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Reproductive condition of an invasive snake in urban, savanna, and forest habitats

Predation by Brown Treesnakes ( Boiga irregularis ) has caused the decline or extinction of all native forest vertebrates on Guam, including birds, lizards, and bats. Loss of the highest-quality prey has caused shifts in Brown Treesnake size and life history. We sought to understand how reproductive condition varies among habitats island-wide, particularly comparing urban and savanna habitats (where some primarily non-native large prey remain) with prey-depauperate forest habitats. We dissected 100 snakes from each of 18 sites stratified by 6 habitat types and calculated a body condition index, relative follicle size (for females; N = 817) and relative testes size (for males; N = 992) for each. Low detection of reproductive females continues to hamper research on their biology, as only 11% of females had secondary vitellogenic follicles (N = 89). Although variation among sites within habitats was high, above-average reproductive development of both males and females was most frequently observed in urban and savanna habitats, emphasizing the potential for prey subsidies in these habitats to fuel population growth and affect invasive snake management objectives. Furthermore, the trade-off between body condition and reproductive condition appeared to be more severe in forested habitats and less severe in urban habitats for both males and females. There were few obvious seasonal patterns, although there was a slightly lower probability of females having secondary vitellogenic follicles in the wet season compared to the dry season.

NeoBiota

Designing biosurveillance using target analysis for a cryptic invasive species, the brown treesnake (Boiga irregularis)

Biosurveillance is especially important for invasive species that are difficult to control or remove once established. Target analysis, or the strategic evaluation of factors that affect or improve detection of organisms, is one approach to improve biosurveillance over time. Due to their ecologically and economically devastating effects, brown treesnakes ( Boiga irregularis ), an invasive snake in the United States territory of Guam, have a dedicated biosecurity program to prevent their spread. Guam has been occupied by brown treesnakes since the 1940s, and they are distributed throughout the island. The neighboring Saipan Island, otherwise known as Saipan, participates in biosecurity programs designed to prevent their establishment. We evaluated biosurveillance protocols within rapid response and detection exercises for brown treesnakes. Our goal was to identify, account for, and reduce potential error when inferring absence from lack of detection using data collected in Guam and Saipan. We evaluated 1) whether brown treesnake detection probability was density dependent; 2) whether using terrestrial environmental DNA ( eDNA ) sampling detected snakes in occupied areas; 3) the likelihood brown treesnakes would have been detected in Saipan, given sampling effort during a biosurveillance exercise, and whether lizards could be used as a proxy on snake-free islands for estimating factors that affect detection; 4) the effect of transect design on pacing and subsequently pacing on lizard counts; and 5) the importance of observer training in surveying for cryptic reptiles and completing broad versus targeted bioinventories. We found that brown treesnake detection probability was density dependent when using visual surveys and declined with abundance in Guam, supporting that greater effort was required to have high confidence that lack of detection supported absence in areas with low snake densities. The terrestrial eDNA sampling protocol we developed was effective at detecting snakes in occupied locations in Guam and may be a complementary tool. However, further validation of eDNA protocols would still be required for early detection application. We also found that among trained observers, the ability to detect lizards was positively predictive of the number of snakes they detected at the standard snake densities in Guam. Thus, lizard detection appeared to be a reasonable proxy for evaluating visual search protocols for snakes on snake-free islands. Overall, creating easy pace-setting relationships between time and distance (i.e., 10 m per min), with flagging that clearly marked location on a transect, greatly reduced variability in search pace across survey nights. Observers that were continuously trained to maintain brown treesnake search profiles detected twice as many, on average, cryptic reptiles compared to individuals who had never received training, which was not true for the non-reptiles (birds and small mammals). Additionally, conducting broad bioinventories with many target species reduced detection of the target species, and this was increasingly true as years since the observer completed training for the target species increased. We thus found that designing biosurveillance exercises to evaluate search protocols can help improve or refine survey methods to reduce noise in the data, and accounting for that can improve the estimated confidence that failure to detect an invasive species infers absence.

NeoBiota

Small but tough: What can ecophysiology of croaking gourami Trichopsis vittatus (Cuvier 1831) tell us about invasiveness of non-native fishes in Florida?

