Transdisciplinary approach to the characterisation and current status of Spanish karstic lakes on gypsum

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They are usually small lakes but often displaying a high relative depth, located in active gypsum karst areas, with a high saturation of Ca 2 + and SO 4 2− in its waters. These lakes can usually stratify from spring to early autumn when the depth is high enough, then a sulphide-rich anoxic hypolimnion can develop in deep layers. So far, neither a comprehensive scientific definition of their ecological characteristics nor an exhaustive catalogue of their occurrence in a particular territory are available. This paper delves on their biotic and abiotic features needed for their identification as an ecosystem type. A proper methodology was also designed and applied for the evaluation of their conservation status, with the definition of reference values and assessment methods following the criteria of the Habitats Directive. The Spanish karstic lakes on gypsum (THCI 3190) were here identified, statistically representative sites were selected, and spatial GIS methods and multimetric indices were applied to assess the range, area, structure and function, and future prospects of this Habitat Type, as requested for the reporting according to Article 17 of the Habitats Directive. Results showed a favourable conservation status of this habitat type in the Alpine and Atlantic regions of Spain, but unfavourable-inadequate status in the Mediterranean, due to the pressures and impacts acting on some specific sites. A critical analysis of the methodologies and the values obtained for its conservation status was carried out. EU Habitats Directive Habitat Types of Community Interest Karstic lakes on gypsum Conservation status Assessment Figures Figure 1 Figure 2 Figure 3 1. Introduction Lentic ecosystems (standing waters) are affected worldwide by anthropogenic pressures, and they are one of the most threatened ecosystem types (Verhoeven 2014 ). Further, they are highly vulnerable to environmental shifts caused by climate change (Reid et al. 2019 ). These impacts may result in a degradation of the ecological structure and an alteration of their functioning (Dodson et al. 2005 ; Camacho et al. 2012 , 2016 ). Proper management and conservation are essential aspects to ensure their ecological integrity, biodiversity, and functioning, for instance, promoting biogeochemical mechanisms that provide ecosystem services such as carbon sequestration (Morant et al. 2020 ). To this end, a proper assessment of their condition is essential, to better define mechanisms to maintain or improve their ecological condition and integrity. But for that purpose, it is also necessary a proper characterisation of the ecosystem type, with the knowledge of their main ecological features both abiotic and biotic. In Europe, the assessment of the status of lentic waterbodies is currently regulated by two European Directives that establish the framework for the management and conservation of aquatic ecosystems and water resources. The Water Framework Directive (WFD; 2000/60/EC) (European Commission 2000 ) is the legal basis for water policy in the European Union (EU), which aims to achieve the good ecological status of aquatic ecosystems. The Habitats Directive (HD; 92/43/EEC) (European Commission 1992 ) is the framework for the EU’s biodiversity policy, aiming to protect, maintain or restore, at a favourable conservation status, the Habitat Types of Community Interest (HTCI) and their characteristic species, as well as, particularly, specific species. Among these HTCI, there are some types that are directly, (Group 31 of Annex 1, standing waters) or indirectly associated with aquatic ecosystems. The "ecological status" (assessed according to the WFD) is an expression of the quality of the structure and functioning of the aquatic ecosystems associated with surface waters. Meanwhile, the "conservation status" (HD) is the set of influences on a natural habitat type and on their typical species that can affect its natural distribution, its structure and function in the long term, as well as the survival of its typical species. There is a partial convergence of the objectives set by both Directives with the achievement of the good status for waterbodies and the conservation of HTCI and species included in Annexes II and IV of the HD as the final goals (Stefanidis et al. 2021 ), by considering management and protection measures to improve water-dependent habitats, species and water quality (European Commission 2011 ). The definition and classification of ecosystems and their associated habitats is essential, first to identify them, then to allow a proper assessment and management. HTCI in general, and habitats associated to aquatic ecosystems in particular, are usually described based on the structuring role of plants in the ecosystems (DG Environment 2013 ). As in other HTCI, in lentic habitats, plant communities can also play a determining role, but other defining elements of the habitat type are also needed for a proper habitat characterisation. Therefore, and especially in the case of HTCI in group 31 (standing waters), attention must be also paid to the global ecological characteristics of the habitat type, knowing that the occurrence of a HTCI is not solely described by the presence or absence of certain taxa, but rather by a sort of biotic and abiotic features. Consequently, the assessment of the conservation status must be based on the consideration of all those abiotic and biotic properties as defining the habitat /ecosystem type. Within the group of lentic ecosystems, which can be associated with the habitat types of standing waters (group 31 of Annex I of the HD), there are some characteristic lake and pond types, very rare worldwide, featured by their karstic character on areas rich in gypsum, providing specific ecological properties being key in their definition as a specific type of ecosystem and HTCI (Camacho et al. 2009 ). In this work we focus on this HTCI 3190, “Karstic lakes on gypsum”, one of the less known HTCI of Annex I of the HD from an ecological point of view. Officially, there are only 19 sites within the N2000 network classified as HTCI 3190 throughout the EU, 9 in Germany, 5 in Spain, 3 in Latvia, and 2 in Lithuania (European Environmental Agency 2022 ). The objectives of this paper were 1) to improve the information and knowledge of this restrictive HTCI (3190), with a detailed ecological description, 2) to identify the habitat and assess its global conservation status in Spain from the four parameters by the established protocols, following a scientific approach (Range of distribution, Area of occupancy, Structure and function, and Future prospects), and 3) to make a critical analysis of the methodologies established for the evaluation and the global value of the conservation status. 2. Material and methods 2.1. Ecological definition Following the Interpretation manual of European Union habitats EUR28 (DG Environment 2013 ), a deeper ecological characterisation of the Spanish karstic lakes on gypsum from the ecological features described in the scientific literature, accompanied with several field campaigns, was carried out. The main ecological features, including abiotic and biotic parameters, as well as the presence of the typical species, were used for the classification of the lentic habitat sites featured as HTCI 3190. 2.2. Validation of sites classified as HTIC 3190: karstic lakes on gypsum A validation of lakes with potential features linked to the HTCI 3190 previously selected from a database was carried out by overlaying the localities to the Geological Map of Spain 1:50,000 (GEODE) to ensure they were located on karstic areas with gypsum content. A deeper study of the rest of abiotic and biotic parameters, from bibliographic references and field data, was done to ensure they fit to the ecological description of the karstic lakes on gypsum. Some small ponds or other localities not included in the database up to the moment of the study could not be included in this assessment, though ongoing updates of the database would slightly increase the number of lakes identified as HTCI 3190 in Spain. 2.3. Selection of a minimum number of sites and data images sources In order to select a representative number of Iberian sites for the assessment of the conservation status among all the sites identified as HTCI 3190, specific criteria were used to avoid biases in the assessment and extrapolation of the conservation status. The minimum number of statistically representative sites (n) was obtained from the total population (total number of lakes on gypsum karst officially catalogued in Spain until the moment of the work) using the following formula (Camacho et al. 2019a ): $$n=\frac{{Z}_{\alpha }^{2}Npq}{{e}^{2}\left(N-1\right)+{Z}_{\alpha }^{2}pq}$$ where: N = population size (total number of lakes on gypsum karst in Spain) Z α = confidence level, expressed in values relative to the desired confidence percentage. The most used values are: 1.28 for a confidence level of 80%, 1.44 for 85%, 1.65 for 90%, 1.69 for 91%, 1.75 for 92%, 1.81 for 93%, 1.88 for 94%, 1.96 for 95% and 2.58 for 99% (Camacho et al. 2019a ) p = probability of success. As any study characteristics for success are defined in this case, by default it is usually assumed that p = q = 0.5 q = proportion of localities that do not have the probability of success, q = 1-p e = desired sampling error (in percentage) Once the minimum number of sites to be assessed was determined, specific criteria to select the sites were defined (Camacho et al. 2019a ) as follows: Extent, covering the largest area possible per site within the type, avoiding evaluating a greater number of different sites. Representativeness in the Natura 2000 Network and other protection figures at the international (Ramsar), national or regional level, also considering sites not officially classified as such. Threat status (or danger of disappearance), with a selection of sites in, a priori, both good and poor condition, indistinctively. Environmental-ecological diversity subtypes, e.g., shallow and deeper lakes, small ponds and bigger lakes). Existing information. Environmental and geographic variability. Accessibility and representativeness of the localities. Each of these criteria were valued and weighted to obtain the final sites to be included in the assessment as representative of the habitat type. Once all the sites classified as HTCI 3190 were designed, and the sites for their assessment were selected, the assessment of the four parameters of the conservation status established for the reporting of the Article 17 of the HD was carried out. The assessment for each site was obtained by determining, for the four parameters (‘Range of distribution’, ‘Area of occupancy’, ‘Structure and function’, and ‘Future prospects’) an evaluation as favourable, unfavourable-inadequate or unfavourable-bad status, by comparing the period 2013–2018 (following the Article 17 of the HD) with Google Earth images collected along this period for the determination of the ‘Range of distribution’ and ‘Area of occupancy’ parameters. Further than the assessment of changes along the period 2013–2018 used for the reporting according to the Article 17 of the HD, a historical recreation based on images collected in several flight missions was build-up. These missions were: American Flight 1956–1957; Interministerial Flight 1973–1986; National Flight 1981–1986; Five-Year Flight 1998–2003, and current (since 2012) high-resolution images series from the Spanish National Aerial Orthophotography Plan (PNOA), available at the Spanish Centre for Geographic Information ( https://centrodedescargas.cnig.es/CentroDescargas/index.jsp ). 2.4. Range of distribution The ‘Range of distribution’ of a habitat type is defined as the outer limits of the general area in which the habitat is found. The range of the HTCI 3190 in Spain was obtained from all the localities designed as 3190, using the Range Tool (EIONET 2022 ) in a GIS environment, following the guidelines of the toolbox (Camacho et al., 2019b ). A gap (or maximum distance between grids of distribution) of 40 km was used, as being karstic lakes on gypsum a non-zonal habitat type, following the unified criteria of the European Commission (DG Environment 2017 ). The status assessment of the parameter ‘Range of distribution’ was defined by comparing the range obtained in 2018 with the range obtained in 2013. The favourable reference range (FRR) was considered as the range when the Directive came into force, in 1994. 2.5. Area of occupancy The parameter 'Area of occupancy’ of a habitat type accounts for the real area it occupies. Each of the sites included as HTCI 3190 were delineated by using the Google Earth Pro® delineation tools as described by Camacho et al. ( 2019b ), differentiating the total area of the wetland, then breaking it down further into its different components: maximum flooded area, current flooded area, marginal vegetation coverage, other land uses or coverages. As said, not only the current flooded area, but also other areas of the wetland that have the characteristics of the habitat type are considered when assessing the area coverage at each site. For the assessment of the parameter 'Area of occupancy’, only the total surface parameter was used, being quantified with images of 2018 (or the closest), and 2013, to assess the differences among the 6-year period established between the assessment reports. As for the ‘Range of distribution’ FRR, the favourable reference area (FRA) was considered as the area when the Directive came into force in 1994. The latter was obtained by viewing and delineating each of the localities with historical images in a GIS (Camacho et al. 2019b ). Alternatively, the reference area was also estimated based on the minimum area that can support all species associated with the habitat type. In this method the cumulative surface area of each site added was correlated with the cumulative species present in the set of sites evaluated. The sorting of these sites when adding the surface areas and species to the curve was done randomly using 999 random iterations. An average curve was drawn showing the response of the coverage of the whole set of species with an increase in the total area of the habitat type when adding sites into the curve build-up. 2.6. Structure and function The parameter ‘Structure and function’ refers to the assessment of the status of the intrinsic ecological characteristics of the habitat types or ecosystems. Structure is understood as the configuration of the different biotic and abiotic elements, while the function refers to the processes linked to those elements. According to the procedures for the monitoring of habitat types, the ECLECTIC multimetric index was used to assess the ‘Structure and function’ (Camacho et al. 2009 , 2019c ). This index considers typical biological, physical-chemical, and hydromorphological variables of the habitat/ecosystem type, whose values are compared with reference values for each variable specific for the habitat type, from which a quantification was obtained. Some of these variables are compulsory, meanwhile others are optional for the final assessment (Camacho et al. 2009 ). The final value of the ECLECTIC index was obtained after the weighting of each of the variables (Camacho et al. 2009 ), being the HTCI at each site assessed as being in a favourable status when ECLECTIC ≥ 70, an unfavourable-inadequate status when 50 ≤ ECLECTIC < 70, and unfavourable-bad status when ECLECTIC < 50. 2.7. Future prospects The evaluation of the parameter ‘Future prospects’ for the conservation of lentic habitat types or ecosystems was associated with the estimation of the pressures and threats. The methodology used was described by Camacho et al. ( 2009 ). A group of pressures and impacts was evaluated in a weighted way, considering hydrological and geomorphological pressures and impacts, water quality, pressures and impacts on the structure of the communities, land uses, or presence of invasive species, accordingly to the potential impact each can have on the conservation status of the HTCI 3190. As calibrated, overall scores between 0–20 points, mean a low level of pressures and threats, thus a favourable conservation status. Scores between 21–50, indicated moderate pressures and threats, meaning an unfavourable-inadequate status, whereas scores higher than 51 points show high or very high level pressures and threats, thus an unfavourable-bad status. 2.8. Conservation status For the assessment of the conservation status at the level of the biogeographical region, the assessments carried out in each of the selected sites for the parameters 'Structure and function' and 'Future prospects' were weighted by its occupied surface area: $$Global parameters assessment= \sum _{i=1}^{n}index value site*\% surface site [up to 18.7\%]$$ Considering the great difference between the surface areas in the different sites due to the nature of THIC 3190, in which there are small ponds of a few square metres, to large lakes with extensions of several hectares, a factor was applied limiting the surface area over which the results could be extrapolated. This factor was three times the percentage of a site in relation to the total, if they all had the same surface area (1/16 selected sites)*3 (%). The results of the assessment of the parameters ‘Range of distribution’ and ‘Area of occupancy’ were obtained by the sum of the values for all sites assessed. Once the four parameters included in the general conservation status assessment matrix were evaluated, the results were provided as: favourable, unfavourable-inadequate, unfavourable-bad and unknown, following the guidelines of DG Environment ( 2017 ). The global value of the conservation status of the HTCI 3190 for each of the three biogeographic regions in Spain (Mediterranean, Atlantic and Alpine, given that this HTCI is not present in the Canary Islands – Macaronesia) was obtained according to the evaluation matrix, using the combination procedure given in Table 1 . Table 1 Procedure for the combination of the results of the conservation status using the four parameters of the evaluation matrix, for the assessment of the conservation status for a HTCI at the biogeographical region scale. Source: DG Environment (2017) Status of the parameters All f avourable, or three f avourable and one u nknown One or more i nadequate, but no bad One or more bad Two or more unknown combined with f avourable, or all unknown Overall assessment conservation status Favourable Unfavourable-inadequate Unfavourable-bad Unknown 3. Results 3.1. Ecological characterisation Karstic lakes on gypsum, mostly feed by groundwater, are formed by the collapse of karstic structures that evolved from the dissolution of materials on a basin lying on gypsum (CaSO 4 ), a highly soluble material that provides the water with a high concentration of sulphates that normally exceed bicarbonates, whose concentration is also high as in other types of karstic lakes. The morphometry of the basin is usually of the sinkhole type, generally with steep slopes walls and a high relative depth, although some basins (ponds) can just display small depressions formed by dissolution when collapse has not yet occurred. The size of the lake is usually small, unless the lake spans throughout several deep sinkholes or even a polje. Their waters are generally subsaline ( sensu Hammer 1986 ), although the conductivity range (generally 1.5-5 mS cm − 1 ) can occasionally overlap with the oligosaline or the low hyposaline range (Camacho et al. 2009 ). In lakes with a high relative depth, the formation of a sulphide-rich hypolimnion