Risks posed by invasive species to the provision of ecosystem services in Europe.

Invasive species significantly impact biodiversity and ecosystem services, yet understanding these effects at large spatial scales remains a challenge. Our study addresses this gap by assessing the current and potential future risks posed by 94 invasive species to seven key ecosystem services in Europe. We demonstrate widespread potential impacts, particularly on outdoor recreation, habitat maintenance, crop provisioning, and soil and nitrogen retention. Exposure to invasive species was higher in areas with lower provision of ecosystem services, particularly for regulating and cultural services. Exposure was also high in areas where ecosystem contributions to crop provision and nitrogen retention were at their highest. Notably, regions vital for ecosystem services currently have low invasion suitability, but face an average 77% increase in potential invasion area. Here we show that, while high-value ecosystem service areas at the highest risk represent a small fraction of Europe (0-13%), they are disproportionally important for service conservation. Our study underscores the importance of monitoring and protecting these hotspots to align management strategies with international biodiversity targets, considering both invasion vulnerability and ecosystem service sustainability.

Using the IUCN Environmental Impact Classification for Alien Taxa to inform decision-making.

The Environmental Impact Classification for Alien Taxa (EICAT) is an important tool for biological invasion policy and management and has been adopted as an International Union for Conservation of Nature (IUCN) standard to measure the severity of environmental impacts caused by organisms living outside their native ranges. EICAT has already been incorporated into some national and local decision-making procedures, making it a particularly relevant resource for addressing the impact of non-native species. Recently, some of the underlying conceptual principles of EICAT, particularly those related to the use of the precautionary approach, have been challenged. Although still relatively new, guidelines for the application and interpretation of EICAT will be periodically revisited by the IUCN community, based on scientific evidence, to improve the process. Some of the criticisms recently raised are based on subjectively selected assumptions that cannot be generalized and may harm global efforts to manage biological invasions. EICAT adopts a precautionary principle by considering a species' impact history elsewhere because some taxa have traits that can make them inherently more harmful. Furthermore, non-native species are often important drivers of biodiversity loss even in the presence of other pressures. Ignoring the precautionary principle when tackling the impacts of non-native species has led to devastating consequences for human well-being, biodiversity, and ecosystems, as well as poor management outcomes, and thus to significant economic costs. EICAT is a relevant tool because it supports prioritization and management of non-native species and meeting and monitoring progress toward the Kunming-Montreal Global Biodiversity Framework (GBF) Target 6.

Uso de la Clasificación de Impacto Ambiental de los Taxones Exóticos de la UICN para la toma de decisiones Resumen La Clasificación de Impacto Ambiental de los Taxones Exóticos (EICAT, en inglés) es una herramienta importante para las políticas y manejo de las invasiones biológicas y ha sido adoptada como un estándar de la Unión Internacional para la Conservación de la Naturaleza (UICN) para medir la seriedad del impacto ambiental causado por los organismos que viven fuera de su extensión nativa. La EICAT ya ha sido incorporada a algunos procedimientos locales y nacionales de toma de decisiones, lo que la vuelve un recurso particularmente relevante para abordar el impacto de las especies no nativas. Algunos principios conceptuales subyacentes de la EICAT han sido cuestionados recientemente, en particular aquellos relacionados con el uso del principio de precaución. Aunque todavía son relativamente nuevas, las directrices para la aplicación e interpretación de la EICAT tendrán una revisión periódica, basada en evidencia científica, por parte de la comunidad de la UICN para mejorar el proceso. Algunas de las críticas recientes están basadas en suposiciones seleccionadas subjetivamente que no pueden generalizarse y podrían perjudicar los esfuerzos globales para manejar las invasiones biológicas. La EICAT adopta un principio de precaución cuando considera el historial de impacto de una especie en cualquier otro lugar ya que algunos taxones tienen características que podrían volverlos más dañinos. Además, las especies no nativas suelen ser factores de pérdida de bidiversidad, incluso bajo otras presiones. Cuando ignoramos el principio de precaución al abordar el impacto de las especies no nativas, hay consecuencias devastadoras para el bienestar humano, la biodiversidad y los ecosistemas, así como resultados pobres de conservación, y por lo tanto con costos económicos importantes. La EICAT es una herramienta relevante porque respalda la priorización y el manejo de las especies no nativas y ayuda con el cumplimiento y monitoreo del progreso para llegar al objetivo 6 del Marco Mundial de Biodiversidad Kunming-Montreal.

