the invader has been removed, it is vital to quantify the impacts of the invasion within an economic context. Here we develop optimal management strategies for biological invasions, which minimize both the direct economic costs of removal and restoration, and the ecological costs of present and future damage caused by the invasion. We demonstrate how this can be formulated as a linear programming problem, enabling the fast and efficient computation of optimal solutions. Using a simple example, we outline some general principles for the optimal control of an invader that damages its environment. Notably, we show that the most effective strategies often switch the priority of removal and restoration over time, and outline the conditions under which restoration is prioritized over removal. The proportion of total funds allocated to restoration will depend on the annual budget, the persistence of damage, and the relative costs of damage, removal and restoration.     

Conflicting values: ecosystem services and invasive tree management

Tree species have been planted widely beyond their native ranges to provide or enhance ecosystem services such as timber and fibre production, erosion control, and aesthetic or amenity benefits. At the same time, non-native tree species can have strongly negative impacts on ecosystem services when they naturalize and subsequently become invasive and disrupt or transform communities and ecosystems. The dichotomy between positive and negative effects on ecosystem services has led to significant conflicts over the removal of non-native invasive tree species worldwide. These conflicts are often viewed in only a local context but we suggest that a global synthesis sheds important light on the dimensions of the phenomenon. We collated examples of conflict surrounding the control or management of tree invasions where conflict has caused delay, increased cost, or cessation of projects aimed at invasive tree removal. We found that conflicts span a diverse range of taxa, systems and countries, and that most conflicts emerge around three areas: urban and near-urban trees; trees that provide direct economic benefits; and invasive trees that are used by native species for habitat or food. We suggest that such conflict should be seen as a normal occurrence in invasive tree removal. Assessing both positive and negative effects of invasive species on multiple ecosystem services may provide a useful framework for the resolution of conflicts.     

Plant-disperser mutualisms in a semi-arid habitat invaded by <Emphasis Type="Italic">Lantana camara</Emphasis> L.

Dispersal is an important ecological process that affects plant population structure and community composition. Invasive plants with fleshy fruits rapidly form associations with native and invasive dispersers, and may affect existing native plant-disperser associations. We asked whether frugivore visitation rate and fruit removal was associated with plant characteristics in a community of fleshy-fruited plants and whether an invasive plant receives more visitation and greater fruit removal than native plants in a semi-arid habitat of Andhra Pradesh, India. Tree-watches were undertaken at individuals of nine native and one invasive shrub species to assess the identity, number and fruit removal by avian frugivores. Network analyses and generalised linear mixed-effects models were used to understand species and community-level patterns. All plants received most number of visits from abundant, generalist avian frugivores. Number of frugivore visits and time spent by frugivores at individual plants was positively associated with fruit crop size, while fruit removal was positively associated with number of frugivore visits and their mean foraging time at individual plants. The invasive shrub, Lantana camara L. (Lantana), had lower average frugivore visit rate than the community of fleshy-fruited plants and received similar average frugivore visits but greater average per-hour fruit removal than two other concurrently fruiting native species. Based on the results of our study, we infer that there is little evidence of competition between native plants and Lantana for the dispersal services of native frugivores and that more data are required to assess the nature of these interactions over the long term. We speculate that plant associations with generalist frugivores may increase the functional redundancy of this frugivory network, buffering it against loss of participating species.     

Does localized control of invasive eastern gambusia (Poeciliidae: Gambusia holbrooki) increase population growth of generalist wetland fishes?

While invasive fish management is heavily focussed on containment measures when introductions occur, examples from invasive species management in terrestrial systems suggest that there may also be considerable conservation benefits in implementing localized control programmes. We conducted a field‐based experiment to assess the effectiveness of removing a globally significant invasive fish, eastern gambusia Gambusia holbrooki, from natural wetland habitats of south‐eastern Australia. With recent work suggesting the impacts of eastern gambusia may be minimal for species with generalist life‐history strategies, we hypothesized that the removal of eastern gambusia will reduce localized population growth of the invasive species, but will have little influence on the population growth of more generalist sympatric wetland fish species. We used a predictive modelling approach to investigate changes in eastern gambusia populations following removal activities, and how sympatric fish species responded to such changes. Although eastern gambusia rapidly populated habitats, we demonstrated that control actions substantially reduced the rate of population increase over the four‐month study period. This suggests that control may be an effective localized strategy to suppress eastern gambusia densities. There was however, no evidence of any response to the removal actions by any of the three sympatric fish species investigated – carp gudgeon (Hypseleotris spp.), Australian smelt (Retropinna semoni) and the invasive common carp (Cyprinus carpio). These results support previous work which suggests that the flexible life‐history strategies and behavioural traits of all three species allow co‐existence with eastern gambusia. The study highlights the importance of understanding the potential outcomes of control options which is particularly pertinent for established aquatic invasive species where information on control effectiveness, population dynamics and/or ecosystem response is currently lacking.     

