Research Article |
Corresponding author: Thomas I. Gibson ( tom.gibson@cefas.gov.uk ) Academic editor: Angela Brandt
© 2025 Thomas I. Gibson, Rebecca S. Millard, Isla MacMillan, Nick Taylor, Mark Thrush, Hannah Tidbury.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gibson TI, Millard RS, MacMillan I, Taylor N, Thrush M, Tidbury H (2025) Application of a theoretical simulator to the optimisation of risk-based invasive species surveillance. NeoBiota 97: 19-46. https://doi.org/10.3897/neobiota.97.121188
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Early detection and rapid response are critical to the successful management of non-indigenous species (NIS) and rely on effective surveillance programmes. Risk-based surveillance, where surveillance targets high risk locations, is the most efficient form of NIS surveillance. However, further research is required on the impact of different levels of emphasis on risk, in sampling designs and on surveillance efficacy. This study implements a theoretical surveillance simulator to model the relative merit of different surveillance strategies with different levels of focus on NIS risk for NIS detection at one or more sites. Three potential surveillance scenarios were modelled: random, risk-based and heavy risk-based surveillance, each with three distributions of combined NIS risks of introduction and establishment: exponential, random and uniform. An example analysis using model derived NIS risk data is also provided. Sensitivity and elasticity analyses were conducted to identify variables which influence model outputs. The interaction between sampling method detection probability and changes in NIS abundance was modelled. It was found that NIS risk distribution influences the relative performance of different surveillance strategies and that risk- and heavy risk-based surveillance have lower times to detections and, generally, higher surveillance probabilities of detection compared to random surveillance at more skewed NIS risk distributions. However, there was a trade-off between short detection time and detection failure in risk-based and particularly heavy risk-based surveillance. Therefore, an over-emphasis on risk-based surveillance could provide suboptimal NIS detection. Sensitivity and elasticity analysis showed that the number of NIS seed sites, mean site visit rate and method detection probability had the largest effects on detection time, highlighting the complexity of designing surveillance programmes. In conclusion, the optimal surveillance strategy is conditional on the risk distribution and this study highlights the value of model-based simulators to guide decision-making in the design of NIS surveillance programmes.
Establishment risk, introduction risk, non-indigenous species, risk-based surveillance, surveillance design, theoretical model
Non-indigenous species (NIS) are species which have spread to new regions outside their natural biogeographical range with the aid of human actions (
NIS introductions occur via various pathways. Five key pathways in the marine environment are commercial shipping, recreational boating, movement of aquaculture stock, the aquarium trade and natural dispersal (
Early detection and rapid response rely on effective surveillance programmes which must be in place prior to NIS arrival to allow early detection (
Computer simulations which compare the effect of using simulated risk-based surveillance designs to random and other surveillance designs, on parameters which are of importance to NIS surveillance, such as time to detection or detection probability, provide a useful method to address this knowledge gap. These models vary in their sophistication and have been used in research into invasive plant pathogens (
This study develops and implements a theoretical model, referred to as a surveillance simulator, to assess the relative merit of different surveillance strategies, which differ in their level of risk focus, for NIS detection. Although developed for early warning monitoring of marine NIS, where the species is established at a relatively small number of sites, the simulator is generic and can be applied to any terrestrial or aquatic organisms while requiring a minimal amount of species-specific data. The simulator calculates the time to NIS detection across multiple simulations, following the introduction and establishment of a NIS at one or more sites. The survey probability of detection, over time, is also calculated across simulations. Differential risk of introduction and establishment between sites is incorporated. Surveillance is simulated under three potential scenarios: random surveillance, risk-based surveillance and heavy risk-based surveillance. For risk-based surveillance, the visit rate is increased by the relative risk of NIS introduction and establishment. For heavy risk-based surveillance, this relative increase is enhanced for the highest risk sites. The simulator also incorporates the interaction between the detection probability of a method and changes in the abundance of NIS. Sensitivity and elasticity analyses are performed to determine the effect of changes in selected parameters on time to NIS detection and the failure to detect NIS. Findings are discussed in the context of optimisation of surveillance for NIS and the operation of the model rather than providing detailed differences between theoretical scenarios. Application of the model is further illustrated using NIS risk scores for 10,249 sites, derived from model predictions based on empirical data, for three scenarios focused on: risk of introduction, risk of spread and risk of impact and representing three different risk distributions.
