A dark side of conservation biology: Protected areas fail in representing subterranean biodiversity

ORIGINAL ARTICLE A dark side of conservation biology: Protected areas fail in representing subterranean biodiversity Raquel Colado 1 | Pedro Abellán 2 | Susana Pallarés 2 | Stefano Mammola 3,4,5 | Roberto Milione 3,6 | Arnaud Faille 7 | Javier Fresneda 8,9 | David Sánchez-Fernández 1 1 Departamento de Ecología e Hidrología, Facultad de Biología, Universidad de Murcia, Murcia, Spain 2 Departamento de Zoología, Universidad de Sevilla, Seville, Spain 3 DarkMEG—Molecular Ecology Group, Water Research Institute (IRSA), National Research Council of Italy (CNR), Verbania Pallanza, Italy 4 LIBRe—Laboratory for Integrative Biodiversity Research, Finnish Museum of Natural History, University of Helsinki, Helsinki, Finland 5 National Biodiversity Future Center, Palermo, Italy 6 Department of Life Sciences and Systems Biology, University of Torino, Torino, Italy 7 Department of Entomology, Stuttgart State Museum of Natural History, Stuttgart, Germany 8 Ca de Massa, 25526 Llesp–El Pont de Suert, Lleida, Spain 9 Museu de Ciències Naturals (Zoología), Barcelona, Spain Correspondence Raquel Colado, Departamento de Ecología e Hidrología, Facultad de Biología, Universidad de Murcia, Murcia, Spain. Email: [email protected] Funding information Consejería de Economía, Conocimiento, Empresas y Universidad de la Junta de Andalucía-Fondo Social Europeo de Andalucía 2014-2020, Grant/Award Number: SP-DOC_01211; Ministerio de Ciencia e Innovaci on; Ministry of Economy and Innovation (Spain), Grant/Award Number: RYC2019-027446-I; Biodiversa+2021-2022 project DarCo, Grant/Award Number: BIODIV21_0006 Editor: Manu E. Saunders and Associate Editor: Raphael K. Didham Abstract 1. Biodiversity conservation is a central imperative of the 21st century. Subterranean ecosystems deliver critical nature’s contributions to people and harbour a broad diversity of poorly understood specialised organisms. However, the subterranean biome is still largely overlooked in global biodiversity targets. 2. We assessed how well subterranean biodiversity is represented in protected areas (Natura 2000 and Emerald networks) in two global hotspots of subterranean biodiversity (the Pyrenees and the Alps). For this, we used two comprehensive databases of terrestrial subterranean taxa, that is, leiodids (beetles) from the Pyrenees and spiders from the Alps, and identified priority areas in each region using both species richness and geographic rarity patterns. 3. Our results show the incapacity of surface-protected area networks to represent subterranean biodiversity, as more than 70% and 90% of the identified priority areas (and 40% and 22% of the species) are not effectively covered by protected areas in the Pyrenees and the Alps, respectively. 4. These findings call for developing an urgent plan for subterranean biodiversity conservation within the European Biodiversity Strategy for 2030. KEYWORDS Alps, gap analysis, hydrographic basin, Natura 2000 Network, Pyrenees, subterranean species Received: 6 October 2022 Accepted: 26 June 2023 DOI: 10.1111/icad.12666 This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. © 2023 The Authors. Insect Conservation and Diversity published by John Wiley & Sons Ltd on behalf of Royal Entomological Society. 674 Insect Conserv Divers. 2023;16:674–683. wileyonlinelibrary.com/journal/icad INTRODUCTION The conservation of biodiversity is a central imperative of the 21st century (Cowie et al., 2022; Loreau et al., 2021). However, biodiversity continues to be lost at an unprecedented rate and thus many ecosystem services are being compromised (IPBES, 2021). The most common conservation strategy for halting biodiversity loss is focused on area-based methods (i.e., the establishment of protected areas) (Fahrig et al., 2022; Maxwell et al., 2020). For example, the Biodiversity Strategy for 2030 by the European Union proposes to transform 30% of Europe’s lands and seas into effectively managed protected areas while bolstering economic development and climate mitigation (European Commission, 2020). Several similar bold protected area targets have been advanced, including the Half-Earth Project (E.O. Wilson Biodiversity Foundation), the ‘Global Safety Net’(Dinerstein et al., 2020) and recently, the UN Biodiversity Conference (COP15) agreement (7–19 December 2022; Montreal, Canada), all aiming to reach between 30% and 50% of protected areas across the globe. While theoretical, protected area networks should strive to achieve a comprehensive coverage of multiple biodiversity components and ecosystem services, the reality is that some habitats and taxa are often overlooked in protected areas designation and other conservation actions (Adamo et al., 2022; Cardoso et al., 2011; Donaldson et al., 2016). A clear example of this is in subterranean ecosystems, whose exceptional forms of life continue to be neglected in conservation plans (Fiˇ ser et al., 2022; Sánchez-Fernández et al., 2021; Wynne et al., 2021). Subterranean ecosystems are likely the most widespread nonmarine environments on Earth, covering at least 19% of the Earth’s surface (Chen et al., 2017; Sánchez-Fernández et al., 2021). Yet, only 6.9% of known subterranean ecosystems overlap with protected areas globally (Sánchez-Fernández et al., 2021). Importantly, most of these protected areas have been established to preserve surface habitats and species, not including any specific characteristics or components of the subterranean habitats underneath among their conservation criteria (Mammola, Cardoso, et al., 2019; Sánchez-Fernández et al., 2021). The reasons why subterranean ecosystems should be better protected are numerous and rapidly grow as research advances. First, subterranean ecosystems harbour a great diversity of specialised and endemic species with distinctive morphologies, physiologies, behaviours, reduced dispersal capacity and as a consequence, a distribution range often limited to the karstic massif in which they occur (Culver & Pipan, 2019; Jeannel, 1926). Subterranean lineages show interesting adaptive convergences (Assmann et al., 2010; Fresneda & Salgado, 2017; Juan et al., 2010; Mammola et al., 2020; Trontelj et al., 2012; Wilkens & Strecker, 2003) and unique evolutionary trajectories that largely match the geological history of mountain ranges and cave habitats (Culver & White, 2005; Ribera et al., 2018). Furthermore, the subterranean environment has a key role in the regulation and provision of