Article
To read the full-text of this research, you can request a copy directly from the authors.

Abstract

In light of recent alarming trends in human population growth, climate change, and other environmental modifications, a "Warning to humanity" manifesto was published in BioScience in 2017. This call reiterated most of the ideas originally expressed by the Union of Concerned Scientists in 1992, including the fear that we are "pushing Earth's ecosystems beyond their capacities to support the web of life. " As subterranean biologists, we take this opportunity to emphasize the global importance and the conservation challenges associated with subterranean ecosystems. They likely represent the most widespread nonmarine environments on Earth, but specialized subterranean organisms remain among the least documented and studied. Largely overlooked in conservation policies, subterranean habitats play a critical role in the function of the web of life and provide important ecosystem services. We highlight the main threats to subterranean ecosystems and propose a set of effective actions to protect this globally important natural heritage.

No full-text available

Request Full-text Paper PDF

To read the full-text of this research,
you can request a copy directly from the authors.

... As the upcoming CBD meeting will be held in the South China Karst, a region supporting the highest diversity of subterranean-adapted fishes, beetles, and millipedes globally (and likely to emerge as a subterranean biodiversity hotspot), we provide this roadmap for conserving the world's subterranean resources. Through a multilateral effort to robustly examine, monitor, and incorporate the subterranean biome into future conservation targets, the CBD will further improve the ecological effectiveness of protected areas by including groundwater resources, subterranean ecosystem services, and the profoundly endemic subsurface biodiversity (Elshall et al., 2020;Mammola et al., 2019Mammola et al., , 2020. To this end, this roadmap embodies five conceptual areas: (1) science gaps and data management needs; (2) anthropogenic stressors; (3) socioeconomic analysis and conflict resolution; (4) environmental education; and (5) national policies and multilateral agreements. ...
... Perhaps even more than surface ecosystems, the terrestrial subterranean biome is riddled by extensive knowledge gaps (Box 1, Figure 1). For example, for most known subterranean-obligate species, we have little more than observational data from a few human-accessible localities (Mammola et al., 2019(Mammola et al., , 2020; this makes assessing species for protective management extremely challenging. While the importance of ecosystem services associated with groundwater quality and cave-roosting bats is relatively well-documented (Elshall et al., 2020;Griebler et al., 2014;Mammola et al., 2019), a substantial effort will be required to quantify the scope, importance, and habitat requirements of subterranean bioindicator and ecosystem service species (Elshall et al., 2020). ...
... For example, for most known subterranean-obligate species, we have little more than observational data from a few human-accessible localities (Mammola et al., 2019(Mammola et al., , 2020; this makes assessing species for protective management extremely challenging. While the importance of ecosystem services associated with groundwater quality and cave-roosting bats is relatively well-documented (Elshall et al., 2020;Griebler et al., 2014;Mammola et al., 2019), a substantial effort will be required to quantify the scope, importance, and habitat requirements of subterranean bioindicator and ecosystem service species (Elshall et al., 2020). ...
Article
Full-text available
The 15th UN Convention on Biological Diversity (CBD) (COP15) will be held in Kunming, China in October 2021. Historically, CBDs and other multilateral treaties have either alluded to or entirely overlooked the subterranean biome. A multilateral effort to robustly examine, monitor, and incorporate the subterranean biome into future conservation targets will enable the CBD to further improve the ecological effectiveness of protected areas by including groundwater resources, subterranean ecosystem services, and the profoundly endemic subsurface biodiversity. To this end, we proffer a conservation roadmap that embodies five conceptual areas: (1) science gaps and data management needs; (2) anthropogenic stressors; (3) socioeconomic analysis and conflict resolution; (4) environmental education; and (5) national policies and multilateral agreements.
... Due to the intrinsic inaccessibility of subterranean ecosystems (Ficetola, Canedoli & Stoch, 2019) and many impediments to research (Mammola et al., 2021a), we currently know too little about subterranean biota to be able to routinely implement costeffective conservation interventions. To date, conservation of subterranean ecosystems has been dominated by problem-based studies focused on identifying the main drivers associated with subterranean biodiversity decline (Mammola et al., 2019a;Gerovasileiou & Bianchi, 2021). For example, we have elucidated the ecological impacts of polluted surface waters percolating underground (Di Lorenzo et al., 2015Manenti et al., 2021), the long-term consequences of climate change on specialised subterranean organisms adapted to thermally stable conditions (Mammola et al., 2019c;Pallarés et al., 2020a,b;Colado et al., 2022), and some of the negative impacts that pathogens and alien species can cause to subterranean ecosystems (Howarth et al., 2007;Wynne et al., 2014;Howarth & Stone, 2020;Hoyt, Kilpatrick & Langwig, 2021). ...
... We determined the main threats to subterranean ecosystems based on recent syntheses (Mammola et al., 2019a(Mammola et al., , 2020a complemented by our expert opinion. We grouped threats into the following eight categories: (i) Alien species & Pathogens (impacts due to alien species or pathogens); (ii) Climate change (impacts related to the alteration of climatic conditions); (iii) Overexploitation & Poaching (indiscriminate collection of species or overexploitation of biological resources); (iv) Pollution (organic and inorganic pollution events); (v) Surface habitat change (habitat alteration at the surface that affect subterranean systems; e.g. ...
... Monitoring was by far the most recommended intervention in our database (Figs 2E, 4C) despite its effects being indirecta pattern that is not exclusive to subterranean conservation (Buxton et al., 2020. This likely relates to a deficit of knowledge about subterranean ecosystems, which are notoriously difficult to explore, study, understand, and ultimately to protect (Ficetola, Canedoli & Stoch, 2019;Mammola et al., 2019aMammola et al., , 2021aGerovasileiou & Bianchi, 2021). Despite their perceived importance, only 11% of the proposed monitoring plans and methods were tested. ...
Article
Full-text available
Subterranean ecosystems are among the most widespread environments on Earth, yet we still have poor knowledge of their biodiversity. To raise awareness of subterranean ecosystems, the essential services they provide, and their unique conservation challenges, 2021 and 2022 were designated International Years of Caves and Karst. As these ecosystems have traditionally been overlooked in global conservation agendas and multilateral agreements, a quantitative assessment of solution-based approaches to safeguard subterranean biota and associated habitats is timely. This assessment allows researchers and practitioners to understand the progress made and research needs in subterranean ecology and management. We conducted a systematic review of peer-reviewed and grey literature focused on subterranean ecosystems globally (terrestrial, freshwater, and saltwater systems), to quantify the available evidence-base for the effectiveness of conservation interventions. We selected 708 publications from the years 1964 to 2021 that discussed, recommended, or implemented 1,954 conservation interventions in subterranean ecosystems. We noted a steep increase in the number of studies from the 2000s while, surprisingly, the proportion of studies quantifying the impact of conservation interventions has steadily and significantly decreased in recent years. The effectiveness of 31% of conservation interventions has been tested statistically. We further highlight that 64% of the reported research occurred in the Palearctic and Nearctic biogeographic regions. Assessments of the effectiveness of conservation interventions were heavily biased towards indirect measures (monitoring and risk assessment), a limited sample of organisms (mostly arthropods and bats), and more accessible systems (terrestrial caves). Our results indicate that most conservation science in the field of subterranean biology does not apply a rigorous quantitative approach, resulting in sparse evidence for the effectiveness of interventions. This raises the important question of how to make conservation efforts more feasible to implement, cost-effective, and long-lasting. Although there is no single remedy, we propose a suite of potential solutions to focus our efforts better towards increasing statistical testing and stress the importance of standardising study reporting to facilitate meta-analytical exercises. We also provide a database summarising the available literature, which will help to build quantitative knowledge about interventions likely to yield the greatest impacts depending upon the subterranean species and habitats of interest. We view this as a starting point to shift away from the widespread tendency of recommending conservation interventions based on anecdotal and expert-based information rather than scientific evidence, without quantitatively testing their effectiveness.
... Logistical difficulties in sampling groundwater ecosystems (Larned, 2012) and the often cryptic morphology of stygofauna (Bradford et al., 2010;Danielopol & Pospisil, 2001;Finston et al., 2007) have meant that biodiversity inventories of subterranean ecosystems are severely lacking in many locations (Ficetola et al., 2019;. Investigating the spatial scales of endemism within groundwater ecosystems is a critical step in understanding the implications of increasing threats, such as water abstraction and contaminant infiltration, as well as the efficacy of different management policies and practices (Boulton, 2020;Mammola et al., 2019). ...
... There have been repeated calls for accelerated scientific work to identify groundwater biodiversity, which is threatened with extinction before being discovered, identified, and ideally assigned a conservation status and protected (Gladstone et al., 2021;Mammola et al., 2019). Like most countries, knowledge of groundwater fauna is exceptionally poor in New Zealand. ...
... Our results support common findings of high biodiversity and short-range endemism in groundwater faunas internationally (e.g., Boulton, 2020;Gladstone et al., 2021) and likewise for the use of genetic data in identifying morphologically cryptic species, which are common in groundwaters (Boulton, 2020;Delic et al., 2017;Eme et al., 2018;Gladstone et al., 2021). By contributing to knowledge of the biodiversity and spatial distributions of groundwater taxa, we hope to help address some of the knowledge gaps inhibiting conservation of groundwater biodiversity (e.g., Boulton, 2020;Mammola et al., 2019Mammola et al., , 2020. ...
Article
Full-text available
We used DNA barcoding to assess the diversity and distribution of New Zealand's groundwater amphipods and isopods (Crustacea) and to determine whether biodiversity and endemism within tectonically active New Zealand are similar to those of more tectonically stable continents. Sixty-five wells were sampled in seven aquifers across four regions within the North and South islands of New Zealand, and resident invertebrates were morphologically identified and then assessed using sequencing of the mitochondrial DNA cytochrome c oxidase subunit one (COI) gene. Invertebrates were found in 54 wells. Of the 228 individual amphipods and isopods found in 36 of the wells, 154 individuals were successfully sequenced for COI (68% success rate) from 25 wells, with at least one well in each aquifer containing sequenced individuals. Of the 45 putative species identified using Barcode Index Numbers (BINs), 30 BINs (78% of all taxa and 83% of amphipods) were previously unrecorded. Substantial morphologically cryptic, species-level diversity was revealed, particularly within the amphipod Family Paraleptamphopidae. Similarly, one isopod taxon morphologically identified as Cruregens fontanus was assigned to five well-separated BINs based on COI sequences. Endemism appeared high, with all taxa regionally endemic; 87% of species were restricted to one aquifer and more than 50% restricted to one well. Non-saturated species accumulation curves indicated that, while additional sampling may increase the range of some currently identified taxa, additional range-restricted taxa are also likely to be discovered. Patterns of diversity and short-range endemism were similar to those found elsewhere, including locations which are more tectonically stable. The predominance of local endemism within New Zealand's groundwater fauna suggests that land-use activities and groundwater extraction require careful evaluation to minimize threats to groundwater biodiversity.
... Using modern technologies, environmental processes and threats on above-ground surface ecosystems (e.g., forest ecosystems) can easily be mapped with remote sensing (Rose et al., 2015). Yet most subterranean ecosystems such as caves are challenging to map and have frequently been overlooked and neglected in prioritisation (Mammola et al., 2019;McClure et al., 2020;Sánchez-Fernández et al., 2021). Up to 90 % of species in a single cave may be undescribed in some countries (Manenti et al., 2018;Whitten, 2009), meaning many thousands of species remain undescribed and potentially at risk (Ficetola et al., 2019). ...
... Up to 90 % of species in a single cave may be undescribed in some countries (Manenti et al., 2018;Whitten, 2009), meaning many thousands of species remain undescribed and potentially at risk (Ficetola et al., 2019). However, most conservation projects and funds are focused on taxa generally considered to be charismatic (Ford et al., 2017), but neglect fragile ecosystems with high endemism such as cave ecosystems (Mammola et al., , 2019Manenti et al., 2018). Subterranean ecosystems are threatened by both immediate threats to the caves themselves, and modifications of the surrounding environment (Phelps et al., 2016;Tanalgo et al., 2018). ...