allows of its colonisation by green and purple sulphur bacteria. Table 2 indicates the main abiotic features of the HTCI “karstic lakes on gypsum”, used for the determination of sites assigned to the HTCI 3190. Communities of Charetea, Lemnetea and Potamogetonion are the dominant elements of the submerged vegetation (DG Environment 2013 ). Table 2 Main abiotic properties characterising THIC 3190, and other characteristic data. Source: (Camacho et al. 2009 ). Variable Feature Trophic status Oligo-mesotrophic Mineralisation Relatively high (freshwaters –oligosaline to subsaline). Conductivity: 1–5 mS/cm Content in bases High Water colour Colourless pH > 7,5 Transparency High Hydroperiod Generally permanent, sometimes temporary Lithology Gypsum and limestones Substrate Marls Depth Generally deep (> 2 m) Distribution in the Iberian Peninsula Gypsum-rich areas The definition of “karstic lakes on gypsum” by the EU Interpretation Manual does not fit to all lakes that lie on gypsum. From the geological and hydrogeological point of view, this name could include a much significant number of wetlands in Spain because gypsum outcrops broadly extend in the Spanish Alpine Orogenes and in some Neogene basins. However, many of these lakes present hypersaline waters; therefore, the Habitats Directive classified them differently as the definition of the HTCI 3190 includes only subsaline lakes. Deeping on the short definition given by the interpretation manual for Habitats of Community Interest (DG Environment 2013 ), Spanish karstic lakes on gypsum, are characterised by a high concentration of sulphate, allowing that, in anaerobic respiration processes in the sediments or in the anoxic hypolimnion (for deep enough lakes), sulphate can be used as an electron acceptor (Atlas and Bartha 2002 ), causing the accumulation of hydrogen sulphide in the deep layers while stratification is maintained, which greatly determines the biological communities in these depths. If altered by increased inputs of nutrients, phytoplankton massive growth in surface waters derived from a hypothetical eutrophication process would considerably reduce light penetration and hinder or prevent the formation of dense deep populations of photosynthetic microorganisms that normally include cryptophyceans, cyanobacteria and/or photosynthetic bacteria, around the oxic-anoxic interface (Camacho 2006 ), which characteristic of this habitat. Under natural (non-altered) conditions, and linked to the arrangement of the populations in the vertical profile mediated by the physical-chemical structuring of the water column, surface waters are usually relatively poor in nutrients, especially in phosphorus. Therefore, phytoplankton growth is controlled by the availability of these nutrients mediated by recirculation. Surface chlorophyll concentrations are naturally in the oligo-mesotrophic range (usually below 10 mg m − 3 ). However, very high concentrations of chlorophyll and/or bacteriochlorophyll of even several hundred mg m − 3 can be found in depth around the oxic-anoxic interface (Camacho 1997 ; Chicote 2004 ). This summer chlorophyll maximum can thus be considered as characteristic of this type of ecosystems in conditions of good functional conservation. In the Iberian stratified karstic sinkholes (both on gypsum and limestone) the formation of these chlorophyll metalimnetic maxima is common (Camacho 2006 , Camacho et al., 2009 ), generally formed by one or a few species of cryptophyceans (Pedrós-Alió et al. 1987 ; Gasol et al. 1992 ; García-Gil et al. 1993 ; Camacho et al. 2001a ; Chicote 2004 ) and/or cyanobacteria (Camacho et al. 1996 ; Camacho et al. 2000a ), which form dense populations at the oxic-anoxic interface coinciding with the bottom of the thermocline. In this type of lakes, the appearance, in anoxic zones, of populations of photosynthetic sulphur bacteria that use hydrogen sulphide as an electronic donor for photosynthesis is also a common feature (Van Gemerden and Mas 1995 ; Camacho et al. 2000b ). In karstic lakes on gypsum with metalimnetic chlorophyll maxima, the highest photosynthetic rates, in absolute terms, can be found in the narrow range of depths occupied by these deep populations of algae, cyanobacteria, and/or photosynthetic bacteria (Planas 1973 , 1990 ; Camacho and Vicente 1998 ), although their productivity (production per biomass unit) is normally lower than that of more superficial layers (Camacho 2006 ). This, together with the narrow range of depths they cover, means that only a part of the primary planktonic production is located in the deep metalimnion, although in certain systems the metalimnetic contribution may account for most of the primary planktonic production of the lake (Camacho et al. 2001b ). Due to the high slope of the shores, the relative extent of the littoral zone is generally small compared to the pelagic zone (open waters), although the saturated zone colonised by helophytes can be quite extensive, especially in non-collapsed shallow ponds. When the subsidence is small and the depth of the basin formed is low, the flooding is maintained only temporarily or occurs in the central area, originating a shallow lake essentially with littoral characteristics, and a higher relative importance of macrophyte communities. The submerged macrophytic vegetation can include both charophytes and higher plants, as well as several species of emergent helophytes in the littoral zone (Cirujano 1990 , 1995 ; Cirujano and Medina 2002 ). The contribution of organic matter from the marginal and riverside vegetation, mainly the remains of leaves from deciduous trees, can generate an important contribution of allochthonous organic materials that, given the often small reduced size of the lakes, can induce an increase in the trophic level and, indirectly, a greater production of hydrogen sulphide. Even some of these lakes can remain completely anoxic during a part of the annual cycle, with high concentrations of hydrogen sulphide and populations of purple sulphur bacteria that can reach up to the surface, giving a characteristic red colour to the waters. In the Iberian Peninsula, karstic lakes on gypsum show zooplankton populations with a heterogeneous vertical distribution, with species of crustaceans and rotifers with a distribution mainly epilimnetic, others with a metalimnetic distribution (Alfonso et al. 1987; Esteve et al. 1988 ; Miracle and Alfonso 1993 ; Gasol et al. 1995 ; Miracle and Armengol 1995) or with migratory capacity, and even populations of anaerobic ciliates (thanks to mutualistic symbiosis with bacteria; Finlay et al. 1991 ; Esteban et al. 1993 ) that colonise the anoxic hypolimnion rich in hydrogen sulphide in a markedly reducing environment. In the systems exclusively fed by groundwater (small sinkholes) without connections to surface watercourses, fish, when they exist, are generally introduced, while in systems with surface water connections, which serve as dispersal, the fish community may be more diverse and more structured by ecological interactions (Granado 2000 ). 3.2 Sites identified as karstic lakes on gypsum in Spain and selection of sites for the assessment of the structure and function, and future prospects Considering the ecological characterisation, and after the validation process following the procedures indicated in the Methods section, up to 79 lakes from the available data set on which we started (UVEG 2022 ), some of them grouped in lake complexes, were finally considered in Spain as belonging to THIC 3190 (Supplementary Table 1). Sites are mostly distributed throughout the East half of the Iberian Peninsula, in three of the four biogeographic regions of Spain. Though they are mostly located in the Mediterranean biogeographic region, they also appear in the Atlantic and Alpine regions, but do not appearing in the Macaronesian Canary Islands. Part of the typified sites were located in lakes complexes, associated with geological areas characterised by the presence of gypsum, and the genesis by karst dissolution. Figure 1 represents the site of the THIC throughout the Spanish territory and their location within the three biogeographic regions of the Spanish part of the Iberian Peninsula and the Balearic Islands (through this HTCI was not identified so far in the Balearic Islands). Following the number of sites needed for a proper assessment of the for the assessment of the structure and function, and future prospects (with the population size of 79 sites, and a defined error of 15%), and considering the amount of information necessary to carry out the relevant evaluation, the selection of a sample size was estimated with a total of 18 lakes representative of the type of habitat/ecosystem, based on the criteria defined in the material and methods section. Table 3 shows the sites finally selected as representative of the heterogeneity and intrinsic characteristics, protection and availability of information on the type of habitat/ecosystem. Table 3 Final selection of sites evaluated as representative from the HTCI 3190 Karstic lakes on gypsum in Spain according to the defined criteria. (ALP: Alpine; ATL: Atlantic; MED: Mediterranean biogeographical region). Codes of the different protection figures (Ramsar, Water Framework Directive, WFD, and Spanish Inventory of Wetlands) are also included in each lake included in each case. Site Region Hydroperiod Province Lake complex Natura 2000 network Ramsar Waterbody (WFD) Code Spanish Inventory of Wetlands Lago de Arreo MED Permanent Álava ES2110007 1258 ES091MSPF1019 IH211002 Laguna de Arbieto ATL Permanent Vizcaya Estanque Grande de Abajo de Estaña MED Permanent Huesca Estaña ES2410072 ES091MSPF1014 Laguna de Tortajada MED Permanent Teruel ES2420131 Laguna de Barcena, MED Permanent Burgos Gayanes Laguna de Alboraj MED Permanent Albacete ES4210011 IH421059 Laguna de La Atalaya MED Permanent Cuenca Fuentes - Río Moscas ES080MSPFL11_a Laguna de los Capellanes MED Temporary Cuenca IH423033 Estany de Banyoles MED Permanent Girona Banyoles ES5120008 1257 ES100MSPF0450401 Estanyol de Vilà MED . Girona Banyoles ES5120008 1257 ES100MSPF0450401 Estanyol de La Coromina MED . Girona ES100MSPFH1040040 Estany de Montcortés ALP Permanent Lleida ES5130019 ES091MSPF1029 Laguna de Zóñar MED Permanent Córdoba Sur de Córdoba ES0000034 446 ES050MSPFES0512000007 IH613017 Laguna Grande de Archidona MED Permanent Málaga Archidona 1911 ES060MSPF0614520 IH617023 Ballesteros-5 MED Permanent Cuenca Arcas ES4230008 ES080MSPFL13 IH423003 (refers to the complex) Arcas-2 MED Permanent Cuenca Arcas ES4230008 ES080MSPFL13 Arcas-4 MED Temporary Cuenca Arcas ES4230008 ES080MSPFL13 Lagunillo de las Tortugas MED Permanent Cuenca ES4230014 ES080MSPFL12 IH423006 (refers to the complex) 3.3. Range and area of distribution of HTCI 3190 in Spain The sites classified as HTCI 3190, karstic lakes on gypsum, showed an uneven distribution throughout the Iberian Peninsula (Fig. 2 ), being located mainly concentrated in specific areas. The Mediterranean region showed sites spread in several areas of its eastern part, with a range of distribution of 3,000 km 2 . The Alpine region showed only one site (Estany de Montcortés, in Lleida), with a range, equivalent to the distribution, of 100 km 2 . Finally, two sites were in the Atlantic region (Laguna de Olandina, in Álava, and Laguna de Arbieto, in Vizcaya), with a range of distribution of 200 km 2 . The area of the HTCI 3190 (considering the 79 sites classified) distributed in the three biogeographic regions in Spain shows a clear predominance of the Mediterranean region, with more than 94% of the total surface of this HTCI in Spain, and with very specific areas located in the Alpine and Atlantic regions (Table 4 ). All the karstic lakes on gypsum occupied an area of 308 ha in Spain. The maximum water flooding area was around 250 ha, meanwhile the emerged vegetation (helophytes) coverage was of 83 ha (Table 4 ). Table 4 Area covered by the HTCI 3190 and its different components at the end of the period 2013–2018 using Google Earth Pro® by biogeographic regions. (ALP: Alpine; ATL: Atlantic; MED: Mediterranean region). Since these are average values, an overlap can occur. Surface (ha) Total Maximum Current Emerged vegetation Others ALP 15.2 12.3 12.3 2.8 0.0 ATL 2.1 1.2 1.1 1.2 0.0 MED 290.9 236.1 203.1 79.0 7.1 Total 308.1 249.6 216.5 83.1 7.2 The assessment of the changes separated by biogeographic regions showed stable values in the Alpine and Atlantic regions, and changes of around 1% in the total surface in the Mediterranean region (Table 5 ) for the six-years evaluation period. The variations in the current flooding surface showed negative trends in contrast with the increases in the marginal vegetation coverage in this evaluation period, especially in lower latitudes of the Mediterranean region, being the “current flooding area” more linked with hydroclimatic features. Table 5 Values of change (in ha) in occupied surface area of HTCI 3190 during the period 2013–2018 using Google Earth Pro® by biogeographic regions in Spain. (ALP: Alpine; ATL: Atlantic; MED: Mediterranean region). Positive values represent an increase in coverage, whereas negative values show a decrease. Assessment period (ha) 2013–2018 Total Maximum Current Vegetation Others ALP 0.2 -0.1 -0.1 0.2 0.0 ATL 0.0 0.1 0.0 0.2 0.0 MED 3.2 1.0 -7.7 8.7 0.8 Total 3.4 1.0 -7.8 9.1 0.8 Both range and area covered by the HTCI 3190 did not significantly decrease when compared to the values when the Habitats Directive came into force (1994), as assessed by historic aerial images (Supplementary Figs. 1 to 26), therefore, they were assessed as favourable for the three biogeographic regions. Using the alternative procedure of the species-area curve, a 'favourable reference area' (FRA) for this THIC at national level could be determined of being around 300 ha, where increases in the total area of the THIC did not associate to large increases in the accumulation of species associated with the habitat type (Fig. 3 ). Thus, the 'Area of occupancy’ of this HTCI in Spain is still above the FRA, and this assessment procedure for the 'favourable reference area' also gives a favourable conservation status when all country is considered for this parameter. 3.4. Structure and function, Future prospects, and overall conservation status When assessing the ‘Structure and function’ parameter using the ECLECTIC index, the results obtained for each of the 18 sites selected as representative of the type, compiled in Table 6 , showed that none of the assessed sites had an unfavourable-bad status (U2). However, notable differences appear between some of the studied sites, with 50% of them in an unfavourable-inadequate status (U1), and another 50% with a favourable status. Table 6 Final value of the application of the ECLECTIC index, both with the evaluation of the compulsory variables, and with the evaluation of all variables (compulsory and optative), based on the available information, for the 18 sites selected as representative. The complete and disaggregated evaluations are compiled in Supplementary Table 2 (FV: favourable status; U1: unfavourable-inadequate status; ALP: Alpine; ATL: Atlantic; MED: Mediterranean region) Site Region ECLECTIC compulsory index value Status ECLECTIC all index value Status Lago de Arreo MED 73.9 FV 78.9 FV Laguna de Arbieto ATL 85.4 FV 79.2 FV Estanque Grande de Abajo de Estaña MED 92.7 FV 86.5 FV Laguna de Tortajada MED 60.4 U1 59.8 U1 Laguna de Barcena MED 87.5 FV 88.1 FV Laguna de Alboraj MED 63.5 U1 63.5 U1 Laguna de La Atalaya MED 57.3 U1 59.8 U1 Laguna de los Capellanes MED 67.7 U1 66.5 U1 Estany de Banyoles MED 65.8 U1 61.6 U1 Estanyol de Vilà MED 63.5 U1 63.5 U1 Estanyol de La Coromina MED 59.4 U1 59.4 U1 Estany de Montcortés ALP 87.5 FV 83.3 FV Laguna de Zóñar MED 56.3 U1 56.3 U1 Laguna Grande de Archidona MED 80.2 FV 84.4 FV Ballesteros-5 MED 95.8 FV 95.8 FV Arcas-2 MED 100 FV 91.7 FV Arcas-4 MED 95.8 FV 95.8 FV Lagunillo de las Tortugas MED 50 U1 52.8 U1 Regarding the ‘Future prospects’ parameter, the results obtained for each of the 18 sites, compiled in Table 7 , showed the values obtained from the pressure and threat evaluation method, and their equivalence to the conservation status class, with up to 10 sites with low levels of pressure, which was assimilated to favourable status for this parameter. In the same way, 7 sites, with moderate pressures, showed an unfavourable-inadequate status, and only one showed high pressure levels and threats, and thus, a bad status. Table 7 Final value of the application of the pressure and threat index, based on the available information, and structure and function for comparison, for the 18 sites selected as representative for the assessment. The complete and disaggregated evaluations are compiled in Supplementary Table 3. FV: favourable status; U1: unfavourable-inadequate status; U2: unfavourable-bad status; ALP: Alpine; ATL: Atlantic; MED: Mediterranean biogeographic region; P&A: Pressures and threats; S&F: Structure and function) Site Region P&T index value P&A Status S&F Status Lago de Arreo MED 10 FV FV Laguna de Arbieto ATL 12 FV FV Estanque Grande de Abajo de Estaña MED 12 FV FV Laguna de Tortajada MED 16 FV U1 Laguna de Barcena MED 13 FV FV Laguna de Alboraj MED 43 U1 U1 Laguna de La Atalaya MED 30 U1 U1 Laguna de los Capellanes MED 33 U1 U1 Estany de Banyoles MED 106 U2 U1 Estanyol de Vilà MED 31 U1 U1 Estanyol de La Coromina MED 28 U1 U1 Estany de Montcortés ALP 8 FV FV Laguna de Zóñar MED 26 U1 U1 Laguna Grande de Archidona MED 8 FV FV Ballesteros-5 MED 17 FV FV Arcas-2 MED 17 FV FV Arcas-4 MED 19 FV FV Lagunillo de las Tortugas MED 31 U1 U1 Supplementary Table 4 collects the references to assess the ‘Structure and function’ and ‘Future prospects’ in each selected site following the indices detailed. Comparing the status of the ‘Structure and function’ parameter, from the ECLECTIC index, with the ‘Future prospects’ parameter, from pressures and threats, almost all the sites showed similar status assessment (Table 7 ). When considering the set of the statistically representative sites, per biogeographic region, and following the weighting and extrapolation method explained above, both the ‘Structure and function’ and ‘Future prospects’ were classified as unfavourable-inadequate in the Mediterranean region, as the weighted final value of the ECLECTIC index was 62.2, slightly below the limit established by the favourable status (70), and the total value for pressures and threats extrapolated from the weighting by area meant a value of 45, meaning moderate pressure levels, and thus, unfavourable inadequate status. Both parameters were classified as favourable in the representative lakes for the Atlantic (Laguna de Arbieto) and the Alpine (Estany de Montcortés) biogeographical regions (Table 8 ), as the values of the ECLECTIC and pressures and threats indices indicated the good condition and lower pressures level in the two sites assessed for these regions. Combining these two parameters with the previous assessment of the ‘Range of distribution’ and ‘Area of occupancy’ in the General Matrix, the final conservation status for the Mediterranean region of Spain of the HTCI 3190, karstic lakes on gypsum, for the period 2013–2018 was assessed as unfavourable-inadequate, whereas for both the Atlantic and Alpine biogeographical regions, the conservation status was favourable (Table 8 ). Table 8 Evaluation of the conservation status of each of the four parameters included in the General Matrix, and global evaluation of the THIC 3190 for the period 2013–2018. FV: favourable status; U1: unfavourable-inadequate status; U2: unfavourable-bad status; ALP: Alpine; ATL: Atlantic; MED: Mediterranean biogeographical region. RANGE OF DISTRIBUTION AREA OF OCCUPANCY STRUCTURE AND