Biodiversity Data Journal :
Species Conservation Profiles
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Corresponding author: Ana Sofia P. S. Reboleira (sreboleira@snm.ku.dk)
Academic editor: Paulo Borges
Received: 16 Apr 2021 | Accepted: 01 Jul 2021 | Published: 20 Aug 2021
© 2021 Ana Sofia Reboleira, Rita Eusébio
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Reboleira ASPS, Eusébio RP (2021) Cave-adapted beetles from continental Portugal. Biodiversity Data Journal 9: e67426. https://doi.org/10.3897/BDJ.9.e67426
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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.
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.
Coleoptera, subterranean habitats, Iberian Peninsula, conservation, rocky habitats, troglobiont, stygobiont
Cave fauna has relatively low specific richness, but high conservation value for the global biodiversity of our planet (
Amongst subterranean fauna, the beetles (Insecta, Coleoptera) stand out, as a group that presents the greatest animal specific richness worldwide (
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 (
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 (
All cave-adapted beetle species from continental Portugal are endemic and exhibit an extremely reduced distribution area. These unique species face major conservation threats and lack of adequate specific management (
Caves of continental Portugal have been intensively sampled over the last 15 years. Sampling was performed by direct searching and baited pitfall traps, both in caves and in the mesovoid shallow substratum. Caves consist of underground spaces where a human can fit, while the mesovoid shallow substratum (MSS) consists of a matrix of unaggregated rock that can be found in scree slopes. Most of this sampling was performed under the framework of Master and Doctoral studies (
Extent of occurence (EOO) and area of occupancy (AOO) were calculated using the Geospatial Conservation Assessment Tool (GeoCAT) with an approximation to the standard IUCN 2 km × 2 km cells (4 km2) and the maps were created in the open source software QGIS 3.14.16, with the natural protected areas of Portugal layer (
Threats were observed in situ in the field and complemented by appropriate literature surveys. Threats, conservation actions, research needed and the type of habitat classification were assigned, based on the IUCN Red List database.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Dytiscidae |
Iberoporus pluto is the largest and widest of all the subterranean species of its genus (
Suppl. materials
The extent of occurrence (EOO) and area of occupancy (AOO) are both 4 km².
Iberoporus pluto is a groundwater-adapted beetle known from a stream in a single cave, located in north-eastern Sicó karst area. The cave stream, where it was found, flows in a subterranean system of approximately 15 km of horizontal implementation (
A decline in EOO is inferred due to the degradation of the cave by anthropogenic impact as Soprador do Carvalho Cave is subject to recreational visitation and direct trampling on the subterranean stream.
No decline in AOO has been observed, but it is inferred due to the decline and vulnerability of the habitat.
Iberoporus pluto occurs in a single cave, Soprador do Carvalho, in the Sicó karst area in central Portugal and it is under moderate disturbance (
This cave is the only known location in the historical range, as several caves in the same karst massif nearby have been sampled and this species has never been found elsewhere. Therefore, the current trend in number of locations is stable.
Only one specimen (female) of this species is known from a single location in central Portugal.
The specimen was collected in the bottom of a clay pool connected to a subterranean stream (
Soprador do Carvalho Cave is located in the vicinity of a village, agricultural fields and a quarry (
The genus Iberoporus Castro & Delgado, 2001 is endemic to the Iberian Peninsula and exclusively stygobiont (
The cave is explored for tourism and visitors step over the habitat where the species was found (
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 (
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.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Carabidae |
This species belongs to the T. fulvus-group of species.
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The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both 4 km2.
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 (
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.
AOO decline has been inferred due to the vulnerability of the habitat.
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 (
Alcobertas is the only known location in the historical range, therefore, the current trend in number of locations is stable.
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 (
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 (
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 (
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 (
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 (
The habitats are protected under the EU “Rede Natura 2000” (
Further investigation is needed about the population size, extent of distribution, ecology and life cycle. It is urgent to develop a management plan for this species, which will consequently improve the conservation of further cave-adapted species of Serra dos Candeeiros subunit of the Estremenho massif.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Carabidae |
This species belongs to the “T. fulvus-group” species complex.
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The extent of occurrence (EOO) is 73.4 km² and the maximum estimated area of occupancy (AOO) is 24 km².
Trechus gamae is a troglobiont beetle, known from five caves and from the mesovoid shallow substrate (scree slopes), all located in the Santo António Plateau, the central subunit of the Estremenho karst massif (
Trechus gamae was found in five caves: Algar de Marradinhas II, Algar das Gralhas VII, Algar do Pena, Algar da Arroteia and Algar do Ladoeiro, all located in the Santo António Plateau, the central subunit of the Estremenho karst massif. Recently, a single specimen was found in the mesovoid shallow substrate at 0.5 m depth in scree slopes of Fórnea, which is also located in the Santo António Plateau, showing that this species may also disperse through more superficial subterranean habitats (
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 (
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.