... Furthermore, contextspecific threats (e.g., vulnerability to religious activities in Buddhist regions, where caves often become temples or religious sites) need to be accounted for explicitly for indices to be effective. Cave ecosystems host both high diversity and site-specific endemism, and are used by up to 30 % of all bat species yet are rarely included in global priorities (Mammola et al., 2019;Sánchez-Fernández et al., 2021). The persistence of high levels of biodiversity in caves is linked to more pristine cave environments with less anthropogenic pressure. ...
Article
Full-text available
Research and media attention is disproportionately focused on taxa and ecosystems perceived as charismatic, while other equally diverse systems such as caves and subterranean ecosystems are often neglected in biodiversity assessments and prioritisations. Highlighting the urgent need for protection, an especially large fraction of cave endemic species may be undescribed. Yet these more challenging systems are also vulnerable, with karsts for example losing a considerable proportion of their area each year. Bats are keystone to cave ecosystems making them potential surrogates to understand cave diversity patterns and identify conservation priorities. On a global scale, almost half (48 %) of known bat species use caves for parts of their life histories, with 32 % endemic to a single country, and 15 % currently threatened. We combined global analysis of cave bats from the IUCN spatial data with site-specific analysis of 1930 bat caves from 46 countries to develop global priorities for the conservation of the most vulnerable subterranean ecosystems. Globally, 28 % of caves showed high bat diversity and were highly threatened. The highest regional concentration of conservation priority caves was in the Palearctic and tropical regions (except the Afrotropical, which requires more intensive cave data sampling). Our results further highlight the importance of prioritising bat caves by incorporating locally collected data and optimising parameter selection (i.e., appropriate landscape features and threats). Finally, to protect and conserve these ecosystems it is crucial that we use frameworks such as this to identify priorities in species and habitat-level and map vulnerable underground habitats with the highest biodiversity and distinctiveness.
... Second, we focus on the analytical and technical aspects of eDNA, including its potential and limitations for subterranean environments, and third, we propose research avenues that will advance molecular-based studies of subterranean ecology. Indeed, the vastness and magnitude of subterranean environments requires effective and standardised sampling strategies to accurately monitor the taxonomically and functionally diverse communities comprising subterranean ecosystems (Mammola et al., 2019a). Beyond its analytical value, eDNA has the potential to focus a spotlight on some of the most essential, but undervalued and threatened, ecosystems in the world, and may dismantle conceptual boundaries (e.g., subterranean vs surface), creating a more holistic approach to studying freshwater and terrestrial ecology. ...
... Subterranean and groundwater ecosystem management strategies in many locations around the world rely on surveys and ongoing biomonitoring to assess and monitor potential changes in biological community composition (Gibson et al., 2019;Mammola et al., 2019a;Steube et al., 2009). Traditional sampling approaches such as haul net surveys and litter traps include sampling the habitat followed by morphological identification and confirmation of species present with DNA sequence data (i.e., Sanger sequences). ...
... As discussed above, the health, viability, and functioning of subterranean communities and ecosystems are increasingly under threat from pollution, climate change, and water extraction that impact the groundwater profile (Mammola et al., 2019a;Mammola et al., 2019c). Therefore, there is an urgent need for research on subterranean communities and the elements within them. ...
Article
Full-text available
Monitoring of biota is pivotal for the assessment and conservation of ecosystems. Environments worldwide are being continuously and increasingly exposed to multiple adverse impacts, and the accuracy and reliability of the biomonitoring tools that can be employed shape not only the present, but more importantly, the future of entire habitats. The analysis of environmental DNA (eDNA) metabarcoding data provides a quick, affordable, and reliable molecular approach for biodiversity assessments. However, while extensively employed in aquatic and terrestrial surface environments, eDNA-based studies targeting subterranean ecosystems are still uncommon due to the lack of accessibility and the cryptic nature of these environments and their species. Recent advances in genetic and genomic analyses have established a promising framework for shedding new light on subterranean biodiversity and ecology. To address current knowledge and the future use of eDNA methods in groundwaters and caves, this review explores conceptual and technical aspects of the application and its potential in subterranean systems. We briefly introduce subterranean biota and describe the most used traditional sampling techniques. Next, eDNA characteristics, application, and limitations in the subsurface environment are outlined. Last, we provide suggestions on how to overcome caveats and delineate some of the research avenues that will likely shape this field in the near future. We advocate that eDNA analyses, when carefully conducted and ideally combined with conventional sampling techniques, will substantially increase understanding and enable crucial expansion of subterranean community characterisation. Given the importance of groundwater and cave ecosystems for nature and humans, eDNA can bring to the surface essential insights, such as study of ecosystem assemblages and rare species detection, which are critical for the preservation of life below, as well as above, the ground.
... In addition, it is difficult to precisely measure important abiotic factors such as gas exchange and temperature that affect insect behavior and survival in soil (Villani and Wright 1990;Hagedorn et al. 2019), partly because soil macrofauna actively create channels, pores, and aggregates, and transform soil nutrients in ways that directly modify the chemical, structural, and acoustic characteristics of the soil in which they live (Brussaard et al. 1997;Veen et al. 2019). Consequently, current technologies are barely in the initial stages of developing more than a fleeting awareness of the diverse environments sustaining the subterranean biota (Mammola et al. 2019). The difficulty of precisely investigating subterranean biotremology has restricted the capability and opportunity to conduct such studies. ...
... Nevertheless, there are fundamentally important reasons to study communication and the incidental movement and feeding vibrations of arthropods in soil, given the ecological importance of subterranean herbivory (Hunter 2001;Lavelle et al. 2006;van Dam 2009;Adhikari and Hartemink 2016;Veen et al. 2019) and the evolutionary processes (Camacho et al. 1992) leading to a high diversity of known subterranean arthropod lineages (Mammola et al. 2019). Subterranean arthropods interact with the environment over a much smaller spatial scale than above-ground arthropods (Veen et al. 2019), and subterranean species sometimes are restricted to small areas such as isolated mountain valleys (van Tol et al. 2004). ...
... Subterranean arthropods interact with the environment over a much smaller spatial scale than above-ground arthropods (Veen et al. 2019), and subterranean species sometimes are restricted to small areas such as isolated mountain valleys (van Tol et al. 2004). Population growth often is small because energy-limited and stable environments have selected for long-lived species with low metabolism and fecundity (Mammola et al. 2019). Consequently, needs for conservation efforts to protect endemic subterranean arthropods are reported often (Harvey et al. 2011), especially in view of recently reported trends of reduced biomass of flying insects in protected areas (Hallmann et al. 2017). ...
... Groundwater is the largest reservoir of liquid freshwater globally that is of direct and major interest to humans (Gibert & Deharveng, 2002;Mammola et al., 2019). Despite its vastness, groundwater is also one of the most endangered habitats. ...
... Despite its vastness, groundwater is also one of the most endangered habitats. Habitat loss, overexploitation, contamination and climate change threaten its inhabitants (Mammola et al., 2019). Groundwater fauna is unique in many aspects. ...
... It comprises phylogenetic and geographic relicts (Humphreys, 2000), disproportionally high number of endemic species (Bregović et al., 2019) and plays a key role in ecosystem services (Griebler & Avramov, 2015). However, conservation biologists have largely neglected groundwater biodiversity, even in comparatively well-studied regions such as Europe and North America (Mammola et al., 2019). Species inventories and mapping of groundwater biodiversity have been lagging behind initiatives on surface systems (Ficetola et al., 2019). ...
Article
Full-text available
Aim Groundwater harbours an exceptional fauna and provides invaluable ecosystem services, yet is among the least explored and consequently least protected ecosystems. Successful protection of its biodiversity depends on complete species inventories, knowledge of species spatial distribution, and quantification of biodiversity patterns, as well as disentanglement of the processes that shaped biodiversity patterns. We studied the hyper‐speciose amphipod genus Niphargus as a model system within a global subterranean biodiversity hotspot. We linked the biodiversity patterns with possible underlying processes and discuss the needs to include information on different origins of biodiversity into conservation approaches. Location Europe, Western Balkans. Methods We analysed biodiversity patterns of Niphargus using two biodiversity metrics, species richness and phylogenetic diversity, on a grid‐based approach. To account for high cryptic diversity, we replaced nominal species with taxonomic units identified in unilocus delimitations (MOTUs). We built a time‐calibrated multilocus phylogeny of 512 Niphargus MOTUs from within and outside the study area, and calculated Faith's phylogenetic diversity, standardized effect sizes of phylogenetic diversity, and residual of phylogenetic diversity regressed onto species richness. Results Within the study area, we recognized 245 MOTUs, belonging to different Niphargus clades. Species richness is highest in a north‐western hotspot, although some species‐rich cells were detected also in the south‐east. High phylogenetic diversity coincides with high species richness in the north‐west, while in the south‐east it is lower than expected. Main conclusions We have shown that species richness does not predictably correlate with phylogenetic diversity. This difference suggests that different processes have led to the formation of species‐rich areas in the Western Balkans: through a combination of dispersal and speciation in the north‐west, and local radiation in the south‐east, respectively. This calls for caution in conservation strategies relying solely on number of species and may change the view on conservation priorities within this region.
... Due to the unique abiotic characteristics and dependencies on organic matter input from the surface, life in subterranean environments is often accompanied by dramatic morphological, anatomical and ecological adaptations, and worldwide many species have evolved to be endemic cave specialists [8]. Furthermore, trophic chains in caves are generally regarded as simpler and with communities being characterised by (i) lacking photosynthetic primary producers, (ii) invertebrates that are typically adapted and confined to all aspects of their life cycle underground, and (iii) metabolically active microbes with important biogeochemical activities [9]. While the species diversity of cave macro-organisms is low [10], the diversity of cave micro-organisms can be as high as in surface communities [11][12][13][14][15][16][17][18][19]. ...
... The transition from surface to subterranean ecosystems involves marked environmental changes, resulting from a combination of lack of light, low nutrient content and more stable conditions [9,58]. Here, we investigated the invertebrate and bacterial communities, along with soil properties of understudied caves in subarctic Northern Norway to understand biotic interactions with nutrient availability in these fragile ecosystems. ...
Article
Full-text available
Background Subarctic regions are particularly vulnerable to climate change, yet little is known about nutrient availability and biodiversity of their cave ecosystems. Such knowledge is crucial for predicting the vulnerability of these ecosystems to consequences of climate change. Thus, to improve our understanding of life in these habitats, we characterized environmental variables, as well as bacterial and invertebrate communities of six subarctic caves in Northern Norway. Results Only a minuscule diversity of surface-adapted invertebrates were found in these caves. However, the bacterial communities in caves were compositionally different, more diverse and more complex than the nutrient-richer surface soil. Cave soil microbiomes were less variable between caves than between surface communities in the same area, suggesting that the stable cave environments with tougher conditions drive the uniform microbial communities. We also observed only a small proportion of cave bacterial genera originating from the surface, indicating unique cave-adapted microbial communities. Increased diversity within caves may stem from higher niche specialization and levels of interdependencies for nutrient cycling among bacterial taxa in these oligotrophic environments. Conclusions Taken together this suggest that environmental changes, e.g., faster melting of snow as a result of global warming that could alter nutrient influx, can have a detrimental impact on interactions and dependencies of these complex communities. This comparative exploration of cave and surface microbiomes also lays the foundation to further investigate the long-term environmental variables that shape the biodiversity of these vulnerable ecosystems.
... It is a worldwide consensus that subterranean habitats, as well as their communities, are highly singular, fragile, and represent one of the most threatened environments in the world (Elliott, 2005;Culver, Pipan, 2019;Mammola et al., 2019). Troglobites, in particular, are intrinsically fragile and vulnerable to environmental changes, given their small populations with low resilience (Culver, Pipan, 2009). ...
... Troglobites, in particular, are intrinsically fragile and vulnerable to environmental changes, given their small populations with low resilience (Culver, Pipan, 2009). All troglobitic species should be protected by law and included in at least the Vulnerable category (VU) of IUCN (International Union for Conservation of Nature), an idea that has been advocated by several authors (e.g., Juberthie, 2000b;, 2021Fernandes et al., 2016;Trajano et al., 2016;Gallão, Bichuette, 2018;Culver, Pipan, 2019;Mammola et al., 2019;Bichuette, 2021). ...
