Cave-adapted beetles from continental Portugal

Abstract Background The cave biodiversity of continental Portugal faces tremendous conservation challenges, mostly linked to their direct destruction and contamination infiltrating from the surface. Beetles are the most diverse insects and one of the most diverse arthropod groups in caves of Portugal. New information We present the IUCN Red List profiles for the cave-adapted beetles from continental Portugal, all endemic to their respective geological units and massifs. Ground beetles (Carabidae) are the most diverse family of cave-adapted beetles in continental Portugal, followed by rove beetles (Staphylinidae). Beetles in caves of Portugal are mostly terrestrial and only one species is known to have evolved to live in groundwater. Trechus is the most diverse genus with four species, followed by Domene with three species and by Speonemadus and Iberoporus, both with one cave-adapted species. The aim of this contribution is to assess all endemic cave-adapted species of beetles from continental Portugal and to support their specific protection, to promote adequate management of surface habitats and the establishment of priority areas for conservation. The main biodiversity erosion drivers that are impacting the conservation of the studied species are pollution infiltrating from the surface, urbaniation, modifications of the natural habitat for touristic purposes and mining, quarrying and energy production infrastructures. This document can be used in spatial planning and territory management in karst, based on the current scientific knowledge.


Introduction
Cave fauna has relatively low specific richness, but high conservation value for the global biodiversity of our planet (Mammola et al. 2019). Subterranean species have unique traits that emerged as a result of isolation and convergent adaptation towards the underground life (Mammola et al. 2020).
Amongst subterranean fauna, the beetles (Insecta, Coleoptera) stand out, as a group that presents the greatest animal specific richness worldwide (Didham et al. 2020). Of the 211 recognised families of Coleoptera (Bouchard et al. 2011), around 10 have cave-adapted species, of which about 92% belong to the families Carabidae (mostly Trechinae) and Leiodidae (mostly Cholevinae) (Deharveng and Bedos 2018).
Portugal is located in the western part of the Iberian Peninsula, has more than 2000 caves identified and is considered a hotspot of subterranean biodiversity (Reboleira et al. 2011b). More than 58 terrestrial and 64 aquatic subterranean-obligate species are described to date (Reboleira et al. 2013a, Reboleira and Enghoff 2018, Ribera and Reboleira 2019. Beetles are the third most diverse order of terrestrial cave-adapted species in continental Portugal, preceeded by pseudoscorpions (13 species) and terrestrial isopods (12 species) (Reboleira et al. 2015, Zaragoza and).
The first beetle species, collected in caves of Portugal, was identified as Trechus fulvus Dejean, 1831, captured by M.L.W. Schaufuss; the caves then explored remain unknown (Putzeys 1870). The first cave-adapted beetle of Portugal, Trechus machadoi Jeannel, 1941, was found more than a century later and described, based on two specimens collected by the pioneer of Portuguese cave biology, António de Barros Machado (Jeannel 1941). Since this time, almost seven decades have passed until the description of further cave-adapted beetles in continental Portugal, also belonging to the genus Trechus Clairville, 1806 (Reboleira et al. 2010). The first Staphylinidae beetle was described in 2010 from a karst cave, belonging to the genus Domene Fauvel, 1873 (Reboleira et al. 2011a), followed more recently by the description of two more species of the same genus from non-karst caves (Serrano et al. 2015, Magrini andCarotti 2019). Despite the high

Basis of EOO and AOO: Taxonomic notes
Iberoporus pluto is the largest and widest of all the subterranean species of its genus (Ribera and Reboleira 2019).

Region for assessment:
-Europe  Yes

Habitat (narrative)
The specimen was collected in the bottom of a clay pool connected to a subterranean stream (Ribera and Reboleira 2019).

Justification for trend
Soprador do Carvalho Cave is located in the vicinity of a village, agricultural fields and a quarry (NEC 2020, Neves et al. 2005. It is also affected by touristic activities as tourists visit the cave and step on the stream (Ribera and Reboleira 2019).

Justification for threats
The cave is explored for tourism and visitors step over the habitat where the species was found (Ribera and Reboleira 2019). Other threats include the proximity to a quarry located at 1.4 km from the cave entrance. It is also 67 m from the closest houses and is surrounded by agricultural lands. The subterranean stream flows below habitational areas and run-off of urban wastewater directly to the stream is observable (Reboleira et al. 2011b).