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) (
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 (
The distribution area of the species is all covered by the Natural Park of Serras d'Aire e Candeeiros. However, intense quarry activity is currently ongoing in the surrounding areas of the known localities. Algar do Pena is located 300 m from a quarry and Algar das Gralhas VII 168 m from the same quarry. Algar do Ladoeiro's entrance is 840 m from the closest urban areas, Algar das Marradinhas II is located 1.5 km from the nearest village and Algar da Arroteia is located 112 m from the closest house and 1.3 km from a quarry. Algar do Pena Cave hosts a laboratory and is open for visits upon previous booking. Fórnea is located 595 m from the closest house and 1.9 km from a quarry. The distribution area faces severe groundwater contamination (
Although the habitat is protected by law under the “Rede Natura 2000” (
Additional investigation about the population size, extent of distribution, ecology and life cycle is required. The development of a management plan that will improve the conservation of this cave-adapted species in the Santo António Plateau subunit of the Estremenho karst massif is fundamental to ensure its conservation and protection.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Carabidae |
This species belongs to the “T. fulvus-group” species complex.
Suppl. materials
The extent of occurrence (EOO) is 4 km2 and the maximum estimated area of occupancy (AOO) is 12 km².
Trechus lunai is a troglobiont Carabidae known from three horizontal caves, located in Serra de Aire/São Mamede Plateau (
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 (
Three populations are known exclusively from the Serra d'Aire/São Mamede Plateau subunit of the Estremenho karst massif (
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.
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 (
Trechus lunai was the only troglobiont species captured in these caves. All known localities are caves that flood seasonally (
Almonda Cave is located 50 m from a factory that conducts the subterranean river into the building for industrial use of the water and at 420 m from the village centre. The surrounding village is also heavily populated by agricultural fields. Contenda and Moinhos Velhos caves are heavily contaminated, as their subterranean network extends below the village of Mira d'Aire. The entrance of Moinhos Velhos Cave is located in the village centre and it has been explored for tourism since the 1960s with several complementary touristic infrastructures built (
The Contenda and Moinhos-Velhos caves develop below the village of Mira d'Aire and infiltration of sewage is observed in the underground. Therefore an effort to improve sewage treatment is necessary in order to prevent wastewater run-off into subterranean gealleries and groundwaters. Almonda Cave is classified as Property of Public Interest (IIP) since 1993 and protected due to archaeological heritage (
Information about the population size, extent of distribution, ecology and life cycle of this species is scarce, therefore, further investigation is required. It is also necessary to develop a management plan that will improve the conservation of the species in the Serra de Aire/S. Mamede Plateau subunits of the Estremenho karst massif.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Carabidae |
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 ground-beetle known from Portugal (
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The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both of 4 km².
Trechus tatai is a cave-adapted hygrophilous Carabidae known only from one small cave, located in Serra do Montejunto (
Trechus tatai only occurs in one cave, Algar do Javali, located in Serra do Montejunto. This species is geographically isolated (
Algar do Javali is the only known location for this species, therefore, the trend in number of locations is stable.
Only one population is known from Algar do Javali Cave, in Montejunto karst massif.
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 (
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 (
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.
Although this cave is protected by law through the “Rede Natura 2000” (
Some crucial steps necessary for the protection of the species are the development of a management plan for the conservation of this cave-adapted species in Serra do Montejunto and the promotion of further studies regarding population size, extent of distribution, ecology and life cycle.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Leiodidae |
This species is part of the "Speonemadus escalerai-group" and is recognisable by the equal/subequal length of the 2nd, 4th, 5th and 7th antennomeres and a slightly transverse and hexagonal pronotum (
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The extent of occurrence (EOO) is 7.4 km2 and the maximum estimated area of occupancy (AOO) is 12 km2. The three caves are located along a 48 km straight line, with Algarão do Remexido Cave being 23 km from Vale Telheiro and Vale Telheiro Cave being 25 km from Senhora Cave.
Speonemadus algarvensis was collected in three caves in the southermost province of Portugal in the Algarve, being most likely endemic to the central and eastern parts of the Algarve karst massif, a region also known as Barrocal Algarvio (
Speonemadus algarvensis is known from three caves (Vale Telheiro, Algarão do Remexido and Senhora) in the Algarve karst massif (
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 (
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.
Specimens were collected in three caves, 15 to 30 m deep, with high humidity levels and average temperatures of 17.8ºC (Vale Telheiro Cave), 18.8ºC (Senhora Cave) and 19.3ºC (Algarão do Remexido Cave). This species is endemic to the Algarve karst massif (
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 (
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.