Identification of potential invasive alien species in Spain through horizon scanning.

Invasive alien species have widespread impacts on native biodiversity and ecosystem services. Since the number of introductions worldwide is continuously rising, it is essential to prevent the entry, establishment and spread of new alien species through a systematic examination of future potential threats. Applying a three-step horizon scanning consensus method, we evaluated non-established alien species that could potentially arrive, establish and cause major ecological impact in Spain within the next 10 years. Overall, we identified 47 species with a very high risk (e.g. Oreochromis niloticus, Popillia japonica, Hemidactylus frenatus, Crassula helmsii or Halophila stipulacea), 61 with high risk, 93 with moderate risk, and 732 species with low risk. Many of the species categorized as very high or high risk to Spanish biodiversity are either already present in Europe and neighbouring countries or have a long invasive history elsewhere. This study provides an updated list of potential invasive alien species useful for prioritizing efforts and resources against their introduction. Compared to previous horizon scanning exercises in Spain, the current study screens potential invaders from a wider range of terrestrial, freshwater, and marine organisms, and can serve as a basis for more comprehensive risk analyses to improve management and increase the efficiency of the early warning and rapid response framework for invasive alien species. We also stress the usefulness of measuring agreement and consistency as two different properties of the reliability of expert scores, in order to more easily elaborate consensus ranked lists of potential invasive alien species.

A horizon scan exercise for aquatic invasive alien species in Iberian inland waters.

As the number of introduced species keeps increasing unabatedly, identifying and prioritising current and potential Invasive Alien Species (IAS) has become essential to manage them. Horizon Scanning (HS), defined as an exploration of potential threats, is considered a fundamental component of IAS management. By combining scientific knowledge on taxa with expert opinion, we identified the most relevant aquatic IAS in the Iberian Peninsula, i.e., those with the greatest geographic extent (or probability of introduction), severe ecological, economic and human health impacts, greatest difficulty and acceptability of management. We highlighted the 126 most relevant IAS already present in Iberian inland waters (i.e., Concern list) and 89 with a high probability of being introduced in the near future (i.e., Alert list), of which 24 and 10 IAS, respectively, were considered as a management priority after receiving the highest scores in the expert assessment (i.e., top-ranked IAS). In both lists, aquatic IAS belonging to the four thematic groups (plants, freshwater invertebrates, estuarine invertebrates, and vertebrates) were identified as having been introduced through various pathways from different regions of the world and classified according to their main functional feeding groups. Also, the latest update of the list of IAS of Union concern pursuant to Regulation (EU) No 1143/2014 includes only 12 top-ranked IAS identified for the Iberian Peninsula, while the national lists incorporate the vast majority of them. This fact underlines the great importance of taxa prioritisation exercises at biogeographical scales as a step prior to risk analyses and their inclusion in national lists. This HS provides a robust assessment and a cost-effective strategy for decision-makers and stakeholders to prioritise the use of limited resources for IAS prevention and management. Although applied at a transnational level in a European biodiversity hotspot, this approach is designed for potential application at any geographical or administrative scale, including the continental one.

The EICAT+ framework enables classification of positive impacts of alien taxa on native biodiversity.

Species introduced through human-related activities beyond their native range, termed alien species, have various impacts worldwide. The IUCN Environmental Impact Classification for Alien Taxa (EICAT) is a global standard to assess negative impacts of alien species on native biodiversity. Alien species can also positively affect biodiversity (for instance, through food and habitat provisioning or dispersal facilitation) but there is currently no standardized and evidence-based system to classify positive impacts. We fill this gap by proposing EICAT+, which uses 5 semiquantitative scenarios to categorize the magnitude of positive impacts, and describes underlying mechanisms. EICAT+ can be applied to all alien taxa at different spatial and organizational scales. The application of EICAT+ expands our understanding of the consequences of biological invasions and can inform conservation decisions.

80 questions for UK biological security.