FUNCTION FUTURE PROSPECTS OVERALL CONSERVATION STATUS MED FV FV U1 U1 U1 ATL FV FV FV FV FV ALP FV FV FV FV FV 4. Discussion Karstic lakes on gypsum are a very peculiar type of lentic ecosystem, with very restrictive characteristics, included by the HD as a habitat type (3190). These ecosystems are strongly linked with the geological and hydrogeological context associated with gypsum areas (Almécija 1997 ; Rodriguez-Rodriguez et al. 2006; Andreo et al. 2016 ), although their ecological definition is also related to their morphology, physical-chemical water features, and biotic processes and components, especially in particular microbial communities (Van Gemerden and Mas 1995 ; Camacho et al. 2000b ). However, the definition included in the interpretation manual for HTCI (DG Environment 2013 ), does not reflect exactly the ecological features observed in Spanish karstic lakes on gypsum. Under natural conditions, the large level fluctuations indicated in the Habitat Interpretation Manual (DG Environment 2013 ) are not so large in the Spanish lakes, being even infrequent in many cases, and highly dependent on the relations with the aquifer. In some cases, these lakes are the origin of more or less important streams or watercourses depending on the upwelling flow and the capacity of the aquifer that feeds them (Camacho et al. 2009 ). In Spain as well, there are lakes with a maximum depth higher than 7 m. Otherwise, the importance of some abiotic features, characteristics of this HTCI, indicated in Table 2 , are good indicators for the identification of sites as 3190, even more than macrophyte communities that can be found in other habitat types (Camacho et al. 2009 ). The particularities of this type of habitat, which is located in specific conditions where the karstification process takes place on materials with a high content in gypsum, means that its distribution is not attributable to bioclimatic factors, rather to specific geological factors. The distribution and range showed how the sites were located in specific areas where the geological characteristics allowed the formation of these systems. Their assessment was determined by comparison with the range when the Directive came into force, defining a reference range. However, the number of 79 lakes identified as HTCI 3190 are still far from the current 5 sites officially classified and protected within the N2000 network for this habitat type in Spain, and just 19 in all the EU (European Environmental Agency 2022 ), which shows that further declaration of Special Conservation Areas within the Natura 2000 Network is needed to enlarge the protection of this habitat type of community interest. Not only at the European level, but other protected figures do not include a large number of these ecosystems either. Of the 18 lakes identified as HTCI 3190 studied here wih detail as representative, only 7 are included in the Spanish Inventory of Wetlands (Table 3 ). Up to now, this inventory did not include some of the most important sites of the type, such as the Estaña or Banyoles complexes, neither the Estany de Montcortés. These and other sites, such as the Arbieto, Atalaya or Bárcena lakes, meet the requirements defined for their inclusion in this inventory, as well as having the peculiarity of being HTCI 3190. However, since the Spanish Inventory of Wetlands is a legal figure that requires the regional governments to submit to the central government the localities to be included, until each regional government sends this information, a lake/wetland cannot be included in the Inventory, thus is not officially recognized. The specific area occupied by the sites classified as 3190 was also calculated by aerial images, following a widespread method in ecology and conservation biology (Camacho et al. 2019b ). This method allows the area to be assessed from the delineation manually, which can be quite accurate, especially with high quality images. However, it is costly when the number of sites to be surveyed is very large, so other methods that automate the identification and delineation of these surfaces may be more suitable, although the accuracy may be reduced (Doña et al. 2021 ). The status assessment related to the habitat distribution and coverage was also carried out on the basis of the differences to the moment when the Directive came into force. Looking at the historic aerial images, an encompassed balance between disappeared sites (due to clogging processes and changes in land use), and those newly appearing by genesis due to karstic collapse during the last decades (Supplementary Figs. 4 and 26), or restored after having disappeared, like Laguna de Santiago (Supplementary Fig. 15), seems evident. This encompassing of newly appearance and disappearance is one of the most outstanding dynamic features of this HTCI, as the time scale of genesis is assimilable to the time scale of natural and anthropogenic processes causing its disappearance (see e.g., Supplementary Figs. 4 and 26, how several sites were formed by karstification processes during the last decades). This is one of the reasons why, despite being a geographically restricted habitat found under specific geologic conditions, its conservation status referring to the range and area was assessed as favourable. Particularly for the Atlantic region, in the area occupied by the Laguna de Arbieto, a karstic lake on gypsum disappeared in the 1980s, as observed in aerial images. However, and following the criteria to determine the status, as this site had disappeared before the moment when the Habitats Directive came into force, there were no implications for the assessment of these two parameters. Therefore, methods other than the comparison to the 1994 range and surface to define favourable reference values should be considered, such as, for example, the species-area curve we proposed, a method that is also used for other ecological applications (Tjørve 2003 ; Ibáñez et al. 2006 ) which here helped to determine a minimum area capable of harbouring the maximum number of plant species characteristic of the habitat type. With respect to the assessment of the area of occupancy for the evaluation period 2013–2018, differences among the surfaces (total, maximum flooded, current flooded, vegetation coverage, others) were also identified, though only the total area was used for the overall conservation status assessment. Results showed how most of the total surface corresponded to the flooded areas (70.6%), followed by 27% of the surface occupied by marginal emerged vegetation, and 2.4% the surface occupied by other uses. Looking at the differences along the period, a trend was found for the increase in the helophyte coverage and a reduction of open waters, especially in lower latitudes of the Spanish Mediterranean biogeographical region. This trend could be attributed to environmental changes, some of them related to climate change, such as reducing water supply thus reducing the water depth, and favouring the helophyte colonisation, especially that of Phragmites australis , a highly invasive, though autochthonous, species in littoral shallow areas of lakes (Cirujano et al. 2010 ), as well as a possible acceleration the lakes siltation in the long term due to increased sediment transport in the agricultural areas surrounding many of these sites. However, the large inter- and intra-annual variability associated with hydro-climatic characteristics in the Mediterranean area, as well as other management actions that may influence the hydrological dynamics of wetlands, should be taken into account. Trends should be framed only on the period they were assessed, as these short-term trends may vary. Therefore, the assessment of the area parameter is only based on the total surface of the wetland, and not on the coverages of its components. It would be necessary to continue this evaluation in future periods to see if these trends continue, which would indicate a change in the structure of the habitat type, at least in shallow systems. The ‘Structure and function’ parameter was assessed by the multimetric ECLECTIC index, obtained from a quantification and weighting of different parameters. Apart from the importance of the Chlorophyll- a concentration (Poikane et al. 2010 ; Carvalho et al. 2013 ), other parameters, such as the biological community composition and coverage of submerged and emerged macrophytes, are relevant in the biological part of this index, given the role of plants in the structure and functioning of the habitats (Camacho et al. 2016 ). Hydro-geomorphological factors and physical-chemical parameters are also included in the index, given the importance of these components in the lentic ecosystems condition (Verhoeven et al. 2006 ; Poikane et al. 2020 ). The evaluation of the parameter 'Structure and function' by the ECLECTIC index is carried out based on a sum of weighted values of different metrics, whose final value defines the parameter “structure and function“ of the conservation status based on the global score obtained. Looking at the results of the ECLECTIC index, and their assimilations to status classes, lakes with impacts that directly or indirectly affect their ecological integrity showed lower index value, and unfavourable status. Estany de Banyoles, with human pressure on its banks and catchment basin, Laguna de los Capellanes located within a cattle farm with a progressive modification of its hydroperiod and morphology, and Laguna de Alboraj as well as Laguna de Zóñar, with intensive agriculture activity in the surroundings, were assessed as unfavourable-inadequate for the ‘Structure and function’ parameter. These sites showed, in general, relatively high concentrations of total phosphorus in water and high values of Chlorophyll- a . In Laguna de Zóñar, the loose of typical species, both plants (submerged macrophytes and helophytes) as well as zooplankton and benthic invertebrates, contributed to the reduction of the value of the conservation status (Junta de Andalucía 2005 ). Contrarily, sites such as Laguna de Arbieto, in the Atlantic region, or Estany de Montcortés, in the Alpine region, showed a favourable status for the structure and function parameter. In both cases, the evaluated variables identified the ecosystem as displaying a good ecological integrity, due to their biological diversity, the maintenance of their natural hydromorphological functioning, and the values of their physical-chemical parameters within the appropriate range for the HTCI. The multimetric ECLECTIC index combines the evaluation and application of different metrics referring to the specific elements of lentic ecosystems and their functioning. Some of the main assessing variables included in the index, such as the Chlorophyll- a concentration, and the nutrients concentrations, might make vary the final value (from favourable to unfavourable) due to their importance in the weighting, especially when not all the variables, both compulsory and optional, are assessed. Some of these variables are also used as the metrics for the assessment of the ecological status according to the WFD (Poikane et al. 2010 ; Carvalho et al. 2013 ; Poikane et al. 2020 ). Both approaches, the ECLECTIC method, and the different metrics according to the WFD, coincide in the determination of some common biological, physical-chemical and hydromorphological features, whose separate values integrated in the multimetric ECLECTIC index, allow the global evaluation of the status of ecological health of these lentic ecosystems. However, the ECLECTIC index differs from those used by the WFD, in which the lowest value of those obtained in the evaluations of biological quality elements, physical-chemical and hydro-morphological factors, is the one that gives the overall status value (one out, all out). Therefore, the proposed method could overestimate the final status value compared to other multimetric methods (European Commission 2000 ). The two legal frameworks for the protection of water (WFD) and nature (HD) converge when evaluating and achieving similar objectives (European Commission 2011 ). However, WFD and HD are currently implemented separately, which can make more difficult to achieve their respective goals (Stefanidis et al. 2021 ). The ‘Future prospects’ parameter was obtained from the assessment of pressures and threats that were previously demonstrated to affect the ecosystem condition, such as hydrology and geomorphology (Jusik and Macioł 2014 ; Evtimova and Donohue 2016 ; Poikane et al. 2020 ), water quality (Verhoeven et al. 2006 ), land uses (Nielsen et al. 2012 ; Morant et al. 2021 ) and exotic species (Reid et al. 2019 ). There was a notable difference in the level of pressure exerted in each of the sites selected as representative of the THIC 3190. Up to 10 from 18 assessed sites showed low levels of pressure, which assimilates to favourable future prospects (e.g., in Laguna de Arbieto, Estanque Grande de Estaña, and Estany de Montcortés). The levels of pressure exerted, both directly on hydrology and on their communities, as well as indirect pressures, were low and did not significantly affect the structure and ecological function of ecosystems, as the same favourable status was generally obtained for the ‘Structure and function’ parameter. In the same way, those sites with moderate pressures, and an unfavourable-inadequate status in the 'Future prospects' parameter, showed an unfavourable-inadequate status in the 'Structure and function' parameter, since the pressures exerted on the components of aquatic systems resulted on damages on their structure and functioning (Camacho et al. 2009 ). This was the case of Laguna de la Atalaya, Laguna de los Capellanes, Laguna de Zóñar, and Lagunillo de las Tortugas, where management actions are needed to reduce or eliminate these sources of pressure affecting their structure and ecological functioning, thus, their conservation status. This multimetric method, like the ECLECTIC index, could overestimate the calculated status, due to the integration of a set of variables, in which only one could reduce the final value of the status in a lake that apparently could be defined as favourable. Additionally to the method used here, alternative methodologies have been proposed for the assessment of pressures and threats have been proposed that can be spatially explicit, thus approaching the localisation of the areas for actuation, for instance, the use the land uses in the catchment area as a proxy to estimate the pressure level over lentic ecosystems by the LUPLES method (Morant et al. 2021 ), which demonstrated to be correlated with ecological indicators of impacts. At global level, the conservation status assessed within the four parameters. for the HTCI 3190 in Spain showed its unfavourable-inadequate status in the Mediterranean region, reflected by the moderate pressures and impacts and their response in the structure and functioning of some of the sites included in the assessment. In this region, the area of occupancy was maintained over the decades, because of an equilibrium between new formed sites that counterbalanced the disappeared lakes. Meanwhile, the status was favourable in the Atlantic and Alpine regions, though this was determined with a low sample size, with only two and one karstic lakes on gypsum assessed, respectively. As indicated above, this HTCI is highly specific and quite particular, thus, the low number of sites classified as HTCI 3190, and the low area occupied was not equivalent to a poor status, as the sites remain over the decades. Thus, by applying the different methods defined in this article, and following the criteria for the definition of the conservation status, it seems the results mostly reflect the real condition of the HTCI in Spain, considering the particularities of the ecosystem type. 5. Conclusions For the ecological characterisation of the lakes, main parameters identifying karstic lakes on gypsum were physical-chemical features associated to the active gypsum karst areas where they are located, conferring particular hydro-morphometric characteristics, as well as the presence of certain types of submerged macrophytes species, though a revision should incorporate additional ecological features as those indicated in this study. Localised in certain areas with specific characteristics, these lakes cover an area of about 300 ha in Spain, being the main catalogued contribution to this habitat type (HTCI 3190) by Western European countries. Their global conservation status was favourable in the Spanish Alpine and Atlantic regions, and unfavourable-inadequate in the Spanish Mediterranean region. This assessment was based on range and area of distribution of the HTCI, plus the assessment and extrapolation of the ‘Structure and function’ and ‘Future prospects’ parameters with multimetric indices, which followed the criteria established by the HD. From the assessment approach defined and applied, management actions of the HTCI should ensure the integrity and protection of the structure and functioning of the ecosystems in current and future scenarios, by minimising threats and maintaining their range and area throughout the biogeographic regions where they are located. However, additional efforts and detailed studies of these systems are needed to identify possible new locations that could be classified as HTCI 3190, especially small temporary ponds associated with geological environments rich in gypsum. In turn, a more detailed evaluation of their characteristics and condition is necessary for a more detailed overall assessment of their conservation status. Further, the methods here developed, as well as the specific results of the assessment, would help for the purposes of the Strategic Plan for Wetlands 2030, published by the Government of Spain in 2023 (MITECO, 2023 ), This plan aims to improve the conservation status and establish restoration targets for Spanish wetlands, which would be especially important for the conservation of this very scarce habitat type both at the Iberian and European level, as Spain plays a prominent role for the preservation of most of its sites in Western Europe. Declarations Competing interests “The authors have no relevant financial or non-financial interests to disclose.” Funding This work has been supported by the MITERD, through a TRAGSATEC contract nº: 65393 to the University of Valencia, as well as by the project CLIMAWET-CONS (PID2019-104742RB-I00), granted to AC (University of Valencia) by the Spanish Research Agency (AEI) of the Spanish Ministry for Science and Innovation. Author Contribution All authors contributed to the study conception and design, as well as to material preparation, data collection and analysis. RH promoted the study on behalf of MITERD, and AC did the experimental design. The first draft of the manuscript was written by DM and AC-S, and the final version was elaborated by AC. 