multiple ecological services and nature’s contribution to people (Canedoli et al., 2022), for example: (i) more than half of the world’s population relies on groundwater for consumption and agriculture (Elshall et al., 2020; Griebler et al., 2014); (ii) the consumption and absorption of greenhouse gasses by subterranean microbiota, resulting in a significant removal of CH 4 (between 65% and 90%) (e.g., Fernandez-Cortes et al., 2015; Martin-Pozas et al., 2022); or (iii) other important ecological services by cave-roosting bats ranging from pest arthropod suppression to plant pollination and seed dispersal (Fujita & Tuttle, 1991; Kunz et al., 2011). Subterranean ecosystems are connected to the soil and surface, and therefore are affected by several human threats such as deforestation, exploitation of freshwater resources, agriculture, urbanisation, pollution, quarrying and mining, which is triggering the loss of subterranean biodiversity (Castaño-Sánchez et al., 2020; Di Lorenzo et al., 2014; Mammola, Cardoso, et al., 2019; Mammola, Piano, & Cardoso, 2019; Wynne et al., 2018), including bacteria and invertebrates that maintain groundwater in good health (Griebler & Avramov, 2015; Lapworth et al., 2012; Schwarzenbach et al., 2010). While incorporating subterranean ecosystems in expanding networks of protected areas may seem straightforward, different knowledge gaps complicate this task. Global and regional subterranean biodiversity patterns are still largely unknown (Ficetola et al., 2019), preventing us from fully incorporating this biome in systematic conservation planning. Indeed, most of these habitats are not mapped and studied entirely because they are inaccessible to humans (e.g., large vertical shafts, meanders, narrow passages and fissure systems) (Culver & Pipan, 2014,2019; Giachino & Vailati, 2010; Mammola et al., 2016,2021). As a consequence, there are few georeferenced records of the presence of subterranean species, making it a challenge to effectively inventory, manage and conserve subterranean organisms (Wynne et al., 2018,2021). Despite a few recent attempts have been conducted to select priority areas for conserving subterranean biodiversity in both groundwater (Iannella et al., 2021; Michel et al., 2009) and terrestrial systems (Borges et al., 2012; Cardoso et al., 2021; Rabelo et al., 2018), there is an urgent need to understand geographical patterns of subterranean biodiversity and to test the effectiveness of current protected areas in covering sites of importance for the conservation of subterranean biodiversity. This would allow researchers to develop conservation guidelines to ensure that the valuable subterranean species and the ecosystem services provided by these habitats are effectively covered in future conservation strategies (Mammola, Meierhofer, et al., 2022; Wynne et al., 2021). Here, we evaluate the effectiveness of the current network of protected areas in covering subterranean biodiversity in two western European regions recognised as two of the main hotspots of subterranean biodiversity worldwide (Culver et al., 2006): The Pyrenees, Basque Country and coastal mountain systems in Catalonia (Spain) (Pyrenees hereafter) as the first study case, and the Alps as a second study case. For each region, we used two comprehensive databases of terrestrial subterranean biodiversity, that is, leiodids (beetles) from the Pyrenees and subterranean spiders from the Alps. The main objective of this study was to assess the extent to which the subterranean biodiversity is represented in protected areas. Specifically, we: (i) study the distribution patterns of subterranean CONSERVATION OF SUBTERRANEAN BIODIVERSITY 675 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License beetles and subterranean spiders in the Pyrenees and the Alps, respectively; (ii) identify priority areas (sites of importance for the conservation of subterranean biodiversity) in each region integrating both richness and rarity patterns; (iii) evaluate the effectiveness of protected areas regarding the representation of both species and conservation priority areas; and finally (iv) identify conservation gaps of subterranean species distribution and sensitive areas, allowing the identification of further conservation priorities. MATERIALS AND METHODS Study area and biological dataset We compiled two distributional databases on subterranean species in south-western Europe. We considered both troglobionts (obligate subterranean species that live and develop their whole life cycle in the subterranean environment) and troglophile (facultative subterranean organisms that complete at least one stage of their life cycle in the subterranean environment, depending so on the subterranean habitat for their survival) (Howarth & Moldovan, 2018). The first database includes distributional data of a clade of troglobiont beetles of the Leptodirini tribe (family Leiodidae) from the Pyrenees. The second database includes distributional records of subterranean spiders (troglobiont and troglophile species) from the Alps (Figure 1; Tables S1 and S2). We obtained distribution data for leiodids by digitising the information provided by Fresneda and Salgado (2017). This information was updated and expanded with additional data from several sampling campaigns (previously unpublished data). For subterranean spiders, we included the entirety of the Alps range from France in the east extending westward through Switzerland, Italy, Liechtenstein, Austria, Germany and Slovenia, by integrating available data in Global Biodiversity Information Facility (GBIF) (Mammola & Cardoso, 2021), Spiders of Europe (Nentwig et al., 2022), Araneae.it (Pantini & Isaia, 2019) and other literature sources (reviewed in Mammola et al., 2018; Mammola, Pavlek, et al., 2022). Both databases were checked and updated by the authors (including at least one specialist for each group) to validate species taxonomy and distributions. These databases encompass a total of 771 records (species/locality/source) for 193 beetle species in the Pyrenees and 2004 records for 115 subterranean spider species in the Alps—representing ca. one fifth of total species of subterranean spiders documented in Europe (Mammola, Pavlek, et al., 2022). Biodiversity patterns and identification of priority areas We generated maps of species richness for each study area using hydrographic basins as spatial units (Figure 1) and the QGIS Desktop 3.22.1 software. For this, we used the HydroBASINS database at level 12 (Lehner & Grill, 2013). HydroBASINS includes datasets of georeferenced hydrographic basins at 12 nested levels of sub-basin polygons, ranging from millions (level 1) to tens of km 2 (level 12). We used hydrographic basins as spatial units since they represent natural units, more biologically relevant for subterranean organisms than other alternatives such as grid cells. In order to prioritise not only the richest areas, but also those harbouring species with restricted distributions, which might be especially sensitive to human activities, we generated maps showing the patterns of geographic rarity as well, also using hydrographic basins as spatial units. For this, we firstly calculated a geographic rarity index FIGURE 1 Studied areas divided in hydrographic basins (a: Pyrenees; b: Alps). Distribution data of subterranean beetles and spiders are marked in red and blue, respectively. 