Justification for conservation actions
Measures should be taken to prevent infiltration of wastewaters from the village into the cave stream. The nearby quarry has been reported in the national media to be the source of the infiltration of small particles of quarry dust that have been deposited all over the gallery of Algarinho Cave by flood events. This type of slurry is known to perniciously impact groundwater quality (Piccini et al. 2019) and Algarinho Cave is part of the Dueça Speleological System, therefore, hydrologically connected to Soprador do Carvalho Cave.

Justification for research needed
Further investigation is needed about the distribution, ecology and life cycle of the species. Developing a management plan for this species is crucial. This plan will aid the conservation of the cave-adapted species of the Sicó karst area.

Taxonomic notes
This species belongs to the T. fulvus-group of species.

Region for assessment:
-Europe

Geographic range
Biogeographic realm:

Basis (narrative)
The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both 4 km .

Range description
Trechus machadoi is a troglobiont beetle known only in the Alcobertas Cave and in a countinuous mesovoid shallow substratum (MSS) located in the Serra dos Candeeiros subunit of the Estremenho karst massif, central Portugal. The cave extends horizontally for approximately 210 m (Reboleira 2007, Reboleira 2012, Reboleira et al. 2009) and the MSS area is located 80 m from the type locality (Eusébio 2020).

Extent of occurrence
4 Decline (observed)

Justification for trend
Despite intensive sampling in the type locality (cave), no specimens have been found in there since the species description. Only recent sampling in the MSS contiguous to the cave retrieved specimens.

Justification for number of locations
Trechus machadoi is known only from the Alcobertas Cave and from the adjacent MSS, located approximately 80 m from each other. Its distribution is likely to be confined to the subterranean habitats of the Serra dos Candeeiros subunit (Reboleira et al. 2009). This population is under intense disturbance.

Justification for trend
Alcobertas is the only known location in the historical range, therefore, the current trend in number of locations is stable. Habitat specialist: Trend in extent, area or quality?:

Justification for trend
The type locality was monitored for six months in 2009, but no specimens were sampled or observed, most likely due to the human impact on this cave described below (Reboleira 2007).

No
Yes Unknown Unknown

Population Information (Narrative)
So far, only one population is known from the Alcobertas Cave, which is also dispersed in the contiguous mesovoid shallow substratum at 0.5 m depth in scree slopes (Reboleira 2007, Reboleira et al. 2009, Reboleira et al. 2010.

Terrestrial
Yes

Habitat (narrative)
The Alcobertas Cave was subject of a large anthropogenic intervention at the beginning of the 1970s, with the intention to transform it to receive mass tourism. During that process, a second entry was opened near the end of the gallery which induced significant changes in the climatology of the cave (Reboleira 2007). Many explosions inside the cave have resulted in large accumulations of powdery residues that can still be observed nowadays, which, associated with the air fluxes between the two entrances, may have displaced the Trechus machadoi population to other parts of the subterranean network (Reboleira et al. 2009). The Alcobertas Cave is currently used for tourism by a local association. Recently, Trechus machadoi was collected during winter at a depth of 50 cm in the mesovoid shallow substratum (MSS) in scree slopes close to the type locality (Eusébio 2020, Eusébio et al. 2021. These scree slopes are classified as western Mediterranean and thermophile deposits and protected by the Natura 2000 network (Eusébio et al. 2021).

Ecology and traits (narrative)
Trechus machadoi is a troglobiont with reduced eyes and body depigmentation. It lives exclusively in subterranean habitats and is only known from a single cave and from scree slopes habitats in the Serra dos Candeeiros subunit of the Estremenho karst massif in central Portugal (Eusébio 2020, Reboleira 2007, Reboleira et al. 2009).

Justification for threats
Since the 1970s, this cave has been intensively explored for touristic activities. During that period, a second entrance has been opened, drastically changing the environment (Reboleira 2007, Reboleira et al. 2009). The scree slope, where it was recently found, is disturbed by herds of grazing goats (Eusébio 2020, Eusébio et al. 2021). Both sites are located 640 m from a field of energy windmills, 1 km from a quarry, 850 m from agricultural lands and 690 m from the nearest village.