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Int Rev Ges Hydrobiol 78:283-294 Gasol JM, Guerrero R, Pedrós-Alió C (1992) Spatial and temporal dynamics of a metalimnetic cryptomonas peak. J Plankton Res 14:1565-1580 Gasol JM, Jürgens K, Massana R, Calderón-Paz JI, Pedrós-Alió C (1995) Mass development of Daphnia pulex in a sulfide-rich pond (Lake Cisó). Arch Hydrobiol 132:279-296 Granado CA (2000) Ecología de Comunidades: el paradigma de los peces de agua dulce. Universidad de Sevilla, Sevilla Hammer UT (1986) Saline lake ecosystems of the world (Vol. 59). Springer Science & Business Media, Boston Ibáñez I, Clark JS, Dietze MC et al (2006) Predicting biodiversity change: outside the climate envelope, beyond the species–area curve. Ecology 87(8):1896-1906. https://doi.org/10.1890/0012-9658(2006)87[1896:PBCOTC]2.0.CO;2 Junta de Andalucía (2005) Caracterización ambiental de humedales en Andalucía. Sevilla: Junta de Andalucía. http://www.juntadeandalucia.es/medioambiente/site/portalweb/menuitem.7e1cf46ddf59bb227a9ebe205510e1ca/?vgnextoid=45e206b116b75010VgnVCM1000000624e50aRCRD&vgnextchannel=1f27dfde043f4310VgnVCM1000001325e50aRCRD Accessed 12 July 2021 Jusik S, Macioł A (2014) The influence of hydromorphological modifications of the littoral zone in lakes on macrophytes. Oceanol Hydrobiol Stud 43(1):66-76. https://doi.org/10.2478/s13545-014-0119-x Miracle MR, Alfonso MT (1993) Rotifer vertical distributions in a meromictic basin of Lake Banyoles (Spain). Hydrobiologia 255/256:371-380 Miracle MR, Armengol-Díaz X (1995) Population dynamics of oxiclinal species in Lake Arcas-2 (Spain). Hydrobiologia 313/314:291-301 MITECO (2023). Plan Estratégico de Humedales a 2030. Ministerio para la Transición Ecológica y el Reto Demográfico, Madrid, 122 p. https://www.miteco.gob.es/content/dam/miteco/es/biodiversidad/planes-y-estrategias/planestrategicodehumedalespublicacionoficial_tcm30-548431.pdf Morant D, Picazo A, Rochera C, Santamans AC, Miralles-Lorenzo J, Camacho A (2020) Influence of the conservation status on carbon balances of semiarid coastal Mediterranean wetlands. Inland Waters 10(4):453-467. https://doi.org/10.1080/20442041.2020.1772033 Morant D, Perennou C, Camacho A (2021) Assessment of the pressure level over lentic waterbodies through the estimation of land uses in the catchment and hydro-morphological alterations: The LUPLES method. Appl Sci 11(4):1633. https://doi.org/10.3390/app11041633 Nielsen A, Trolle D, Søndergaard M et al (2012) Watershed land use effects on lake water quality in Denmark. Ecol Appl 22:1187–1200. https://doi.org/10.1890/11-1831.1 Pedrós-Alió C, Gasol JM, Guerrero R (1987) On the ecology of a Cryptomonas phaseolus population forming a metalimnetic bloom in Lake Cisó, Spain: annual distribution and loss factors. Limnol Oceanogr 32:285-298 Planas MD (1973) Composición, ciclo y productividad del fitoplancton del lago de Banyoles. Oecol Aquat 1:3-106 Planas MD (1990) Factores de control de la distribución espacial y temporal de la producción primaria del fitoplancton del Lago de Banyoles. Scientia Gerundensis 16:193-204 Poikane S, Alves MH, Argillier C et al (2010) Defining chlorophyll- a reference conditions in European lakes. Environ Manage 45(6):1286-1298. https://doi.org/10.1007/s00267-010-9484-4 Poikane S, Zohary T, Cantonati M (2020) Assessing the ecological effects of hydromorphological pressures on European lakes. Inland Waters 10(2):241-255. https://doi.org/10.1080/20442041.2019.1654800 Reid AJ, Carlson AK, Creed IF, et al (2019) Emerging threats and persistent conservation challenges for freshwater biodiversity. Biol Rev 94(3):849-873. https://doi.org/10.1111/brv.12480 Rodríguez-Rodríguez M, Benavente J, Cruz-San Julián JJ, Martos FM (2006) Estimation of ground-water exchange with semi-arid playa lakes (Antequera region, southern Spain). J Arid Environ 66(2):272-289. https://doi.org/10.1016/j.jaridenv.2005.10.018 Stefanidis K, Oikonomou A, Stoumboudi M, Dimitriou E, Skoulikidis NT (2021) Do water bodies show better ecological status in Natura 2000 protected areas than non-protected ones? The case of Greece. Water 13(21):3007. https://doi.org/10.3390/w13213007 Tjørve E (2003) Shapes and functions of species–area curves: a review of possible models. J Biogeogr 30(6):827-835. https://doi.org/10.1046/j.1365-2699.2003.00877.x UVEG (2022) Meta base de datos de lagos, lagunas y humedales de España. Valencia: Universitat de València Estudi General Van Gemerden H, Mas J (1995) Ecology of phototrophic sulfur bacteria. In: Blankenship RE, Madigan MT Bauer CE (eds) Anoxygenic photosynthetic bacteria. Springer, Dordrech, pp 49-85 t. Verhoeven JT (2014) Wetlands in Europe: perspectives for restoration of a lost paradise. Ecol Eng 66:6-9. https://doi.org/10.1016/j.ecoleng.2013.03.006 Verhoeven JT, Arheimer B, Yin C, Hefting MM (2006) Regional and global concerns over wetlands and water quality. Trends Ecol Evol 21:96–103. https://doi.org/10.1016/j.tree.2005.11.015 Supplementary Figures Supplementary Figures are not available with this version. Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 02 Aug, 2024 Read the published version in Environmental Earth Sciences → Version 1 posted Editorial decision: Accepted 06 Jun, 2024 Reviews received at journal 03 Jun, 2024 Reviewers agreed at journal 21 May, 2024 Reviews received at journal 21 May, 2024 Reviewers agreed at journal 21 May, 2024 Reviewers invited by journal 21 May, 2024 Submission checks completed at journal 02 May, 2024 Editor assigned by journal 02 May, 2024 First submitted to journal 30 Apr, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4350343","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":299588553,"identity":"b465f43a-dd31-4268-9788-f136b839afbc","order_by":0,"name":"Daniel Morant","email":"","orcid":"","institution":"Universitat de València","correspondingAuthor":false,"prefix":"","firstName":"Daniel","middleName":"","lastName":"Morant","suffix":""},{"id":299588556,"identity":"793ac95c-193c-4591-af07-3244fdb65db7","order_by":1,"name":"Alba Camacho-Santamans","email":"","orcid":"","institution":"Ecologia i Ciències Ambientals, Universitat de Barcelona","correspondingAuthor":false,"prefix":"","firstName":"Alba","middleName":"","lastName":"Camacho-Santamans","suffix":""},{"id":299588558,"identity":"0067e88c-b1d2-4c24-85cc-98db158af7d9","order_by":2,"name":"Rafael Hidalgo","email":"","orcid":"","institution":"Gobierno de España","correspondingAuthor":false,"prefix":"","firstName":"Rafael","middleName":"","lastName":"Hidalgo","suffix":""},{"id":299588560,"identity":"02fb7bdb-3930-4ec5-9166-c7c17ebf9f57","order_by":3,"name":"Antonio Camacho","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA3UlEQVRIiWNgGAWjYNACAyjN2MAgR4RyZlQtxkRqgQGglsQGQhr4Z/cffFxQYMPA33468cPPHTbpG24kMD78gUeLxJ3DzMYzDNIYJM7kbpbsPZOWC9TCbMyDz5obyWzSPAaHgd7J3SDN2HY4d8OZA2zS+HTIw7Xwv938G6gl3eDMAfaf+BxmANcikbsNZEuCwfEGNgZ8DjO8kWxszGOQxiNx4+02y962NMOZxxubpfFpkbuR+PAxzx8bOf7+3M03frbZyPMdZj74EZ/DYADZWGDsjIJRMApGwSigDAAAIIlJuLO6taoAAAAASUVORK5CYII=","orcid":"","institution":"Universitat de València","correspondingAuthor":true,"prefix":"","firstName":"Antonio","middleName":"","lastName":"Camacho","suffix":""}],"badges":[],"createdAt":"2024-04-30 16:06:33","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4350343/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4350343/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s12665-024-11700-4","type":"published","date":"2024-08-02T15:57:30+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":56105350,"identity":"59777bc5-051c-4ccc-9dc4-3831dd99bfd4","added_by":"auto","created_at":"2024-05-08 15:30:15","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":3118361,"visible":true,"origin":"","legend":"\u003cp\u003eLocation of the lakes or lake complexes assigned as THIC 3190 in Spain, with the delimitation of the biogeographic regions\u003c/p\u003e","description":"","filename":"MorantetalFigure1.png","url":"https://assets-eu.researchsquare.com/files/rs-4350343/v1/498a9d94f09b65e16358244e.png"},{"id":56105351,"identity":"da3df714-eeb3-4757-9290-122c9ea8a8b1","added_by":"auto","created_at":"2024-05-08 15:30:15","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":2445764,"visible":true,"origin":"","legend":"\u003cp\u003eMap of distribution and range of HTCI 3190 Karstic lakes on gypsum in Spain, generated from the Range tool in a GIS environment.\u003c/p\u003e","description":"","filename":"MorantetalFigure2.png","url":"https://assets-eu.researchsquare.com/files/rs-4350343/v1/425d8a54be413dab7770d118.png"},{"id":56105942,"identity":"5a375f4a-91ae-4342-8eda-b5ce820eaed2","added_by":"auto","created_at":"2024-05-08 15:38:15","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1091879,"visible":true,"origin":"","legend":"\u003cp\u003eExample of the results of one of the iterations for the species-area curve for THIC 3190, obtained by a random sorting iteration method.\u003c/p\u003e","description":"","filename":"MorantetalFigure3.png","url":"https://assets-eu.researchsquare.com/files/rs-4350343/v1/35bf8560e09279c9e9eabede.png"},{"id":61793916,"identity":"7f57ecae-f302-416f-9b1e-a0519e69b34a","added_by":"auto","created_at":"2024-08-05 16:16:49","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":8324220,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4350343/v1/f69ed3bf-6426-459c-bfe6-a4f44dd7bda6.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Transdisciplinary approach to the characterisation and current status of Spanish karstic lakes on gypsum","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eLentic ecosystems (standing waters) are affected worldwide by anthropogenic pressures, and they are one of the most threatened ecosystem types (Verhoeven \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Further, they are highly vulnerable to environmental shifts caused by climate change (Reid et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). These impacts may result in a degradation of the ecological structure and an alteration of their functioning (Dodson et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Camacho et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Proper management and conservation are essential aspects to ensure their ecological integrity, biodiversity, and functioning, for instance, promoting biogeochemical mechanisms that provide ecosystem services such as carbon sequestration (Morant et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). To this end, a proper assessment of their condition is essential, to better define mechanisms to maintain or improve their ecological condition and integrity. But for that purpose, it is also necessary a proper characterisation of the ecosystem type, with the knowledge of their main ecological features both abiotic and biotic.\u003c/p\u003e \u003cp\u003eIn Europe, the assessment of the status of lentic waterbodies is currently regulated by two European Directives that establish the framework for the management and conservation of aquatic ecosystems and water resources. The Water Framework Directive (WFD; 2000/60/EC) (European Commission \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) is the legal basis for water policy in the European Union (EU), which aims to achieve the good ecological status of aquatic ecosystems. The Habitats Directive (HD; 92/43/EEC) (European Commission \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1992\u003c/span\u003e) is the framework for the EU\u0026rsquo;s biodiversity policy, aiming to protect, maintain or restore, at a favourable conservation status, the Habitat Types of Community Interest (HTCI) and their characteristic species, as well as, particularly, specific species. Among these HTCI, there are some types that are directly, (Group 31 of Annex 1, standing waters) or indirectly associated with aquatic ecosystems.\u003c/p\u003e \u003cp\u003eThe \"ecological status\" (assessed according to the WFD) is an expression of the quality of the structure and functioning of the aquatic ecosystems associated with surface waters. Meanwhile, the \"conservation status\" (HD) is the set of influences on a natural habitat type and on their typical species that can affect its natural distribution, its structure and function in the long term, as well as the survival of its typical species. There is a partial convergence of the objectives set by both Directives with the achievement of the good status for waterbodies and the conservation of HTCI and species included in Annexes II and IV of the HD as the final goals (Stefanidis et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), by considering management and protection measures to improve water-dependent habitats, species and water quality (European Commission \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe definition and classification of ecosystems and their associated habitats is essential, first to identify them, then to allow a proper assessment and management. HTCI in general, and habitats associated to aquatic ecosystems in particular, are usually described based on the structuring role of plants in the ecosystems (DG Environment \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). As in other HTCI, in lentic habitats, plant communities can also play a determining role, but other defining elements of the habitat type are also needed for a proper habitat characterisation. Therefore, and especially in the case of HTCI in group 31 (standing waters), attention must be also paid to the global ecological characteristics of the habitat type, knowing that the occurrence of a HTCI is not solely described by the presence or absence of certain taxa, but rather by a sort of biotic and abiotic features. Consequently, the assessment of the conservation status must be based on the consideration of all those abiotic and biotic properties as defining the habitat /ecosystem type.\u003c/p\u003e \u003cp\u003eWithin the group of lentic ecosystems, which can be associated with the habitat types of standing waters (group 31 of Annex I of the HD), there are some characteristic lake and pond types, very rare worldwide, featured by their karstic character on areas rich in gypsum, providing specific ecological properties being key in their definition as a specific type of ecosystem and HTCI (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). In this work we focus on this HTCI 3190, \u0026ldquo;Karstic lakes on gypsum\u0026rdquo;, one of the less known HTCI of Annex I of the HD from an ecological point of view. Officially, there are only 19 sites within the N2000 network classified as HTCI 3190 throughout the EU, 9 in Germany, 5 in Spain, 3 in Latvia, and 2 in Lithuania (European Environmental Agency \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe objectives of this paper were 1) to improve the information and knowledge of this restrictive HTCI (3190), with a detailed ecological description, 2) to identify the habitat and assess its global conservation status in Spain from the four parameters by the established protocols, following a scientific approach (Range of distribution, Area of occupancy, Structure and function, and Future prospects), and 3) to make a critical analysis of the methodologies established for the evaluation and the global value of the conservation status.\u003c/p\u003e"},{"header":"2. Material and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Ecological definition\u003c/h2\u003e \u003cp\u003eFollowing the Interpretation manual of European Union habitats EUR28 (DG Environment \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), a deeper ecological characterisation of the Spanish karstic lakes on gypsum from the ecological features described in the scientific literature, accompanied with several field campaigns, was carried out. The main ecological features, including abiotic and biotic parameters, as well as the presence of the typical species, were used for the classification of the lentic habitat sites featured as HTCI 3190.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Validation of sites classified as HTIC 3190: karstic lakes on gypsum\u003c/h2\u003e \u003cp\u003eA validation of lakes with potential features linked to the HTCI 3190 previously selected from a database was carried out by overlaying the localities to the Geological Map of Spain 1:50,000 (GEODE) to ensure they were located on karstic areas with gypsum content. A deeper study of the rest of abiotic and biotic parameters, from bibliographic references and field data, was done to ensure they fit to the ecological description of the karstic lakes on gypsum. Some small ponds or other localities not included in the database up to the moment of the study could not be included in this assessment, though ongoing updates of the database would slightly increase the number of lakes identified as HTCI 3190 in Spain.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Selection of a minimum number of sites and data images sources\u003c/h2\u003e \u003cp\u003eIn order to select a representative number of Iberian sites for the assessment of the conservation status among all the sites identified as HTCI 3190, specific criteria were used to avoid biases in the assessment and extrapolation of the conservation status. The minimum number of statistically representative sites (n) was obtained from the total population (total number of lakes on gypsum karst officially catalogued in Spain until the moment of the work) using the following formula (Camacho et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019a\u003c/span\u003e):\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$n=\\frac{{Z}_{\\alpha }^{2}Npq}{{e}^{2}\\left(N-1\\right)+{Z}_{\\alpha }^{2}pq}$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003ewhere:\u003c/p\u003e \u003cp\u003eN\u0026thinsp;=\u0026thinsp;population size (total number of lakes on gypsum karst in Spain)\u003c/p\u003e \u003cp\u003eZ\u003csub\u003eα\u003c/sub\u003e= confidence level, expressed in values relative to the desired confidence percentage. The most used values are: 1.28 for a confidence level of 80%, 1.44 for 85%, 1.65 for 90%, 1.69 for 91%, 1.75 for 92%, 1.81 for 93%, 1.88 for 94%, 1.96 for 95% and 2.58 for 99% (Camacho et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019a\u003c/span\u003e)\u003c/p\u003e \u003cp\u003ep\u0026thinsp;=\u0026thinsp;probability of success. As any study characteristics for success are defined in this case, by default it is usually assumed that p\u0026thinsp;=\u0026thinsp;q\u0026thinsp;=\u0026thinsp;0.5\u003c/p\u003e \u003cp\u003eq\u0026thinsp;=\u0026thinsp;proportion of localities that do not have the probability of success, q\u0026thinsp;=\u0026thinsp;1-p\u003c/p\u003e \u003cp\u003ee\u0026thinsp;=\u0026thinsp;desired sampling error (in percentage)\u003c/p\u003e \u003cp\u003eOnce the minimum number of sites to be assessed was determined, specific criteria to select the sites were defined (Camacho et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019a\u003c/span\u003e) as follows:\u003c/p\u003e \u003cp\u003e \u003cul\u003e \u003cli\u003e \u003cp\u003eExtent, covering the largest area possible per site within the type, avoiding evaluating a greater number of different sites.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eRepresentativeness in the Natura 2000 Network and other protection figures at the international (Ramsar), national or regional level, also considering sites not officially classified as such.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eThreat status (or danger of disappearance), with a selection of sites in, a priori, both good and poor condition, indistinctively.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eEnvironmental-ecological diversity subtypes, e.g., shallow and deeper lakes, small ponds and bigger lakes).\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eExisting information.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eEnvironmental and geographic variability.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eAccessibility and representativeness of the localities.\u003c/p\u003e \u003c/li\u003e \u003c/ul\u003e \u003c/p\u003e \u003cp\u003eEach of these criteria were valued and weighted to obtain the final sites to be included in the assessment as representative of the habitat type.