Multiple national and international trends and drivers are radically changing what biological security means for the United Kingdom (UK). New technologies present novel opportunities and challenges, and globalisation has created new pathways and increased the speed, volume and routes by which organisms can spread. The UK Biological Security Strategy (2018) acknowledges the importance of research on biological security in the UK. Given the breadth of potential research, a targeted agenda identifying the questions most critical to effective and coordinated progress in different disciplines of biological security is required. We used expert elicitation to generate 80 policy-relevant research questions considered by participants to have the greatest impact on UK biological security. Drawing on a collaboratively-developed set of 450 questions, proposed by 41 experts from academia, industry and the UK government (consulting 168 additional experts) we subdivided the final 80 questions into six categories: bioengineering; communication and behaviour; disease threats (including pandemics); governance and policy; invasive alien species; and securing biological materials and securing against misuse. Initially, the questions were ranked through a voting process and then reduced and refined to 80 during a one-day workshop with 35 participants from a variety of disciplines. Consistently emerging themes included: the nature of current and potential biological security threats, the efficacy of existing management actions, and the most appropriate future options. The resulting questions offer a research agenda for biological security in the UK that can assist the targeting of research resources and inform the implementation of the UK Biological Security Strategy. These questions include research that could aid with the mitigation of Covid-19, and preparation for the next pandemic. We hope that our structured and rigorous approach to creating a biological security research agenda will be replicated in other countries and regions. The world, not just the UK, is in need of a thoughtful approach to directing biological security research to tackle the emerging issues.

Priority setting for invasive species management by the water industry.

The expansion of Invasive Alien Species (IAS) is a growing concern to the UK water industry because of their diverse impacts on water quality, infrastructure and eradication costs. New regulations reinforcing the industry's responsibilities beyond operational costs, coupled with continued range expansion and establishment of new IAS will increase damages. To tackle IAS effectively, the water industry requires reliable information about which species pose the greatest risk to operations and which areas are most vulnerable to invasion. Here we assess potential biosecurity threats for the 24 water companies in the UK using well-established modelling research techniques such as risk assessment and distribution modelling. Using a consensus approach with environmental managers and water companies, we identified 11 IAS of concern for the UK water industry, including five plants, three crustaceans, two molluscs and one fish. These invaders pose important hazards in terms of water quality, flood protection, human health, integrity of infrastructures, recreational and aesthetic values, amongst others. We used distribution models to predict their potential expansion under current and future 2050 climate scenarios within each of the 24 water companies in the UK. Water companies in the South East of England (Cambridge Water, Anglian Water, Affinity Water and Thames Water) are under the highest risk of invasional meltdown from multiple IAS, both now and under future scenarios. The quagga mussel poses the most serious risk of immediate spread and may exacerbate the impacts of the widespread zebra mussel for the water industry. The information generated in this study can support the prioritization of species and regions at risk, so that funds for prevention and eradication of invasions are well allocated. Ultimately, this study demonstrates that scientific risk assessments, usually restricted to the academic and public sectors, can be extremely useful to guide decision-making by the private sector.

Small hydropower plants as a threat to the endangered pearl mussel Margaritifera margaritifera.

Freshwater mussels are undergoing rapid global declines due to habitat loss and fragmentation, among other factors, but little is known about the effects of small hydropower plants. Here we assessed the impact of small hydropower plants on the abundance and size structure of the imperilled pearl mussel Margaritifera margaritifera. For this, we sampled 66 sites in three Portuguese rivers (Mente, Rabaçal and Tuela) located upstream and downstream of dams and within the reservoirs. Pearl mussels were significantly more abundant upstream than downstream of dams (97.4% more) or within reservoirs (98.5% more). In addition, juveniles were mostly found upstream of dams. The most significant environmental alterations that explained the observed patterns were related to changes in sediment characteristics (accumulation of fine sediments and organic matter in reservoirs) and water chemistry, most notably suspended solids (highest values in reservoirs) and dissolved oxygen (lowest values in reservoirs). Overall, results show that small hydropower plants can deeply affect pearl mussel populations: specimens almost disappeared from the areas within the reservoirs and sites located downstream only retained adults without signs of recent recruitment. Future management measures devoted to the conservation of pearl mussels should take into account the results reported here to avoid the construction of new dams in pearl mussel rivers; improve management of the river flow in downstream areas; and consider the decommissioning of dams in pearl mussel rivers with a particular attention devoted to the re-naturalization of river sections under the influence of the reservoir and downstream areas.

Author Correction: A global database for metacommunity ecology, integrating species, traits, environment and space.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A global database for metacommunity ecology, integrating species, traits, environment and space.