Trichopsis vittata (Cuvier, 1831) is a small, freshwater gourami (Fam: Osphronemidae ) native to southeast Asia. It was first detected in Florida in the 1970s and seems to have persisted for decades in a small area. In this study, we documented T. vittata ’s ecophysiological tolerances (salinity and low-temperature) and qualitatively compared them to published values for other sympatric non-native species that have successfully invaded much of the Florida peninsula. Trichopsis vittata survived acute salinity shifts to 16 psu and was able to survive up to 20 psu when salinity was raised more slowly (5 psu per week). In a cold-tolerance experiment, temperature was lowered from 24 °C at 1 °C hr -1 until fish died. Mean temperature at death (i.e., lower lethal limit) was 7.2 °C. Trichopsis vittata seems as tolerant or more tolerant than many other sympatric non-native fishes for the variables we examined. However, T. vittata is the only species that has not dispersed since its introduction. Species other than T. vittata have broadly invaded ranges, many of which include the entire lower third of the Florida peninsula. It is possible that tolerance to environmental parameters serves as a filter for establishment, wherein candidate species must possess the ability to survive abiotic extremes as a first step. However, a species’ ability to expand its geographic range may ultimately rely on a secondary set of criteria including biotic interactions and life-history variables.

NeoBiota

Evaluating simplistic methods to understand current distributions and forecast distribution changes under climate change scenarios: An example with coypu ( Myocastor coypus )

Invasive species provide a unique opportunity to evaluate factors controlling biogeographic distributions; we can consider introduction success as an experiment testing suitability of environmental conditions. Predicting potential distributions of spreading species is not easy, and forecasting potential distributions with changing climate is even more difficult. Using the globally invasive coypu ( Myocastor coypus [Molina, 1782]), we evaluate and compare the utility of a simplistic ecophysiological based model and a correlative model to predict current and future distribution. The ecophysiological model was based on winter temperature relationships with nutria survival. We developed correlative statistical models using the Software for Assisted Habitat Modeling and biologically relevant climate data with a global extent. We applied the ecophysiological based model to several global circulation model ( GCM ) predictions for mid-century. We used global coypu introduction data to evaluate these models and to explore a hypothesized physiological limitation, finding general agreement with known coypu distribution locally and globally and support for an upper thermal tolerance threshold. Global circulation model based model results showed variability in coypu predicted distribution among GCMs , but had general agreement of increasing suitable area in the USA. Our methods highlighted the dynamic nature of the edges of the coypu distribution due to climate non-equilibrium, and uncertainty associated with forecasting future distributions. Areas deemed suitable habitat, especially those on the edge of the current known range, could be used for early detection of the spread of coypu populations for management purposes. Combining approaches can be beneficial to predicting potential distributions of invasive species now and in the future and in exploring hypotheses of factors controlling distributions.

NeoBiota

Provenance of invaders has scale-dependent impacts in a changing wetland ecosystem

Exotic species are associated with a variety of impacts on biodiversity, but it is unclear whether impacts of exotic species differ from those of native species with similar growth forms or native species invading disturbed sites. We compared presence and abundance of native and exotic invaders with changes in wetland plant species diversity over a 28-year period by re-surveying 22 ponds to identify factors correlated with observed changes. We also compared communities found within dense patches of native and exotic emergent species with similar habits. Within patches, we found no categorical diversity differences between areas dominated by native or exotic emergent species. At the pond scale, the cover of the exotic grass Phragmites australis best predicted change in diversity and evenness over time, likely owing to its significant increase in coverage over the study period. These changes in diversity and evenness were strongest in younger, less successionally-advanced ponds. Changes associated with cover of P. australis in these ponds were not consistent with expected diversity decreases, but instead with a dampening of diversity gains, such that the least-invaded ponds increased in diversity the most over the study period. There were more mixed effects on evenness, ranging from a reduction in evenness gains to actual losses of evenness in the ponds with highest invader cover. In this wetland complex, the habit, origin and invasiveness of species contribute to diversity responses in a scale- and context-dependent fashion. Future efforts to preserve diversity should focus on preventing the arrival and spread of invaders that have the potential to cover large areas at high densities, regardless of their origin. Future studies should also investigate more thoroughly how changes in diversity associated with species invasions are impacted by other ongoing ecosystem changes.