\u003c/p\u003e \u003cp\u003eOnce all the sites classified as HTCI 3190 were designed, and the sites for their assessment were selected, the assessment of the four parameters of the conservation status established for the reporting of the Article 17 of the HD was carried out. The assessment for each site was obtained by determining, for the four parameters (\u0026lsquo;Range of distribution\u0026rsquo;, \u0026lsquo;Area of occupancy\u0026rsquo;, \u0026lsquo;Structure and function\u0026rsquo;, and \u0026lsquo;Future prospects\u0026rsquo;) an evaluation as favourable, unfavourable-inadequate or unfavourable-bad status, by comparing the period 2013\u0026ndash;2018 (following the Article 17 of the HD) with Google Earth images collected along this period for the determination of the \u0026lsquo;Range of distribution\u0026rsquo; and \u0026lsquo;Area of occupancy\u0026rsquo; parameters. Further than the assessment of changes along the period 2013\u0026ndash;2018 used for the reporting according to the Article 17 of the HD, a historical recreation based on images collected in several flight missions was build-up. These missions were: American Flight 1956\u0026ndash;1957; \u003cem\u003eInterministerial\u003c/em\u003e Flight 1973\u0026ndash;1986; National Flight 1981\u0026ndash;1986; Five-Year Flight 1998\u0026ndash;2003, and current (since 2012) high-resolution images series from the Spanish National Aerial Orthophotography Plan (PNOA), available at the Spanish Centre for Geographic Information (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://centrodedescargas.cnig.es/CentroDescargas/index.jsp\u003c/span\u003e\u003cspan address=\"https://centrodedescargas.cnig.es/CentroDescargas/index.jsp\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4. Range of distribution\u003c/h2\u003e \u003cp\u003eThe \u0026lsquo;Range of distribution\u0026rsquo; of a habitat type is defined as the outer limits of the general area in which the habitat is found. The range of the HTCI 3190 in Spain was obtained from all the localities designed as 3190, using the Range Tool (EIONET \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) in a GIS environment, following the guidelines of the toolbox (Camacho et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2019b\u003c/span\u003e). A gap (or maximum distance between grids of distribution) of 40 km was used, as being karstic lakes on gypsum a non-zonal habitat type, following the unified criteria of the European Commission (DG Environment \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe status assessment of the parameter \u0026lsquo;Range of distribution\u0026rsquo; was defined by comparing the range obtained in 2018 with the range obtained in 2013. The favourable reference range (FRR) was considered as the range when the Directive came into force, in 1994.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.5. Area of occupancy\u003c/h2\u003e \u003cp\u003eThe parameter 'Area of occupancy\u0026rsquo; of a habitat type accounts for the real area it occupies. Each of the sites included as HTCI 3190 were delineated by using the Google Earth Pro\u0026reg; delineation tools as described by Camacho et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2019b\u003c/span\u003e), differentiating the total area of the wetland, then breaking it down further into its different components: maximum flooded area, current flooded area, marginal vegetation coverage, other land uses or coverages. As said, not only the current flooded area, but also other areas of the wetland that have the characteristics of the habitat type are considered when assessing the area coverage at each site.\u003c/p\u003e \u003cp\u003eFor the assessment of the parameter 'Area of occupancy\u0026rsquo;, only the total surface parameter was used, being quantified with images of 2018 (or the closest), and 2013, to assess the differences among the 6-year period established between the assessment reports. As for the \u0026lsquo;Range of distribution\u0026rsquo; FRR, the favourable reference area (FRA) was considered as the area when the Directive came into force in 1994. The latter was obtained by viewing and delineating each of the localities with historical images in a GIS (Camacho et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2019b\u003c/span\u003e). Alternatively, the reference area was also estimated based on the minimum area that can support all species associated with the habitat type. In this method the cumulative surface area of each site added was correlated with the cumulative species present in the set of sites evaluated. The sorting of these sites when adding the surface areas and species to the curve was done randomly using 999 random iterations. An average curve was drawn showing the response of the coverage of the whole set of species with an increase in the total area of the habitat type when adding sites into the curve build-up.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e2.6. Structure and function\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe parameter \u0026lsquo;Structure and function\u0026rsquo; refers to the assessment of the status of the intrinsic ecological characteristics of the habitat types or ecosystems. Structure is understood as the configuration of the different biotic and abiotic elements, while the function refers to the processes linked to those elements. According to the procedures for the monitoring of habitat types, the ECLECTIC multimetric index was used to assess the \u0026lsquo;Structure and function\u0026rsquo; (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2019c\u003c/span\u003e). This index considers typical biological, physical-chemical, and hydromorphological variables of the habitat/ecosystem type, whose values are compared with reference values for each variable specific for the habitat type, from which a quantification was obtained. Some of these variables are compulsory, meanwhile others are optional for the final assessment (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The final value of the ECLECTIC index was obtained after the weighting of each of the variables (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), being the HTCI at each site assessed as being in a favourable status when ECLECTIC\u0026thinsp;\u0026ge;\u0026thinsp;70, an unfavourable-inadequate status when 50\u0026thinsp;\u0026le;\u0026thinsp;ECLECTIC\u0026thinsp;\u0026lt;\u0026thinsp;70, and unfavourable-bad status when ECLECTIC\u0026thinsp;\u0026lt;\u0026thinsp;50.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e2.7. Future prospects\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe evaluation of the parameter \u0026lsquo;Future prospects\u0026rsquo; for the conservation of lentic habitat types or ecosystems was associated with the estimation of the pressures and threats. The methodology used was described by Camacho et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). A group of pressures and impacts was evaluated in a weighted way, considering hydrological and geomorphological pressures and impacts, water quality, pressures and impacts on the structure of the communities, land uses, or presence of invasive species, accordingly to the potential impact each can have on the conservation status of the HTCI 3190. As calibrated, overall scores between 0\u0026ndash;20 points, mean a low level of pressures and threats, thus a favourable conservation status. Scores between 21\u0026ndash;50, indicated moderate pressures and threats, meaning an unfavourable-inadequate status, whereas scores higher than 51 points show high or very high level pressures and threats, thus an unfavourable-bad status.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e2.8. Conservation status\u003c/h2\u003e \u003cp\u003eFor the assessment of the conservation status at the level of the biogeographical region, the assessments carried out in each of the selected sites for the parameters 'Structure and function' and 'Future prospects' were weighted by its occupied surface area:\u003cdiv id=\"Equb\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equb\" name=\"EquationSource\"\u003e\n$$Global parameters assessment= \\sum _{i=1}^{n}index value site*\\% surface site [up to 18.7\\%]$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eConsidering the great difference between the surface areas in the different sites due to the nature of THIC 3190, in which there are small ponds of a few square metres, to large lakes with extensions of several hectares, a factor was applied limiting the surface area over which the results could be extrapolated. This factor was three times the percentage of a site in relation to the total, if they all had the same surface area (1/16 selected sites)*3 (%). The results of the assessment of the parameters \u0026lsquo;Range of distribution\u0026rsquo; and \u0026lsquo;Area of occupancy\u0026rsquo; were obtained by the sum of the values for all sites assessed.\u003c/p\u003e \u003cp\u003eOnce the four parameters included in the general conservation status assessment matrix were evaluated, the results were provided as: favourable, unfavourable-inadequate, unfavourable-bad and unknown, following the guidelines of DG Environment (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The global value of the conservation status of the HTCI 3190 for each of the three biogeographic regions in Spain (Mediterranean, Atlantic and Alpine, given that this HTCI is not present in the Canary Islands \u0026ndash; Macaronesia) was obtained according to the evaluation matrix, using the combination procedure given in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eTable 1\u003c/strong\u003e Procedure for the combination of the results of the conservation status using the four parameters of the evaluation matrix, for the assessment of the conservation status for a HTCI at the biogeographical region scale. Source: DG Environment (2017)\u003c/p\u003e\n\u003ctable style=\"width: 4.3e+2pt;border-collapse:collapse;border:none;\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width:0in;border:solid windowtext 1.0pt;background:#F2F2F2;padding:0in 5.4pt 0in 5.4pt;height:55.0pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:black;'\u003eStatus of the parameters\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:0in;border:solid windowtext 1.0pt;border-left:none;background:#F2F2F2;padding:0in 5.4pt 0in 5.4pt;height:55.0pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height: 115%;font-family:\"Times New Roman\",serif;color:black;'\u003eAll\u0026nbsp;\u003c/span\u003e\u003c/strong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:black;'\u003ef\u003cstrong\u003eavourable, or\u003c/strong\u003e \u003cstrong\u003ethree\u0026nbsp;\u003c/strong\u003ef\u003cstrong\u003eavourable and one\u0026nbsp;\u003c/strong\u003eu\u003cstrong\u003enknown\u003c/strong\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:0in;border:solid windowtext 1.0pt;border-left:none;background:#F2F2F2;padding:0in 5.4pt 0in 5.4pt;height:55.0pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:black;'\u003eOne or more\u0026nbsp;\u003c/span\u003e\u003c/strong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:black;'\u003ei\u003cstrong\u003enadequate, but no\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003ebad\u003c/strong\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:0in;border:solid windowtext 1.0pt;border-left:none;background:#F2F2F2;padding:0in 5.4pt 0in 5.4pt;height:55.0pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height: 115%;font-family:\"Times New Roman\",serif;color:black;'\u003eOne or more\u0026nbsp;\u003c/span\u003e\u003c/strong\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family: \"Times New Roman\",serif;color:black;'\u003ebad\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:0in;border:solid windowtext 1.0pt;border-left:none;background:#F2F2F2;padding:0in 5.4pt 0in 5.4pt;height:55.0pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:black;'\u003eTwo or more\u0026nbsp;\u003c/span\u003e\u003c/strong\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:black;'\u003eunknown\u003c/span\u003e\u003c/strong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:black;'\u003e\u0026nbsp;\u003cstrong\u003ecombined with\u0026nbsp;\u003c/strong\u003ef\u003cstrong\u003eavourable, or all\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eunknown\u003c/strong\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width:78.0pt;border:solid windowtext 1.0pt;border-top: none;padding:0in 5.4pt 0in 5.4pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;'\u003eOverall assessment\u0026nbsp;conservation status\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:92.1pt;border-top:none;border-left:none;border-bottom:solid windowtext 1.0pt;border-right:solid windowtext 1.0pt;padding:0in 5.4pt 0in 5.4pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height: 115%;font-family:\"Times New Roman\",serif;color:#00B050;'\u003eFavourable\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:85.05pt;border-top:none;border-left:none;border-bottom:solid windowtext 1.0pt;border-right:solid windowtext 1.0pt;padding:0in 5.4pt 0in 5.4pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height:115%;font-family:\"Times New Roman\",serif;color:#FF6600;'\u003eUnfavourable-inadequate\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:85.05pt;border-top:none;border-left:none;border-bottom:solid windowtext 1.0pt;border-right:solid windowtext 1.0pt;padding:0in 5.4pt 0in 5.4pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height: 115%;font-family:\"Times New Roman\",serif;color:red;'\u003eUnfavourable-bad\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width:85.05pt;border-top:none;border-left:none;border-bottom:solid windowtext 1.0pt;border-right:solid windowtext 1.0pt;padding:0in 5.4pt 0in 5.4pt;\"\u003e\n \u003cp style='margin-top:3.0pt;margin-right:0in;margin-bottom:3.0pt;margin-left:0in;font-size:11.0pt;font-family:\"Calibri\",sans-serif;text-align:center;line-height:115%;'\u003e\u003cstrong\u003e\u003cspan style='font-size:12px;line-height: 115%;font-family:\"Times New Roman\",serif;color:gray;'\u003eUnknown\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"},{"header":"3. Results","content":"\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e3.1. Ecological characterisation\u003c/h2\u003e \u003cp\u003eKarstic lakes on gypsum, mostly feed by groundwater, are formed by the collapse of karstic structures that evolved from the dissolution of materials on a basin lying on gypsum (CaSO\u003csub\u003e4\u003c/sub\u003e), a highly soluble material that provides the water with a high concentration of sulphates that normally exceed bicarbonates, whose concentration is also high as in other types of karstic lakes. The morphometry of the basin is usually of the sinkhole type, generally with steep slopes walls and a high relative depth, although some basins (ponds) can just display small depressions formed by dissolution when collapse has not yet occurred. The size of the lake is usually small, unless the lake spans throughout several deep sinkholes or even a polje. Their waters are generally subsaline (\u003cem\u003esensu\u003c/em\u003e Hammer \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1986\u003c/span\u003e), although the conductivity range (generally 1.5-5 mS cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) can occasionally overlap with the oligosaline or the low hyposaline range (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). In lakes with a high relative depth, the formation of a sulphide-rich hypolimnion allows of its colonisation by green and purple sulphur bacteria. Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e indicates the main abiotic features of the HTCI \u0026ldquo;karstic lakes on gypsum\u0026rdquo;, used for the determination of sites assigned to the HTCI 3190. Communities of Charetea, Lemnetea and Potamogetonion are the dominant elements of the submerged vegetation (DG Environment \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eMain abiotic properties characterising THIC 3190, and other characteristic data.\u003c/p\u003e \u003cdiv class=\"Credit\"\u003e\u003cp\u003eSource: (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"2\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVariable\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFeature\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrophic status\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eOligo-mesotrophic\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMineralisation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRelatively high (freshwaters \u0026ndash;oligosaline to subsaline). Conductivity: 1\u0026ndash;5 mS/cm\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eContent in bases\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHigh\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater colour\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eColourless\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026gt;\u0026thinsp;7,5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTransparency\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHigh\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHydroperiod\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGenerally permanent, sometimes temporary\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLithology\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGypsum and limestones\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSubstrate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMarls\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDepth\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGenerally deep (\u0026gt;\u0026thinsp;2 m)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDistribution in the Iberian Peninsula\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGypsum-rich areas\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe definition of \u0026ldquo;karstic lakes on gypsum\u0026rdquo; by the EU Interpretation Manual does not fit to all lakes that lie on gypsum. From the geological and hydrogeological point of view, this name could include a much significant number of wetlands in Spain because gypsum outcrops broadly extend in the Spanish Alpine Orogenes and in some Neogene basins. However, many of these lakes present hypersaline waters; therefore, the Habitats Directive classified them differently as the definition of the HTCI 3190 includes only subsaline lakes.\u003c/p\u003e \u003cp\u003eDeeping on the short definition given by the interpretation manual for Habitats of Community Interest (DG Environment \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), Spanish karstic lakes on gypsum, are characterised by a high concentration of sulphate, allowing that, in anaerobic respiration processes in the sediments or in the anoxic hypolimnion (for deep enough lakes), sulphate can be used as an electron acceptor (Atlas and Bartha \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), causing the accumulation of hydrogen sulphide in the deep layers while stratification is maintained, which greatly determines the biological communities in these depths. If altered by increased inputs of nutrients, phytoplankton massive growth in surface waters derived from a hypothetical eutrophication process would considerably reduce light penetration and hinder or prevent the formation of dense deep populations of photosynthetic microorganisms that normally include cryptophyceans, cyanobacteria and/or photosynthetic bacteria, around the oxic-anoxic interface (Camacho \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), which characteristic of this habitat.