Article
Full-text available
The present work brings information on threats to the subterranean fishes in Brazil. Currently, at least 36 species are known, 22 of which are already formally described. Endemism is the rule for most of them. Regarding their conservation, these fishes are in general considered threatened: and most of the already formally described species are included in national lists of threatened fauna, and only four of them are included in the global list of the IUCN. Regarding habitats, Brazilian subterranean fishes occur in alluvial sediments (part of the hyporheic zone), shallow base-level streams, flooded caves, lakes in the water table, upper vadose tributaries, and epikarst aquifers. We detected 11 main threats, mainly related to agriculture, pasture, and hydroelectric plans, but unmanaged tourism and pollution are also significant threats. Two threats affect a high number of species (physical change of the habitat and food restriction). The river basins with the higher number of identified threats are the upper Tocantins (eight) followed by the upper Paraguaçu (six). Effective proposals to protect this neglected component of the Brazilian biodiversity are still scarce, such as monitoring projects and their function in the subterranean communities, besides education projects aiming to develop public awareness.
... Alternatively, these demographic differences may indicate environmental changes that are detrimental to the cave-adapted G. subterraneus but not to G. porphyriticus. Obligate cave species are expected to be more susceptible to environmental changes because of their specialized troglobitic adaptations (Mammola et al. 2019) and some cave-associated salamanders in urbanizing watersheds have suffered declines (Bendik et al. 2014). Deforestation, sedimentation, and water contamination are principal threats to karst systems (Harley et al. 2011;Mammola et al. 2019), but these threats are not apparent in the General Davis Cave watershed. ...
... Obligate cave species are expected to be more susceptible to environmental changes because of their specialized troglobitic adaptations (Mammola et al. 2019) and some cave-associated salamanders in urbanizing watersheds have suffered declines (Bendik et al. 2014). Deforestation, sedimentation, and water contamination are principal threats to karst systems (Harley et al. 2011;Mammola et al. 2019), but these threats are not apparent in the General Davis Cave watershed. ...
Article
Full-text available
Due to their limited geographic distributions and specialized ecologies, cave species are often highly endemic and can be especially vulnerable to habitat degradation within and surrounding the cave systems they inhabit. We investigated the evolutionary history of the West Virginia Spring Salamander (Gyrinophilus subterraneus), estimated the population trend from historic and current survey data, and assessed the current potential for water quality threats to the cave habitat. Our genomic data (mtDNA sequence and ddRADseq-derived SNPs) reveal two, distinct evolutionary lineages within General Davis Cave corresponding to G. subterraneus and its widely distributed sister species, Gyrinophilus porphyriticus, that are also differentiable based on morphological traits. Genomic models of evolutionary history strongly support asymmetric and continuous gene flow between the two lineages, and hybrid classification analyses identify only parental and first generation cross (F1) progeny. Collectively, these results point to a rare case of sympatric speciation occurring within the cave, leading to strong support for continuing to recognize G. subterraneus as a distinct and unique species. Due to its specialized habitat requirements, the complete distribution of G. subterraneus is unresolved, but using survey data in its type locality (and currently the only known occupied site), we find that the population within General Davis Cave has possibly declined over the last 45 years. Finally, our measures of cave and surface stream water quality did not reveal evidence of water quality impairment and provide important baselines for future monitoring. In addition, our unexpected finding of a hybrid zone and partial reproductive isolation between G. subterraneus and G. porphyriticus warrants further attention to better understand the evolutionary and conservation implications of occasional hybridization between the species.
... Concern about cave ecosystems, their species, uniqueness, and vulnerability has increased in the past two decades. There is a scientific consensus that caves are globally threatened environments and need more protection (Furey and Racey 2015;Phelps et al. 2016;Mammola et al. 2019). For environmental licensing of the mineral sector in Brazil, bat caves in areas currently subject to mining have legal protection. ...
Article
Bat caves harbor exceptional populations of insectivorous bats. Those bats play an important role as insect suppressors and produce large quantities of guano, which is essential for maintaining cave ecosystems since entire highly specialized cave biotas may heavily rely on bat guano as their main energy input. Although ecologically relevant, few studies have estimated insect consumption and guano input in Neotropical bat caves. We provide estimates for five bat caves used by Pteronotus gymnonotus and P. personatus (Mormoopidae) in northeastern Brazil. Using a non-invasive automated system, we counted bats, then captured and weighted individuals to estimate insect consumption, and, with collectors and rulers, estimated the amount and speed of guano accumulation in cave sectors. Bat abundance varied between and within caves, up to 158,884 bats, indicating highly dynamic occupation patterns. Insect consumption varied from 0.6 to 2.5 g/bat for P. gymnonotus (~5 to 20% of their body weight) and 0.8 to 2.0 g/bat for P. personatus (~10 to 28% of their body weight). Guano deposition was spatially and temporally heterogeneous (from 0 to 738 g/m2/96h). Some caves showed a 15-cm increase in guano deposits on the cave floor in 7 months. Bulky guano deposits in those caves stressed the bat role as insect suppressors. The present study provides baseline quantitative data on the contributions of bats to cave ecosystems and valuable data for estimates of ecosystem services provided by bats.
... Mammola et al. 2019). Isto faz com o número de interações seja menor, facilitando a sua análise ecológica, mas aumentando também a sua vulnerabilidade e sensibilidade ambiental(Castaño-Sánchez et al. 2020). ...
Article
Full-text available
O Planalto das Cesaredas é formado por calcários marinhos do Jurássico Inferior a Superior (Toarciano ao Titoniano, ~180 a 144 milhões de anos) e alberga várias cavidades naturais com condições propícias à vida subterrânea. Localizado a baixa altitude na transição entre o maciço de Montejunto e vale Tifónico das Caldas da Rainha, este afloramento cársico inclui-se no distrito bioespeleológico Lusitânico, unidade biogeográfica que inclui os maciços da Arrábida, Estremenho, Sicó-Condeixa-Alvaiázere, Outil-Cantanhede e os afloramentos calcários da Península de Lisboa. O interesse científico sobre a biologia das grutas das Cesaredas teve o seu despertar há sete décadas e investigações recentes revelaram que a sua fauna estritamente subterrânea é composta sobretudo por invertebrados: moluscos, aracnídeos, miriápodes, crustáceos e insectos. A descoberta de espécies endémicas, i.e., cuja única área de distribuição mundial se encontra confinada ao Planalto das Cesaredas, lança grandes desafios em termos de conservação e urge estabelecer medidas de proteção para salvaguardar a sua biodiversidade subterrânea. Este artigo oferece uma perspectiva sobre a geologia, composição faunística e a ecologia das grutas das Cesaredas.
... Worldwide, increasing over-extraction and contamination of groundwater resources coupled with direct and indirect climate change impacts (including but not limited to droughts, altered aquifer recharge, reduced surface productivity, changing dynamics of OM flux, and functional cascade effects) are threatening the complex matrix of habitats and biota associated with GDEs (Mammola et al., 2019;Sánchez-Fernández et al., 2021). It has been estimated that almost two thirds of the world's largest aquifers (21 out of 37) have exceeded their sustainable limits (Richey et al., 2015). ...
Article
Full-text available
Groundwater environments interact with and support subterranean biota as well as superficial aquatic and terrestrial ecosystems. However, knowledge of subterranean energy flows remains incomplete. Cross-boundary investigations are needed to better understand the trophic structures of groundwater ecosystems and their reliance on carbon inputs from aboveground. In this study we used carbon and nitrogen stable isotope analyses combined with radiocarbon fingerprints to characterise organic flows in groundwater ecosystems. We coupled these data with DNA metabarcoding of the gut contents of stygofauna to further elucidate organic matter (OM) sources and shifts in diet preferences. Samples were collected from the arid zone Sturt Meadows calcrete aquifer under low rainfall (LR) and high rainfall (HR) conditions. Bayesian modelling of 14 C, δ 13 C, and δ 15 N data indicated that primary consumers (copepods) incorporated mainly particulate organic carbon (POC) under LR but during HR shifted to root derived material (either exudates or direct root grazing). By contrast, diets of secondary consumers (amphipods) were dominated by root material under both LR and HR. Our DNA metabarcoding-based results indicate that amphipods relied primarily on root inputs from perennial trees (likely Eucalyptus and Callitris) during the dry season (LR). Under HR, diets of both amphipods and copepods also included organic material derived from a broad range of more shallow rooted shrubs, and ephemeral herbs and grasses. Our findings illustrate the complexity of functional linkages between groundwater biota and surface terrestrial ecosystems in environments where aboveground productivity, diversity and OM flux to groundwater are intimately linked to often episodic rainfall.
... 16.11 and 12.10 mg/kg, respectively. The average contents of Hg, Cd, Pb and Cr in soil within the research area are 2.03, 1.36, 1.11 and 1.23 times of the soil background values in Shaanxi Province, respectively. ...
Article
Full-text available
The ecological environment in Loess Plateau of Northern Shaanxi is fragile, so the soil pollution caused by the exploitation of coal resources cannot be ignored. With Shigetai Coal Mine in Loess Plateau of Northern Shaanxi as the object of study for field survey and sampling, the content of heavy metals in soil is analyzed, the environmental pollution in the research area is evaluated by the single factor pollution index method, comprehensive pollution index method and potential ecological risk index method, and the spatial distribution characteristics of heavy metals are discussed by the geostatistics method. According to the study results, the average contents of heavy metals Hg, Cd, Pb and Cr are 2.03, 1.36, 1.11 and 1.23 times of the soil background values in Shaanxi Province respectively and the average contents of other heavy metals are lower than the soil background values in Shaanxi Province; Hg and Cd show moderate variation while As, Pb, Cr, Zn, Ni and Cu show strong variation; the skewness coefficients and kurtosis coefficient of Cd, As and Cu in the soil within the research area are relatively high, and these elements are accumulated in large amounts. Single factor pollution index (Pi) and potential ecological risk index (E) indicate that heavy metal Hg is the main pollution factor and mainly distributed in the east and north of the research area. The comprehensive index of potential ecological risk (RI) of the research area is 1336.49, showing an extremely high ecological risk, and the distribution characteristics of potential ecological risk are consistent with that of potential ecological risk index (E) of Hg. The results of ecological risk warning show that Hg is in a slight warning status, while Cd, Pb and Cr are in a warning status. The areas with high ecological risk warning values are mainly distributed in the east and north, and the whole research area shows relatively obvious zonal distribution law. The soil is disturbed greatly during the coal mining, so the ecological governance of the mine area shall adapt to the local natural conditions and regional environmental characteristics and follow the principle of “adjusting governance measures based on specific local conditions and classifications”. An environmentally sustainable governance manner shall be adopted to realize the protection of the ecological environment and high-quality development of coal resources.
... Their special nature has long been noted by the various Pacific Islander cultures that confer sacred or supernatural properties on anchialine shrimps (Gordon, 1936;Wear & Holthuis, 1977;Choy, 1987). Nonetheless, anchialine environments are highly threatened by human activities (e.g., Bucol & Alcala, 2013;Iliffe, 2002;Sakihara, 2012;Husana & Yamamuro, 2013;Mammola et al., 2019). ...
Article
Full-text available
We document the current status of the enigmatic bright-red anchialine shrimps in Tiniguiban Islet in the western Visayas region of the Philippines. A second epigeal anchialine pool was discovered in nearby Hinlaran Islet, and two shrimp species were recorded in the pools, the large barbouriid Parhippolyte uveae Borradaile, 1900 as previously reported and a much smaller atyid Antecaridina lauensis (Edmondson, 1935). The shrimps move in and out of the pools with tidal fluctuations, since both pools completely dry out at low tide. The shrimps remain very abundant in Tiniguiban despite extensive modifications made by its owners and an apparent decrease in dissolved oxygen levels. Magico-religious beliefs regarding these unusual red shrimps persist, yet a subtle shift in values is noted, particularly with the growing popularity of the Tiniguiban pool with local tourists. Further modernization and coastal development may rapidly threaten these rare shrimp populations. Our study provides baseline data for systematic study and future protection of these rare pools and their shrimps that uniquely dwell at the interface of cave and marine habitats.
... The anthropogenic impact has hit all earth ecosystems. Even the somewhat hidden and isolated aquifers are threatened by contamination and over-exploitation (Mammola et al., 2019). Especially, contamination with aromatic hydrocarbons and halogenated organic compounds poses a long-lasting hazard. ...