676 COLADO ET AL. 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License for each species, defined as the inverse of the number of basins in which each species appears (1/n; being nthe number of basins where a species appears). Then, we calculated an average rarity for each basin, which was defined by the sum of the rarity values of all species that appear in that basin divided by the total number of species occurring there (i.e., (Σ1/n)/N; being Nthe total of species that appear in that basin). We ranked basins according to their values of richness and rarity, selecting the top 50 basins for each criterion and obtaining two lists with the highest richness and rarity values. We then identified as priority areas for subterranean biodiversity conservation (PACSB) in each region those basins that met both criteria, that is, basins with the highest values for both richness and rarity, ensuring that we are selecting the sites of importance for the conservation of subterranean biodiversity. Effectiveness assessment of protected areas We evaluated the effectiveness of the Natura 2000 Network for the conservation of subterranean beetles in the Pyrenees, and both the Natura 2000 Network and the Emerald Network for protecting subterranean spiders in the Alps. The Natura 2000 Network is the main tool to promote the long-term survival of both species and habitats in Europe. It consists of ‘Special Areas of Conservation’(SAC) and ‘Special Protection Areas’(SPA), designed under the European Commission Habitats Directive (EU Council Directive 92/43/EEC) and the Birds Directive (79/409/EEC), respectively. Importantly, as the Natura 2000 was not implemented in Switzerland and a number of sampling localities in the Alps are located in this country, we also considered as protected areas the pan-European Emerald Network in this territory. This network includes protected areas that aim to conserve wildlife and their associated natural habitats. It was launched in 1989 by the Council of Europe as part of its work under the Convention on the Conservation of European Wildlife and Natural Habitats (Bern Convention), which was started in 1982. Despite the fact that there are other types of protection tools in both study areas (e.g., National Parks, Natural Reserves, etc.), we have focused on the Natura 2000 Network because: (i) it is the main pillar to preserve biodiversity in Europe; (ii) it was created under a common legislative framework in the whole European Union; and (iii) many of the other protection tools are also included within the Natura 2000 Network. Firstly, we overlapped sampling localities with protected areas to identify ‘gap species’as those species distributed exclusively in caves located in unprotected areas. In addition, we overlapped the previously identified PACSB (basins) with the considered protected area networks in both study cases, classifying these priority areas into three types as a function of the calculated overlapping values: PACSB ‘covered or protected’by protected areas (overlapping value ≥50%), PACSB ‘marginally represented’(<50%) and PACSB ‘outside’protected areas (0%). These analyses were conducted using QGIS Desktop 3.22.1 software. In order to identify those basins that better complement the current network of protected areas in representing subterranean species, we computed the minimum set of basins that represent all the gap species at least once in our study areas. For this, we used the R package ‘prioritizr’v.7.1.1 with Gurobi solver (Hanson et al., 2023), which uses mixed integer linear programming techniques to solve conservation planning problems. Because there were multiple optimal solutions, we computed conservation importance (irreplaceability) scores for each basin following Ferrier et al. (2000)asimplementedin‘prioritizr’.The analysis of irreplaceability is a standard methodology of systematic conservation planning that has been used for multiple types of organisms, areas or environments (e.g., Lombard et al., 1999; Pressey, 1998,1999; Pressey et al., 1994). The obtained irreplaceability scores were used to quantify the relative importance of each basin for representing all the gap species in the fewest number of basins. RESULTS Biodiversity patterns and priority areas We recorded a total of 193 species of subterranean beetles in 175 hydrographic basins in the Pyrenees and 115 species of subterranean spiders in 472 hydrographic basins in the Alps (Table 1; Figure 1). Ninety-nine and 123 species of Pyrenean subterranean beetles, as well as 19 and 24 species of Alpine subterranean spiders, occur at a single locality and basin, respectively (Tables S1 and S2). In addition, all the beetles (193) of the Pyrenees and 62% of the 115 subterranean spider species are endemic to each study area. TABLE 1 Summary of study areas (Py: Pyrenees; A: Alps). Species Basins Gap species Irreplaceability PACSB Py 193 175 78 (40.4%) 49 17 TB TP NA P MR O 78 0 0 5 10 2 A 115 472 26 (22.6%) 22 11 TB TP NA P MR O 8171 1100 Abbreviations: MR, marginally represented; NA, no accurate information on the habitat/biology of the species; O, outside protected areas; P, covered or protected areas; PACSB, Priority areas for the conservation of subterranean biodiversity; TB, Troglobionts; TP, troglophiles. CONSERVATION OF SUBTERRANEAN BIODIVERSITY 677 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License Maps of species richness and rarity are shown in Figure S1.The basins with the highest subterranean beetle richness (4–7 species) were medium-altitude basins located at both northern and southern slopes of the Pyrenees (Figure S1), and a similar pattern was observed for the