Conservation actions:
Use type: Ecosystem service type:

Justification for conservation actions
The habitats are protected under the EU "Rede Natura 2000" (Directive 1992, ICNB 2000, but the species lacks a formal protection framework. Monitoring of the population trends and extent of distribution in the underground in the nearby areas is crucial to understand the species' abundance patterns and life cycle. Improvements are needed to reduce the impacts of the quarry and windmills on the habitat.  Map of records (Google Earth):

Taxonomic notes
This species belongs to the "T. fulvus-group" species complex.

Region for assessment:
-Europe

Geographic range
Biogeographic realm:

Basis (narrative)
The extent of occurrence (EOO) is 73.4 km² and the maximum estimated area of occupancy (AOO) is 24 km².

Population
Unknown Unknown Unknown Unknown Unknown Unknown

Population Information (Narrative)
Amongst the six known localities with populations of T. gamae, Algar das Gralhas VII Cave had the largest population, followed by Algar do Pena Cave, Algar das Marradinhas Cave and Algar da Arroteia Cave (Reboleira 2007, Reboleira andOrtuño 2011). Algar do Ladoeiro Cave and the mesovoid shallow substrate in scree slopes in Fórnea (contiguous to Algar da Arroteia Cave) had the smallest population (Reboleira et al. 2009;Eusébio et al. 2021). All the known populations are within the Santo António Plateau.

Justification for trend
All the subpopulations face threats derived from intensive quarrying activity, which changes land use and disturbs the natural processes of the habitat. The subpopulations from Algar do Ladoeiro, Algar das Marradinhas II, Algar da Arroteia and Fórnea face threats of pollution and land use disturbance due to the proximity of urbanised areas.

System:
Habitat specialist: Trend in extent, area or quality?: Habitat importance:

Generation length (yr):
Dependency of single sp?: Threat type:

Habitat
Terrestrial Yes

Habitat (narrative)
Trechus gamae was found in the deepest parts of the caves, from 50 to 95 m depth, all with high humidity levels (> 98%) and temperatures ranging from 13.5ºC (in Algar do Pena) to 17ºC (in Algar de Marradinhas II) (Reboleira 2007, Reboleira 2012, Reboleira and Ortuño 2011, Reboleira et al. 2009). The caves where the species were found are located at an altitude ranging from 250 to 485 m a.s.l. (Reboleira and Ortuño 2011). More recently, a single specimen was collected in MSS traps in scree slopes in Fórnea, which is located between the Algar da Arroteia and the Algar do Ladoeiro Caves. The MSS specimen was collected in the winter, in a pitfall trap placed at 50 cm deep into the scree slope (Eusébio et al. 2021).

Ecology and traits (narrative)
Trechus gamae was the only cave-adapted beetle collected in the caves and MSS of the Santo António Plateau and it shows a strict subterranean lifestyle (Eusébio et al. 2021, Reboleira 2007, Reboleira 2012, Reboleira and Ortuño 2011, Reboleira et al. 2009, Reboleira et al. 2010. Both adults and larvae were found in the sampled caves and some seasonal abundance variation was recorded, being more abundant in spring and summer (Reboleira and Ortuño 2011).

Threats
Ongoing Threats: Conservation action type:

Justification for conservation actions
Although the habitat is protected by law under the "Rede Natura 2000" (Directive 1992, ICNB 2000), the species is not. Population trends need to be monitored in order to better understand the species' abundance patterns and life cycle. Measures should be taken to prevent infiltration of wastewaters from the nearby town into the soil and underground habitats and to minimise the negative effects of the quarry in the habitat.

Other
International Very important

Taxonomic notes
This species belongs to the "T. fulvus-group" species complex.

Region for assessment:
-Europe Suppl. materials 1, 5 Known habitat extent

Basis (narrative)
The extent of occurrence (EOO) is 4 km and the maximum estimated area of occupancy (AOO) is 12 km².

Justification for number of locations
Trechus lunai was found in three caves of the Serra de Aire/São Mamede Plateau subunits. The southernmost distribution is the Almonda Cave, but this species is also known from the Contenda and Moinhos Velhos Caves' system (Reboleira et al. 2009).

Justification for trend
The subpopulation in Almonda cave is subject to wastewater and pollution infiltration and the subpopulations of the Contenda and Moinhos Velhos Caves face heavy contamination derived from the village under which they are located.