Chapter
This chapter discusses the relevance of microbial biodiversity in the groundwater environment. We provide information on the general methodological approaches that are used to estimate the microbial diversity. The importance of biodiversity as an inherent value of an ecosystem and its implication for ecosystem functioning is highlighted. We provide insight on the spatial and temporal scales of microbial diversity, which can span several orders of magnitude in aquifers. Relevant groundwater taxa and groundwater habitats are presented in terms of case studies that illustrate the breadth of ongoing research. Finally, the chapter discusses the knowledge gaps and potential for microbial diversity research in groundwater. Main concepts covered: Microbial diversity connects genetic variance, species biodiversity up to the ecosystem level. We delineate the biotic and abiotic conditions, which are the driving and limiting factors of biodiversity. Consequently, we discuss the effect of varying spatial scales and how to compare intra-and inter-community diversity as well as total diversity of a landscape. The intrinsic and utilitarian value of microbial biodiversity in groundwater habitats is presented in terms of ecosystem functioning. We cover the concepts of ecosystem stability and functional redundancy. Main methods covered: The methods covered in this chapter include a brief discussion of the molecular tools that are used to characterize the microbial diversity in groundwater. We highlight the main advantages of new sequencing techniques which work at unprecedented sensitivity and speed. Therefore, we stress the importance of bioinformatic and statistical analysis combined with a thoughtful experimental design. Conclusion/Outlook: The groundwater environment is a vital link for terrestrial and aquatic ecosystems. The microbial biodiversity in the subsurface is a critical factor for ecosystem services, but remains a frontier of ecological research.
... Metazoans play a potentially important, albeit insufficiently documented role, in maintaining goods and services delivered by groundwater ecosystems to humankind (Boulton et al., 2008;Griebler and Avramov, 2015). Yet, they are neither recognized as conservation priorities, nor integrated into management policies of groundwater resources (Mammola et al., 2019). In Europe, the assessment of groundwater resources is still exclusively based upon criteria of water quality and aquifer productivity, whereas that of surface water bodies is largely based upon the concept of ecological status. ...
Chapter
Aim: Groundwater hosts a high diversity of metazoans that dwell in open spaces within the rock. That diversity makes an important, yet underestimated, contribution to freshwater biodiversity. Here, we briefly describe groundwater habitats, the composition and spatial distribution of groundwater metazoan assemblages, the characteristic traits of specialist groundwater organisms, and their role in groundwater ecosystems. Main concepts covered/Main methods covered: Groundwater habitats all have in common that they are dark and hence they lack photosynthetic production. They are nevertheless highly diverse in terms of void space available to metazoans, interconnectedness of the voids and hydrological linkages to surface ecosystems. Groundwater metazoan assemblages are made of species living exclusively in groundwater, which we refer to as specialist groundwater species, and species that complete their life cycle both in groundwater and in surface water, which we refer to as generalists. Assemblages of specialist groundwater metazoans show a high spatial turnover in species composition over short distances and low local species diversity relative to regional diversity. A major evolutionary hypothesis is that selective pressures specific to groundwater shape converging phenotypes among specialist groundwater species, which differ strikingly from those observed in surface species. Phenotypes of specialist groundwater metazoans are nevertheless extremely diverse, even though the combination of selective pressures causing that diversity is yet to be elucidated. Groundwater metazoans have direct and indirect effect on hydraulic attributes of aquifers and ecosystem processes, which contribute to delivering important ecosystem services such as water provision and purification. Conclusion/outlook: Groundwater metazoans belong to the invisible component of Earth's biodiversity. Documenting and understanding their compositional and functional diversity has revealed the fascinating content of a Pandora's Box of which much still remains unknown. Much of the progress achieved in understanding mechanisms shaping phenotypes, biodiversity distribution patterns and organisms' function is yet to translate into policies that frame the management of groundwater ecosystems and the important services they deliver to humans.
... This information, combined with evaluation of exposure to climatic changes at their present locations, is needed to identify species or populations at greatest risk and to preserve this fragile and valuable component of global biodiversity, especially when it is systematically neglected by global climate change conservation strategies and agendas (Arneth et al., 2020;S anchez-Fern andez et al., 2021). In the era of climate change, when insect populations are going extinct at an unprecedented peace (Cardoso et al., 2020;Eisenhauer et al., 2019), it is important not to forget the biodiversity that dwells below our feet (Mammola et al., 2019c;S anchez-Fern andez et al., 2021;Wynne et al., 2021). It will be affected the same, if not more so, by a global temperature increase. ...
Article
The climatic variability hypothesis predicts the evolution of species with wide thermal tolerance ranges in environments with variable temperatures, and the evolution of thermal specialists in thermally stable environments. In caves, the extent of spatial and temporal thermal variability experienced by taxa decreases with their degree of specialization to deep subterranean habitats. We use Phylogenetic Generalized Least Squares to model the relationship between thermal tolerance (upper lethal limits), subterranean specialization (estimated using ecomorphological traits) and habitat temperature in sixteen beetle species of the tribe Leptodirini (Leiodidae). We found a significant, negative relationship between thermal tolerance and the degree of subterranean specialization. Conversely, habitat temperature had only a marginal effect on lethal limits. In agreement with the climatic variability hypothesis and under a climate change context, we show that the specialization process to live in deep subterranean habitats involves a reduction of upper lethal limits, but not an adjustment to habitat temperature. Thermal variability seems to exert a higher evolutionary pressure than mean habitat temperature to configure the thermal niche of subterranean species. Our results provide novel insights on thermal physiology of species with poor dispersal capabilities and on the evolutionary process of adaptation to subterranean environments. We further emphasize that the pathways determining vulnerability of subterranean species to climate change greatly depend on the degree of specialization to deep subterranean environments.
... In light of current concern about the fate of subterranean ecosystems (Mammola et al., 2019), investigating biodiversity pattern and local genetic differentiation of groundwater species is essential. Central questions are: What factors determine subterranean dispersal and biodiversity? ...
Article
Groundwater is one of our most important resources, however groundwater ecosystems are among the most understudied habitats of the planet earth. Studies on groundwater organisms are hampered by the difficult accessibility of species, the lack of morphological differentiation and the limitation for laboratory cultures. One important approach to overcome these shortcomings is to provide sensitive genetic methods to unravel patterns of biodiversity, population structure and gene flow in natural populations. In this study we present five sets of microsatellite markers developed for the isopods Asellus aquaticus and Proasellus slavus, the cyclopoides Paracyclops fimbriatus and Acanthocyclops sensitivus and the harpacticoide Bryocamptus echinatus (Crustacea). Two of these species were subjected to detailed population genetic analyses: We studied 501 specimens of Asellus aquaticus from four different regions in Northern Germany using nine microsatellite markers and 70 specimens of Bryocamptus echinatus using nine microsatellite markers from three different sampling sites in South-Western Germany. Our results show that genetic diversity is high (A. aquaticus: 10 to 20 and B. echinatus: 4 to 18 alleles per locus) among populations of aquatic invertebrates, populations are highly differentiated (FST > 0.2) and genetic differentiation was associated with geographic patterns. Applications of molecular genetic methods and their use for the detection of hydrological exchange processes relevant for drinking water suppliers are demonstrated and discussed.
... Many of these species occur in biodiversity hotspots that are threatened by both anthropogenic and natural threats 6 . Caves are key habitats for bats 12 but are nonetheless threatened and in need of conservation; despite hosting high endemism, cave ecosystems receive little attention in terms of fund allocation for scientific studies and conservation compared to their surface counterparts such as agricultural and forest ecosystems [11][12][13][14] . Cave taxa are adapted to light-limited underground environments and most of them are dependent on mobile species such as bats to transport organic nutrients into these environments 15,16 . ...
Preprint
Full-text available
Understanding biodiversity patterns as well as drivers of population declines, and range losses provides crucial baselines for monitoring and conservation. However, the information needed to evaluate such trends remains unstandardised and sparsely available for many taxonomic groups and habitats, including the cave-dwelling bats and cave ecosystems. Here, we present the DarkCideS 1.0 , a global database of bat caves and bat species based on curated data from the literature, personal collections, and existing datasets. The database contains information for geographical distribution, ecological status, species traits, and parasites and hyperparasites for 679 bat species known to occur in caves or use caves in their life-histories. The database contains 6746 georeferenced occurrences for 402 cave-dwelling bat species from 2002 cave sites in 46 countries and 12 terrestrial biomes. The database has been developed to be a collaborative, open-access, and user-friendly platform, allowing continuous data-sharing among the community of bat researchers and conservation biologists. The database has a range of potential applications in bat research and enables comparative monitoring and prioritisation for conservation.
... These areas are currently one of the most popular tourist destinations and require special protection. Groundwater, which created picturesque karst forms, is in great danger when it comes to anthropogenic impacts; therefore, sustainable development in the creation of new geoattractions in this area must be based on the principles of geoethics, because humanity in the future may depend much more on these water reserves (Gorelick et al., 1983;Kačaroğlu, 1999;Mammola et al., 2019). ...
Article
Full-text available
Geodiversity is the natural diversity of features of geological structure, relief, and soil cover, including the relationships between these features, their properties, and their impact on other elements of the natural and cultural environment. It is described and analyzed using various types of quantitative, qualitative, or quantitative–qualitative methods. The concept of a geodiversity map presented in this article belongs to the third of these groups of methods. Despite the use of optimization methods in the form of a hexagon grid or the analytic hierarchy process calculator, it still remains partially subjective. The use of this method to calculate the geodiversity of an entire province (the Western Carpathians) gives a general view of the natural diversity of this area and allows regions to be selected for more detailed analyses or comparisons to be made between them. The geodiversity map is also a very good background on which to illustrate geotourist potential, which is expressed in terms of the number and distribution of geosites. However, in the case of the Western Carpathians, these two variables do not correlate with each other.
... With this study, we take advantage of the potential that the subterranean environment offers to develop the field of conservation physiology (Cooke et al., 2013) to provide a physiologically-based vulnerability assessment of an endemic species. However, much more effort is needed to (i) increase our knowledge on the thermal tolerance of subterranean fauna (Mammola et al., 2019a) and (ii) consider the neglected subterranean ecosystems in conservation and management policies. Complementary approaches to estimate thermal tolerance not only in subterranean species but also in other poor dispersal species could provide more accurate predictions of their capacity to face climate change. ...
Article
Full-text available
1. Scientists are renewing their efforts to predict the impact of climate change on biodiversity. Subterranean environments represent ideal systems to study the effect of global change in species with poor dispersal capabilities. 2. We assess the vulnerability to climate change of the subterranean pseudoscorpion Neobisium (Blothrus) vasconicum vasconicum (Nonídez, 1925) (Neobisiidae). 3. Thermal tolerance was measured using two complementary estimates of upper thermal limits: (i) from thermal conditions of the localities in which the species occurs (realised upper thermal limit, RUTL), and (ii) from experimentally determined thermal tolerance data (physiological upper thermal limit, PhUTL). Then, thermal safety margins (TSM) were calculated for all known localities for current and future climatic conditions, using the thermal limits from both approaches. 4. The physiological thermal limit (PhUTL = 17.57 C) was 3.27 C higher than that obtained from the distributional and climate data (i.e., the hottest cave in which the species occurs; RUTL = 14.3 C). Regarding TSM, the future temperature (2070; RCP 8.5) of a half of the caves will be higher than the RUTL and in none of them, it would exceed the average PhUTL. This indicates that the species could have some physiological capacity to cope with warming temperatures in situ. 5. We hypothesize that the most realistic upper thermal limit of the species could be between the RUTL and PhUTL. This study shows that complementary approaches to estimate thermal tolerance could provide more accurate predictions of the capacity to face climate change, not only in subterranean species, but also in poor dispersal species.
... Caves are important habitats for bats and other unique species but are nonetheless threatened and in need of urgent conservation 10 . Despite hosting high endemism, cave ecosystems receive little attention in terms of fund allocation and appropriate priorities for scientific studies and conservation compared to their surface counterparts such as agricultural and forest ecosystems 10,13,[15][16][17][18] . Cave taxa are adapted to light-limited underground environments and most of them are dependent on mobile species such as bats to transport organic nutrients into these environments [19][20][21] . ...