The use of functional information in the form of species traits plays an important role in explaining biodiversity patterns and responses to environmental changes. Although relationships between species composition, their traits, and the environment have been extensively studied on a case-by-case basis, results are variable, and it remains unclear how generalizable these relationships are across ecosystems, taxa and spatial scales. To address this gap, we collated 80 datasets from trait-based studies into a global database for metaCommunity Ecology: Species, Traits, Environment and Space; "CESTES". Each dataset includes four matrices: species community abundances or presences/absences across multiple sites, species trait information, environmental variables and spatial coordinates of the sampling sites. The CESTES database is a live database: it will be maintained and expanded in the future as new datasets become available. By its harmonized structure, and the diversity of ecosystem types, taxonomic groups, and spatial scales it covers, the CESTES database provides an important opportunity for synthetic trait-based research in community ecology.

Developing a list of invasive alien species likely to threaten biodiversity and ecosystems in the European Union.

The European Union (EU) has recently published its first list of invasive alien species (IAS) of EU concern to which current legislation must apply. The list comprises species known to pose great threats to biodiversity and needs to be maintained and updated. Horizon scanning is seen as critical to identify the most threatening potential IAS that do not yet occur in Europe to be subsequently risk assessed for future listing. Accordingly, we present a systematic consensus horizon scanning procedure to derive a ranked list of potential IAS likely to arrive, establish, spread and have an impact on biodiversity in the region over the next decade. The approach is unique in the continental scale examined, the breadth of taxonomic groups and environments considered, and the methods and data sources used. International experts were brought together to address five broad thematic groups of potential IAS. For each thematic group the experts first independently assembled lists of potential IAS not yet established in the EU but potentially threatening biodiversity if introduced. Experts were asked to score the species within their thematic group for their separate likelihoods of i) arrival, ii) establishment, iii) spread, and iv) magnitude of the potential negative impact on biodiversity within the EU. Experts then convened for a 2-day workshop applying consensus methods to compile a ranked list of potential IAS. From an initial working list of 329 species, a list of 66 species not yet established in the EU that were considered to be very high (8 species), high (40 species) or medium (18 species) risk species was derived. Here, we present these species highlighting the potential negative impacts and the most likely biogeographic regions to be affected by these potential IAS.

Inter-basin water transfers and the expansion of aquatic invasive species.

Inter-basin Water Transfers (IBWT) are recognized as one of the major pathways of freshwater invasion. They provide a direct link between previously isolated catchments and may modify the habitat conditions of the receiving waters such that they become more favourable for the establishment of invasive species. Combined, IBWT and invasive species will intensify the stress upon native species and ecosystems. Using the Severn and Thames Rivers -two of the largest river systems in Great Britain-as a case study, here we assess the potential influence of IBWT on the expansion of invasive species and thus their impact on biodiversity conservation. The Thames Valley is subject to extensive water abstraction, and an increasing population means that supplemented flow from the River Severn is being considered. Multi-scale Suitability Models, based on climate and water chemistry respectively, provided novel evidence that there is serious risk for further spread of invasive species in the focus area, particularly of the quagga mussel, a recent invader of the Thames River. Native freshwater mussels are particularly vulnerable to changing environmental conditions, and may suffer the decrease in alkalinity and increase in sedimentation associated with an IBWT from the lower Severn to the upper Thames. Regional models suggest considerable overlap between the areas suitable for three vulnerable native freshwater mussels and the expansion of invasive species that negatively impact upon the native mussels. This study illustrates the use of novel spatially-explicit techniques to help managers make informed decisions about the risks associated with introducing aquatic invasive species under different engineering scenarios. Such information may be especially important under new legislation (e.g. EU Invasive Species Regulation No 1143/2014) which increases the responsibility of water managers to contain and not transfer invasive species into new locations.

Current and future effects of global change on a hotspot's freshwater diversity.