Large-Scale Removal of Invasive Honeysuckle Decreases Mosquito and Avian Host Abundance

Invasive species rank second only to habitat destruction as a threat to native biodiversity. One consequence of biological invasions is altered risk of exposure to infectious diseases in human and animal populations. The distribution and prevalence of mosquito-borne diseases depend on the complex interactions between the vector, the pathogen, and the human or wildlife reservoir host. These interactions are highly susceptible to disturbance by invasive species, including terrestrial plants. We conducted a 2-year field experiment using a Before–After/Control–Impact design to examine how removal of invasive Amur honeysuckle (Lonicera maackii) in a forest fragment embedded within a residential neighborhood affects the abundance of mosquitoes, including two of the most important vectors of West Nile virus, Culex pipiens and Cx. restuans. We also assessed any potential changes in avian communities and local microclimate associated with Amur honeysuckle removal. We found that (1) removal of Amur honeysuckle reduces the abundance of both vector and non-vector mosquito species that commonly feed on human hosts, (2) the abundance and composition of avian hosts is altered by honeysuckle removal, and (3) areas invaded with honeysuckle support local microclimates that are favorable to mosquito survival. Collectively, our investigations demonstrate the role of a highly invasive understory shrub in determining the abundance and distribution of mosquitoes and suggest potential mechanisms underlying this pattern. Our results also give rise to additional questions regarding the general impact of invasive plants on vector-borne diseases and the spatial scale at which removal of invasive plants may be utilized to effect disease control.     

Maximum sustainable yield of populations with continuous age-structure.

We address the problem of finding the harvesting policy that will maximize the yield and maintain a population in a steady state. The population is characterized by continuous age classes and therefore follows differential equations. Here, we assume that the equations are linear (no density dependence). Two possible constraints are considered: either recruitment or total population are fixed to a constant. Under these conditions, the optimal policy is to harvest the fraction theta of a younger age class ? and to harvest totally an older age class b. The optimal solution (theta, ?, b) can be calculated explicitly if the fecundity and mortality schedules are given. The solution is compared to the simpler strategy of harvesting all individuals beyond a single age class a. It is shown that the latter strategy can be much less profitable than harvesting two age classes because it cannot take account of the different values of individuals according to their age.     

Age-structured non-pharmaceutical interventions for optimal control of COVID-19 epidemic

In an epidemic, individuals can widely differ in the way they spread the infection depending on their age or on the number of days they have been infected for. In the absence of pharmaceutical interventions such as a vaccine or treatment, non-pharmaceutical interventions (e.g. physical or social distancing) are essential to mitigate the pandemic. We develop an original approach to identify the optimal age-stratified control strategy to implement as a function of the time since the onset of the epidemic. This is based on a model with a double continuous structure in terms of host age and time since infection. By applying optimal control theory to this model, we identify a solution that minimizes deaths and costs associated with the implementation of the control strategy itself. We also implement this strategy for three countries with contrasted age distributions (Burkina-Faso, France, and Vietnam). Overall, the optimal strategy varies throughout the epidemic, with a more intense control early on, and depending on host age, with a stronger control for the older population, except in the scenario where the cost associated with the control is low. In the latter scenario, we find strong differences across countries because the control extends to the younger population for France and Vietnam 2 to 3 months after the onset of the epidemic, but not for Burkina Faso. Finally, we show that the optimal control strategy strongly outperforms a constant uniform control exerted over the whole population or over its younger fraction. This improved understanding of the effect of age-based control interventions opens new perspectives for the field, especially for age-based contact tracing.     

Optimal eradication: when to stop looking for an invasive plant

The notion of being sure that you have completely eradicated an invasive species is fanciful because of imperfect detection and persistent seed banks. Eradication is commonly declared either on an ad hoc basis, on notions of seed bank longevity, or on setting arbitrary thresholds of 1% or 5% confidence that the species is not present. Rather than declaring eradication at some arbitrary level of confidence, we take an economic approach in which we stop looking when the expected costs outweigh the expected benefits. We develop theory that determines the number of years of absent surveys required to minimize the net expected cost. Given detection of a species is imperfect, the optimal stopping time is a trade-off between the cost of continued surveying and the cost of escape and damage if eradication is declared too soon. A simple rule of thumb compares well to the exact optimal solution using stochastic dynamic programming. Application of the approach to the eradication programme of Helenium amarum reveals that the actual stopping time was a precautionary one given the ranges for each parameter.     