NeoBiota

Contact rates with nesting birds before and after invasive snake removal: Estimating the effects of trap-based control

Invasive predators are responsible for almost 60% of all vertebrate extinctions worldwide with the most vulnerable faunas occurring on islands. The brown treesnake ( Boiga irregularis ) is a notorious invasive predator that caused the extirpation or extinction of most native forest birds on Guam. The success of avian reintroduction efforts on Guam will depend on whether snake-control techniques sufficiently reduce contact rates between brown treesnakes and reintroduced birds. Mouse-lure traps can successfully reduce brown treesnake populations at local scales. Over a 22-week period both with and without active snake removal, we evaluated snake-trap contact rates for mouse- and bird-lure traps. Bird-lure traps served as a proxy for reintroduced nesting birds. Overall, mouse-lure traps caught more snakes per trap night than did bird-lure traps. However, cameras revealed that bird-lure traps had a snake contact rate almost 15 times greater than the number of successfully captured snakes. Snakes that entered bird-lure traps tended to be larger and in better body condition and were mostly captured in bird-lure traps, despite numerous adjacent mouse-lure traps. Traps placed along grid edges caught more snakes than interior traps, suggesting continuous immigration into the trapping grid within which bird-lure traps were located. Contact between snakes and bird-lure traps was equivalent before and after snake removal, suggesting mouse-lure traps did not adequately reduce the density of snakes that posed a risk to birds, at least at the timescale of this project. This study provides evidence that some snakes exhibit prey selectivity for live birds over live mouse lures. Reliance on a single control tool and lure may be inadequate for support of avian reintroductions and could lead to unintended harvest-driven trait changes of this invasive predator.

NeoBiota

The impact is in the details: Evaluating a standardized protocol and scale for determining non-native insect impact

Assessing the ecological and economic impacts of non-native species is crucial to providing managers and policymakers with the information necessary to respond effectively. Most non-native species have minimal impacts on the environment in which they are introduced, but a small fraction are highly deleterious. The definition of ‘damaging’ or ‘high-impact’ varies based on the factors determined to be valuable by an individual or group, but interpretations of whether non-native species meet particular definitions can be influenced by the interpreter’s bias or level of expertise, or lack of group consensus. Uncertainty or disagreement about an impact classification may delay or otherwise adversely affect policymaking on management strategies. One way to prevent these issues would be to have a detailed, nine-point impact scale that would leave little room for interpretation and then divide the scale into agreed upon categories, such as low, medium, and high impact. Following a previously conducted, exhaustive search regarding non-native, conifer-specialist insects, the authors independently read the same sources and scored the impact of 41 conifer-specialist insects to determine if any variation among assessors existed when using a detailed impact scale. Each of the authors, who were selected to participate in the working group associated with this study because of their diverse backgrounds, also provided their level of expertise and uncertainty for each insect evaluated. We observed 85% congruence in impact rating among assessors, with 27% of the insects having perfect inter-rater agreement. Variance in assessment peaked in insects with a moderate impact level, perhaps due to ambiguous information or prior assessor perceptions of these specific insect species. The authors also participated in a joint fact-finding discussion of two insects with the most divergent impact scores to isolate potential sources of variation in assessor impact scores. We identified four themes that could be experienced by impact assessors: ambiguous information, discounted details, observed versus potential impact, and prior knowledge. To improve consistency in impact decision-making, we encourage groups to establish a detailed scale that would allow all observed and published impacts to fall under a particular score, provide clear, reproducible guidelines and training, and use consensus-building techniques when necessary.

NeoBiota

Tools for increasing visual encounter probabilities for invasive species removal: A case study of brown treesnakes