Article
Full-text available
Understanding biodiversity patterns as well as drivers of population declines, and range losses provides crucial baselines for monitoring and conservation. However, the information needed to evaluate such trends remains unstandardised and sparsely available for many taxonomic groups and habitats, including the cave-dwelling bats and cave ecosystems. We developed the DarkCideS 1.0 (https://darkcides.org/), a global database of bat caves and species synthesised from publicly available information and datasets. The DarkCideS 1.0 is by far the largest database for cave-dwelling bats, which contains information for geographical location, ecological status, species traits, and parasites and hyperparasites for 679 bat species are known to occur in caves or use caves in part of their life histories. The database currently contains 6746 georeferenced occurrences for 402 cave-dwelling bat species from 2002 cave sites in 46 countries and 12 terrestrial biomes. The database has been developed to be collaborative and open-access, allowing continuous data-sharing among the community of bat researchers and conservation biologists to advance bat research and comparative monitoring and prioritisation for conservation.
... To date, seven caves have been surveyed, five of which are in the same locality of Imi Ougoug, without the species being observed. If a study was to be carried out on Ifri Ouado, it could, on the one hand, help assess threats to this species using the International Union for Conservation of Nature (IUCN) red List criteria (see also recommendations in Mammola et al. 2019). Furthermore, such a study would also contribute to the discussion undertaken by Mammola et al. (2018) on understanding how climate change may impact troglobiont species. ...
Article
Full-text available
The female of Agraecina agadirensis was discovered during a scientific internship in autumn 2019 in a cave system of the Moroccan High Atlas. Since then, several surveys were needed to find a male of the species, which was finally collected in April 2021. Based on this specimen, a detailed description, an extended taxonomic diagnosis, photos of the habitus, as well as drawings of the male pedipalpus are presented. Furthermore, we draw attention to anthropogenic threats to this endemic troglobionte species and propose the legal protection of the Imi Ougoug cave system.
... Despite their recognized value, academic research on and formal conservation of GDEs lags behind research and conservation efforts focused on ecosystems with substantial surface water input (e.g., streams; Mammola et al. 2019). Moreover, the majority of peer-reviewed GDE research focuses on hydrological and ecological characteristics (Eamus and Froend 2006, Kløve et al. 2011, Adams et al. 2015, Rohde et al. 2019, rather than their social value and characteristics. ...
Article
Full-text available
Groundwater dependent ecosystems (GDEs) are increasingly recognized as important conservation targets with linked ecological and social value. However, the social uses and values of GDEs have received relatively little research attention in the peer-reviewed literature, precluding their greater inclusion in policy and management decisions. To help fill this gap, we provide a case study from Kona, Hawaiʻi, where multiple types of GDEs are abundant, to illustrate the diversity of social uses and values of GDEs. To explore these uses and values, we combined a literature review, archival analysis, and key-informant interviews with resource managers and lineal descendants connected to three prominent GDEs: Indigenous aquaculture systems, anchialine pools, and nearshore ecosystems. Interviews focused on current and historical uses and values of GDEs, contemporary management challenges and strategies, and desired visions for the future. Interviewees expressed a range of uses and values associated with GDEs, which we categorized using a Hawaiʻi-based cultural ecosystem service framework focused on social connections, physical and mental health, spirituality, and knowledge. Importantly, results suggest that the historical value of these systems directly informs current social value, and that restoration efforts are largely carried out through biocultural approaches, which emphasize the mutually reinforcing restoration of ecology and culture. We found that interviewees seek to restore ecosystem functions, cultural practice, and connection to place, and in some cases, local food production. Achieving these goals requires addressing multiple and interacting threats to these systems including invasive species, land-based sources of pollution, groundwater pumping, and climate change. Importantly, effective and equitable restoration also rests on recognition and amplification of Indigenous rights, knowledge, practice, and governance. These results provide important lessons for land and water management and policy in Hawaiʻi as well as other islands and coastal areas where GDEs have important linked social and ecological value.
... Ninety-five percent of global freshwater (excluding the polar ice caps) is stored in the continental subsurface constituting a major source of ecosystem services (Griebler et al. 2014). However, at the same time, subterranean ecosystems are threatened by different anthropogenic impacts and by a general inadequacy of protection policies, conditions that seriously endanger their biological diversity and the ecosystem services they provide (Boulton et al. 2008;Mammola et al. 2019). Improving our knowledge of groundwater fauna and its origin is thus of paramount importance for implementing effective conservation practices. ...
Article
Full-text available
Groundwater ecosystems host a rich and unique, but still largely unexplored and undescribed, biodiversity. Several lineages of ostracod crustaceans have subterranean representatives or are exclusively living in groundwaters. The stygobitic genus Pseudolimnocythere Klie, 1938 has a West Palearctic distribution, and includes few living and fossil species of marine origin. Through a comprehensive literature review and the description of the two new living species, Pseudolimnocythere abdita sp. nov. and Pseudolimnocythere sofiae sp. nov. , from springs in the Northern Apennines, Italy, a morphological analysis was carried out with the aim of comparing the valve morphology of living and fossil species, and to discuss previous hypotheses about time and mode of colonization of inland waters. Pseudolimnocythere species show a low variability in valve morphology, with a remarkable stasis over geological times. The distribution of extant and fossil species is consistent with a scenario of multiple and independent events of colonization of continental habitats linked to sea level variations starting from Middle Miocene in the Paratethys and, later, in the Mediterranean. The most common colonization routes of inland waters have taken place through karst formations along ancient coastlines, although we cannot exclude some minor active migration through the hyporheic zone of streams. Available distribution data suggest a poor dispersal ability of Pseudolimnocythere species after they had colonized continental waters.
... A trait dataset such as the one released in this work is a first, necessary step towards the goal of obtaining a multi-pronged prioritization that accounts for multiple biodiversity facets 97 . This is of the utmost importance given the current threats on subterranean ecosystems, and the unique conservation challenges associated with these biota [98][99][100] . ...
Article
Full-text available
Species traits are an essential currency in ecology, evolution, biogeography, and conservation biology. However, trait databases are unavailable for most organisms, especially those living in difficult-to- access habitats such as caves and other subterranean ecosystems. We compiled an expert-curated trait database for subterranean spiders in Europe using both literature data (including grey literature published in many different languages) and direct morphological measurements whenever specimens were available to us. We started by updating the checklist of European subterranean spiders, now including 512 species across 20 families, of which at least 192 have been found uniquely in subterranean habitats. For each of these species, we compiled 64 traits. The trait database encompasses morphological measures, including several traits related to subterranean adaptation, and ecological traits referring to habitat preference, dispersal, and feeding strategies. By making these data freely available, we open up opportunities for exploring different research questions, from the quantification of functional dimensions of subterranean adaptation to the study of spatial patterns in functional diversity across European caves.
... We have speculated that interspecific trait variability among groundwater invertebrates is low in most situations as a likely consequence of the intense selective pressures of the groundwater environment. Models of traits and ecosystem stability suggest that low trait variability makes groundwater ecosystems particularly vulnerable to change, with limited capacity for recovery, emphasising the recent and urgent calls for the improved recognition, conservation and management of groundwater ecosystems (Mammola et al., 2019a;Boulton, 2020;Fattorini et al., 2020;Iannella et al., 2021;Sánchez-Fernández et al., 2021;Wynne et al., 2021). ...
Article
Full-text available
Groundwater comprises the largest freshwater ecosystem on the planet. It has a distinct regime of extreme, yet stable environmental conditions that have favoured the development of similar morphological and functional traits in the resident invertebrate fauna (stygofauna). The analysis of community traits is increasingly used as an alternative to taxonomy‐based assessments of biodiversity, especially for monitoring ecosystem status and linking the functions of organisms to ecological processes, yet it has been rarely applied to stygofauna and groundwater ecosystems. In this paper, we review the variation in functional traits among the invertebrate fauna of this important ecosystem. We focus on the stygofauna and processes of alluvium and fractured rock aquifers that are typified by small voids and fissures that constrain the habitats and environmental conditions. As a first step, we compare trait variability between groundwater and surface water invertebrate communities and then examine the significance of the ranges of these traits to the vulnerability of the ecosystem to change. Fifteen potentially useful functional traits are recognised, with ten of these having narrower ranges (i.e., exhibiting fewer states, or attributes, of a particular trait) in groundwater than they do in surface water. Our synthesis suggests that the relative stability of groundwater environments has led to low trait variability. The low biomass and low reproductive rate of stygofauna suggest that recovery potential following disturbance is likely to be low. For the purposes of both improved understanding and effective management, further work is needed to document additional functional traits and their states in groundwater fauna, enabling a better understanding of the relationship between response and effect traits in these ecosystems.
... In Scotland, the reefs of Serpula vermicularis, which is presumably native there (Ten Hove and , are deemed of conservation importance and worth of restoration where degraded (Cook et al., 2021). Marifugia cavatica is one of the specialized subterranean organisms that would justify the conservation of subterranean ecosystems (Mammola et al., 2019). Protection and restoration have been also envisaged for the reefs of Hydroides dianthus in the eastern USA (Palmer et al., 2022), where the species is possibly native (Otani and Yamanishi, 2010;Sandonnini et al., 2021); however, anthropogenic transport has introduced it to Europe, West Africa, Brazil, Japan and China (Dong et al., 2018;Link et al., 2008;Otani and Yamanishi, 2010;Sun et al., 2017): that the reefs of the same species are considered a benefit in a region and a nuisance in others may seem contradictory. ...
Chapter
The Serpulidae are a large family of sedentary polychaetes, characterized by a calcareous habitation tube, which they cannot leave. The calcium carbonate tube is in the form of both aragonite and calcite, in fairly constant ratio for each taxon. Tubes are cemented firmly to any hard substrate (in only few species tubes are free). Although in the majority of the species the tubes encrust the substrate for all their length, the distal part may eventually detach and grow erectly. Certain species in dense populations build tubes vertical to the substrate in clumps and cement the tubes to each other. This gives serpulids the capability of forming reef-life structures when densely settling. Despite the relative smallness of the individual tubes (rarely longer than 15 cm and wider than 1 cm), such reef-like structures may cover tens of m², with a layer more than 1 m thick. Serpulid reefs can be divided roughly into seven groups, according to the building modality and the type of habitat they occupy: (i) pseudocolonies; (ii) littoral belts; (iii) subtidal to deep-water reefs; (iv) reefs in coastal lakes and harbours; (v) brackish water reefs; (vi) tapestries in freshwater caves; (vii) biostalactites inside marine caves. The role of serpulid reefs in the ecosystems they inhabit is multifarious and may be distinguished in functions (biomass and production, benthic pelagic coupling, resistance and resilience, reproductive and survivorship strategies, trophodynamics, bioconstruction, living space and refuge, nursery, sediment formation and retention, food for other species, carbonate deposition and storage) and services (water clearance, reef associated fishery, cultural benefits). On the other hand, many serpulids are important constituents of biological fouling, and their calcareous masses damage submerged artefacts, causing huge economic costs. Positive and negative roles of serpulid reefs need to be compared with common metrics; the overall balance, however, is still to be assessed.
... Caves are energy-poor ecosystems characterized by high humidity, low natural light, spatial confinement, climatic stability, and low biodiversity (Lamprinou et al., 2012;Culver and Pipan, 2019). For this reason, these ecosystems are very sensitive to anthropogenic pressures (Mammola et al., 2019), which can promote the development of microflora with peculiar characteristics. Some caves are used as tourist attractions or as in situ scientific laboratories. ...
Article
Full-text available
This study aimed to estimate the green formation lampenflora of “Stiffe” caves in order to evaluate their suitability as an isolation source of cyanobacteria useful for the production of polyhydroxyalkanoates (PHAs). The cave system was chosen as the sampling site due to its touristic use and the presence of high-impact illuminations. The biofilms and the mats of the illuminated walls were sampled. Samples were investigated by 16S rRNA gene analysis and culturable cyanobacteria isolation. The isolated strains were then screened for the production of PHAs under typical culturing and nutritional starvation. Cultures were checked for PHA accumulation, poly-β-hydroxybutyrate (PHB) presence (infrared spectroscopy), and pigment production. The 16S rRNA gene metabarcoding. Highlighted a considerable extent of the pressure exerted by anthropogenic activities. However, the isolation yielded eleven cyanobacteria isolates with good PHA (mainly PHB)-producing abilities and interesting pigment production rates (chlorophyll a and carotenoids). Under normal conditions (BG110), the accumulation abilities ranged from 266 to 1,152 ng mg dry biomass–1. The optimization of bioprocesses through nutritional starvation resulted in a 2.5-fold increase. Fourier transform infrared (FTIR) studies established the occurrence of PHB within PHAs extracted by cyanobacteria isolates. The comparison of results with standard strains underlined good production rates. For C2 and C8 strains, PHA accumulation rates under starvation were higher than Azospirillum brasilense and similar to Synechocystis cf. salina 192. This study broadened the knowledge of the microbial communities of mats and biofilms on the lightened walls of the caves. These findings suggested that these structures, which are common in tourist caves, could be used to isolate valuable strains before remediation measures are adopted.