The influence of time since introduction on the population growth of introduced species and the consequences for management

Several processes likely act to change the demographic rates of introduced species over time, and this changing demography could influence the optimal management of invasive populations. Optimal management strategies should be derived based on the demography. However, we have a poor understanding of the degree to which the demography of introduced species changes following initial introduction. We used published matrix population models of introduced plant populations to test how population growth rate and elasticity change with time since introduction. We did not find a significant relationship between population growth rate and time since introduction. However, elasticity to stasis increased while elasticity to growth decreased with time since introduction. Broadly, as time since introduction progressed the elasticities of the introduced plant populations became more similar to those that have been reported for native species. These results suggest that the optimal management strategy should be derived incorporating elasticity through time, especially when the time scope of management is long or the available demographic data were obtained in the past.     

入侵种薇甘菊防治措施及策略评估

薇甘菊Mikania micrantha H.B.K.是入侵中国南部地区的一种危害极大的外来入侵种。防治薇甘菊的措施和策略,应根据薇甘菊的物种特性及所能支配的防除资源,综合考虑、统筹安排。薇甘菊能快速生长覆盖其他植物,其残株断枝易成为独立生长的新个体、种子量巨大且极易扩散,喜入侵次生植被。物理清除适用于小面积,对大面积出现的薇甘菊不应采用。化学防除主要用于应急,但未能改变适于薇甘菊生长的生境,薇甘菊仍可能在数年后再次为害。田野菟丝子Cuscuta campestris控制薇甘菊适用于大面积危害区域,当只以控制而不以根除自然群落的薇甘菊为目标时效果很好。群落改造能一劳永逸地解除薇甘菊的危害,但只适用于宜林地,且成本较高。天敌控制和植物化感防治都尚在探索阶段。未来还应发掘有效天敌,研发综合防治措施,揭示宏观分布规律。薇甘菊防治的正确策略是:对于已形成区域性危害的薇甘菊,根除已经不可能,应采取持久战而非速决战,防止薇甘菊入侵新区域是防治的重中之重,必须优先治理轻度入侵地和能向周边区域甚至是遥远区域扩散的传播源,建立防治示范区或样地。     

Simulating control strategies for a spatially structured weed invasion: <Emphasis Type="Italic">Spartina alterniflora</Emphasis> (Loisel) in Pacific Coast estuaries

Patterns in the spatial arrangement of invasive plant populations can provide opportunity for strategic placement of control efforts. Smooth cordgrass (Spartina alterniflora) is rapidly invading the intertidal mudflats of Pacific Coast estuaries. Its pattern of spread is distinctive. Seedlings establish in open mud and then grow vegetatively to form expanding circular patches, which dot the mudflats and eventually coalesce into a contiguous monospecific meadow. The invasion typically begins in the upper tide zone and then moves down the tidal gradient. A spatially explicit model was used to simulate the spread of S. alterniflora and compare various strategies for control in a situation where only a fraction of the total infestation could be controlled each year. A strategy of killing outlying patches first and then attacking the dense meadows (moving up the tidal gradient) led to eradication with up to 44% less time and effort than a strategy of attacking the dense meadows first and outlying patches second (moving down the tidal gradient). In the control of contiguous meadows located adjacent to the shoreline, the best strategy was to approach one end of the infestation, moving across the meadow to the other end. Suppression of seeds was not an effective control strategy. In general, effective control strategies were those that first eliminate the plant in areas where current or future vegetative growth is greatest. Application of these results in control programs for S. alterniflora and similar invasive species could greatly reduce the costs of control work and improve the likelihood of local or complete eradication.     

Impacts of invasive fish removal through angling on population characteristics and juvenile growth rate

Exploitation can modify the characteristics of fish populations through the selective harvesting of individuals, with this potentially leading to rapid ecological and evolutionary changes. Despite the well‐known effects of invasive fishes on aquatic ecosystems generally, the potential effects of their selective removal through angling, a strategy commonly used to manage invasive fish, are poorly understood. The aim of this field‐based study was to use the North American pumpkinseed Lepomis gibbosus as the model species to investigate the consequences of selective removal on their population characteristics and juvenile growth rates across 10 populations in artificial lakes in southern France. We found that the maximal individual mass in populations decreased as removal pressure through angling increased, whereas we did not observed any changes in the maximal individual length in populations as removal pressure increased. Total population abundance did not decrease as removal pressure increased; instead, here was a U‐shaped relationship between removal pressure and the abundance of medium‐bodied individuals. In addition, population biomass had a U‐shaped curve response to removal pressure, implying that invasive fish populations can modulate their characteristics to compensate for the negative effects of selective removals. In addition, individual lengths at age 2 and juvenile growth rates decreased as removal pressure through angling increased, suggesting a shift toward an earlier size at maturity and an overall slower growing phenotype. Therefore, these outputs challenge the efficiency of selective management methods, suggesting the use of more proactive strategies to control invasive populations, and the need to investigate the potential ecological and evolutionary repercussions of nonrandom removal.     