Early detection and rapid response (EDRR) are essential to identifying and decisively responding to the introduction or spread of an invasive species, thus avoiding population establishment and improving the probability of achieving eradication. However, detection can be challenging at the onset of a species invasion as low population densities can reduce detection probability and conceal the true extent of the situation until the species is well established. This is doubly challenging if the invading species displays cryptic behavior or is nocturnal, thus further limiting opportunities for its discovery. Survey methods that maximize a searcher’s ability to detect an incipient population are therefore critical for successful EDRR. Brown treesnakes (Boiga irregularis) on Guåhan are a classic cautionary example of the dangers of not detecting an invasion early on, and the risk of their introduction to other islands within the Marianas, Hawai’i and beyond remains. Nocturnal visual surveys are known to detect brown treesnakes of all sizes and are the primary detection tool used by the Brown Treesnake Rapid Response Team, but detection probability remains low in complex forest habitats. As such, we investigated the use of two potential enhancements to nocturnal visual surveys – a live mouse lure and spray scent attractant – that may create hotspots of increased detection probability during surveys. We found that, while brown treesnake detection probabilities were low for all surveys, visual surveys conducted on transects with live mouse lures resulted in detection probabilities that were 1.3 times higher than on transects without live mouse lures. Conversely, the spray scent attractant did not increase the probability of detecting brown treesnakes compared to transects without scent, and in fact had detection probabilities that were 0.66 times lower, though the reasons for this phenomenon are unclear. Unlike scent attractants, live mouse lures likely provide both visual and olfactory cues that attract brown treesnakes to transects and thus provide more opportunities to detect and capture them. These enhancements were trialed on Guåhan, where prey populations are depressed. It remains unclear whether live mouse lures will be as effective for EDRR applications in prey-rich settings.

Guam

Chew-cards can accurately index invasive rat densities in Mariana Island forests

Rats ( Rattus spp.) are likely established on 80–90% of the world’s islands and represent one of the most damaging and expensive biological invaders. Effective rat control tools exist but require accurate population density estimates or indices to inform treatment timing and effort and to assess treatment efficacy. Capture-mark-recapture data are frequently used to produce robust density estimates, but collecting these data can be expensive, time-consuming, and labor-intensive. We tested a potentially cheaper and easier alternative, chew-cards, as a count-based (quantitative) index of invasive rat densities in tropical forests in the Mariana Islands, an archipelago in the western North Pacific Ocean. We trialed chew-cards in nine forest grids on two Mariana Islands by comparing the proportion of cards chewed to capture-mark-recapture density estimates and manipulated rat densities to test whether the relationship was retained. Chew-card counts were positively correlated with rat capture-mark-recapture density estimates across a range of rat densities found in the region. Additionally, the correlation between the two sampling methods increased with the number of days chew-cards were deployed. Specifically, when chew-cards were deployed for five nights, a 10% increase in the proportion of cards chewed equated to an estimated increase in rat density of approximately 2.4 individuals per ha (R 2 = 0.74). Chew-cards can provide a valid index of rat densities in Mariana Island forests and are a cheaper alternative to capture-mark-recapture sampling when relative differences in density are of primary interest. New cost-effective monitoring tools can enhance our understanding and management of invaded islands while stretching limited resources further than some conventional approaches, thus improving invasive species management on islands.

NeoBiota

Scanning the horizon for invasive plant threats using a data-driven approach

Early detection and eradication of invasive plants are more cost-effective than managing well-established invasive plant populations and their impacts. However, there is high uncertainty around which taxa are likely to become invasive in a given area. Horizon scanning that combines a data-driven approach with rapid risk assessment and consensus building among experts can help identify invasion threats. We performed a horizon scan of potential invasive plant threats to Florida, USA—a state with a high influx of introduced species, conditions that are generally favorable for plant establishment, and a history of negative impacts from invasive plants. We began with an initial list of 2128 non-native plant taxa that are known invaders or crop pests. We built on previous invasive species horizon scans by developing data-based criteria to prioritize 100 taxa for rapid risk assessment. The semi-automated prioritization process included selecting taxa “on the horizon” (i.e., not yet in the target location and not on a noxious weed list) with climate matching, naturalization history, “weediness” record, and global commonness. We derived overall invasion risk scores with rapid risk assessment by evaluating the likelihood of each of the taxa arriving, establishing, and having an impact in Florida. Then, following a consensus-building discussion, we identified six plant taxa as high risk, with overall risk scores ranging from 75 to 100 out of a possible 125. The six taxa are globally distributed, easily transported to new areas, found in regions with climates similar to Florida’s, and can impact native plant communities, human health, or agriculture. Finally, we evaluated our initial and final lists for potential biases. Assessors tended to assign higher risk scores to taxa that had more available information. In addition, we identified biases towards four plant families and certain geographical regions of origin. Our horizon scan approach identified taxa conforming to metrics of high invasion risk and used a methodology refined for plants that can be applied to other locations.