\u003c/p\u003e \u003cp\u003eUnder natural (non-altered) conditions, and linked to the arrangement of the populations in the vertical profile mediated by the physical-chemical structuring of the water column, surface waters are usually relatively poor in nutrients, especially in phosphorus. Therefore, phytoplankton growth is controlled by the availability of these nutrients mediated by recirculation. Surface chlorophyll concentrations are naturally in the oligo-mesotrophic range (usually below 10 mg m\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e). However, very high concentrations of chlorophyll and/or bacteriochlorophyll of even several hundred mg m\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e can be found in depth around the oxic-anoxic interface (Camacho \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Chicote \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). This summer chlorophyll maximum can thus be considered as characteristic of this type of ecosystems in conditions of good functional conservation. In the Iberian stratified karstic sinkholes (both on gypsum and limestone) the formation of these chlorophyll metalimnetic maxima is common (Camacho \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, Camacho et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), generally formed by one or a few species of cryptophyceans (Pedr\u0026oacute;s-Ali\u0026oacute; et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Gasol et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Garc\u0026iacute;a-Gil et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Camacho et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2001a\u003c/span\u003e; Chicote \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) and/or cyanobacteria (Camacho et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Camacho et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2000a\u003c/span\u003e), which form dense populations at the oxic-anoxic interface coinciding with the bottom of the thermocline. In this type of lakes, the appearance, in anoxic zones, of populations of photosynthetic sulphur bacteria that use hydrogen sulphide as an electronic donor for photosynthesis is also a common feature (Van Gemerden and Mas \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Camacho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2000b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn karstic lakes on gypsum with metalimnetic chlorophyll maxima, the highest photosynthetic rates, in absolute terms, can be found in the narrow range of depths occupied by these deep populations of algae, cyanobacteria, and/or photosynthetic bacteria (Planas \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e1973\u003c/span\u003e, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Camacho and Vicente \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1998\u003c/span\u003e), although their productivity (production per biomass unit) is normally lower than that of more superficial layers (Camacho \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). This, together with the narrow range of depths they cover, means that only a part of the primary planktonic production is located in the deep metalimnion, although in certain systems the metalimnetic contribution may account for most of the primary planktonic production of the lake (Camacho et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2001b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDue to the high slope of the shores, the relative extent of the littoral zone is generally small compared to the pelagic zone (open waters), although the saturated zone colonised by helophytes can be quite extensive, especially in non-collapsed shallow ponds. When the subsidence is small and the depth of the basin formed is low, the flooding is maintained only temporarily or occurs in the central area, originating a shallow lake essentially with littoral characteristics, and a higher relative importance of macrophyte communities. The submerged macrophytic vegetation can include both charophytes and higher plants, as well as several species of emergent helophytes in the littoral zone (Cirujano \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1990\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Cirujano and Medina \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2002\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe contribution of organic matter from the marginal and riverside vegetation, mainly the remains of leaves from deciduous trees, can generate an important contribution of allochthonous organic materials that, given the often small reduced size of the lakes, can induce an increase in the trophic level and, indirectly, a greater production of hydrogen sulphide. Even some of these lakes can remain completely anoxic during a part of the annual cycle, with high concentrations of hydrogen sulphide and populations of purple sulphur bacteria that can reach up to the surface, giving a characteristic red colour to the waters.\u003c/p\u003e \u003cp\u003eIn the Iberian Peninsula, karstic lakes on gypsum show zooplankton populations with a heterogeneous vertical distribution, with species of crustaceans and rotifers with a distribution mainly epilimnetic, others with a metalimnetic distribution (Alfonso et al. 1987; Esteve et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Miracle and Alfonso \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Gasol et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Miracle and Armengol 1995) or with migratory capacity, and even populations of anaerobic ciliates (thanks to mutualistic symbiosis with bacteria; Finlay et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Esteban et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1993\u003c/span\u003e) that colonise the anoxic hypolimnion rich in hydrogen sulphide in a markedly reducing environment. In the systems exclusively fed by groundwater (small sinkholes) without connections to surface watercourses, fish, when they exist, are generally introduced, while in systems with surface water connections, which serve as dispersal, the fish community may be more diverse and more structured by ecological interactions (Granado \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2000\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e3.2 Sites identified as karstic lakes on gypsum in Spain and selection of sites for the assessment of the structure and function, and future prospects\u003c/p\u003e\u003cp\u003eConsidering the ecological characterisation, and after the validation process following the procedures indicated in the Methods section, up to 79 lakes from the available data set on which we started (UVEG \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), some of them grouped in lake complexes, were finally considered in Spain as belonging to THIC 3190 (Supplementary Table\u0026nbsp;1). Sites are mostly distributed throughout the East half of the Iberian Peninsula, in three of the four biogeographic regions of Spain. Though they are mostly located in the Mediterranean biogeographic region, they also appear in the Atlantic and Alpine regions, but do not appearing in the Macaronesian Canary Islands. Part of the typified sites were located in lakes complexes, associated with geological areas characterised by the presence of gypsum, and the genesis by karst dissolution. Figure\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e represents the site of the THIC throughout the Spanish territory and their location within the three biogeographic regions of the Spanish part of the Iberian Peninsula and the Balearic Islands (through this HTCI was not identified so far in the Balearic Islands).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFollowing the number of sites needed for a proper assessment of the for the assessment of the structure and function, and future prospects (with the population size of 79 sites, and a defined error of 15%), and considering the amount of information necessary to carry out the relevant evaluation, the selection of a sample size was estimated with a total of 18 lakes representative of the type of habitat/ecosystem, based on the criteria defined in the \u003cspan refid=\"Sec2\" class=\"InternalRef\"\u003ematerial and methods\u003c/span\u003e section. Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e shows the sites finally selected as representative of the heterogeneity and intrinsic characteristics, protection and availability of information on the type of habitat/ecosystem.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFinal selection of sites evaluated as representative from the HTCI 3190 Karstic lakes on gypsum in Spain according to the defined criteria. (ALP: Alpine; ATL: Atlantic; MED: Mediterranean biogeographical region). Codes of the different protection figures (Ramsar, Water Framework Directive, WFD, and Spanish Inventory of Wetlands) are also included in each lake included in each case.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRegion\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHydroperiod\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eProvince\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLake complex\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eNatura 2000 network\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eRamsar\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eWaterbody (WFD)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eCode Spanish Inventory of Wetlands\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLago de Arreo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026Aacute;lava\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES2110007\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1258\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES091MSPF1019\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIH211002\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Arbieto\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eATL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVizcaya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanque Grande de Abajo de Esta\u0026ntilde;a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eHuesca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEsta\u0026ntilde;a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES2410072\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES091MSPF1014\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Tortajada\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTeruel\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES2420131\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Barcena,\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBurgos\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGayanes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Alboraj\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAlbacete\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES4210011\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIH421059\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de La Atalaya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCuenca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFuentes - R\u0026iacute;o Moscas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES080MSPFL11_a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de los Capellanes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTemporary\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCuenca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIH423033\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstany de Banyoles\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGirona\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBanyoles\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES5120008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1257\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES100MSPF0450401\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanyol de Vil\u0026agrave;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGirona\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBanyoles\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES5120008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1257\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES100MSPF0450401\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanyol de La Coromina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGirona\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES100MSPFH1040040\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstany de Montcort\u0026eacute;s\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eALP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eLleida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES5130019\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES091MSPF1029\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Z\u0026oacute;\u0026ntilde;ar\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eC\u0026oacute;rdoba\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSur de C\u0026oacute;rdoba\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES0000034\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e446\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES050MSPFES0512000007\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIH613017\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna Grande de Archidona\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u0026aacute;laga\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eArchidona\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1911\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES060MSPF0614520\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIH617023\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBallesteros-5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCuenca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eArcas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES4230008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES080MSPFL13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIH423003 (refers to the complex)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArcas-2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCuenca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eArcas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES4230008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES080MSPFL13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArcas-4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTemporary\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCuenca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eArcas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES4230008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES080MSPFL13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLagunillo de las Tortugas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePermanent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCuenca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eES4230014\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eES080MSPFL12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIH423006 (refers to the complex)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.3. Range and area of distribution of HTCI 3190 in Spain\u003c/h2\u003e \u003cp\u003eThe sites classified as HTCI 3190, karstic lakes on gypsum, showed an uneven distribution throughout the Iberian Peninsula (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), being located mainly concentrated in specific areas.\u003c/p\u003e\u003cp\u003eThe Mediterranean region showed sites spread in several areas of its eastern part, with a range of distribution of 3,000 km\u003csup\u003e2\u003c/sup\u003e. The Alpine region showed only one site (Estany de Montcort\u0026eacute;s, in Lleida), with a range, equivalent to the distribution, of 100 km\u003csup\u003e2\u003c/sup\u003e. Finally, two sites were in the Atlantic region (Laguna de Olandina, in \u0026Aacute;lava, and Laguna de Arbieto, in Vizcaya), with a range of distribution of 200 km\u003csup\u003e2\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe area of the HTCI 3190 (considering the 79 sites classified) distributed in the three biogeographic regions in Spain shows a clear predominance of the Mediterranean region, with more than 94% of the total surface of this HTCI in Spain, and with very specific areas located in the Alpine and Atlantic regions (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). All the karstic lakes on gypsum occupied an area of 308 ha in Spain. The maximum water flooding area was around 250 ha, meanwhile the emerged vegetation (helophytes) coverage was of 83 ha (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eArea covered by the HTCI 3190 and its different components at the end of the period 2013\u0026ndash;2018 using Google Earth Pro\u0026reg; by biogeographic regions. (ALP: Alpine; ATL: Atlantic; MED: Mediterranean region). Since these are average values, an overlap can occur.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSurface (ha)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMaximum\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCurrent\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEmerged vegetation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eOthers\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eALP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e15.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e12.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e12.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eATL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e290.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e236.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e203.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e79.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e7.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e308.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e249.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e216.5\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e83.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e7.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe assessment of the changes separated by biogeographic regions showed stable values in the Alpine and Atlantic regions, and changes of around 1% in the total surface in the Mediterranean region (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e) for the six-years evaluation period. The variations in the current flooding surface showed negative trends in contrast with the increases in the marginal vegetation coverage in this evaluation period, especially in lower latitudes of the Mediterranean region, being the \u0026ldquo;current flooding area\u0026rdquo; more linked with hydroclimatic features.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eValues of change (in ha) in occupied surface area of HTCI 3190 during the period 2013\u0026ndash;2018 using Google Earth Pro\u0026reg; by biogeographic regions in Spain. (ALP: Alpine; ATL: Atlantic; MED: Mediterranean region). Positive values represent an increase in coverage, whereas negative values show a decrease.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAssessment period (ha) 2013\u0026ndash;2018\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMaximum\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCurrent\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVegetation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eOthers\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eALP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eATL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-7.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e3.4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e-7.8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e9.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eBoth range and area covered by the HTCI 3190 did not significantly decrease when compared to the values when the Habitats Directive came into force (1994), as assessed by historic aerial images (Supplementary Figs.