Terrestrial
Yes

Habitat (narrative)
Trechus lunai only occurs in the deepest parts of the caves, from 50 to 80 m depth. The three caves have high humidity levels and average temperature of 18ºC (Reboleira et al. 2009. In Moinhos Velhos Cave, the species was found in a subterranean stream that has high input of sewage from surface and is located below the village of Mira d'Aire in the connection between the Gruta de Mira d'Aire Show Cave and the Pena Spring. Moinhos Velhos is contiguous to Contenda Cave and are hydrologically connected (Reboleira et al. 2009). The other locality is Almonda Cave, which is located 8.4 km in a straight line from the Moinhos Velhos-Contenda cave system. Almonda Cave, also known as "Olho do Moinho da Fonte", is the largest cave of Portugal with more than 10 km of horizontal development mapped (Thomas 1991).

Ecology and traits (narrative)
Trechus lunai was the only troglobiont species captured in these caves. All known localities are caves that flood seasonally (Reboleira et al. 2009).

Taxonomic notes
This species belongs to the "T. fulvus-group" species complex. It is recognisable by the shape of the aedeagus and has a slim body, rudimentary wings, reduced eyes (microphthalmia) and depigmentation. This species is the most troglomorphic groundbeetle known from Portugal (Reboleira et al. 2010).

Region for assessment:
-Europe

Fig. 5
Map of records (Google Earth):

Basis of EOO and AOO:
Min Elevation/Depth (m):

Geographic range
Biogeographic realm:

Basis (narrative)
The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both of 4 km².

Range description
Trechus tatai is a cave-adapted hygrophilous Carabidae known only from one small cave, located in Serra do Montejunto (Reboleira et al. 2010).

Justification for number of locations
Trechus tatai only occurs in one cave, Algar do Javali, located in Serra do Montejunto. This species is geographically isolated (Reboleira et al. 2010).

Justification for trend
Algar do Javali is the only known location for this species, therefore, the trend in number of locations is stable.

Population Information (Narrative)
Only one population is known from Algar do Javali Cave, in Montejunto karst massif.

Habitat
Terrestrial Yes

Habitat (narrative)
Trechus tatai was only collected in the deep oligotrophic areas of the cave. It was never found in areas with high organic material content (bat guano accumulation zones). The cave is 10 m deep and extends for 80 m. Temperatures in the deepest zone of the cave ranged from 14.2 ºC in winter to 15 ºC in summer (Reboleira et al. 2010).

Ecology and traits (narrative)
Trechus tatai is the most troglomorphic carabid beetle from continental Portugal. Other caves of the area were sampled, but T. tatai was never collected elsewhere. The seasonal activity pattern of the beetle was studied during one year in the deepest zone of the cave and specimens were collected during winter, autumn and spring (Reboleira et al. 2010). The absence of T. tatai during summer is most likely caused by the low humidity in the cave during this season. It is very likely that, during summer, this species escapes from the cave into the MSS in search of smaller gaps that retain humidity (Reboleira et al. 2010).

Justification for threats
Algar do Javali Cave is located 1.6 km from a quarry with intensive extraction activity and 2.9 km from the closest village, which induces deep changes in land use at the surface and potential biotic exchange, such as introduction of invasive alien species. The cave entrance is located 50 m from a road and surrounded by Eucalyptus intensive plantation, with direct impact on land use at the surface, pollution and groundwater depletion.

Justification for conservation actions
Although this cave is protected by law through the "Rede Natura 2000" (Directive 1992, ICNB 2000), the species is not. This species is rare, a single cave endemic and it is considered the most troglomorphic carabid beetle of Portugal; therefore, a conservation plan for this cave area is crucial to ensure its environmental sustainability.

Taxonomic notes
This species is part of the "Speonemadus escalerai-group" and is recognisable by the equal/subequal length of the 2 , 4 , 5 and 7 antennomeres and a slightly transverse and hexagonal pronotum (Reboleira et al. 2017).

Region for assessment:
-Europe

Geographic range
Biogeographic realm: There are three populations known from Portugal, all from caves in the Algarve karst massif. The largest number of individuals was collected in Vale Telheiro Cave, followed by Senhora Cave and Algarão do Remexido Cave (Reboleira et al. 2017).

Justification for trend
All populations are under risk due to wastewater infiltration derived from the urbanised areas in the region. The subpopulation of Algarão do Remexido cave is threatened by agricultural pollution infiltration and the subpopulation from Senhora cave is threatened by industrial residue pollution.