rarity index (0.7–1) (Figure S1). In the Alps, the basins with the highest number of subterranean spider species (4–14 species) were found in the Western Alps, near the Mediterranean Sea, in the central area and in some isolated basins in the Eastern Alps (Figure S1). The highest rarity index values (between 0.106 and 1) were found in the Western and Eastern Alps and in some basins of the Central Alps (Figure S1). We selected 17 priority areas for subterranean species in the Pyrenees, mainly located on Ariège, the Catalan pre-coastal range near the Padres mountains and the massifs of the Penedes region (Table 1; Figure 2a; Figure S2). We selected 11 priority areas, mainly located in the South-western (Mercantour National Park), Western (Lanzo valleys and Monte Fenera Natural Park) and Eastern Alps (Table 1; Figure 2b; Figure S2). Effectiveness of protected areas We identified 78 gap species in the Pyrenees (40.4% of the total) and 26 in the Alps (22.6%) (Table 1). Additionally, by superimposing the protected areas with the PACSB, we found that only 5 out of the 17 PACSB in the Pyrenees can be considered as protected, while 10 were marginally protected (i.e., <50% of the surface covered by protected areas) and 2 were completely outside protected areas (Table 1; Figure 2a). In the case of the Alps, of the 11 identified PACSB, 10 were marginally protected and 1 totally unprotected (Table 1; Figure 2b). Finally, the minimum number of basins that represent all the gap species previously identified in our study area was 49 (irreplaceability values from 1 to 4) in the Pyrenees (Table 1; Figure 3a) and 22 (irreplaceability values from 0.306 to 3) in the Alps (Table 1; Figure 3b). In the Pyrenees, the basins with highest irreplaceability values and therefore those to be prioritised to complement the current network of protected areas were located on the border massifs between Saint- Jean-Pied-de-Port and Orbaiceta-Roncesvalles, while those in the Alps were primarily located in subterranean systems in the provinces of Bergamo and Como (Lombardia). DISCUSSION Our analyses showed that more than 70% of the identified PACSB in the Pyrenees were not effectively protected (i.e., they were marginally FIGURE 2 Priority areas for the conservation of subterranean biodiversity in orange with protected areas marked in green for the Natura 2000 Network and in brown for the Emerald Network (a: Pyrenees; b: Alps). FIGURE 3 Unprotected basins that would complement the Natura 2000 Network and pan-European Emerald Network (a: Pyrenees; b: Alps). The colour scale represents irreplaceability values ranging from the lowest (blue) to the highest (red). The protected areas are marked in green for the Natura 2000 Network and in brown for the Emerald Network for the conservation. 678 COLADO ET AL. 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License protected or outside protected areas), and 40% of the species did not have any population under protected areas. In the case of the Alps, there were different basins in the western, central and eastern areas identified as PACSB that were not effectively protected (i.e., more than 90% of the PACSB were marginally protected) and more than 22% of the species were totally outside of protected areas. This reflects the incapacity of the current surface-protected area networks to represent subterranean fauna, which is highly dependent on the surface ecosystems because water, matter and energy inputs to the subterranean environment come from the surface through the water that infiltrates through the network of fissures (Delay & Juberthie, 1981). Therefore, the unprotected belowground biodiversity could be highly vulnerable to potential surface land use changes, and some human activities such as agriculture, mining, damming or logging can directly or indirectly impact subterranean ecosystems and their biodiversity (Ferreira et al., 2022;Fi ˇ ser et al., 2022; Prieto et al., 2015). These impacts are expected to be aggravated by climate change, which affects multiple subterranean ecological processes at different spatial and temporal scales (Mammola, Piano, et al., 2019; Sánchez-Fernández et al., 2021). According to our results, 49 and 22 basins in the Pyrenees and the Alps, respectively, should be included in the network of protected areas in order to cover all species. However, we are aware that it is almost impossible that any of these areas would be incorporated as new Natura 2000 sites, as none of these subterranean species are included in the Annex II of the habitats directive. Indeed, some species of cave-roosting bats and a handful of specialised subterranean species [e.g., Congeria kusceri (Bole) (Mollusca: Bivalvia), Leptodirus hochenwartii Schmidt (Coleoptera: Leiodidae), and Proteus anguinus Laurenti (Amphibia: Proteidae)] are the sole representatives of the subterranean environment included in the Habitat Directive. We proffer the following recommendations (or alternatives) to integrate the conservation of subterranean invertebrate species into protected area management. One alternative would be to propose new conservation focal points such as ‘microreserves for subterranean biodiversity conservation’, also considering the management of the whole karst (including epikarst and soil). In this case, we refer to areas mainly devoted to the protection of small range species or habitats that are outside of current protected area networks and under identified threat (e.g., microreserves protecting lepidopteran species in the Sierra de Guadarrama, bats in the Cueva Secreta del Sagreo in the province of Jaén and the caves of Morciguillos and Judia in the province of Cuenca in Spain). Another alternative would be to use information from the International Union for Conservation of Nature (IUCN) about extinction risk of species, which is slowly becoming available even for subterranean invertebrates (e.g., Milano et al., 2022). It is interesting to note that the new Biodiversity Strategy for 2030, proposing to protect 30% of Europe surface (European Commission, 2020) would be efficient to protect the biodiversity of all subterranean spiders from the Alps (as protecting 10% of the basins of this study area would cover the whole diversity of known species). However, to protect the whole of subterranean leiodid species known so far in the Pyrenees, 54% of the basins would be required (i.e., 94 out of 175). Therefore, the data provided will pose challenges to conservation managers for the effective management and conservation of