\u0026nbsp;1 to 26), therefore, they were assessed as favourable for the three biogeographic regions. Using the alternative procedure of the species-area curve, a 'favourable reference area' (FRA) for this THIC at national level could be determined of being around 300 ha, where increases in the total area of the THIC did not associate to large increases in the accumulation of species associated with the habitat type (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Thus, the 'Area of occupancy\u0026rsquo; of this HTCI in Spain is still above the FRA, and this assessment procedure for the 'favourable reference area' also gives a favourable conservation status when all country is considered for this parameter.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e3.4. Structure and function, Future prospects, and overall conservation status\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eWhen assessing the \u0026lsquo;Structure and function\u0026rsquo; parameter using the ECLECTIC index, the results obtained for each of the 18 sites selected as representative of the type, compiled in Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e, showed that none of the assessed sites had an unfavourable-bad status (U2). However, notable differences appear between some of the studied sites, with 50% of them in an unfavourable-inadequate status (U1), and another 50% with a favourable status.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFinal value of the application of the ECLECTIC index, both with the evaluation of the compulsory variables, and with the evaluation of all variables (compulsory and optative), based on the available information, for the 18 sites selected as representative. The complete and disaggregated evaluations are compiled in Supplementary Table\u0026nbsp;2 (FV: favourable status; U1: unfavourable-inadequate status; ALP: Alpine; ATL: Atlantic; MED: Mediterranean region)\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRegion\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eECLECTIC compulsory index value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eStatus\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eECLECTIC all index value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eStatus\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLago de Arreo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e73.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e78.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Arbieto\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eATL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e85.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e79.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanque Grande de Abajo de Esta\u0026ntilde;a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e92.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e86.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Tortajada\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e60.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e59.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Barcena\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e87.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e88.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Alboraj\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e63.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e63.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de La Atalaya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e57.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e59.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de los Capellanes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e67.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e66.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstany de Banyoles\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e65.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e61.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanyol de Vil\u0026agrave;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e63.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e63.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanyol de La Coromina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e59.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e59.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstany de Montcort\u0026eacute;s\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eALP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e87.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e83.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Z\u0026oacute;\u0026ntilde;ar\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e56.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e56.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna Grande de Archidona\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e80.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e84.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBallesteros-5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e95.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e95.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArcas-2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e91.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArcas-4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e95.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e95.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLagunillo de las Tortugas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e52.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eRegarding the \u0026lsquo;Future prospects\u0026rsquo; parameter, the results obtained for each of the 18 sites, compiled in Table\u0026nbsp;\u003cspan refid=\"Tab7\" class=\"InternalRef\"\u003e7\u003c/span\u003e, showed the values obtained from the pressure and threat evaluation method, and their equivalence to the conservation status class, with up to 10 sites with low levels of pressure, which was assimilated to favourable status for this parameter. In the same way, 7 sites, with moderate pressures, showed an unfavourable-inadequate status, and only one showed high pressure levels and threats, and thus, a bad status.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab7\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 7\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFinal value of the application of the pressure and threat index, based on the available information, and structure and function for comparison, for the 18 sites selected as representative for the assessment. The complete and disaggregated evaluations are compiled in Supplementary Table\u0026nbsp;3. FV: favourable status; U1: unfavourable-inadequate status; U2: unfavourable-bad status; ALP: Alpine; ATL: Atlantic; MED: Mediterranean biogeographic region; P\u0026amp;A: Pressures and threats; S\u0026amp;F: Structure and function)\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRegion\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eP\u0026amp;T index value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eP\u0026amp;A Status\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u0026amp;F Status\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLago de Arreo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Arbieto\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eATL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanque Grande de Abajo de Esta\u0026ntilde;a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Tortajada\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Barcena\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Alboraj\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de La Atalaya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de los Capellanes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstany de Banyoles\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e106\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanyol de Vil\u0026agrave;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstanyol de La Coromina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEstany de Montcort\u0026eacute;s\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eALP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna de Z\u0026oacute;\u0026ntilde;ar\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLaguna Grande de Archidona\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBallesteros-5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArcas-2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArcas-4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLagunillo de las Tortugas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eSupplementary Table\u0026nbsp;4 collects the references to assess the \u0026lsquo;Structure and function\u0026rsquo; and \u0026lsquo;Future prospects\u0026rsquo; in each selected site following the indices detailed.\u003c/p\u003e \u003cp\u003eComparing the status of the \u0026lsquo;Structure and function\u0026rsquo; parameter, from the ECLECTIC index, with the \u0026lsquo;Future prospects\u0026rsquo; parameter, from pressures and threats, almost all the sites showed similar status assessment (Table\u0026nbsp;\u003cspan refid=\"Tab7\" class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWhen considering the set of the statistically representative sites, per biogeographic region, and following the weighting and extrapolation method explained above, both the \u0026lsquo;Structure and function\u0026rsquo; and \u0026lsquo;Future prospects\u0026rsquo; were classified as unfavourable-inadequate in the Mediterranean region, as the weighted final value of the ECLECTIC index was 62.2, slightly below the limit established by the favourable status (70), and the total value for pressures and threats extrapolated from the weighting by area meant a value of 45, meaning moderate pressure levels, and thus, unfavourable inadequate status. Both parameters were classified as favourable in the representative lakes for the Atlantic (Laguna de Arbieto) and the Alpine (Estany de Montcort\u0026eacute;s) biogeographical regions (Table\u0026nbsp;\u003cspan refid=\"Tab8\" class=\"InternalRef\"\u003e8\u003c/span\u003e), as the values of the ECLECTIC and pressures and threats indices indicated the good condition and lower pressures level in the two sites assessed for these regions.\u003c/p\u003e \u003cp\u003eCombining these two parameters with the previous assessment of the \u0026lsquo;Range of distribution\u0026rsquo; and \u0026lsquo;Area of occupancy\u0026rsquo; in the General Matrix, the final conservation status for the Mediterranean region of Spain of the HTCI 3190, karstic lakes on gypsum, for the period 2013\u0026ndash;2018 was assessed as unfavourable-inadequate, whereas for both the Atlantic and Alpine biogeographical regions, the conservation status was favourable (Table\u0026nbsp;\u003cspan refid=\"Tab8\" class=\"InternalRef\"\u003e8\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab8\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 8\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEvaluation of the conservation status of each of the four parameters included in the General Matrix, and global evaluation of the THIC 3190 for the period 2013\u0026ndash;2018. FV: favourable status; U1: unfavourable-inadequate status; U2: unfavourable-bad status; ALP: Alpine; ATL: Atlantic; MED: Mediterranean biogeographical region.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRANGE OF DISTRIBUTION\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAREA OF OCCUPANCY\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSTRUCTURE AND FUNCTION\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFUTURE PROSPECTS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eOVERALL CONSERVATION STATUS\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMED\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eU1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003eU1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eATL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003eFV\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eALP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003eFV\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eKarstic lakes on gypsum are a very peculiar type of lentic ecosystem, with very restrictive characteristics, included by the HD as a habitat type (3190). These ecosystems are strongly linked with the geological and hydrogeological context associated with gypsum areas (Alm\u0026eacute;cija \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Rodriguez-Rodriguez et al. 2006; Andreo et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), although their ecological definition is also related to their morphology, physical-chemical water features, and biotic processes and components, especially in particular microbial communities (Van Gemerden and Mas \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Camacho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2000b\u003c/span\u003e). However, the definition included in the interpretation manual for HTCI (DG Environment \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), does not reflect exactly the ecological features observed in Spanish karstic lakes on gypsum. Under natural conditions, the large level fluctuations indicated in the Habitat Interpretation Manual (DG Environment \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) are not so large in the Spanish lakes, being even infrequent in many cases, and highly dependent on the relations with the aquifer. In some cases, these lakes are the origin of more or less important streams or watercourses depending on the upwelling flow and the capacity of the aquifer that feeds them (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). In Spain as well, there are lakes with a maximum depth higher than 7 m. Otherwise, the importance of some abiotic features, characteristics of this HTCI, indicated in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, are good indicators for the identification of sites as 3190, even more than macrophyte communities that can be found in other habitat types (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe particularities of this type of habitat, which is located in specific conditions where the karstification process takes place on materials with a high content in gypsum, means that its distribution is not attributable to bioclimatic factors, rather to specific geological factors. The distribution and range showed how the sites were located in specific areas where the geological characteristics allowed the formation of these systems. Their assessment was determined by comparison with the range when the Directive came into force, defining a reference range. However, the number of 79 lakes identified as HTCI 3190 are still far from the current 5 sites officially classified and protected within the N2000 network for this habitat type in Spain, and just 19 in all the EU (European Environmental Agency \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), which shows that further declaration of Special Conservation Areas within the Natura 2000 Network is needed to enlarge the protection of this habitat type of community interest. Not only at the European level, but other protected figures do not include a large number of these ecosystems either. Of the 18 lakes identified as HTCI 3190 studied here wih detail as representative, only 7 are included in the Spanish Inventory of Wetlands (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Up to now, this inventory did not include some of the most important sites of the type, such as the Esta\u0026ntilde;a or Banyoles complexes, neither the Estany de Montcort\u0026eacute;s. These and other sites, such as the Arbieto, Atalaya or B\u0026aacute;rcena lakes, meet the requirements defined for their inclusion in this inventory, as well as having the peculiarity of being HTCI 3190. However, since the Spanish Inventory of Wetlands is a legal figure that requires the regional governments to submit to the central government the localities to be included, until each regional government sends this information, a lake/wetland cannot be included in the Inventory, thus is not officially recognized.\u003c/p\u003e \u003cp\u003eThe specific area occupied by the sites classified as 3190 was also calculated by aerial images, following a widespread method in ecology and conservation biology (Camacho et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2019b\u003c/span\u003e). This method allows the area to be assessed from the delineation manually, which can be quite accurate, especially with high quality images. However, it is costly when the number of sites to be surveyed is very large, so other methods that automate the identification and delineation of these surfaces may be more suitable, although the accuracy may be reduced (Do\u0026ntilde;a et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The status assessment related to the habitat distribution and coverage was also carried out on the basis of the differences to the moment when the Directive came into force. Looking at the historic aerial images, an encompassed balance between disappeared sites (due to clogging processes and changes in land use), and those newly appearing by genesis due to karstic collapse during the last decades (Supplementary Figs.\u0026nbsp;4 and 26), or restored after having disappeared, like Laguna de Santiago (Supplementary Fig.\u0026nbsp;15), seems evident. This encompassing of newly appearance and disappearance is one of the most outstanding dynamic features of this HTCI, as the time scale of genesis is assimilable to the time scale of natural and anthropogenic processes causing its disappearance (see e.g., Supplementary Figs.\u0026nbsp;4 and 26, how several sites were formed by karstification processes during the last decades). This is one of the reasons why, despite being a geographically restricted habitat found under specific geologic conditions, its conservation status referring to the range and area was assessed as favourable. Particularly for the Atlantic region, in the area occupied by the Laguna de Arbieto, a karstic lake on gypsum disappeared in the 1980s, as observed in aerial images. However, and following the criteria to determine the status, as this site had disappeared before the moment when the Habitats Directive came into force, there were no implications for the assessment of these two parameters. Therefore, methods other than the comparison to the 1994 range and surface to define favourable reference values should be considered, such as, for example, the species-area curve we proposed, a method that is also used for other ecological applications (Tj\u0026oslash;rve \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Ib\u0026aacute;\u0026ntilde;ez et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) which here helped to determine a minimum area capable of harbouring the maximum number of plant species characteristic of the habitat type.