The simulator was developed in the statistical software R v.4.1.2 (
Schematic of the overall simulator structure showing key inputs and outputs and the role of the elasticity and sensitivity analysis, around the core surveillance simulation. The detailed structure of the surveillance simulations is given in Fig.
Parameter/ Term | Description* |
---|---|
Risk-Based Surveillance | Surveillance strategy where the site visit rate is biased towards higher risk sites. |
Heavy Risk-Based Surveillance | Surveillance strategy where the site visit rate is heavily biased towards higher risk sites. |
Introduction risk probability distribution | The statistical distribution which determines the probability of NIS introduction at a site. |
Establishment risk probability distribution | The statistical distribution which determines the probability of NIS establishment at a site. |
NIS risk | The probability of NIS introduction and establishment at a site, calculated by multiplying the introduction and establishment probability together. |
Surveillance time period | The maximum time period (in years) over which a simulation may run. |
Seed site(s) | A site(s) into which a NIS becomes introduced and established based on its relative NIS risk during a simulation. |
Mean site visit rate | The mean number of times which a site is visited in a year. |
Method detection probability | The probability of detecting a NIS at a site when it is searched during a simulation. |
Survey probability of detection | The probability of detecting NIS at a site(s), at a given time point by a simulation, as calculated using all simulations in a simulator run. |
Detection dynamic | The relationship between method detection probability and the abundance of a NIS. Either fixed, threshold or linear. |
Detection summary | Method used to summarise the time to detection if multiple seed sites are used in a simulation. |
The input parameters are controlled via the config_sim.yaml file. The user specifies the number of sites and a probability of introduction (getIntroProbability) and establishment per site (getEstablishProbability). The distributions from which to randomly draw probabilities of introduction and establishment are either: an equal uniform distribution which requires a user specified probability value, random uniform distribution, truncated normal distribution (bounded by 0 and 1), truncated exponential (bounded by 0 and 1) or lognormal distribution (bounded by 0 and 1). Example distributions, used in the later simulator application example, are shown in Fig.
NIS risk distributions of the probability of NIS becoming introduced and established at a site, showing exponential (A), random uniform (B) and equal uniform (C) risk distributions used in the simulator application example.
A mean site visit rate is defined by the user and used to calculate the visit rate for each individual site. Under random surveillance, the visit rate for each individual site is identical. Under risk-based surveillance, the risk-based visit rate for each site (Vrs) is calculated as: Vrs = Vs ∙ (Nrs / Nrx-), where Vs is the visit rate per site and Nrx- is the overall mean NIS risk probability across sites. Under heavy risk-based surveillance, the visit rate for each site (Vhrs) is calculated in the same manner, but the site NIS risk and mean NIS risk across all sites are raised to the power of three: Vhrs = Vs ∙ (Nrs3 / Nrx-3). Therefore, under the risk-based surveillance scenarios, the simulator assigns a relatively higher visit rate to those sites with greater NIS risk. Higher visit rates at high-risk sites are further enhanced under heavy risk-based surveillance. See Fig.
A conceptual example of the relationship between the relative NIS risk at each site (numbers within hexagons, assuming an exponential distribution) and the site visit rate assuming random, risk-based and heavy risk-based surveillance, over three site visits (blue outline) during a model run. The highest risk hexagons in this example represent two port sites, one seeded with a NIS at the beginning of the simulation (orange fill). Under random surveillance three sites are visited with no relationship to risk, under risk-based, three high risk sites are visited and under heavy risk-based surveillance, the highest risk site, only, is visited three times.
The method detection probability defines the probability of the sampling method detecting the NIS during a site visit. The method detection probability may be fixed or vary with NIS abundance linearly or in a threshold manner. Under a linear relationship, the user defines the abundance required to change the detection probability by 0.01. Under a threshold relationship, the user defines a threshold abundance value and two detection probabilities to use when abundance is below, above or equal to the threshold value.
Introductions at multiple seed sites, up to the number of sites in the simulation, may also be selected by the user. If abundance is required, values for multiple sites are either set by the user or randomly drawn from a Poisson distribution with a user specified mean. For multiple sites, the user must select the detection summary method, i.e. how the time to detection is summarised over multiple seed sites in that simulation (ProcessMultipleResults). Time to detection may be taken from the first seed site to be detected or the last seed site.