subterranean species. We are aware that our dataset has intrinsic limitations, including the uneven sampling effort and the coarse scale of the spatial units considered. For example, the exhaustiveness of the data on the French slope of the Pyrenees is low. Despite this, it seems that the area of Saint Girons (the subterranean laboratory of Moulis) was much more intensively sampled than other areas of the Pyrenees. Likewise, knowledge on the subterranean spider fauna in the Western Italian Alps is much greater than in other areas, due to intensive sampling by local researchers (e.g., Arnò & Lana, 2005; Isaia et al., 2011). Despite working with the best information available, it is likely that some sites of importance for the conservation of subterranean biodiversity have been missed owing to uneven sampling effort. It is also possible that some important locations in priority basins are included within protected areas even with a low overlapping value (10%–50%). Thus, an urgent and immediate task will be to increase basic knowledge on species biodiversity and distribution across the full breadth of subterranean environments (Sendra et al., 2011; Trajano & de Carvalho, 2017), including those subterranean habitats not directly accessible to humans (Mammola et al., 2021). The best ways to achieve this could be to limit administratively inefficient constraints and develop cooperative projects with local speleological groups, as significant discoveries of the last decades were made by speleologists who took the time to repeatedly visit caves and collect specimens of species of scientific and/or conservation interest (Garcia et al., 2020; Hernando & Castro, 2022; Pérez Fernández & Pérez Ruiz, 2018). Practical conservation of subterranean biodiversity Most of the known subterranean diversity is found in a zone around 45 latitude in Europe and around 35latitude in North America (Culver et al., 2006). The analysis approach used in this study has been applied in other groundwater conservation studies (Iannella et al., 2021;Michel et al., 2009). Therefore, the methodology used could be perfectly applicable to other subterranean areas, although it would be necessary to explore which groups are bioindicators in the subterranean environment and to improve knowledge on their distribution ranges. Although the study of the two specific groups considered here is certainly interesting by itself, as they represent important components of the subterranean fauna, their richness and rarity patterns and sensitivity to human activities do not necessarily correlate with those of other groups or the subterranean fauna as a whole. We found that, in general, an alarming percentage of species are outside of protected areas. This worrying result may not be as problematic if there are no direct threats to their habitat. In some cases, the addition of protected areas in non-threatened areas could increase administrative burdens for those working to increase knowledge of cave fauna. An effective conservation programme must therefore be compatible with monitoring and scientific activity, which would in fact provide the best basis for informing conservation action, and must always be linked to the documentation of threats to cave fauna. CONSERVATION OF SUBTERRANEAN BIODIVERSITY 679 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License Conservation efforts should first target the areas under intense pressure (despite having potentially less diversity as a result of threats to species). Therefore, the next steps to improve conservation efforts in priority areas would be: (i) the identification of potential threats (e.g., quarrying, damming of water courses, urbanisation, infrastructure such as roads, highways and railways, groundwater withdrawal and/or the overexploitation of aquifers, deforestation and land cover change) in order to prioritise areas whose protection is most urgent; (ii) listing of subterranean habitats, exhaustive sampling, further taxonomic work, and improving monitoring of subterranean fauna; and (iii) improving effective practical conservation measures, as needed. Finally, the effectiveness of conservation actions should be quantitatively tested, and for this, quantitative studies should be carried out to provide references to other scientists and conservation agents, with the aim of improving conservation interventions (Hedrick et al., 2020; Mammola, Meierhofer, et al., 2022; Sendra et al., 2011; Wilson et al., 2005). Overall, these findings will be key to frame a coherent plan for subterranean biodiversity conservation within the European Biodiversity Strategy for 2030. AUTHOR CONTRIBUTIONS Raquel Colado: Conceptualization; investigation; methodology; formal analysis; writing –review and editing; writing –original draft; supervision; data curation; validation; visualization; software. Pedro Abellán: Conceptualization; investigation; writing –review and editing; methodology; formal analysis; data curation; validation; visualization; software. Susana Pallarés: Conceptualization; investigation; writing –original draft; methodology; validation; visualization; writing –review and editing; formal analysis; data curation; supervision. Stefano Mammola: Investigation; validation; writing –review and editing; data curation. Roberto Milione: Data curation; validation; writing –review and editing; investigation. Arnaud Faille: Investigation; validation; writing –review and editing; data curation. Javier Fresneda: Investigation; writing –review and editing; validation; methodology; supervision; data curation. David Sánchez-Fernández: Funding acquisition; investigation; conceptualization; writing –original draft; methodology; validation; visualization; writing –review and editing; project administration; supervision; data curation. ACKNOWLEDGEMENTS This work is supported by the FRAGGLE project (PID2021-124640NB-I00) funded by MCIN/AEI/10.13039/501100011033 and by ‘ERDF A way of making Europe’, and from the Biodiversa+2021 project DarCo, Grant PCI2022-135076-2 funded by MCIN/AEI/10.13039/501100011033 and by the ‘European Union NextGenerationEU/PRTR’.DavidSánchez- Fernández is funded by a postdoctoral grant (RYC2019-027446-I funded by MCIN/AEI/10.13039/501100011033 and by ‘ESF Investing in your future’). Raquel Colado is funded by a predoctoral grant of the Spanish Ministry of Science and Innovation (FPI). Susana Pallarés is funded by a postdoctoral contract from the ‘Consejería de Economía, Conocimiento, Empresas y Universidad de la Junta de Andalucía-Fondo Social