NeoBiota

Regional models do not outperform continental models for invasive species

Aim : Species distribution models can guide invasive species prevention and management by characterizing invasion risk across space. However, extrapolation and transferability issues pose challenges for developing useful models for invasive species. Previous work has emphasized the importance of including all available occurrences in model estimation, but managers attuned to local processes may be skeptical of models based on a broad spatial extent if they suspect the captured responses reflect those of other regions where data are more numerous. We asked whether species distribution models for invasive plants performed better when developed at national versus regional extents. Location : Continental United States. Methods : We developed ensembles of species distribution models trained nationally, on sagebrush habitat, or on sagebrush habitat within three ecoregions (Great Basin, eastern sagebrush, and Great Plains) for nine invasive plants of interest for early detection and rapid response at local or regional scales. We compared the performance of national versus regional models using spatially independent withheld test data from each of the three ecoregions. Results : We found that models trained using a national spatial extent tended to perform better than regionally trained models. Regional models did not outperform national ones even when considerable occurrence data were available for model estimation within the focal region. Information was often unavailable to fit informative regional models precisely in those areas of greatest interest for early detection and rapid response. Main conclusions : Habitat suitability models for invasive plant species trained at a continental extent can reduce extrapolation while maximizing information on species’ responses to environmental variation. Standard modeling methods can capture spatially varying limiting factors, while regional or hierarchical models may only be advantageous when populations differ in their responses to environmental conditions, a condition expected to be relatively rare at the expanding boundaries of invasive species’ distributions.

NeoBiota

Size distribution and reproductive phenology of the invasive Burmese python (Python molurus bivittatus) in the Greater Everglades Ecosystem, Florida, USA

The design of successful invasive species control programs is often hindered by the absence of basic demographic data on the targeted population. Establishment of invasive Burmese pythons ( Python molurus bivittatus ) in the Greater Everglades Ecosystem, Florida USA has led to local precipitous declines (> 90%) of mesomammal populations and is also a major threat to native populations of reptiles and birds. Efforts to control this species are ongoing but are hampered by the lack of access to and information on the expected biological patterns of pythons in southern Florida. We present data from more than 4,000 wild Burmese pythons that were removed in southern Florida over 26 years (1995–2021), the most robust dataset representing this invasive population to date. We used these data to characterize Burmese python size distribution, size at maturity, clutch size, and seasonal demographic and reproductive trends. We broadened the previously described size ranges by sex and, based on our newly defined size-stage classes, showed that males are smaller than females at sexual maturity, confirmed a positive correlation between maternal body size and potential clutch size, and developed predictive equations to facilitate demographic predictions. We also refined the annual breeding season (approx.100 days December into March), oviposition timing (May), and hatchling emergence and dispersal period (July through October) using correlations of capture morphometrics with observations of seasonal gonadal recrudescence (resurgence) and regression. Determination of reproductive output and timing can inform population models and help managers arrest population growth by targeting key aspects of python life history. These results define characteristics of the species in Florida and provide an enhanced understanding of the ecology and reproductive biology of Burmese pythons in their invasive Everglades range.

Florida

Burmese pythons in Florida: A synthesis of biology, impacts, and management tools

Burmese pythons ( Python molurus bivittatus ) are native to southeastern Asia, however, there is an established invasive population inhabiting much of southern Florida throughout the Greater Everglades Ecosystem. Pythons have severely impacted native species and ecosystems in Florida and represent one of the most intractable invasive-species management issues across the globe. The difficulty stems from a unique combination of inaccessible habitat and the cryptic and resilient nature of pythons that thrive in the subtropical environment of southern Florida, rendering them extremely challenging to detect. Here we provide a comprehensive review and synthesis of the science relevant to managing invasive Burmese pythons. We describe existing control tools and review challenges to productive research, identifying key knowledge gaps that would improve future research and decision making for python control.