Deforestation, climate change and invasive species constitute three global threats to biodiversity that act synergistically. However, drivers and rates of loss of freshwater biodiversity now and in the future are poorly understood. Here we focus on the potential impacts of global change on freshwater mussels (Order Unionida) in Sundaland (SE Asia), a vulnerable group facing global declines and recognized indicators of overall freshwater biodiversity. We used an ensemble of distribution models to identify habitats potentially suitable for freshwater mussels and their change under a range of climate, deforestation and invasion scenarios. Our data and models revealed that, at present, Sundaland features 47 and 32 Mha of habitat that can be considered environmentally suitable for native and invasive freshwater mussels, respectively. We anticipate that by 2050, the area suitable for palm oil cultivation may expand between 8 and 44 Mha, representing an annual increase of 2-11%. This is expected to result in a 20% decrease in suitable habitat for native mussels, a drop that reaches 30% by 2050 when considering concomitant climate change. In contrast, the habitat potentially suitable for invasive mussels may increase by 44-56% under 2050 future scenarios. Consequently, native mussels may compete for habitat, food resources and fish hosts with invasive mussels across approximately 60% of their suitable range. Our projections can be used to guide future expeditions to monitor the conservation status of freshwater biodiversity, and potentially reveal populations of endemic species on the brink of extinction. Future conservation measures-most importantly the designation of nature reserves-should take into account trends in freshwater biodiversity generally, and particularly species such as freshwater mussels, vital to safeguard fundamental ecosystem services.

Protected areas offer refuge from invasive species spreading under climate change.

Protected areas (PAs) are intended to provide native biodiversity and habitats with a refuge against the impacts of global change, particularly acting as natural filters against biological invasions. In practice, however, it is unknown how effective PAs will be in shielding native species from invasions under projected climate change. Here, we investigate the current and future potential distributions of 100 of the most invasive terrestrial, freshwater, and marine species in Europe. We use this information to evaluate the combined threat posed by climate change and invasions to existing PAs and the most susceptible species they shelter. We found that only a quarter of Europe's marine and terrestrial areas protected over the last 100 years have been colonized by any of the invaders investigated, despite offering climatically suitable conditions for invasion. In addition, hotspots of invasive species and the most susceptible native species to their establishment do not match at large continental scales. Furthermore, the predicted richness of invaders is 11%-18% significantly lower inside PAs than outside them. Invasive species are rare in long-established national parks and nature reserves, which are actively protected and often located in remote and pristine regions with very low human density. In contrast, the richness of invasive species is high in the more recently designated Natura 2000 sites, which are subject to high human accessibility. This situation may change in the future, since our models anticipate important shifts in species ranges toward the north and east of Europe at unprecedented rates of 14-55 km/decade, depending on taxonomic group and scenario. This may seriously compromise the conservation of biodiversity and ecosystem services. This study is the first comprehensive assessment of the resistance that PAs provide against biological invasions and climate change on a continental scale and illustrates their strategic value in safeguarding native biodiversity.

Global ecological impacts of invasive species in aquatic ecosystems.

The introduction of invasive species, which often differ functionally from the components of the recipient community, generates ecological impacts that propagate along the food web. This review aims to determine how consistent the impacts of aquatic invasions are across taxa and habitats. To that end, we present a global meta-analysis from 151 publications (733 cases), covering a wide range of invaders (primary producers, filter collectors, omnivores and predators), resident aquatic community components (macrophytes, phytoplankton, zooplankton, benthic invertebrates and fish) and habitats (rivers, lakes and estuaries). Our synthesis suggests a strong negative influence of invasive species on the abundance of aquatic communities, particularly macrophytes, zooplankton and fish. In contrast, there was no general evidence for a decrease in species diversity in invaded habitats, suggesting a time lag between rapid abundance changes and local extinctions. Invaded habitats showed increased water turbidity, nitrogen and organic matter concentration, which are related to the capacity of invaders to transform habitats and increase eutrophication. The expansion of invasive macrophytes caused the largest decrease in fish abundance, the filtering activity of filter collectors depleted planktonic communities, omnivores (including both facultative and obligate herbivores) were responsible for the greatest decline in macrophyte abundance, and benthic invertebrates were most negatively affected by the introduction of new predators. These impacts were relatively consistent across habitats and experimental approaches. Based on our results, we propose a framework of positive and negative links between invasive species at four trophic positions and the five different components of recipient communities. This framework incorporates both direct biotic interactions (predation, competition, grazing) and indirect changes to the water physicochemical conditions mediated by invaders (habitat alteration). Considering the strong trophic links that characterize aquatic ecosystems, this framework is relevant to anticipate the far-reaching consequences of biological invasions on the structure and functionality of aquatic ecosystems.