Europeo de Andalucía 2014–2020’(SP-DOC_01211). David Sánchez-Fernández is funded by a postdoctoral contract from the Spanish Ministry of Science and Innovation (Ram on y Cajal programs, RYC2019-027446-I). Further support comes from the project Biodiversa+2021-2022 project DarCo (BIODIV21_0006). We express our gratitude to all biospeleologists, speleologists and taxonomists who are dedicated to investigating the subterranean environment: without them we would not have enough information to be able to study and preserve this unique ecosystem. We would especially like to thank Charles Bourdeau, Pau Balart García and Ignacio Ribera Galán for their help and dedication to the study of subterranean biodiversity. CONFLICT OF INTEREST STATEMENT There are no conflicts of interest to be declared among the authors of the manuscript. DATA AVAILABILITY STATEMENT Data availability statement is in zenoro. The data of the supplementary material is in the manuscript. ORCID Raquel Colado https://orcid.org/0000-0002-1525-800X Pedro Abellán https://orcid.org/0000-0001-5566-5083 Susana Pallarés https://orcid.org/0000-0001-8677-7475 Stefano Mammola https://orcid.org/0000-0002-4471-9055 Arnaud Faille https://orcid.org/0000-0003-3274-5915 Javier Fresneda https://orcid.org/0000-0003-0701-6016 David Sánchez-Fernández https://orcid.org/0000-0003-1766-0761 REFERENCES Adamo, M., Sousa, R., Wipf, S., Correia, R.A., Lumia, A., Mucciarelli, M. et al. (2022) Dimension and impact of biases in funding for species and habitat conservation. Biological Conservation, 272, 109636. Available from: https://doi.org/10.1016/j.biocon.2022.109636 Borges, P.A.V., Cardoso, P., Amorim, I.R. et al. (2012) Volcanic caves: priorities for conserving the Azorean endemic troglobiont species. International Journal of Speleology, 41, 101–112. https://doi.org/10.5038/ 1827-806X.41.1.11 Elshall, A.S., Arik, A.D., El-Kadi, A.I., Pierce, S., Ye, M., Burnett, K.M. et al. (2020) Groundwater sustainability: a review of the interactions between science and policy. Environmental Research Letters,15(9), 093004. Available from: https://doi.org/10.1088/1748-9326/ab8e8c Arnò, C. & Lana, E. (2005) Ragni cavernicoli del Piemonte e della Valle d’Aosta. In: La Grafica Nuova. Torino: Associazione Gruppi Speleologici Piemontesi Regione Piemonte, p. 265. Assmann, T., Casale, A., Drees, C., Habel, J.C., Matern, A. & Schuldt, A. (2010) The dark side of relict species biology: cave animals as ancient lineages. In: Relict species. Berlin, Heidelberg: Springer, pp. 91–103. Available from: https://doi.org/10.1007/978-3-540-92160-8_4 Canedoli, C., Ficetola, G.F., Corengia, D., Tognini, P., Ferrario, A. & Padoa- Schioppa, E. (2022) Integrating landscape ecology and the assessment of ecosystem services in the study of karst areas. Landscape Ecology, 37(1), 347–365. Available from: https://doi.org/10.1007/ s10980-021-01351-2 Cardoso, P., Erwin, T.L., Borges, P.A. & New, T.R. (2011) The seven impediments in invertebrate conservation and how to overcome them. Biological Conservation, 144(11), 2647–2655. Available from: https:// doi.org/10.1016/j.biocon.2011.07.024 Cardoso, R.C., Ferreira, R.L. & Souza-Silva, M. (2021) Priorities for cave fauna conservation in the Iuiú karst landscape, northeastern Brazil: a threatened spot of troglobitic species diversity. Biodiversity and 680 COLADO ET AL. 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License Conservation, 30, 1433–1455. Available from: https://doi.org/10. 1007/s10531-021-02151-5 Castaño-Sánchez, A., Hose, G.C. & Reboleira, A.S.P. (2020) Ecotoxicological effects of anthropogenic stressors in subterranean organisms: a review. Chemosphere, 244, 125422. Available from: https://doi.org/ 10.1016/j.chemosphere.2019.125422 Chen, Z., Goldscheider, N., Auler, A., Bakalowicz, M., Broda, S., Drew, D. et al. (2017) World karst aquifer map (WHYMAP WOKAM) BGR, IAH, KIT. UNESCO, 10, b2. https://doi.org/10.25928/b2.21_sfkq- r406 Cowie, R.H., Bouchet, P. & Fontaine, B. (2022) The sixth mass extinction: fact, fiction or speculation? Biological Reviews, 97, 640–663. Available from: https://doi.org/10.1111/brv.12816 Culver, D.C., Deharveng, L., Bedos, A., Lewis, J.J., Madden, M., Reddell, J.R. et al. (2006) The mid-latitude biodiversity ridge in terrestrial cave fauna. Ecography, 29(1), 120–128. Available from: https:// doi.org/10.1111/j.2005.0906-7590.04435.x Culver, D.C. & Pipan, T. (2014) Shallow subterranean habitats. In: Ecology, evolution, and conservation. Oxford, New York: Oxford University Press. Culver, D.C. & Pipan, T. (2019) The biology of caves and other subterranean habitats. Oxford: Oxford University Press. Available from: https:// doi.org/10.1093/oso/9780198820765.001.0001 Culver, D.C. & White, W.B. (2005) Encyclopedia of caves. Amsterdam: Elsevier. Delay, B. & Juberthie, C. (1981) Les problemes d’introduction d’énergie dans le milieu souterrain superficial. Memoires de Biospeologie,8, 45–52. Di Lorenzo, T., Di Marzio, W.D., Sáenz, M.E., Baratti, M., Dedonno, A.A., Iannucci, A. et al. (2014) Sensitivity of hypogean and epigean freshwater copepods to agricultural pollutants. Environmental Science and Pollution Research, 21(6), 4643–4655. Available from: https://doi. org/10.1007/s11356-013-2390-6 Dinerstein, E., Joshi, A.R., Vynne, C., Lee, A.T.L., Pharand-Deschênes, F., França, M. et al. (2020) A “global safety net”to reverse biodiversity loss and stabilize Earth’s climate. Science Advances, 6(36), eabb2824. Available from: https://doi.org/10.1126/sciadv.abb2824 Donaldson, M.R., Burnett, N.J., Braun, D.C., Suski, C.D., Hinch, S.G., Cooke, S.J. et al. (2016) Taxonomic bias and international biodiversity conservation research. Facets, 1(1), 105–113. Available from: https://doi.org/10.1139/facets-2016-0011 European Commision. (2020) Europe 2020 European Commision. Available from: https://ec.europa.eu/eu2020/pdf/COMPLET%20EN% 20BARROSO%20%20%20007%20-%20Europe%202020%20-%20EN% 20version.pdf [Accessed 6th Sep 2022]. Fahrig, L., Watling, J.I., Arnillas, C.A., Arroyo-Rodríguez, V., Jörger- Hickfang, T., Müller, J. et al. (2022) Resolving the SLOSS dilemma for biodiversity conservation: a research agenda. Biological Reviews, 97(1), 99–114. Available from: https://doi.org/10.1111/brv.12792 Fernandez-Cortes, A., Cuezva, S., Alvarez-Gallego, M., Garcia-Anton, E., Pla, C., Benavente, D. et al. (2015) Subterranean atmospheres may act as daily methane sinks. Nature Communications, 6(1), 1–11. Available from: https://doi.org/10.1038/ncomms8003 Ferreira, R.L., Bernard, E., Da Cruz Júnior, F.W., Pil o, L.B., Calux, A., Souza- Silva, M. et al. (2022) Brazilian cave heritage under siege. Science, 375(6586), 1238–1239. Available from: https://doi.org/10.1126/ science.abo1973 Ferrier, S., Pressey, R.L. & Barrett, T.W. (2000) A new predictor of the irreplaceability of areas for achieving a conservation goal, its application to real-world planning, and a research agenda for further refinement. Biological Conservation, 93(3), 303–325. Available from: https://doi. org/10.1016/S0006-3207(99)00149-4 Ficetola, G.F., Canedoli, C. & Stoch, F. (2019) The Racovitzan impediment and the hidden biodiversity of unexplored environments. Conservation Biology, 33(1), 214–216. Available from: https://doi.org/10. 