Florida

Capsaicin-treated bait is ineffective in deterring non-target mammals from trap disturbance during invasive lizard control

Excluding non-target species from invasive species control efforts can be challenging due to non-target attraction to trap structure, baits, and lures. Various methods have been used to deter non-target species from entering or disturbing traps including altered features (e.g., mesh size, trip mechanism, or entrances), staking traps, and chemical deterrents. Invasive populations of Argentine Black and White Tegu lizards ( Salvator merianae ) occur in several locations across Florida and Georgia, and there are ongoing trapping efforts to control them. At sites in Georgia, non-target mammals disturb most of the lizard traps (>80%), consume egg bait/lures, and thus reduce trap efficacy. In contrast, our Florida site has fewer problems with non-target mammals. Our goal was to quantify the efficacy of capsaicin-coated eggs, a known distasteful irritant to mammals, as a non-target bait deterrent in live traps set for tegus in both Georgia and Florida. We conducted feeding assays on three tegus and found that individuals readily consumed food coated in capsaicin. We then conducted a three-part, live trapping experiment to test 1) if trap disturbance by mammals habituated to eggs without capsaicin decreased when capsaicin-coated eggs were deployed in Georgia, 2) if mammals not habituated to eggs as bait (treated or untreated) disturbed live traps at the same rate as those habituated to eggs in Georgia, and 3) if tegu capture rates were different when capsaicin treated eggs were deployed in Florida. In Georgia, we found that trap disturbance by non-target mammals did not decrease when capsaicin was applied to eggs in an area previously habituated to trapping with this bait nor when applied in a novel area. In Florida, we found no significant difference in tegu captures using capsaicin-treated vs. untreated bait. Tegus were tolerant of capsaicin, but capsaicin treated eggs did not reduce non-target mammal disturbance to traps. Therefore, removal of invasive populations could be problematic if methods to reduce trap disturbance by non-targets are not identified and deployed.

NeoBiota

Limitations of invasive snake control tools in the context of a new invasion on an island with abundant prey

In October 2020, a new population of invasive brown treesnakes ( Boiga irregularis ) was discovered on the 33-ha Cocos Island, 2.5 km off the south coast of Guam. Cocos Island is a unique conservation resource, providing refuge for many lizards and birds, including endangered species, which were extirpated from mainland Guam by invasive predators including brown treesnakes. We sought to evaluate the usefulness of toxic baiting with acetaminophen-treated carrion baits and cage trapping, common tools for the control of brown treesnakes on mainland Guam, as potential eradication tools on Cocos Island. We evaluated multiple bait types and bait presentations: on the ground, suspended in the canopy emulating aerial bait applications and in four plastic-tube bait station configurations intended to exclude non-target species. We monitored all baits with time-lapse cameras. Despite improved exclusion of non-targets by bait station design, most baits were quickly removed by non-target species, particularly coconut crabs ( Birgus latro ) and Mariana monitors ( Varanus tsukamotoi ). Monitoring of 1,250 baits available for 2,427 bait nights resulted in no observations of brown treesnakes taking any bait. Subsequently, we tested two trap types commonly used on Guam and compared trapping success with live versus dead mouse lures. In 10,553 trap nights using live and dead mouse lures, we only captured one brown treesnake, in a trap with a live mouse lure. These baiting and trapping rates are so low as to be ineffectual for all practical purposes. Concurrent visual searching and hand capture of brown treesnakes during initial rapid response efforts demonstrates that these low baiting and trapping success rates are not a result of low snake density. We make a case for our assumption that the ineffectiveness of these tools on Cocos Island is due to the context of extremely high abundance of preferred live prey, primarily large geckos and birds. Our results have profound conservation ramifications, because any future island invasions by brown treesnakes are likely to occur within similarly prey-rich environments where these baiting and trapping methods might be similarly ineffective.

NeoBiota

Outcomes of control and monitoring of a widespread riparian invader (Tamarix spp.): A comparison of synthesis approaches