The simulation is run by the function runSurveillanceSimulation (Fig.
Schematic of the surveillance simulation showing key steps and outputs, as defined by the runSurveillanceSimulation function.
At each time step, a single site is selected to be visited dependent on the mean visit rate (Fig.
Sensitivity analysis determines the impact that absolute changes in each model parameter have on the output, i.e. time taken to detect NIS and forms a component of the surveillance simulator. Sensitivity analysis can be implemented for the number of sites, number of years, mean visit rate, method detection probability and number of seed sites (makeSensitivityParamsTable). The simulator runs iteratively (runSurveillanceSensitivity; results formatted by formatSensitivityResults), incrementally altering input parameters, one at a time, by a user-defined interval within a specified range and plotting the results. Summary statistics such as number of times a NIS was detected/not detected and the mean, maximum and minimum time to detection are output for each parameter. Outputs are generated by the report-NIS-intro-detect-sensitivity.Rmd R Markdown file and other helper functions.
Elasticity analysis is also included in the simulator. Elasticity (ξ) is proportional sensitivity, it estimates the effect of a proportional change in a parameter on the proportional change in the output, i.e. time taken to detect NIS (
Three different NIS introduction and establishment risk distributions were implemented for each surveillance strategy (random, risk-based and heavy risk-based). These risk distributions were equal uniform (probability: 0.8), random uniform and exponential. The equal uniform distribution was selected to provide a default example with no variation in risk. The random uniform distribution provided a scenario where risk varied between sites, whereas the exponential distribution was used to represent a situation where most sites are of no or low risk and a small number of sites are of high risk (
For sensitivity and elasticity analysis, the exponential risk distribution was used as, under this risk distribution, the largest differences between sampling programmes were seen. For the sensitivity analysis, the number of seed sites, survey sites, years, mean visit rate and detection probability were run with selected parameters defined, based on the authors’ knowledge of sampling programmes (Table
Parameters | Sensitivity Values | Elasticity Values | ||||
---|---|---|---|---|---|---|
Minimum | Maximum | Interval | Default | 25% Decrease | 25% Increase | |
Number of Seed Sites | 1 | 100 | 10 | |||
Number of Survey Sites | 50 | 200 | 25 | 100 | 75 | 125 |
Number of Years | 10 | 50 | 5 | 30 | 22.5 | 37.5 |
Mean Visit Rate | 0.25 | 4 | 0.25 | 1 | 0.75 | 1.25 |
Method Detection Probability | 0.1 | 1 | 0.1 | 0.8 | 0.6 | 1.0 |
The effect of dynamic detection (where the method detection probability is linked to NIS abundance) was explored using an exponential risk distribution and with seed site set to 1 and 10. The abundance model parameters assumed a starting population of 1, intrinsic growth rate of 1.5 with logistic growth and a population carrying capacity at each site of 100,000 individuals. For a linear relationship between abundance and detection method sensitivity, the starting detection method sensitivity was set to 0.1 with an increase of 0.01 per abundance increase of 500, up to 0.8. For a threshold relationship between abundance and detection method sensitivity, an abundance threshold of 10,000 was set such that method detection sensitivity below and above this threshold was 0.1 and 0.8, respectively. Parameter values were arbitrarily selected to demonstrate the functionality of the simulator.
As a practical example, the surveillance simulator was used to assess the relative performance of random, risk-based and heavy risk-based surveillance for three marine NIS introduction and spread scenarios created by a model to prioritise surveillance activities for NIS species for the UK coastline (the ‘Site Prioritisation Tool’ or SPT model, Cefas, in prep.). This hierarchical model was developed to provide information for surveillance programmes by scoring and ranking 10,249/5 km × 5 km grid squares representing the UK coastline. Empirical data for a range of risk parameters was grouped into pathways, distributed amongst four risk categories: Introduction risk (pathways: intentional introduction, shipping, recreational boating, fishery and aquaculture release), Establishment risk (temperature, salinity and substrate), Impact risk (environment and industry) and Spread risk (recreational boating, fishery and aquaculture release; Suppl. material
Risk distributions of the combined probability of marine NIS becoming introduced and established at a site, spread from that site and the site being negatively impacted. Scores generated using the SPT model (Cefas, in prep.) for Scenario A shipping risk weighted, Scenario B spread risk weighted and Scenario C impact risk weighted.