\u003c/p\u003e \u003cp\u003eWith respect to the assessment of the area of occupancy for the evaluation period 2013\u0026ndash;2018, differences among the surfaces (total, maximum flooded, current flooded, vegetation coverage, others) were also identified, though only the total area was used for the overall conservation status assessment. Results showed how most of the total surface corresponded to the flooded areas (70.6%), followed by 27% of the surface occupied by marginal emerged vegetation, and 2.4% the surface occupied by other uses. Looking at the differences along the period, a trend was found for the increase in the helophyte coverage and a reduction of open waters, especially in lower latitudes of the Spanish Mediterranean biogeographical region. This trend could be attributed to environmental changes, some of them related to climate change, such as reducing water supply thus reducing the water depth, and favouring the helophyte colonisation, especially that of \u003cem\u003ePhragmites australis\u003c/em\u003e, a highly invasive, though autochthonous, species in littoral shallow areas of lakes (Cirujano et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), as well as a possible acceleration the lakes siltation in the long term due to increased sediment transport in the agricultural areas surrounding many of these sites. However, the large inter- and intra-annual variability associated with hydro-climatic characteristics in the Mediterranean area, as well as other management actions that may influence the hydrological dynamics of wetlands, should be taken into account. Trends should be framed only on the period they were assessed, as these short-term trends may vary. Therefore, the assessment of the area parameter is only based on the total surface of the wetland, and not on the coverages of its components. It would be necessary to continue this evaluation in future periods to see if these trends continue, which would indicate a change in the structure of the habitat type, at least in shallow systems.\u003c/p\u003e \u003cp\u003eThe \u0026lsquo;Structure and function\u0026rsquo; parameter was assessed by the multimetric ECLECTIC index, obtained from a quantification and weighting of different parameters. Apart from the importance of the Chlorophyll-\u003cem\u003ea\u003c/em\u003e concentration (Poikane et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), other parameters, such as the biological community composition and coverage of submerged and emerged macrophytes, are relevant in the biological part of this index, given the role of plants in the structure and functioning of the habitats (Camacho et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Hydro-geomorphological factors and physical-chemical parameters are also included in the index, given the importance of these components in the lentic ecosystems condition (Verhoeven et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Poikane et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The evaluation of the parameter 'Structure and function' by the ECLECTIC index is carried out based on a sum of weighted values of different metrics, whose final value defines the parameter \u0026ldquo;structure and function\u0026ldquo; of the conservation status based on the global score obtained.\u003c/p\u003e \u003cp\u003eLooking at the results of the ECLECTIC index, and their assimilations to status classes, lakes with impacts that directly or indirectly affect their ecological integrity showed lower index value, and unfavourable status. Estany de Banyoles, with human pressure on its banks and catchment basin, Laguna de los Capellanes located within a cattle farm with a progressive modification of its hydroperiod and morphology, and Laguna de Alboraj as well as Laguna de Z\u0026oacute;\u0026ntilde;ar, with intensive agriculture activity in the surroundings, were assessed as unfavourable-inadequate for the \u0026lsquo;Structure and function\u0026rsquo; parameter. These sites showed, in general, relatively high concentrations of total phosphorus in water and high values of Chlorophyll-\u003cem\u003ea\u003c/em\u003e. In Laguna de Z\u0026oacute;\u0026ntilde;ar, the loose of typical species, both plants (submerged macrophytes and helophytes) as well as zooplankton and benthic invertebrates, contributed to the reduction of the value of the conservation status (Junta de Andaluc\u0026iacute;a \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Contrarily, sites such as Laguna de Arbieto, in the Atlantic region, or Estany de Montcort\u0026eacute;s, in the Alpine region, showed a favourable status for the structure and function parameter. In both cases, the evaluated variables identified the ecosystem as displaying a good ecological integrity, due to their biological diversity, the maintenance of their natural hydromorphological functioning, and the values of their physical-chemical parameters within the appropriate range for the HTCI.\u003c/p\u003e \u003cp\u003eThe multimetric ECLECTIC index combines the evaluation and application of different metrics referring to the specific elements of lentic ecosystems and their functioning. Some of the main assessing variables included in the index, such as the Chlorophyll-\u003cem\u003ea\u003c/em\u003e concentration, and the nutrients concentrations, might make vary the final value (from favourable to unfavourable) due to their importance in the weighting, especially when not all the variables, both compulsory and optional, are assessed. Some of these variables are also used as the metrics for the assessment of the ecological status according to the WFD (Poikane et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Poikane et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Both approaches, the ECLECTIC method, and the different metrics according to the WFD, coincide in the determination of some common biological, physical-chemical and hydromorphological features, whose separate values integrated in the multimetric ECLECTIC index, allow the global evaluation of the status of ecological health of these lentic ecosystems. However, the ECLECTIC index differs from those used by the WFD, in which the lowest value of those obtained in the evaluations of biological quality elements, physical-chemical and hydro-morphological factors, is the one that gives the overall status value (one out, all out). Therefore, the proposed method could overestimate the final status value compared to other multimetric methods (European Commission \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). The two legal frameworks for the protection of water (WFD) and nature (HD) converge when evaluating and achieving similar objectives (European Commission \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). However, WFD and HD are currently implemented separately, which can make more difficult to achieve their respective goals (Stefanidis et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe \u0026lsquo;Future prospects\u0026rsquo; parameter was obtained from the assessment of pressures and threats that were previously demonstrated to affect the ecosystem condition, such as hydrology and geomorphology (Jusik and Macioł \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Evtimova and Donohue \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Poikane et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), water quality (Verhoeven et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), land uses (Nielsen et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Morant et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and exotic species (Reid et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). There was a notable difference in the level of pressure exerted in each of the sites selected as representative of the THIC 3190. Up to 10 from 18 assessed sites showed low levels of pressure, which assimilates to favourable future prospects (e.g., in Laguna de Arbieto, Estanque Grande de Esta\u0026ntilde;a, and Estany de Montcort\u0026eacute;s). The levels of pressure exerted, both directly on hydrology and on their communities, as well as indirect pressures, were low and did not significantly affect the structure and ecological function of ecosystems, as the same favourable status was generally obtained for the \u0026lsquo;Structure and function\u0026rsquo; parameter. In the same way, those sites with moderate pressures, and an unfavourable-inadequate status in the 'Future prospects' parameter, showed an unfavourable-inadequate status in the 'Structure and function' parameter, since the pressures exerted on the components of aquatic systems resulted on damages on their structure and functioning (Camacho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). This was the case of Laguna de la Atalaya, Laguna de los Capellanes, Laguna de Z\u0026oacute;\u0026ntilde;ar, and Lagunillo de las Tortugas, where management actions are needed to reduce or eliminate these sources of pressure affecting their structure and ecological functioning, thus, their conservation status. This multimetric method, like the ECLECTIC index, could overestimate the calculated status, due to the integration of a set of variables, in which only one could reduce the final value of the status in a lake that apparently could be defined as favourable. Additionally to the method used here, alternative methodologies have been proposed for the assessment of pressures and threats have been proposed that can be spatially explicit, thus approaching the localisation of the areas for actuation, for instance, the use the land uses in the catchment area as a proxy to estimate the pressure level over lentic ecosystems by the LUPLES method (Morant et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), which demonstrated to be correlated with ecological indicators of impacts.\u003c/p\u003e \u003cp\u003eAt global level, the conservation status assessed within the four parameters. for the HTCI 3190 in Spain showed its unfavourable-inadequate status in the Mediterranean region, reflected by the moderate pressures and impacts and their response in the structure and functioning of some of the sites included in the assessment. In this region, the area of occupancy was maintained over the decades, because of an equilibrium between new formed sites that counterbalanced the disappeared lakes. Meanwhile, the status was favourable in the Atlantic and Alpine regions, though this was determined with a low sample size, with only two and one karstic lakes on gypsum assessed, respectively. As indicated above, this HTCI is highly specific and quite particular, thus, the low number of sites classified as HTCI 3190, and the low area occupied was not equivalent to a poor status, as the sites remain over the decades. Thus, by applying the different methods defined in this article, and following the criteria for the definition of the conservation status, it seems the results mostly reflect the real condition of the HTCI in Spain, considering the particularities of the ecosystem type.\u003c/p\u003e"},{"header":"5. Conclusions","content":"\u003cp\u003eFor the ecological characterisation of the lakes, main parameters identifying karstic lakes on gypsum were physical-chemical features associated to the active gypsum karst areas where they are located, conferring particular hydro-morphometric characteristics, as well as the presence of certain types of submerged macrophytes species, though a revision should incorporate additional ecological features as those indicated in this study. Localised in certain areas with specific characteristics, these lakes cover an area of about 300 ha in Spain, being the main catalogued contribution to this habitat type (HTCI 3190) by Western European countries. Their global conservation status was favourable in the Spanish Alpine and Atlantic regions, and unfavourable-inadequate in the Spanish Mediterranean region. This assessment was based on range and area of distribution of the HTCI, plus the assessment and extrapolation of the \u0026lsquo;Structure and function\u0026rsquo; and \u0026lsquo;Future prospects\u0026rsquo; parameters with multimetric indices, which followed the criteria established by the HD.\u003c/p\u003e \u003cp\u003eFrom the assessment approach defined and applied, management actions of the HTCI should ensure the integrity and protection of the structure and functioning of the ecosystems in current and future scenarios, by minimising threats and maintaining their range and area throughout the biogeographic regions where they are located. However, additional efforts and detailed studies of these systems are needed to identify possible new locations that could be classified as HTCI 3190, especially small temporary ponds associated with geological environments rich in gypsum. In turn, a more detailed evaluation of their characteristics and condition is necessary for a more detailed overall assessment of their conservation status. Further, the methods here developed, as well as the specific results of the assessment, would help for the purposes of the Strategic Plan for Wetlands 2030, published by the Government of Spain in 2023 (MITECO, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), This plan aims to improve the conservation status and establish restoration targets for Spanish wetlands, which would be especially important for the conservation of this very scarce habitat type both at the Iberian and European level, as Spain plays a prominent role for the preservation of most of its sites in Western Europe.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting interests\u003c/h2\u003e \u003cp\u003e\u0026ldquo;The authors have no relevant financial or non-financial interests to disclose.\u0026rdquo;\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis work has been supported by the MITERD, through a TRAGSATEC contract n\u0026ordm;: 65393 to the University of Valencia, as well as by the project CLIMAWET-CONS (PID2019-104742RB-I00), granted to AC (University of Valencia) by the Spanish Research Agency (AEI) of the Spanish Ministry for Science and Innovation.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAll authors contributed to the study conception and design, as well as to material preparation, data collection and analysis. RH promoted the study on behalf of MITERD, and AC did the experimental design. The first draft of the manuscript was written by DM and AC-S, and the final version was elaborated by AC. All authors commented on previous versions of the manuscript. and read and approved the final manuscript\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eAuthors are grateful to the colleagues of the Spanish Ministry for the Ecological Transition and the Demographic Challenge (MITERD), TRAGSATEC, and the University of Valencia who collaborated in this work. This work has been supported by the MITERD, through a TRAGSATEC contract n\u0026ordm;: 65393 granted to the University of Valencia, as well as by the project CLIMAWET-CONS (PID2019-104742RB-I00), granted to AC (University of Valencia) by the Spanish Research Agency (AEI) of the Spanish Ministry for Science and Innovation.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAlfonso MT, Miracle MR (1987) Estudio comparativo del zooplancton de tres ullales del Parque Natural de la Albufera de Valencia. Limnetica 3:263-272\u003c/li\u003e\n\u003cli\u003eAlm\u0026eacute;cija C (1997) Hydrological and hydrochemical study of the wetland complexes at the north of the province of Malaga. 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Trends Ecol Evol 21:96\u0026ndash;103. https://doi.org/10.1016/j.tree.2005.11.015 \u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Supplementary Figures","content":"\u003cp\u003eSupplementary Figures are not available with this version.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"environmental-earth-sciences","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"enge","sideBox":"Learn more about [Environmental Earth Sciences](https://www.springer.com/journal/12665)","snPcode":"12665","submissionUrl":"https://submission.nature.com/new-submission/12665/3","title":"Environmental Earth Sciences","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"EU Habitats Directive, Habitat Types of Community Interest, Karstic lakes on gypsum, Conservation status, Assessment","lastPublishedDoi":"10.21203/rs.3.rs-4350343/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4350343/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eKarstic lakes on gypsum are a very peculiar type of ecosystem declared as a Habitat Type of Community Interest (Type 3190) by the European Habitats Directive. They are usually small lakes but often displaying a high relative depth, located in active gypsum karst areas, with a high saturation of Ca\u003csub\u003e2\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e and SO\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e2\u0026minus;\u003c/sup\u003e in its waters. These lakes can usually stratify from spring to early autumn when the depth is high enough, then a sulphide-rich anoxic hypolimnion can develop in deep layers. So far, neither a comprehensive scientific definition of their ecological characteristics nor an exhaustive catalogue of their occurrence in a particular territory are available. This paper delves on their biotic and abiotic features needed for their identification as an ecosystem type. A proper methodology was also designed and applied for the evaluation of their conservation status, with the definition of reference values and assessment methods following the criteria of the Habitats Directive. The Spanish karstic lakes on gypsum (THCI 3190) were here identified, statistically representative sites were selected, and spatial GIS methods and multimetric indices were applied to assess the range, area, structure and function, and future prospects of this Habitat Type, as requested for the reporting according to Article 17 of the Habitats Directive. Results showed a favourable conservation status of this habitat type in the Alpine and Atlantic regions of Spain, but unfavourable-inadequate status in the Mediterranean, due to the pressures and impacts acting on some specific sites. A critical analysis of the methodologies and the values obtained for its conservation status was carried out.\u003c/p\u003e","manuscriptTitle":"Transdisciplinary approach to the characterisation and current status of Spanish karstic lakes on gypsum","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-05-08 15:30:10","doi":"10.21203/rs.3.rs-4350343/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Accepted","date":"2024-06-06T07:53:15+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-06-03T10:42:05+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"314067643454583229339229358871734273226","date":"2024-05-21T18:21:55+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-05-21T13:55:56+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"58823210796850310412891418697205710088","date":"2024-05-21T13:41:45+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-05-21T12:02:37+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-05-02T08:06:47+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-05-02T08:06:47+00:00","index":"","fulltext":""},{"type":"submitted","content":"Environmental Earth Sciences","date":"2024-04-30T16:04:59+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"environmental-earth-sciences","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"enge","sideBox":"Learn more about [Environmental Earth Sciences](https://www.springer.com/journal/12665)","snPcode":"12665","submissionUrl":"https://submission.nature.com/new-submission/12665/3","title":"Environmental Earth Sciences","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"0aee34cf-c44f-45d5-8ec6-29935383f9c9","owner":[],"postedDate":"May 8th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-08-05T16:08:47+00:00","versionOfRecord":{"articleIdentity":"rs-4350343","link":"https://doi.org/10.1007/s12665-024-11700-4","journal":{"identity":"environmental-earth-sciences","isVorOnly":false,"title":"Environmental Earth Sciences"},"publishedOn":"2024-08-02 15:57:30","publishedOnDateReadable":"August 2nd, 2024"},"versionCreatedAt":"2024-05-08 15:30:10","video":"","vorDoi":"10.1007/s12665-024-11700-4","vorDoiUrl":"https://doi.org/10.1007/s12665-024-11700-4","workflowStages":[]},"version":"v1","identity":"rs-4350343","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4350343","identity":"rs-4350343","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}