... Caves are energy-poor ecosystems characterized by high humidity, low natural light, spatial confinement, climatic stability, and low biodiversity (Lamprinou et al., 2012;Culver and Pipan, 2019). For this reason, these ecosystems are very sensitive to anthropogenic pressures (Mammola et al., 2019), which can promote the development of microflora with peculiar characteristics. Some caves are used as tourist attractions or as in situ scientific laboratories. ...
... Subterranean ecosystems are likely the most widespread nonmarine environments on Earth, yet specialized subterranean organisms are among the least documented and studied groups (Mammola et al. 2019). Within stygobitic communities (i.e., restricted to aquatic subterranean habitats), snails are one of the least studied taxa and face elevated risk of extinction due to their limited geographic distribution and habitat degradation, including nutrient enrichment and groundwater contamination (Gladstone et al. 2021). ...
Article
Full-text available
The Enigmatic Cavesnail, Fontigens antroecetes (Hubricht, 1940), is a cave adapted hydrobioid snail listed as state endangered in Illinois. It is known from only one cave in Illinois, Stemler Cave, and from several caves in the eastern Ozark ecoregion of Missouri. Little is known about the snail’s reproductive habits, embryological development, or growth rates. I attempted to gain basic life history information by breeding Enigmatic Cavesnails under simulated cave conditions in the laboratory. Six adult snails were collected from Stemler Cave and held in aerated containers of cave water with one or two cobbles from the cave stream. Containers of snails were housed in incubators set at the average cave water temperature of 13 °C. The snails produced 49 embryos in captivity over the course of 34 weeks. Eggs were deposited singly, attached to the underside of rocks within small pits or crevices. Nearly 82% of embryos developed to hatching. Mean estimated development time of embryos was 70.7 days. Survival of hatchling snails was poor. Limited data available from surviving hatchling snails suggests slow growth rates. The process was replicated with nine Enigmatic Cavesnails collected from Cliff Cave in St. Louis County MO. Captive Cliff Cave snails produced 34 embryos over 46 weeks and varied from the Stemler population in their oviposition behavior, with a majority of eggs deposited on the top surface of rocks. Cliff Cave snail embryos also had longer mean estimated development times (82.17 days).
... Since the time of their exploitation, quarries have been regularly visited and transformed. In natural cavities, the vulnerability of subterranean habitats and their fauna to disturbance or pollution is generally accepted (MAMMOLA et al. 2019). We wonder whether the latter factors lead to malacofauna impoverishment through the degradation of habitats. ...
Conference Paper
Full-text available
Since the birth of biospeleology in the 19 th century, researchers have studied all groups of invertebrates living in subterranean ecosystems. Similarly to natural caves, artificial cavities provide suitable habitats for a distinct underground fauna, a big part of it still being poorly understood. Mollusc investigations in underground quarries are indeed limited because of their lower malacological biodiversity. At 10-40 m below Paris, kilometers of passageways are the result of previous exploitation of limestone, chalk and gypsum, establishing an underground network used and visited by underground workers and explorers for centuries. This work presents the first large-scale inventory of the malacological biodiversity in the Parisian underground quarries. Seven eutroglophiles, one endogean and seven trogloxenes taxa were identified. Oxychilus species' are the most common in quarries and the occurrence of Zonitoides arboreus is a new mention for the Paris region. The inventory of molluscan taxa in the Parisian quarries considers the relationship between human activities and the environment in this particular ecosystem to better understand quarries' biodiversity. The organic allochthonous material introduced by man and the molluscan carnivorous occasional feeding behaviour seem to be two conditions that favour the presence of eutroglophile mollusc. Résumé Depuis l'avènement de la biospéléologie au XIXe siècle, les chercheurs ont étudié tous les groupes d'invertébrés vivant dans les écosystèmes souterrains. Tout comme les grottes naturelles, les cavités artificielles sont des habitats propices à une faune souterraine particulière, dont une grande partie demeure mal connue. En effet, elles sont peu étudiées en raison de leur plus faible biodiversité malacologique. Entre 10 et 40 mètres de profondeur sous Paris et dans les environs, des kilomètres de galeries sont le résultat de l'exploitation antérieure du calcaire, de la craie et du gypse. Pendant des siècles, l'ensemble du réseau souterrain a été utilisé et fréquenté par les travailleurs et les explorateurs. Cette étude présente le premier inventaire à large échelle de la diversité malacologique des carrières parisiennes. Sept taxons eutroglophiles, un endogé et sept trogloxènes ont été identifiés. Les espèces d'Oxychilus sont les plus courantes dans les carrières et la présence de Zonitoides arboreus est une mention inédite pour la région parisienne. L'inventaire des mollusques dans les carrières parisiennes met en évidence l'importance d'appréhender les relations entre l'homme et l'environnement dans cet écosystème particulier pour mieux comprendre sa biodiversité. La matière organique allochtone apportée par l'homme et le régime carnivore occasionel de certains mollusques semblent être deux conditions favorables au développement des mollusques eutroglophiles.
... Many karst regions are classified as hotspots of biodiversity because of the variety of species observed in different habitats in surface water (ponds and lakes), epikarst water (cavities, fractures, and conduits), unsaturated fissures (vadose zones), and deep groundwater fractures and conduits [4]. Zagmajster et al. [5], according to the Linnean biodiversity shortfall [6,7], suggest that the majority of troglobites and stygobites have not yet been described, with a prevalence of known macroscopic subterranean animals concerning the meiofauna, i.e., with a size between 0.06 and 1 mm [8]. ...
Article
Full-text available
We predicted the global warming effects on the stygofauna of Murgia–Salento karstic groundwaters in Italy for 2050, which contribute to a biodiversity loss assessment in the climate change context. For quantitative impact estimations, we defined a local resilience score (LRS) for sampled species between 2018 and 2021. A resilience model equation of the stygobiont species conservation was obtained from a surface best-fit of the assigned LRS and the corresponding values of independent variables describing the environmental quality of monitored habitats and LRS. The principal components of the correlation between the monitored variables and LRS were obtained via factor analysis. Three-dimensional surface maps of stygofauna species resilience (SSR) were constructed to visualize and quantitatively compare the biodiversity loss of species assemblages owing to environmental and habitat quality modifications. The proposed SSR model was applied to the sampled stygofauna, and the decrease in local species resilience for 2050 was predicted. Independent variable factors were updated for 2050 to consider increases of up to 2 °C and 0.04 mS/cm in groundwater temperature and electric conductance observed for 2021. The SSR model results predicted a high impact on the resilience of Parastenocaris cf. orcina (80%), newly retrieved Crustacea Copepod Cyclopidae gen 1 sp 1, and three other stygobites (~50%). The resilience of Metacyclops stammeri had minor impacts.
Preprint
Full-text available
Protected areas are a key tool for conserving biodiversity, sustaining ecosystem services and improving human well-being. Global initiatives that aim to expand and connect protected areas generally focus on controlling ‘above ground’ impacts such as land use, overlooking the potential for human actions in adjacent areas to affect protected areas through groundwater flow. Here, we assess the potential footprint of these impacts by mapping groundwatersheds, a groundwater-modified watershed delineation. We find that most groundwatersheds (83%) of the world’s protected areas are partially unprotected and are overall only 52% protected by surface area. These findings highlight a widespread potential risk to protected areas if activities affecting groundwater are uncontrolled within their groundwatersheds, underscoring the need for groundwatershed-focused protection measures. Delineating groundwatersheds can catalyze needed discussions about protected area connectivity and effectiveness, and investments in groundwatershed conservation and management that can help ensure groundwater-dependent ecosystems are uncompromised by avoidable external underground threats.
Article
Full-text available
Context A landscape is defined as a “system of ecosystems” and this is a model in which karst areas can easily be integrated. In karst areas, much of the connectivity between the units of the landscape is underground, with aquifers and caves forming a continuous layered tissue. However, underground environments are among the least studied landscapes on Earth because of limited accessibility and the difficulty of performing surveys. Objectives The aim of this paper is to provide a conceptual framework for applying principles of landscape ecology to research on karst environments. Methods By adapting the standard patch-corridor-matrix model to a 3d model, the main issues that need to be addressed were identified. These include identifying the main morphological (surface and underground) karst features; determining the landscape structure through its features, composition, and configuration; and developing adequate indices. Results The landscape spatial structure of different karst areas influences fundamental ecological functions and biodiversity patterns. Determining how structure, biodiversity, and functions relate reveals important insights into the functioning of karst systems. Emphasizing the provisioning of ecosystem services is essential in supporting the concept that karst regions are vital for human well-being because they host valuable resources and fundamental ecosystem processes. The paper discusses how this framework helps address anthropogenic impacts and conservation issues on karst. Conclusions The potential of applying a landscape approach to karst systems lies in developing models that provide ecological information relevant to understanding karst systems and understanding their implications for natural resources management.
Article
Full-text available
This review is devoted to the problem of the development of “lampenflora” - phototrophic fouling communities in karst caves’ areas with artificial lighting used for tourism purposes. The experience of domestic and foreign colleagues helps to understand the causes of its occurrence, the conditions for the formation and development of communities; describes the mechanisms of adaptation of individual species and communities to the complex conditions of underground ecosystems. Lampenflora is not typical for the cave environment, and in this regard, the adverse consequences of its presence are found in the form of an impact on the local biota and abiotic parameters of the environment. The review compares lampenflora and natural communities of phototrophs in the entrance zones of caves illuminated by sunlight.
Article
Full-text available
1. Coastal aquifers are vital water sources for humanity. Their quality and the ecosystem services they provide depend on the integrity of their subterranean biota. However, current anthropogenic impacts such as climate change effects and coastal population growth place enormous pressure on the sustainability of these environments. 2. Despite the significance of subterranean biota to ecosystem function and the delivery of ecosystem services, stygofauna-groundwater-dwelling aquatic animals-have until recently been largely ignored in aquifer monitoring and management. This issue is of importance in both coastal and inland zones. Common threats in inland and coastal areas are water extraction, reduced recharge caused by aridification, and pollution, while, in coastal zones, additional complications arise from sea-level change and salt water ingress. 3. This review examines stygofaunal diversity, impacts, and future conservation challenges in coastal aquifers. Focussing on Australia, we provide a summary of the available data on stygofaunal communities and distributions; identify and describe potential threats to these communities across the diverse coastal regions of the continent; and propose future research priorities with the goal of facilitating the long-term preservation of these ecosystems on the Australian continent. While we focus this review on Australia, the threats and management issues discussed are relevant globally. 4. Recent subterranean studies in Australia have been primarily undertaken in inland areas, and while coastal data exist, ecological assessment of coastal subterranean ecosystems is incomplete, compromising the efficacy of conservation plans. This review indicates that the Australian continent hosts five major coastal stygofaunal biodiversity areas characterised by heterogeneous community assemblages, involving a total of 17 taxonomic groups spanning microscopic.
Preprint
Full-text available
Research and media attention is disproportionately focused on taxa and ecosystems perceived as charismatic, while other systems with high levels of endemism, are often under-protected and overlooked such as caves and subterranean ecosystems. Yet these more challenging systems are also threatened, with karsts for example losing around 6% of their area each year, highlighting the urgent need for protection, especially as up to 90% of cave endemic species may be undescribed. Bats are keystone to cave ecosystems making them potential surrogates to understand cave diversity patterns and assay conservation priorities. Almost half (48%) of known bat species use caves for parts of their life histories, with 32% endemic to a single country, and 15% currently threatened. We combine global analysis of cave bats from the IUCN with site specific analysis of 1930 bat caves from 46 countries to develop global priorities for the conservation of the most vulnerable cave ecosystems. Globally, 28% of caves showed high diversity and were highly threatened and 4% had high diversity but not currently threatened. Amongst regions, the highest concentration of conservation priority caves were in the Palearctic, and tropical regions except the Afrotropics, which requires more intensive data sampling. Our results further highlight the importance of prioritising bat caves using locally collected data, and parameter selection is optimised (i.e., appropriate landscape features and threats). Finally, to protect and conserve these ecosystems it is crucial that we identify priorities in species and habitat-level, and map vulnerable habitats with the highest biodiversity and distinctiveness.