Comparison of the time to detection between different risk distributions showed that results varied with surveillance strategy (Table
The overall survey detection probability of NIS over time, calculated across 10,000 simulations, assuming an exponential (A), random uniform (B) and equal uniform (C) risk distribution.
Model Run | Distribution | Detection Dynamic | Number of Seed Sites | Scenario | Detection Time (Years) | Detection Failure (%) | Survey Probability of Detection at Time (Years) | ||||
---|---|---|---|---|---|---|---|---|---|---|---|
Median Detection Time | Interquartile Range | 1 | 5 | 10 | 30 | ||||||
Run 1 | Exponential | Constant | 1 | Random | 0.85 | 1.34 | 0.00 | 0.56 | 0.98 | 1.00 | 1.00 |
Risk-Based | 0.41 | 0.85 | 0.28 | 0.75 | 0.96 | 0.99 | 1.00 | ||||
Heavy Risk-Based | 0.47 | 1.86 | 10.68 | 0.57 | 0.78 | 0.83 | 0.89 | ||||
Run 2 | Random Uniform | 1 | Random | 0.89 | 1.39 | 0.00 | 0.54 | 0.98 | 1.00 | 1.00 | |
Risk-Based | 0.56 | 1.01 | 0.15 | 0.68 | 0.97 | 0.99 | 1.00 | ||||
Heavy Risk-Based | 0.52 | 1.45 | 7.49 | 0.61 | 0.84 | 0.88 | 0.93 | ||||
Run 3 | Equal Uniform | 1 | Random | 0.87 | 1.39 | 0.00 | 0.54 | 0.98 | 1.00 | 1.00 | |
Risk-Based | 0.86 | 1.36 | 0.00 | 0.56 | 0.98 | 1.00 | 1.00 | ||||
Heavy Risk-Based | 0.88 | 1.35 | 0.00 | 0.55 | 0.98 | 1.00 | 1.00 | ||||
Run 4 | Exponential | Linear | 1 | Random | 6.03 | 4.39 | 0.00 | 0.09 | 0.40 | 0.97 | 1.00 |
Risk-Based | 3.34 | 5.03 | 0.50 | 0.20 | 0.63 | 0.95 | 0.99 | ||||
Heavy Risk-Based | 3.55 | 6.70 | 12.48 | 0.24 | 0.49 | 0.76 | 0.88 | ||||
Run 5 | 10 | Random | 8.79 | 1.78 | 0.00 | 0.00 | 0.00 | 0.78 | 1.00 | ||
Risk-Based | 8.31 | 3.51 | 3.02 | 0.00 | 0.02 | 0.68 | 0.97 | ||||
Heavy Risk-Based | 14.52 | 11.25 | 68.87 | 0.00 | 0.00 | 0.08 | 0.31 | ||||
Run 6 | Exponential | Threshold | 1 | Random | 6.29 | 4.23 | 0.00 | 0.09 | 0.39 | 0.97 | 1.00 |
Risk-Based | 3.20 | 5.20 | 0.29 | 0.21 | 0.63 | 0.96 | 1.00 | ||||
Heavy Risk-Based | 3.63 | 6.54 | 12.39 | 0.24 | 0.49 | 0.76 | 0.88 | ||||
Run 7 | 10 | Random | 8.61 | 1.79 | 0.00 | 0.00 | 0.00 | 0.81 | 1.00 | ||
Risk-Based | 8.14 | 3.54 | 3.16 | 0.00 | 0.01 | 0.69 | 0.97 | ||||
Heavy Risk-Based | 14.71 | 11.45 | 67.67 | 0.00 | 0.00 | 0.09 | 0.32 |
Assuming an exponential risk distribution, an increase in the number of seed sites from 1 to 20 led to an increase in median time to detection across surveillance scenarios: from 0.87 to 3.6 years for random surveillance, 0.35 to 4.11 years for risk-based surveillance and 0.43 to 17.61 years for heavy risk-based surveillance (Fig.
The results of the sensitivity analysis, assuming an exponential risk distribution, for the parameters: number of seed sites (A), number of sampling sites (B), number of sampling years (C), mean visit rate (D) and method detection probability (E), showing their effect on median time to detection (left hand column) and the percentage of simulations in each model run where no NIS was detected (right hand column).