Ecology and traits (narrative)
Speonemadus algarvensis occurs exclusively in the Algarve and does not exhibit the typical troglomorphism found in other cave-adapted species, such as evident eye reduction, severe depigmentation and extreme body and appendages elongation, although it has only been collected in caves and never at the surface. This species was found to carry the ectoparasitic fungus of the order Laboulbeniales (Stichomyces conosomatis Thaxt., 1901) attached to the cuticle and foresic acari (Reboleira et al. 2017).

Threats
Ongoing Threats: Conservation action type:

Justification for threats
Algarão do Remexido is located under agricultural lands, 370 m from the closest house and 1.7 km from the closest village. Vale Telheiro is located 290 m from the closest house and 745 m from the closest urbanisation. Senhora Cave is located 168 m from the closest house and 900 m from an industrial complex.

Justification for conservation actions
Although the habitat is protected under legislation by the "Rede Natura 2000" (Directive 1992, ICNB 2000, the species is not. Population trends need to be monitored in order to better understand the species' abundance patterns and life cycle. Measures should be taken to prevent infiltration of wastewaters and to ensure conservation of the natural landscape and plant communities on the surface, which are necessary to maintain nutrient inflow to the subterranean ecosystem.

Justification for research needed
The development of a management plan for the conservation of this cave-adapted species in the Algarve karst massif and the encouragement of more studies regarding population size, extent of distribution, ecology and life cycle are essential measures for the protection of the species.

Taxonomic notes
Individuals display troglomorphism, such as microphthalmia, lack of wings and body elongation (Reboleira et al. 2011a).

Region for assessment:
-Europe  Known habitat extent

Basis (narrative)
The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both of 4 km².

Range description
Domene lusitanica was found in a single cave located in the Sicó karstic massif (Reboleira et al. 2011a). Threat type:

Extent of occurrence
Threats:

Habitat (narrative)
Specimens were exclusively collected in the deepest zones of the cave (10 m deep), in high humidity levels and with average temperatures of 16.4ºC (Reboleira et al. 2011a).

Ecology and traits (narrative)
Domene lusitanica is included in the subgenus Lathromene, together with the other two Portuguese species of cave-adapted Domene: D. viriatoi and D. darinkae. This species is a predator troglobiont rove beetle, with reduced eyes, apterous, depigmented and elongated body and appendages (Reboleira et al. 2011a). It is only known from one cave in the Sicó karst area and is a rare species, as the type locality and several other caves nearby have been monitored for more than a decade and only eight specimens have been observed so far: seven collected in 2010 (type material) and one collected in December 2019.  Reboleira & Taiti, 2015and Porcellio cavernicolus Vandel, 1946, Reboleira et al. 2013c.

Justification for conservation actions
The habitat is located in an "Rede Natura 2000" area (Directive 1992, ICNB 2000.
Population trends need to be monitored in order to better understand the species' abundance patterns and life cycle. Measures should be taken to prevent infiltrations from agricultural lands and livestock farms and to prevent the pernicious effects of the quarry activity on the surrounding habitats.

Justification for research needed
In order to build a sustainable conservation plan for the species in the Sicó karst area, more information about population size, extent of distribution, ecology and life cycle is needed. The threats also need to be addressed and minimised, if possible, in order to improve the habitat quality.

Taxonomic notes
This species displays body, leg and antennae elongation, microphthalmia and lack of wings (Serrano et al. 2015).

Region for assessment:
-Europe

Geographic range
Biogeographic realm:

Basis (narrative)
The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both 4 km .

Range description
Domene viriatoi was collected in two galleries of the Buraco da Moura Cave, located in the Serra da Estrela Mountain foothills (Serrano et al. 2015).

Ecology and traits (narrative)
Domene viriatoi is included in the subgenus Lathromene. Both adults and larvae of this species were observed foraging for preys in bat guano on the cave substrate (Serrano et al. 2015

Justification for threats
The cave entrance is located 127 m from the closest house, 530 m from a hydroelectric power station and 1.2 km from the closest village and is under anthropogenic disturbance due to tourism.