Although the habitat is protected under legislation by the “Rede Natura 2000” (
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.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Staphylinidae |
Individuals display troglomorphism, such as microphthalmia, lack of wings and body elongation (
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The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both of 4 km².
Domene lusitanica was found in a single cave located in the Sicó karstic massif (
Domene lusitanica was collected in Cerâmica Cave, located in the Sicó karst area in central Portugal (Reboleira et al. 2011a). This Ccave extends for 355 m (
This species is known from a single population in central Portugal (
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 (
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 (
Cerâmica Cave is located 550 m from an animal farm, 3.5 km from the nearest village and 3.6 km from a quarry. It is surrounded by agricultural lands and Eucalyptus plantations.
The habitat is located in an “Rede Natura 2000” area (
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.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Staphylinidae |
This species displays body, leg and antennae elongation, microphthalmia and lack of wings (
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The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both 4 km2.
Domene viriatoi was collected in two galleries of the Buraco da Moura Cave, located in the Serra da Estrela Mountain foothills (
Domene viriatoi was collected from the Buraco da Moura Cave at the edge of the Estrela Mountain chain (
This species is known from a single population in the western border of the Estrela Mountain chain, the highest mountain of continental Portugal (
The cave is formed by granite blocks in the margins of the Caniça stream at an elevation of 677 m. It extends for 150 m of underground passages (
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 (
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.
Buraco da Moura Cave was classified as a “National Important Underground Shelter for Bats” therefore a decrease in human disturbance is expected (
Although the habitat is protected under legislation by the “Rede Natura 2000” (
A sustainable conservation plan for the species is only possible if more information about population size, extent of distribution, ecology and life cycle is collected.
Kingdom | Phylum | Class | Order | Family |
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Animalia | Arthropoda | Insecta | Coleoptera | Staphylinidae |
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The extent of occurrence (EOO) and the maximum estimated area of occupancy (AOO) are both 4 km2.
Domene darinkae is a cave-adapted rove beetle known from an abandoned mine in northern Portugal (
Domene darinkae is known from a single horizontal artificial cave, Santa Isabel mine, located in the Marão Mountain chain in north Portugal (
Only one specimen of this species is known from a single location in northern Portugal (
The only known specimen was collected in the rocky debris along the main tunnel of the Santa Isabel mine (
Domene darinkae is included in the subgenus Lathromene. It is a predator and exhibits troglomorphisms, such as depigmentation, elongation of body and antennae and accentuated microphthalmia (
Undisturbed areas in the surface of the mine need to be defined and established.
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.
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-the-year) 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
Beetles are the most diverse insects in Portuguese caves (
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 (
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
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.
This work was supported by a research grant (15471) from the VILLUM FONDEN and by Portuguese National Funds through Fundação para a Ciência e a Tecnologia (FCT) within the cE3c Unit funding UIDB/00329/2020.
Both authors contributed equally.
Villum Young Investigator grant (15471).
HiddenRisk – Understanding the impact of human activities on subterranean biodiversity
Natural History Museum of Denmark
Authors declare no conflict of interest.
1: Trechus machadoi (blue circle); 2: T. gamae (yellow circle); 3:T. lunai (pink circle); 4: T. tatai (red circle); 5: Iberoporus pluto (blue star); 6: Domene lusitanica (yellow diamond); 7: D. viriatoi (pink diamond); 8: D. darinkae (blue diamond); and 9: Speonemadus algarvensis (pink triangle). (A) Detail of northern distribution, (B) Detail of central distribution and (C) Detail of southern distribution. In green are protected areas.
Iberoporus pluto distribution: Soprador do Carvalho Cave, Penela, Coimbra District.
Trechus machadoi distribution: Alcobertas Cave and mesovoid shallow substratum, Rio Maior.
Trechus gamae distribution: (1) Algar da Arroteia Cave; (2) Fórnea (MSS); (3) Algar do Ladoeiro Cave; (4) Algar de Marradinhas II Cave; (5) Algar do Pena Cave; and (6) Algar das Gralhas VII Cave. All caves and MSS are located in the Santo António Plateau, the central subunit of the Estremenho karst massif. (A) Detail of distribution.
Trechus lunai distribution: (1) Contenda and Moinhos Velhos Cave system; and (2) Almonda Cave, both located in the Estremenho karst massif.
Trechus tatai distribution: Algar do Javali Cave, Montejunto karst massif.
Speonemadus algarvensis distribution: (1) Algarão do Remexido Cave; (2) Vale Telheiro Cave; and (3) Senhora cavea, all located in the Algarve karst massif.
Domene lusitanica distribution: Cerâmica Cave, Sicó karst area.
Domene viriatoi distribution: Buraco da Moura Cave, Estrela Mountain chain.
Domene darinkae distribution: Santa Isabel mine, Marão Mountain chain.