1111/cobi.13179 Fiˇ ser, C., Borko, ˇ S., Deli c, T., Kos, A., Premate, E., Zagmajster, M. et al. (2022) The European Green Deal misses Europe’s subterranean biodiversity hotspots. Nature Ecology & Evolution,1–2, 1403–1404. Available from: https://doi.org/10.1038/s41559-022-01859-z Fresneda, J. & Salgado, J.M. (2017) Catálogo de los Cole opteros Leiodidae Cholevinae Kirby, 1837 de la Península Ibérica e Islas Baleares. Monografies del Museu de Ciències Naturals. Consorci del Museu de Ciències Naturals de Barcelona,7,1–308. Fujita, M.S. & Tuttle, M.D. (1991) Flying foxes (Chiroptera: Pteropodidae): threatened animals of key ecological and economic importance. Conservation Biology, 5(4), 455–463. Garcia, L., Miralles-Nunez, A. & Pérez-Fernéndez, T. (2020) A new genus and species of cave-dwelling terrestrial isopod (Crustacea: Oniscidea: Trichoniscidae) from southern Spain. Zootaxa, 4822(2), zootaxa- 4822. Available from: https://doi.org/10.11646/zootaxa.4822.2.7 Giachino, P.M. & Vailati, D. (2010) Subterranean Environments: Hypogean Life. In: Concepts and Collecting Techniques. Verona: World Biodiversity Association. Griebler, C. & Avramov, M. (2015) Groundwater ecosystem services: a review. Freshwater Science, 34, 355–367. Available from: https://doi. org/10.1086/679903 Griebler, C., Malard, F. & Lefébure, T. (2014) Current developments in groundwater ecology—from biodiversity to ecosystem function and services. Current Opinion in Biotechnology, 27, 159–167. Available from: https://doi.org/10.1016/j.copbio.2014.01.018 Hanson, J.O., McCune, J.L., Chadès, I., Proctor, C.A., Hudgins, E.J., & Bennett, J.R. (2023). Optimizing ecological surveys for conservation. Journal of Applied Ecology, 60(1), 41-51. Hedrick, B.P., Heberling, J.M., Meineke, E.K., Turner, K.G., Grassa, C.J., Park, D.S. et al. (2020) Digitization and the future of natural history collections. Bioscience, 70, 243–251. Available from: https://doi.org/ 10.1093/biosci/biz163 Hernando, C. & Castro, A. (2022) Description of the world’s first troglobitic Pselaphini: Geopselaphus bullonorum sp. nov. from southern Spain (Coleoptera, Staphylinidae, Pselaphinae). Subterranean Biology, 43, 1– 10. Available from: https://doi.org/10.3897/subtbiol.43.83592 Howarth, F.G. & Moldovan, O.T. (2018) The ecological classification of cave animals and their adaptations. Cave Ecology, 235, 41–67. Iannella, M., Fiasca, B., Di Lorenzo, T., Di Cicco, M., Biondi, M., Mammola, S. et al. (2021) Getting the ‘Most out of the hotspot’for practical conservation of groundwater biodiversity. Global Ecology and Conservation, 31, e01844.55–e01844.367. Available from: https://doi.org/10.1016/j.gecco.2021.e01844 IPBES. (2021) Summary for policymakers of the global assessment report on biodiversity and ecosystem services of the intergovernmental science-policy platform on biodiversity and ecosystem services. Isaia, M., Paschetta, M., Lana, E., Pantini, P., Schönhofer, A.L., Christian, E. et al. (2011) Subterranean arachnids of the western Italian Alps (Arachnida: Araneae, Opiliones, Palpigradi, Pseudoscorpiones). In: Monografie di Museo Regionale di Scienze Naturali, Vol. 47. Torino: Museo Regionale di Scienze Naturali, pp. 1–325. Jeannel, R. (1926) Faune cavernicole de la France avec une étude des conditions d’existence dans le domaine souterrain. Paris: Lechevalier. Juan, C., Guzik, M.T., Jaume, D. & Cooper, S.J.B. (2010) Evolution in caves: Darwin’s‘wrecks of ancient life’in the molecular era. Molecular Ecology, 19, 3865–3880. Available from: https://doi.org/10.1111/j. 1365-294X.2010.04759.x Kunz, T.H., Braun de Torrez, E., Bauer, D., Lobova, T. & Fleming, T.H. (2011) Ecosystem services provided by bats. Annals of the New York Academy of Sciences, 1223(1), 1–38. Available from: https://doi.org/ 10.1111/j.1749-6632.2011.06004.x Lapworth, D.J., Baran, N., Stuart, M.E. & Warda, R.S. (2012) Emerging organic contaminants in groundwater: a review of sources, fate and occurrence. Environmental Pollution, 163, 287–303. Available from: https://doi.org/10.1016/j.envpol.2011.12.034 CONSERVATION OF SUBTERRANEAN BIODIVERSITY 681 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License Lehner, B. & Grill, G. (2013) Global river hydrography and network routing: baseline data and new approaches to study the world’s large river systems. Hydrological Processes, 27(15), 2171–2186. www. hydrosheds.org. Available from: https://doi.org/10.1002/hyp.9740 Lombard, A.T., Hilton-Taylor, C., Rebelo, A.G., Pressey, R.L. & Cowling, R.M. (1999) Reserve selection in the Succulent Karoo, South Africa: coping with high compositional turnover. Plant Ecology, 142, 35–55. Loreau, M., Barbier, M., Filotas, E., Gravel, D., Isbell, F., Miller, S.J. et al. (2021) Biodiversity as insurance: from concept to measurement and application. Biological Reviews, 96(5), 2333–2354. Available from: https://doi.org/10.1111/brv.12756 Mammola, S., Arnedo, M.A., Fiˇ ser, C., Cardoso, P., Dejanaz, A.J. & Isaia, M. (2020) Environmental filtering and convergent evolution determine the ecological specialization of subterranean spiders. Functional Ecology, 34, 1064–1077. Available from: https://doi.org/10.1111/1365- 2435.13527 Mammola, S. & Cardoso, P. (2021) Cave_dwelling_spiders_europe. Biodiversity Data Journal. Occurrence Dataset accessed via GBIF.org on 2021-08-31. Available from: https://doi.org/10.15468/ygocko Mammola, S., Cardoso, P., Culver, D.C., Deharveng, L., Ferreira, R.L., Fiˇ ser, C. et al. (2019) Scientists’warning on the conservation of subterranean ecosystems. Bioscience, 69(8), 641–650. Available from: https://doi.org/10.1093/biosci/biz064 Mammola, S., Cardoso, P., Ribera, C., Pavlek, M. & Isaia, M. (2018) A synthesis on cave-dwelling spiders in Europe. Journal of Zoological Systematics and Evolutionary Research, 56(3), 301–316. Available from: https://doi.org/10.1111/jzs.12201 Mammola, S., Giachino, P.M., Piano, E., Jones, A., Barberis, M., Badino, G. et al. (2016) Ecology and sampling techniques of an understudied subterranean habitat: the Milieu Souterrain Superficiel (MSS). The Science of Nature, 103(11), 1–24. Available from: https://doi.org/10. 