Conservation
In Place -

Justification for conservation actions
Buraco da Moura Cave was classified as a "National Important Underground Shelter for Bats" therefore a decrease in human disturbance is expected (Serrano et al. 2015). However, specific monitoring plans for this species are required in order to understand its ecology and distribution.
Although the habitat is protected under legislation by the "Rede Natura 2000" (Directive 1992, ICNB 2000, the species is not. Population trends need to be monitored in order to better understand the species' abundance patterns and life cycle. Measures should be taken to prevent infiltration of wastewaters from the nearby town into the soil and underground habitats and to minimise the effects of the hydroelectric power station on the surrounding habitats. Trend:

Basis (narrative)
The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both 4 km .

Range description
Domene darinkae is a cave-adapted rove beetle known from an abandoned mine in northern Portugal (Magrini and Carotti 2019). Only one specimen of this species is known from a single location in northern Portugal (Magrini and Carotti 2019).

Unknown
Unknown Unknown

Terrestrial
Yes

Habitat (narrative)
The only known specimen was collected in the rocky debris along the main tunnel of the Santa Isabel mine (Magrini and Carotti 2019). The geological matrix of this mine is quartzite with iron.

Justification for research needed
Further information about population size, extent of distribution, ecology and life cycle is needed to better protect the species and the habitat. The mine, where the species was found, is an anthropogenic construction that clearly adversely affected the natural habitat of the species that should be the deep fissures and the mesovoid shallow substrate of the area. Therefore, it is recommended to sample these habitats in the area to understand the distribution of this species and to define new conservation priorities.

Discussion
The year 2021 is the International Year of Caves and Karst (http://iyck2021.org), an event organised by the International Union of Speleology to promote the awareness for the importance of caves and their habitats. Under this framework, a global initiative created the International Cave Animal of the Year (http://iyck2021.org/index.php/cave-animal-of-theyear) devoted to cave beetles. Within this initiative, different countries selected their own endemic species as a flag for advocating the conservation of subterranean ecosystems.
Here, we offer information about the distribution (Suppl. material 1), habitat, species ecology, current threats and conservation measures for the nine cave-adapted beetles of continental Portugal. This information is essential to raise the awareness about the threats faced by subterranean ecosystems and to establish conservation measures needed specifically for each country.
Beetles are the most diverse insects in Portuguese caves (Reboleira et al. 2011a, Reboleira et al. 2017. Cave-adapted beetles from continental Portugal are highly endemic with a very reduced extent of occurrence (EOO) and area of occupancy (AOO). This occurs because they are all endemic from their correspondent karst massif unit or collected in artificial mines in granite or quartzite rock. Four of the known species are endemic to single caves, while the other five species can be found in more than one cave. The use of molecular methods may shed further light on the evolutionary relationships and species delimitation, especially for the genus Trechus, which shows a radiation in contiguous areas of karst massifis in central Portugal (Reboleira et al. 2010).
Most of the cave-adapted species of beetles are extremely rare, appearing only once or twice per decade of constant sampling. For example, Iberoporus pluto, the only groundwater-adapted (stygobiont) beetle from Portugal, was described, based on a female specimen and no further specimens have been found in a cave that has been constantly monitored for more than a decade. Its habitat, the Soprador do Carvalho Cave is under serious anthropogenic threats, such as groundwater contamination and touristic pressure (Ribera and Reboleira 2019).
Some of the major threats that cave-adapted species face are habitat destruction due to the intensive quarrying activity that occurs near the majority of the localities and the severe groundwater contamination caused by the lack of proper wastewater treatment in most villages of the central Portugal karst massifs. Some of these threats have been identified previously by Reboleira et al. 2011b, but no specific conservation measures have been taken hitherto. The infiltration of contaminants and fertilisers originating from agricultural practices and industry on the surface, also pose major threats to the integrity of subterranean organisms (Castaño-Sánchez et al. 2020). Therefore, it is urgent to generate ecotoxicological data on the sensitivity of cave-adapted beetles to understand if the pernicious impacts that have been already evaluated for some endemic groundwateradapted species in Portugal (Reboleira et al. 2013b) are also impacting the terrestrial cave species.
In order to create protection strategies for cave-adapted species in continental Portugal, it is necessary to improve our knowledge about their population size, extent of distribution, ecology and life cycle. We hope this contribution may help to support decision-making on territory planning and to establish conservation measures for these highly endemic species. These will act as umbrella species for the conservation of other cave-adapted species that share the same subterranean habitats.