Chapter
This article reports the current knowledge of trophic dynamics in both aquatic and terrestrial subterranean communities. Scarcity of nutrients characterizes subterranean food web interactions and drives opportunistic adaptive strategies. However, recent research contrasts the archetype of poorly structured food chains against emerging convoluted mechanisms sustaining a great range of biotic diversity and functional complexity. Novel analytical designs described in this work provide important new perspectives into the investigation of these key ecological dynamics, which, together with future advances, will enable the understanding of the immense conservational value of these often overlooked ecosystems.
Article
Full-text available
The genus Troglohyphantes Joseph, 1882 (Araneae, Linyphiidae) includes 131 species, mainly distributed across the main European mountain ranges. The Alps and the north-western Dinarides account for 66 species, most of them showing narrow or even point-like distributions. The majority of Troglohyphantes spiders dwell in subterranean habitats including caves, mines, soil litter, rocky debris and other moist and shaded retreats. Despite being intensively studied from taxonomic, ecological and biogeographic standpoints, knowledge on the status of conservation and on the potential risk of extinction of these spiders is lagging. To date, only three species have been included in the global IUCN Red List, but their status has not been updated ever since their last assessment in 1996. The aim of this contribution is to assess the Alpine and north-western Dinaric species of the genus Troglohyphantes and to re-assess the species previously evaluated, according to the last version of the IUCN Red List Categories and Criteria. Amongst the 66 species here considered, 62 had sufficient data to allow the quantification of their Extent Of Occurrence (EOO) and Area Of Occupancy (AOO). Most of the species have a narrow distribution range, with an estimated EOO < 20,000 km ² and AOO < 2,000 km ² , meeting the thresholds for the inclusion in the threatened categories. Five species have a more widespread distribution (EOO > 20,000 km ² ), extending across multiple countries. The quality of the data on distribution of four species was not sufficient to provide a reliable estimation of the distribution range. A continuing decline in EOO, AOO and habitat quality was inferred for 30 species. The majority of them were subterranean specialised species, with a reduced thermal tolerance and a low dispersal ability. Accordingly, changes in subterranean microclimatic conditions due to climate change represent a major threat for these species. Land-use change and habitat alteration were identified as additional relevant threats for several species. A considerable proportion of the species here assessed was found in protected areas and in sites of the Natura 2000 network. In addition, 14 species are formally protected by national and sub-national legislation. At present, 25 species are listed in the regional Red Lists. Long-term monitoring programmes, management plans for both the species and their habitats, expansion of the extant protected areas and designation of new ones, should be considered as the most effective approaches to species conservation.
Preprint
Full-text available
The conservation of biodiversity is a central imperative of the 21st century. Subterranean ecosystems deliver critical nature's contributions to people and harbour a broad diversity of poorly-understood specialized organisms that are of interest from both a conservation and evolutionary perspective. However, the subterranean biome is still systematically overlooked in global biodiversity targets and conservation agendas. The main objective of this study was to assess how far subterranean biodiversity is represented in protected areas (Natura 2000 and Emerald networks) in two global hotspots of subterranean biodiversity (the Pyrenees and the Alps). For this, we used the most complete databases of terrestrial subterranean biodiversity known to us, i.e., leiodids (beetles) from the Pyrenees and spiders from the Alps, and identi ed priority areas in each region using both species richness and geographic rarity patterns. Our results show the incapacity of surface protected area networks to represent subterranean fauna, as more than 70 and 90% of the identi ed priority areas (and the 40 and 22% of the species) are not effectively covered by protected areas in the Pyrenees and the Alps, respectively. These ndings call for urgent policies and would be key to developing a coherent plan for subterranean biodiversity conservation within the European Biodiversity Strategy for 2030.
Article
Full-text available
Terrestrial isopods (Oniscidea) are the most diverse group of troglobionts in caves of continental Portugal. They occur in all karst regions of Portugal, play a major role in decomposition of organic matter in caves and may act as umbrella species for the conservation of all other cave-adapted invertebrates. We present the IUCN Red List profiles for the cave-adapted terrestrial isopods from continental Portugal, based on recent distribution data from caves.
Article
Full-text available
ABSTRACT Insect pollination establishes an ecosystem service around the globe, providing compelling budgetary and creative profits along with developmental values to humans and vital eco�friendly measures for the environment. It is, therefore, essential to understand how insect pollinator populations and communities respond to rapidly changing environments if we are to maintain healthy and effective pollinator services. Although insect pollinators are known to provide ecosystem services to more than 80% of the world’s flowering plants (including cultivated crops), a steep decline (�20–40%) in their population has created an alarming situation for global biodiversity. Threats to bee populations in recent years have increased awareness about the critical role of pollinators for life on earth, as pollinators are predicted to persist only when all animal-pollinated plant species persist. Additionally, increased usage of chemical pesticides may result in the collapse of pollinators which leads to a decrease in food resource density and also facilitates the increasing isolation of natural habitats. So, to overcome pollinators’ decline, joint efforts of all stakeholders are needed to increase their numbers on the planet. We have to cut down the use of synthetic pesticides, ban highly toxic pesticides, tackle problems related to colony collapse disorder (CCD), climate change, habitat loss and provide much-needed help to the native pollinator species to revive their natural habitats. So, this paper aims to focus on appreciating the services of insect pollina�tors and rescuing them from the threats leading to their extinctions which in turn will help in enhancing global food production
Article
Full-text available
Caves are laboratories for many disciplines that work in natural sciences including mineralogy, biology, hydrogeology, and archaeology. In this study, bi-monthly samplings were carried out from three sampling locations within and around the Güvercinkaya Cave, a high-altitude cave located in northwestern Turkey, to evaluate the hydrochemical and microbiological properties and the aquatic macroinvertebrates of the cave stream. Some parameters of the water including pH, electrical conductivity, temperature, oxidation-reduction potential, and dissolved oxygen were measured in-situ, while elemental (70 in total) and ionic composition of water were analyzed in the laboratory. Microbiological analyses of the cave stream were examined through analyses of total bacteria, total coliforms, fecal coliforms, fecal Streptococcus, and Escherichia coli. According to the Piper diagram of hydrochemical data, the cave stream had mainly Ca-Mg-HCO3 character, on the other hand, the Schoeller diagram indicated a common water source in Güvercinkaya cave due to the similar components of the main ionic components of the water. As a result of microbiological analysis, fecal contamination was determined, indicating active wildlife in the cave. Additionally, several aquatic macroinvertebrates taxa, Rhynchelmis limosella, Dugesia sp., Gammarus uludagi which have non-troglobiont character were found in the cave stream. Rhynchelmis limosella detected in this study is the first record for the Turkish fauna.
Article
Full-text available
Scientists’ warnings of a climate and ecological emergency have been published recently. They have been criticised as being unattractive to non-scientists. Here, the criticisms are reviewed and comments presented. The path is long between primary research and the daily concerns of hard-to-reach people (e.g., those who are impoverished). It is enough that expert scientists express their findings accurately and intelligibly to all who are receptive. Outside the ranks of the specialist experts, there are many – intellectuals of all kinds, journalists, politicians, business people, and concerned citizens – who are well placed to contribute to the generation of a worldwide groundswell of practical action. The full range of discourse on the ecological issues is divided into four registers: used in primary research; dissemination of specialists’ thinking to non-specialists; discussion with those engaged in public affairs; and discussion with those who face obstacles to becoming engaged with the issues.
Article
Full-text available
For humans, caves are dark, moist, and often dangerous places. However, caves are also home to many animals that live strange lives: blind beetles that eat bat poop and appreciate it; tiny spiders that spin webs to catch insects, also known as flying food; and white salamanders that swim in cave ponds without getting lost, at least not all the time. Most of these animals do not have eyes—why would you need eyes when you live in the dark? Instead, they have developed other special senses to “see” in the dark. These animals look nothing like the ones you know, and many of them are endangered because they can only live in caves. Mining, pollution, and climate change are threatening cave animals and all their weirdness. Anyone can study and help them, even you!
Article
Full-text available
We report a functional switching valve within the female genitalia of the Brazilian cave insect Neotrogla. The valve complex is composed of two plate-like sclerites, a closure element, and in-and-outflow canals. Females have a penis-like intromittent organ to coercively anchor males and obtain voluminous semen. The semen is packed in a capsule, whose formation is initiated by seminal injection. It is not only used for fertilization but also consumed by the female as nutrition. The valve complex has two slots for insemination so that Neotrogla can continue mating while the first slot is occupied. In conjunction with the female penis, this switching valve is a morphological novelty enabling females to compete for seminal gifts in their nutrient-poor cave habitats through long copulation times and multiple seminal injections. The evolution of this switching valve may have been a prerequisite for the reversal of the intromittent organ in Neotrogla.
Article
Full-text available
The use of semi‐isolated habitats such as oceanic islands, lakes and mountain summits as model systems has played a crucial role in the development of evolutionary and ecological theory. Soon after the discovery of life in caves, different pioneering authors similarly recognized the great potential of these peculiar habitats as biological model systems. In their 1969 paper in Science, ‘The cave environment’, Poulson and White discussed how caves can be used as natural laboratories in which to study the underlying principles governing the dynamics of more complex environments. Together with other seminal syntheses published at the time, this work contributed to establishing the conceptual foundation for expanding the scope and relevance of cave‐based studies. Fifty years after, the aim of this review is to show why and how caves and other subterranean habitats can be used as eco‐evolutionary laboratories. Recent advances and directions in different areas are provided, encompassing community ecology, trophic‐webs and ecological networks, conservation biology, macroecology, and climate change biology. Special emphasis is given to discuss how caves are only part of the extended network of fissures and cracks that permeate most substrates, and thus their ecological role as habitat islands is critically discussed. Numerous studies have quantified the relative contribution of abiotic, biotic and historical factors in driving species distributions and community turnovers in space and time, from local to regional scales. Conversely, knowledge of macroecological patterns of subterranean organisms at a global scale remains largely elusive, due to major geographical and taxonomical biases. Also, knowledge regarding subterranean trophic webs and the effect of anthropogenic climate change on deep subterranean ecosystems is still limited. In these research fields, the extensive use of novel molecular and statistical tools may hold promise for quickly producing relevant information not accessible hitherto.
Article
Full-text available
The Second Warning to Humanity provides a clarion call for wetland researchers and practitioners given the loss and degradation of wetlands, the declining availability of fresh water, and the likely consequences of climate change. A coordinated response and approach to policies has the potential to prevent further degradation and support resilient wetlands that can provide a range of ecosystem services, including buffering wetlands from climate impacts, and avoiding major climate amplification from temperature-induced release of additional carbon dioxide and methane while addressing the causes and consequences of global climate change. The Warning to Humanity also provides an opportunity for organisations such as the Society of Wetland Scientists to raise the profile of wetlands and to initiate a discussion on how to respond and change direction from the destructive development trajectory that led to wetland loss and degradation. It also provides a signal for a reappraisal of the effectiveness of the implementation of the Ramsar Convention on Wetlands as an international mechanism for ensuring the sustainability of wetlands.
Article
Full-text available
https://rdcu.be/ZBWl (Direct link for BMC Evol Biol) Background: An essential question in evolutionary biology is whether shifts in a set of polygenic behaviors share a genetic basis across species. Such a behavioral shift is seen in the cave-dwelling Mexican tetra, Astyanax mexicanus. Relative to surface-dwelling conspecifics, cavefish do not school (asocial), are hyperactive and sleepless, adhere to a particular vibration stimulus (imbalanced attention), behave repetitively, and show elevated stress hormone levels. Interestingly, these traits largely overlap with the core symptoms of human autism spectrum disorder (ASD), raising the possibility that these behavioral traits are underpinned by a similar set of genes (i.e. a repeatedly used suite of genes).