Effective ecological restoration requires empirical assessment to determine outcomes of projects, but conclusions regarding the effects of restoration treatments on the whole ecosystem remain rare. Control of invasive shrubs and trees in the genus Tamarix and associated riparian restoration in the American Southwest has been of interest to scientists and resource managers for decades; dozens of studies have reported highly variable outcomes of Tamarix control efforts, as measured by a range of response variables, temporal and spatial scales and monitoring strategies. We conducted a literature search and review, meta-analysis and vote count (comparison of numerical outcomes lacking reported variances and/or sample sizes) on published papers that quantitatively measured a variety of responses to control of Tamarix . From 96 publications obtained through a global search on terms related to Tamarix control, we found 52 publications suitable for a meta-analysis (n = 777 comparisons) and 63 publications suitable for two vote counts (n = 1,460 comparisons total; 622 comparisons reported as statistically significant) of response to Tamarix control. We estimated responses to control by treatment type (e.g. cut-stump treatment, burning, biocontrol) and ecosystem component (e.g. vegetation, fauna, fluvial processes). Finally, we compared results of the various synthesis methods to determine whether the increasingly stringent requirements for inclusion led to biased outcomes. Vegetation metrics, especially measures of Tamarix response, were the most commonly assessed. Ecosystem components other than vegetation, such as fauna, soils and hydrogeomorphic dynamics, were under-represented. The meta-analysis showed significantly positive responses by vegetation overall to biocontrol, herbicide and cut-stump treatments. This was primarily due to reduction of Tamarix cover; impacts on replacement vegetation were highly variable. We found concordance amongst our varied synthesis approaches, indicating that increased granularity from stricter quantitative techniques does not come at the cost of a biased sample. Overall, our results indicate that common control methods are generally effective for reducing Tamarix , but the indirect effects on other aspects of the ecosystem are variable and remain understudied. Given that this is a relatively well-studied invasive plant species, our results also illustrate the limitations of not only individual studies, but also of reviews for measuring the impact of invasive species control. We call on researchers to investigate the less commonly studied responses to Tamarix control and riparian restoration including the effects on fauna, soil and hydrogeomorphic characteristics.

NeoBiota

Integrating social-ecological outcomes into invasive species management: The Tamarix case

Incorporating societal considerations into decisions related to invasive species management is desirable, but can be challenging because it requires a solid understanding of the ecological functions and socio-cultural and economic benefits and values of the invaded environment before and after invasion. The ecosystem service ( ES ) concept was designed to facilitate such decision-making by establishing direct connections between ecosystem properties and human well-being, but its application in invasive species management has not been systematic. In this Discussion paper, we propose the adoption of the ES cascade model as a framework for understanding the environmental effects, costs and benefits associated with controlling an invasive shrub ( Tamarix spp.) in riparian systems of the western United States. The cascade model has the advantage of explicitly dissecting social-ecological systems into five components: ecosystem structure and processes, ecological functions, ecosystem services, benefits and the economic and socio-cultural valuation of these services and benefits. The first two have received significant attention in the evaluation of Tamarix control effectiveness. The last three have long been implicitly acknowledged over decades of Tamarix management in the region, but have not been formally accounted for, which we believe would increase the effectiveness, accountability and transparency of management efforts.

NeoBiota

Evaluating spatially explicit management alternatives for an invasive species in a riverine network

Invasive species have substantial ecological and economic costs and removing them can require large investments by management agencies. Optimal spatial allocation of removal effort is critical for efficient and effective management of invasive species. Using a series of ecologically informed model simulations, we evaluated and compared different spatially explicit removal strategies for invasive rusty crayfish ( Faxonius rusticus ) in the John Day River, USA. We assessed strategies in terms of their performance on three likely management objectives: suppression (minimise overall population abundance), containment (minimise the spatial extent of invasion) and prevention (minimise spread into a specific area). We developed five spatial removal strategies to achieve those objectives, denoted as: Target Abundance (removal at locations with the highest population abundance), Target Growth (removal at locations with the highest population growth), Target Edges (removal at the most distant locations in the river), Target Downstream (removal at the most downstream invaded segments on the Mainstem), and Target Random (removal at randomly selected locations). Each strategy was assessed at various effort levels, referring to the number of spatial segments in the river in which removals were conducted, after seven years of management. We identified the alternative that best achieved each objective, based on decision criteria for risk-neutral and risk-averse decision-makers and further evaluated strategies based on Pareto efficiency, which identifies the set of alternatives for which an improvement on one objective cannot be had without a decline in performance on another. We found that Target Abundance and Target Growth strategies best achieved the suppression objective, for risk neutral and risk averse decision-makers, respectively and Target Downstream was always best in achieving the prevention objective across both types of decision-makers. No single strategy consistently performed best in terms of the containment objective. In terms of all three objectives, Target Downstream was consistently Pareto efficient across all levels of management effort and both decision criteria. The modelling framework we provided is adaptable to a variety of riverine invasive species to help assess and compare spatial management strategies.

Oregon