The number of sampling sites had little impact on the median time to detection or the detection failure of random or risk-based surveillance (Fig.
Assuming an exponential distribution, the elasticity of median time to detection and detection failure varied between surveillance scenarios, parameters and the direction of change in parameter values (Fig.
The elasticity of the median time to detection (years; A) and the detection failure (%; B) to a 25% increase and decrease in the default values of mean visit rate, number of sampling sites, number of years and method detection probability in each model run, assuming an exponential risk distribution.
Assuming an exponential risk distribution, inclusion of a linear relationship between method detection probability and NIS abundance (i.e. a linear detection dynamic), which grew logistically, resulted in a similar pattern of median time to detection and detection failure, between surveillance scenarios, for one seed site (model run 4; Table
The survey probability of detection of NIS at a site, or all sites, over time, assuming an exponential NIS risk distribution, calculated across 10,000 simulations, for a linear (panels A and B) and threshold detection dynamic (panels C and D) between NIS abundance and method detection probability for one (left hand panels) and ten seed sites (right hand panels).
The overall survey detection probability of NIS over time, calculated across 1000 simulations, assuming the risk distribution in the combined probability of NIS becoming introduced and established at a site, spread from that site and the site being negatively impacted. Scores generated using the ‘Site Prioritisation Tool’ (Cefas, in prep.) for Scenario A shipping risk weighted, Scenario B spread risk weighted and Scenario C impact risk weighted.
Assuming a threshold detection dynamic between method detection probability and NIS abundance, simulations with one seed site (model run 6) produced similar median times to detection and detection failure to a linear detection dynamic with one site (model run 4), across surveillance scenarios (Table
The time to detection and other outputs varied between surveillance strategies for each SPT modelled site risk distribution (Table
Model Run | Surveillance Scenario | Detection Time (Years) | Detection Failure (%) | Survey Probability of Detection at Time (Years) | ||||
---|---|---|---|---|---|---|---|---|
Median Detection Time | Interquartile Range | 1 | 5 | 10 | 30 | |||
Scenario A | Random | 0.93 | 1.47 | 0.00 | 0.53 | 0.98 | 1.00 | 1.00 |
Shipping Risk Weighted | Risk-Based | 0.25 | 0.47 | 0.00 | 0.90 | 0.99 | 1.00 | 1.00 |
Heavy Risk-Based | 0.37 | 0.94 | 4.40 | 0.70 | 0.89 | 0.93 | 0.96 | |
Scenario B | Random | 0.95 | 1.36 | 0.00 | 0.52 | 0.98 | 1.00 | 1.00 |
Spread Risk Weighted | Risk-Based | 0.34 | 0.66 | 0.30 | 0.81 | 0.97 | 0.99 | 1.00 |
Heavy Risk-Based | 0.52 | 1.85 | 8.10 | 0.56 | 0.83 | 0.87 | 0.92 | |
Scenario C | Random | 0.78 | 1.33 | 0.00 | 0.58 | 0.99 | 1.00 | 1.00 |
Impact Risk Weighted | Risk-Based | 0.34 | 0.68 | 0.10 | 0.80 | 0.96 | 0.98 | 1.00 |
Heavy Risk-Based | 0.27 | 0.63 | 7.70 | 0.74 | 0.86 | 0.89 |
Variation in the risk of NIS introduction and establishment between survey sites (NIS risk distribution) and the level of risk focus which the surveillance strategy adopts have important implications for optimising NIS detection. This study shows that the relative performance of surveillance strategies changes with NIS risk distributions derived both theoretically and with model estimates from the SPT model. Generally, under risk- and heavy risk-based surveillance, time to detection was shorter and survey probability of detection was greater than random surveillance for sites with random or exponential NIS risk distributions. For example, assuming an exponential risk distribution, risk-based surveillance detected NIS twice as fast as random surveillance. Risk-based surveillance also had a substantially higher survey probability of detection after 1 year compared to random and heavy risk-based surveillance. This observation generally held for risk and heavy risk-based surveillance for the marine NIS risk distributions derived from the SPT model, with risk-based and heavy risk-based surveillance having the shortest detection times and highest detection probabilities after 1 year across all three scenarios. This is comparable to the performance between risk-based and random surveillance in other studies (
An over-emphasis on the highest risk sites can, in some instances, lead to a failure to detect NIS with little benefit in terms of reduced detection time. Concentrating on a small number of sites has also been shown to be detrimental by a spatially-explicit plant pathogen model (