1007/s00114-016-1413-9 Mammola, S., Lunghi, E., Bilandˇzija, H., Cardoso, P., Grimm, V., Schmidt, S.I. et al. (2021) Collecting eco-evolutionary data in the dark: impediments to subterranean research and how to overcome them. Ecology and Evolution, 11(11), 5911–5926. Available from: https://doi.org/ 10.1002/ece3.7556 Mammola, S., Meierhofer, M.B., Borges, P.A., Colado, R., Culver, D.C., Deharveng, L. et al. (2022) Towards evidence-based conservation of subterranean ecosystems. Biological Reviews, 97, 1476–1510. Available from: https://doi.org/10.1111/brv.12851 Mammola, S., Pavlek, M., Huber, B.A., Isaia, M., Ballarin, F., Tolve, M. et al. (2022) A trait database and updated checklist for European subterranean spiders. Scientific Data, 9(1), 1–13. Available from: https://doi. org/10.1038/s41597-022-01316-3 Mammola, S., Piano, E. & Cardoso, P. (2019) Climate change going deep: the effects of global climatic alterations on cave ecosystems. The Anthropocene Review, 6(1–2), 98–116. Available from: https://doi. org/10.1177/20530196198515 Martin-Pozas, T., Cuezva, S. & Fernandez-Cortes, A. (2022) Role of subterranean microbiota in the carbon cycle and greenhouse gas dynamics. Science of the Total Environment, 831, 154921. Available from: https://doi.org/10.1016/j.scitotenv.2022.154921 Maxwell, S.L., Cazalis, V. & Dudley, N. (2020) Area-based conservation in the twenty-first century. Nature, 586(7828), 217–227. Available from: https://doi.org/10.1038/s41586-020-2773-z Michel, G., Malard, F., Deharveng, L., Di Lorenzo, T., Sket, B. & De Broyer, C. (2009) Reserve selection for conserving groundwater biodiversity. Freshwater Biology, 54, 861–876. Available from: https:// doi.org/10.1111/j.1365-2427.2009.02192.x Milano, F., Borio, L., Komposch, C., Mammola, S., Pantini, P., Pavlek, M. et al. (2022) Species conservation profiles of the endemic spiders Troglohyphantes (Araneae, Linyphiidae) from the Alps and the Northwestern Dinarides. ARPHA Preprints, 3, e87298. Available from: https://doi.org/10.3897/arphapreprints.e87298 Nentwig, W., Blick, T., Gloor, D., Hänggi, A. & Christian, K. (2022) Araneae—spiders of Europe. http://www.araneae.nmbe.ch/ Pantini, P. & Isaia, M. (2019) Araneae.it: the online catalog of Italian spiders, with addenda on other arachnid orders occurring in Italy (Arachnida: Araneae, Opiliones, Palpigradi, Pseudoscorpionida, Scorpiones, Solifugae). Fragmenta entomol ogica, 51(2), 127–152. Available from: https://doi.org/10.4081/fe.2019.374 Pérez Fernández, T. & Pérez Ruiz, A. (2018) La Cueva Secreta del Sagreo (La Iruela, Jaén). In: Grupo de Espeleología de Villacarrillo. Jaén, Spain: Grupo de Espeleología de Villacarrillo, p. 88. Pressey, R.L. (1998) Algorithms, politics and timber: an example of the role of science in a public, political negotiation process over new conservation areas in production forests. In: Wills, R. & Hobbs, R. (Eds.) Ecology for everyone: communicating ecology to scientists, the public and the politicians. Sydney: Surrey Beatty and Sons, pp. 73–87. Pressey, R.L. (1999) Applications of irreplaceability analysis to planning and management problems. Parks, 9(1), 42–51. Pressey, R.L., Johnson, I.R. & Wilson, P.D. (1994) Shades of irreplaceability: towards a measure of the contribution of sites to a reservation goal. Biodiversity and Conservation, 3, 242–262. Prieto, M., Agull o, J., Fadrique, F. & Mas o, G. (2015) Cole opteros hipogeos protegidos o que requieren medidas de conservaci on en Cataluña. Arxius de Miscellània Zoològica, 13, 164–188. Available from: https:// doi.org/10.32800/amz.2015.13.0164 Rabelo, L.M., Souza-Silva, M. & Ferreira, R.L. (2018) Priority caves for biodiversity conservation in a key karst area of Brazil: comparing the applicability of cave conservation indices. Biodiversity and Conservation, 27, 2097–2129. Available from: https://doi.org/10.1007/ s10531-018-1554-6 Ribera, I., Cieslak, A., Faille, A. & Fresneda, J. (2018) Historical and ecological factors determining cave diversity. In: Moldovan, O.T., Kovac, L. & Halse, S. (Eds.) Cave ecology. Switzerland: Springer Nature Switzerland AG, pp. 229–252. Sánchez-Fernández, D., Galassi, D.M.P., Wynne, J.J., Cardoso, P. & Mammola, S. (2021) Don’t forget subterranean ecosystems in climate change agendas. Nature Climate Change, 11, 458–459. Available from: https://doi.org/10.1038/s41558-021-01057-y Schwarzenbach, R., Egli, T., Hofstetter, T.B., Von, Gunten, U. & Wehrli, B. (2010) Global water pollution and human health. Annual Review of Environment and Resources, 35, 109–136. Available from: https://doi. org/10.1146/annurev-environ-100809-125342 Sendra, A., Achurra, A., Barranco, P., Beruete, E., Borges, P.A., Herrero- Borgoñ on, J.J. et al. (2011) Biodiversidad, regiones biogeográficas y conservaci on de la fauna subterránea hispano-lusa. Boletín de la Sociedad Entomol ogica Aragonesa, 49, 365–400. Available from: http:// hdl.handle.net/10400.3/1870 Trajano, E. & de Carvalho, M.R. (2017) Towards a biologically meaningful classification of subterranean organisms: a critical analysis of the Schiner-Racovitza system from a historical perspective, difficulties of its application and implications for conservation. Subterranean Biology, 22, 1–26. Available from: https://doi.org/10.3897/subtbiol.22. 9759 Trontelj, P., Blejec, A. & Fiˇ ser, C. (2012) Ecomorphological convergence of cave communities. Evolution, 66(12), 3852–3865. Available from: https://doi.org/10.1111/j.1558-5646.2012.01734.x Wilkens, H. & Strecker, U. (2003) Convergent evolution of the cavefish Astyanax (Characidae, Teleostei): genetic evidence from reduced eye-size and pigmentation. Biological Journal of the Linnean Society, 80(4), 545–554. Available from: https://doi.org/10.1111/j.1095- 8312.2003.00230.x Wilson,K.,Pressey,R.L.,Newton,A.,Burgman,M.,Possingham,H.& Weston, C. (2005) Measuring and incorporating vulnerability into conservation planning. Environmental Management, 35(5), 527–543. Available from: https://doi.org/10.1007/s00267-004- 0095-9 682 COLADO ET AL. 17524598, 2023, 5, Downloaded from https://resjournals.onlinelibrary.wiley.com/doi/10.1111/icad.12666 by Universidad De Sevilla, Wiley Online Library on [25/04/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License