Biodiversity influences invasion success of a facultative epiphytic seaweed in a marine forest

The biotic resistance hypothesis predicts that more diverse communities should have greater resistance to invasions than species-poor communities. However for facultative and obligate epiphytic invaders a high native species richness, abundance and community complexity might provide more resources for the invader to thrive to. We conducted surveys across space and time to test for the influence of native algal species abundance and richness on the abundance of the invasive facultative epiphytic filamentous alga Lophocladia lallemandii in a Mediterranean Cystoseira balearica seaweed forest. By removing different functional groups of algae, we also tested whether these relationships were dependent on the complexity and abundance of the native algal community. When invasion was first detected, Lophocladia abundance was positively related to species richness, but the correlation became negative after two years of invasion. Similarly, a negative relationship was also observed across sites. The removal experiment revealed that more complex native communities were more heavily invaded, where also a positive relationship was found between native algal richness and Lophocladia, independently of the native algal abundance. Our observational and experimental data show that, at early stages of invasion, species-rich seaweed forests are not more resistant to invasion than species-poor communities. Higher richness of native algal species may increase resource availability (i.e. substrate) for invader establishment, thus facilitating invasion. After the initial invasion stage, native species richness decreases with time since invasion, suggesting negative impacts of invasive species on native biodiversity.     

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.     

Recovery of Native Grasslands after Removing Invasive Pines

The advance of exotic tree and shrub species is one of the main threats to conservation of the last relicts of natural grassland in South America; however, control actions in the region are still scarce and there are almost no evaluations of the recovery of natural ecosystems after removing invasive plants. Monitoring of the vegetation during the years after removal of invasive trees is critical in order to decide whether an active restoration strategy is necessary. The recovery of montane grassland four years after the control of a dense invasion of Aleppo pine (Pinus halepensis) is described in this study. Experimental clearing areas were followed during four years and compared to grassland controls. Variation was seen in the levels of recovery in function of the proximity of sectors of grassland that are free of invasive species and/or the density of invasive trees before control. Native species slowly replaced many exotic herbs that had appeared as pioneers, there was low recruitment of pine seedlings in spite of the quantity of seeds from trees that surrounded the clearings, and species richness and diversity were restored, including cover of the typical grasses in the controls. Recovery of grassland after felling was shown to be successful and does not seem to be seed limited if tree removal occurs early in the invasion process.     

Competing for crevices: interspecific conflict influences retreat-site selection in montane lizards

Direct interference competition between sympatric taxa affects habitat use and shelter-site selection in species within most major vertebrate lineages. However, studies on interspecific social interference in reptiles largely have been confined to research on interactions between non-native (invasive) species and native fauna. Does interspecific interference also influence habitat use within natural assemblages of reptiles? We studied five broadly sympatric species of viviparous montane skinks within the genera Egernia and Eulamprus in southeastern Australia. Previous work has shown strong interspecific overlap in abiotic attributes of shelter-sites for these taxa, but no joint occupancy of retreats. Laboratory trials in which we manipulated the identity of co-occurring lizards revealed frequent displacement from “preferred” (hotter) shelters, with interspecific interactions more intense than intraspecific conflicts. The five species displayed a linear interspecific dominance hierarchy, with larger species displacing smaller taxa. Field manipulations confirmed that interspecific interactions between these species affect shelter-site use. We conclude that direct agonistic encounters between individuals of different species strongly influence habitat use by lizards within this assemblage.     

Optimal schedule for monitoring a plant incursion when detection and treatment success vary over time

Management of an invasive plant species can be viewed as two separate and successive processes. The first, survey, aims to find infested areas and remove individuals. The second, monitoring, consists of repeated visits to these areas in order to prevent possible re-emergence. As detection probability may vary over time, the timing and number of monitoring visits can dramatically impact monitoring efficacy. We explore the optimal timing and number of monitoring visits, by focusing on one infested site. Our decision-analysis framework defines an optimal monitoring schedule which accounts for a time-dependent probability of detection, based on the presence/absence of a flower. We use this framework to investigate the optimal monitoring schedule for Hieracium aurantiacum, an invasive species in the Australian Alps and many other countries. We also perform a sensitivity analysis to draw more general conclusions. For H. aurantiacum eight monitoring visits (compared to 12 visits in the current program) are sufficient to obtain a 99% monitoring efficacy. When four or fewer visits to a site are allowed, it is optimal to visit during the high season, when the weed is likely to initiate flowering. Any extra visits should be scheduled in the early season, before the plants flower. The sensitivity analysis shows that increasing the detection probability early in the season has a greater impact than increasing it late in the season. An effective treatment method increases the value of site visits late in the season, when the detection probability is higher. Our decision-analysis framework can assist invasive species managers to reduce or reallocate management resources by determining the minimum number of monitoring visits required to satisfy an acceptable risk of re-emergence.