The model assumes that the risk distribution of sites can be effectively quantified to guide surveillance. Typically, this information is uncertain, particularly for newly-recorded and poorly-understood NIS. However, there is often enough data for effective survey design (
Sensitivity and elasticity analyses were performed in parallel to allow both the absolute effect of parameter changes on outputs to be examined and the impact of parameter changes to be compared across parameters. These analyses highlighted the key factors which should be considered when designing a surveillance strategy and the utility of the simulator to explore different approaches. For an exponential risk distribution, the number of seed sites, mean visit rate and method detection probability had the strongest effect on detection time, whereas the effects of all parameters on detection failure were more variable. Differential responses of surveillance strategies occurred between risk distributions. When seed site numbers were greater than one, heavy risk-based surveillance performed poorly for time to detection and detection failure, relative to risk-based and random surveillance. It is possible for NIS to establish at multiple sites early in an invasion (
The simulator is an efficient and valuable tool for planning surveillance programmes. While outside the scope of the current study, several opportunities exist for further development into the future. For example, a spatially-explicit model, incorporating NIS distribution and spread of NIS over time, would allow study of how the spatial distribution of sampled sites influences the utility of risk-based surveillance. Spatially-explicit models of pathogen entry and spread have shown that spatial correlations in risk can make it suboptimal to focus on the highest risk sites and a geographic spread of resources to cover all areas of risk is desirable (
In conclusion, variation in the risk of NIS introduction and establishment and the level of risk focus of surveillance programmes interact to influence the efficacy of surveillance regimes. Assuming a skewed risk distribution, an over-emphasis on sampling high risk sites will be outperformed by a more balanced focus on high as well as lower risk sites. However, the optimum approach is dependent on the NIS risk distribution. The relative risk of sites and other survey parameters, has to be quantified for the optimal surveillance design to be selected. Overall, this study highlights the utility of model-based simulators to guide decision-making in the design of the surveillance of NIS and other hazards.
The authors would like to acknowledge our colleague at CEFAS, Mickael Teixeira Alves, for informative discussions on elasticity analysis and quality controlling the manuscript and code. In addition, we would like to thank the two reviewers and editor for their insightful comments on a previous version of this manuscript.
The authors have declared that no competing interests exist.
The authors see no ethical implications of this research. All data created by the simulator was synthetic. For the data used by the SPT model, open source data or data available on request was used. No personal data was used in this study.
This research was funded by The Department for Food Agriculture and Rural Affairs (UK Government). Grant number: C8389 (R&D NIS Surveillance).
Conceptualisation: HT. Methodology: HT, NT, TG, RM, MT, IM. Software: RM, TG, NT. Investigation: TG, RM, MT, IM. Writing - Original draft: TG. Writing - Review and Editing: TG, HT, RM, MT, IM. Visualisation: TG, RM, IM. Supervision: HT. Funding Acquisition: HT.
Thomas I. Gibson https://orcid.org/0000-0002-8732-8716
Rebecca S. Millard https://orcid.org/0000-0002-3896-322X
Isla MacMillan https://orcid.org/0000-0002-8654-6068
Mark Thrush https://orcid.org/0000-0001-7335-406X
Hannah Tidbury https://orcid.org/0000-0001-8549-3518
The simulator code is located in Cefas’ reproducible research account on GitHub in the “C8389-NIS-surveillance-simulator” repository (https://github.com/CefasRepRes/C8389-NIS-surveillance-simulator). The data and outputs, including example mark-down files from theoretical and SPT model runs used in this publication, are available via Zenodo (https://zenodo.org/doi/10.5281/zenodo.10355963). For the SPT model runs, the adjusted code is included along with the data.
Abundance models, exponential growth model, logistic growth model, elasticity analysis, table and figure
Data type: docx
Explanation note: Abundance Models; Exponential Growth Model; Logistic Growth Model; Elasticity Analysis; fig. S1. Spatial representation of the site risk scores output from the Site Prioritisation Tool; table S1. Data sources for each parameter used in the Site Prioritisation Tool.