The importance of the human footprint in shaping the global distribution of terrestrial, freshwater and marine invaders.

Human activities such as transport, trade and tourism are likely to influence the spatial distribution of non-native species and yet, Species Distribution Models (SDMs) that aim to predict the future broad scale distribution of invaders often rely on environmental (e.g. climatic) information only. This study investigates if and to what extent do human activities that directly or indirectly influence nature (hereafter the human footprint) affect the global distribution of invasive species in terrestrial, freshwater and marine ecosystems. We selected 72 species including terrestrial plants, terrestrial animals, freshwater and marine invasive species of concern in a focus area located in NW Europe (encompassing Great Britain, France, The Netherlands and Belgium). Species Distribution Models were calibrated with the global occurrence of species and a set of high-resolution (9×9 km) environmental (e.g. topography, climate, geology) layers and human footprint proxies (e.g. the human influence index, population density, road proximity). Our analyses suggest that the global occurrence of a wide range of invaders is primarily limited by climate. Temperature tolerance was the most important factor and explained on average 42% of species distribution. Nevertheless, factors related to the human footprint explained a substantial amount (23% on average) of species distributions. When global models were projected into the focus area, spatial predictions integrating the human footprint featured the highest cumulative risk scores close to transport networks (proxy for invasion pathways) and in habitats with a high human influence index (proxy for propagule pressure). We conclude that human related information-currently available in the form of easily accessible maps and databases-should be routinely implemented into predictive frameworks to inform upon policies to prevent and manage invasions. Otherwise we might be seriously underestimating the species and areas under highest risk of future invasions.

Priority setting for invasive species management: risk assessment of Ponto-Caspian invasive species into Great Britain.

Invasive species drive important ecological and economic losses across wide geographies, with some regions supporting especially large numbers of nonnative species and consequently suffering relatively high impacts. For this reason, integrated risk assessments able to screen a suite of multiple invaders over large geographic areas are needed for prioritizing the allocation of limited resources. A total of 16 Ponto-Caspian aquatic species (10 gammarids, one isopod, two mysids, and three fishes) have been short-listed as recent or potential future invaders of British waters, whose introduction and spread is of high concern. In this study, we use multiple modeling techniques to assess their risk of establishment and spread into Great Britain. Climate suitability maps for these 16 species differed depending on the eastern and western distribution of species in continental Europe, which was related to their respective migration corridor: southern (Danube-Rhine rivers), and northern (Don and Volga rivers and Baltic lakes). Species whose suitability was high across large parts of Great Britain included four gammarids (Cheliorophium robustum, Dikerogammarus bispinosus, D. villosus, and Echinogammarus trichiatus) and a mysid (Hemimysis anomala). A climatic "heat map" combining the results of all 16 species together pointed to the southeast of England as the area most vulnerable to multiple invasions, particularly the Thames, Anglian, Severn, and Humber river basin districts. Regression models further suggested that alkalinity concentration > 120 mg/L in southeast England may favor the establishment of Ponto-Caspian invaders. The production of integrated risk maps for future invaders provides a means for the scientifically informed prioritization of resources toward particular species and geographic regions. Such tools have great utility in helping environmental managers focus efforts on the most effective prevention, management, and monitoring programs.

Using continuous surface water level and temperature data to characterize hydrological connectivity in riparian wetlands.

Methods to characterize hydrological connectivity at riparian wetlands are necessary for ecosystem management given its importance over ecosystem structure and functioning. In this paper, we aimed to describe hydrological connectivity at one Ebro River reach (NE Spain) and test a method to perform such characterization. Continuous surface water level and temperature data were recorded at five riparian wetlands during the period October 2006-June 2007. Combining water level and temperature, we classified the examined wetlands in three groups, which mainly differed in the dominant water source during different flood stages. Firstly, a comparison of water level fluctuations in riparian wetlands with those in the river channel during events with different characteristics was used to describe hydrological connectivity. Such comparison was also used to extract quantitative hydrological connectivity descriptors as the wetland response initiation time. Secondly, water temperature series were divided in phases with different average, range and daily oscillation, and these parameters were interpreted for each phase to identify dominant flowpaths. By doing so, a more complete description of hydrological connectivity was achieved. Our method provided useful insights to describe hydrological connectivity using a qualitative approach that can be expanded if required to include quantitative parameters for studies of biotic assemblages or ecosystem processes.