Article
Full-text available
Landscapes in tropical regions have been greatly altered by human activities, as a product of growing demands for mineral and agricultural production, as well as those related to the generation of energy (e.g., hydroelectric, wind). In this scenario, caves have suffered several impacts, sometimes irreversible, as they are generally associated with rocks of high economic value and are closely related to epigean systems. Several indices have been proposed to guide conservation policies for the world’s speleological heritage, although few of them consider cave biodiversity as a criterion. To address this knowledge gap, we tested the applicability of four newly proposed indices to assist researchers and policy-makers select priority areas for global cave biodiversity conservation. To compare indices, we used data from 48 caves of the largest carbonate region of South America (Bambui geological group), all found within the Cerrado, a global biodiversity hotspot. Each of the four indices considered cave biodiversity as a criterion, although only three adequately evaluated this attribute. Based on results of Simões index and CCPi, which were the most appropriate in relation to indicate priority caves for biodiversity conservation in regions where the fauna and its distribution are not fully known, 15 of the 48 caves were identified as conservation priorities.
Article
Full-text available
Worldwide, there are at least 12 ILTER sites with an emphasis on karst, landforms arising from the combination of high rock solubility and well-developed solutional channel porosity underground, but the study of cave ecosystems has been largely neglected. Only two ILTER sites, both in Slovenia, are primarily caves. Caves are under-represented for several reasons, but especially because of the overall difficulty of access and the lack of a clear research agenda for cave ecosystem studies. We review several aspects of long-term studies in Postojna Planina Cave System (PPCS), proposing our approach as a model for ILTER research in caves. In PPCS, analysis of short-term temperature data shows a muted daily cycle and seasonality, and analysis of long-term temperature data shows an increase, largely the result of climate change. Changes in drip rate of epikarst aquifers above the cave are correlated with rainfall but with lags and complications resulting from differences in longer term rainfall patterns. Analysis of discharge rates indicates a rapid response to precipitation not only in the Pivka River at its sinking, but also at Unica Spring, where discharge is augmented from other parts of the aquifer, including epikarst. Quantitative analysis of the obligate epikarst-dwelling copepod community shows that, unlike most cave communities, complete sampling of the fauna is possible. Finally, organic carbon levels in PPCS indicate likely carbon limitation in the system. These five factors (temperature, drip rate, river discharge, epikarst copepod fauna, and organic carbon) are the appropriate variables for capturing the essential long-term trends in cave ecosystems and their causes.
Article
Full-text available
Periodic food shortages are a major challenge faced by organisms in natural habitats. Cave-dwelling animals must withstand long periods of nutrient deprivation, as-in the absence of photosynthesis-caves depend on external energy sources such as seasonal floods. Here we show that cave-adapted populations of the Mexican tetra, Astyanax mexicanus, have dysregulated blood glucose homeostasis and are insulin-resistant compared to river-adapted populations. We found that multiple cave populations carry a mutation in the insulin receptor that leads to decreased insulin binding in vitro and contributes to hyperglycaemia. Hybrid fish from surface-cave crosses carrying this mutation weigh more than non-carriers, and zebrafish genetically engineered to carry the mutation have increased body weight and insulin resistance. Higher body weight may be advantageous in caves as a strategy to cope with an infrequent food supply. In humans, the identical mutation in the insulin receptor leads to a severe form of insulin resistance and reduced lifespan. However, cavefish have a similar lifespan to surface fish and do not accumulate the advanced glycation end-products in the blood that are typically associated with the progression of diabetes-associated pathologies. Our findings suggest that diminished insulin signalling is beneficial in a nutrient-limited environment and that cavefish may have acquired compensatory mechanisms that enable them to circumvent the typical negative effects associated with failure to regulate blood glucose levels.
Article
Full-text available
Viewpoint article World Scientists’ Warning to Humanity: A Second Notice WILLIAM J. RIPPLE, CHRISTOPHER WOLF, THOMAS M. NEWSOME, MAURO GALETTI, MOHAMMED ALAMGIR, EILEEN CRIST, MAHMOUD I. MAHMOUD, WILLIAM F. LAURANCE, and 15,364 scientist signatories from 184 countries. http://scientistswarning.forestry.oregonstate.edu/signatories
Article
Full-text available
Bats are unique among mammals, possessing some of the rarest mammalian adaptations, including true self-powered flight, laryngeal echolocation, exceptional longevity, unique immunity, contracted genomes, and vocal learning. They provide key ecosystem services, pollinating tropical plants, dispersing seeds, and controlling insect pest populations, thus driving healthy ecosystems. They account for more than 20% of all living mammalian diversity, and their crown-group evolutionary history dates back to the Eocene. Despite their great numbers and diversity, many species are threatened and endangered. Here we announce Bat1K, an initiative to sequence the genomes of all living bat species (n∼1,300) to chromosome-level assembly. The Bat1K genome consortium unites bat biologists (>132 members as of writing), computational scientists, conservation organizations, genome technologists, and any interested individuals committed to a better understanding of the genetic and evolutionary mechanisms that underlie the unique adaptations of bats. Our aim is to catalog the unique genetic diversity present in all living bats to better understand the molecular basis of their unique adaptations; uncover their evolutionary history; link genotype with phenotype; and ultimately better understand, promote, and conserve bats. Here we review the unique adaptations of bats and highlight how chromosome-level genome assemblies can uncover the molecular basis of these traits. We present a novel sequencing and assembly strategy and review the striking societal and scientific benefits that will result from the Bat1K initiative. Expected final online publication date for the Annual Review of Animal Biosciences Volume 6 is February 15, 2018. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
Article
Full-text available
Twenty-five years ago, the Union of Concerned Scientists and more than 1700 independent scientists, including the majority of living Nobel laureates in the sciences, penned the 1992 “World Scientists’ Warning to Humanity” (see supplemental file S1). These concerned professionals called on humankind to curtail environmental destruction and cautioned that “a great change in our stewardship of the Earth and the life on it is required, if vast human misery is to be avoided.” In their manifesto, they showed that humans were on a collision course with the natural world. They expressed concern about current, impending, or potential damage on planet Earth involving ozone depletion, freshwater availability, marine life depletion, ocean dead zones, forest loss, biodiversity destruction, climate change, and continued human population growth. They proclaimed that fundamental changes were urgently needed to avoid the consequences our present course would bring. The authors of the 1992 declaration feared that humanity was pushing Earth's ecosystems beyond their capacities to support the web of life. They described how we are fast approaching many of the limits of what the ­biosphere can tolerate ­without ­substantial and irreversible harm. The scientists pleaded that we stabilize the human population, describing how our large numbers—swelled by another 2 billion people since 1992, a 35 percent increase—exert stresses on Earth that can overwhelm other efforts to realize a sustainable future (Crist et al. 2017). They implored that we cut greenhouse gas (GHG) emissions and phase out fossil fuels, reduce deforestation, and reverse the trend of collapsing biodiversity.
Article
Full-text available
Over the last two decades there has been an exponential increase in the use of correlative species distribution models (SDMs) to address a variety of topics in ecology, biogeography, evolution, and conservation biology. Conversely, the use of these statistical methods to study the potential distribution of subterranean organisms has lagged behind, relative to their above-ground (epigean) counterparts. The reason for this is possibly related to a number of peculiarities of subterranean systems, which pose important limits, but also opportunities, for these correlative models. The aim of this forum is to explore the caveats that need to be made when generalizing these statistical techniques to caves and other subterranean habitats. We focus on the typical bias in spatial datasets of cave-dwelling species, and provide advice for selecting the model calibration area. In parallel, we discuss the potential use of different large scale surface variables to represent the subterranean condition. A more widespread adoption of these statistical techniques in subterranean biology is highly attractive and has great potential in broadening our knowledge on a variety of ecological topics, especially in the fields of climate change and biodiversity conservation. Their use would especially benefit the study of the biogeographic patterns of subterranean fauna and the impact of past and future climate change on subterranean ecosystems.
Article
Full-text available
Molecular taxonomy often uncovers cryptic species, reminding us that taxonomic incompleteness is even more severe than previous thought. The importance of cryptic species for conservation is poorly understood. Although some cryptic species may be seriously threatened or otherwise important, they are rarely included in conservation programs as most of them remain undescribed. We analysed the importance of cryptic species in conservation by scrutinizing the South European cryptic complex of the subterranean amphipod Niphargus stygius sensu lato. Using uni- and multilocus delineation methods we show that it consists of 15 parapatric and sympatric species, which we describe using molecular diagnoses. The new species are not mere “taxonomic inflation” as they originate from several distinct branches within the genus and coexist with no evidence of lineage sharing. They are as evolutionarily distinct as average nominal species of the same genus. Ignoring these cryptic species will underestimate the number of subterranean endemics in Slovenia by 12 and in Croatia by four species, although alpha diversity of single caves remains unchanged. The new taxonomy renders national Red Lists largely obsolete, as they list mostly large-ranged species but omit critically endangered single-site endemics. Formal naming of cryptic species is critical for them to be included in conservation policies and faunal listings.
Article
Full-text available
The genus Agave is one of the most diverse and rich groups of plants of Mexico. Mexican people have developed several technologies to extract products from Agave, and for many years they have consumed five different alcoholic beverages derived from Agave: Tequila, Mezcal, Bacanora, Raicilla, and Pulque. Additionally, Agave has coevolved with nectar-feeding bats, and in several cases, bats play the main role as functional pollinators in this ecological relationship. But with growth in the demand of agave derived products, management practices have reduced dependence on bat pollination, using instead clonal shoots to replant fields and harvesting plants before flowering, thereby negatively affecting both bats (by decreasing food availability) and agaves (by lowering their genetic diversity). We explore the possibility that bat-friendly practices may be incorporated into the production system. We compiled data about the pollination biology of Agave to infer how many bats could use the available resources, if Mezcal and Tequila producers allowed 5-10% of agave crop inflorescences to flower based on a linear projection using Agave angustifolia (a sister group of A. tequilana). If only 5% of the plants in one hectare were allowed to flower (approximately 222 individuals), then, depending on nectar concentration and total volume, a minimum of 89 individual bats could feed every night during flowering period. This means that allowing 5% of the current total population of A. tequilana reproductive agaves to flower could feed a total of 2,336,250 nectar feeding bats per month.
Chapter
Full-text available
Caves and other subterranean sites such as mines are critical to the survival of hundreds of bat species worldwide, since they often provide shelter for most of a nation’s bat fauna. In the temperate zone, caves provide roosts for hibernation and for some species, breeding in summer, whereas in warmer regions, they support high species richness year round and enormous colonies that maintain substantial ecosystem services. Due to the solubility of the substrate, the highest densities of caves occur in karst landscapes. Given their importance for bats, relatively few studies have investigated factors involved in cave selection, although current evidence suggests that the density and size of caves are the best predictors of species diversity and population sizes. Thermal preferences have been established for some cave-dwelling species as well as their vulnerability to disturbance, particularly during hibernation and reproduction. Growth in limestone quarrying and cave tourism industries worldwide severely threatens cave-dwelling bats, in addition to loss of foraging habitat, hunting for bushmeat, incidental disturbance and disruptive guano harvesting. Apparent declines of cave bats in Europe and North America also pose serious concerns, as do global climate change predictions. The main conservation response to threats to cave bats in these continents has been gating, but this remains relatively untested as a means of protecting colonies in other regions. Research on sustainable harvesting of bats as bushmeat and their responses to different types of human disturbance at caves and loss of surrounding foraging habitats is required. More caves of outstanding importance for bats at national and international levels also require protection.
Article
Full-text available
Ecologists and evolutionary biologists are increasingly using big-data approaches to tackle questions at large spatial, taxonomic, and temporal scales. However, despite recent efforts to gather two centuries of biodiversity inventories into comprehensive databases, many crucial research questions remain unanswered. Here, we update the concept of knowledge shortfalls and review the tradeoffs between generality and uncertainty. We present seven key shortfalls of current biodiversity data. Four previously proposed shortfalls pinpoint knowledge gaps for species taxonomy (Linnean), distribution (Wallacean), abundance (Prestonian), and evolutionary patterns (Darwinian). We also redefine the Hutchinsonian shortfall to apply to the abiotic tolerances of species and propose new shortfalls relating to limited knowledge of species traits (Raunkiæran) and biotic interactions (Eltonian). We conclude with a general framework for the combined impacts and consequences of shortfalls of large-scale biodiversity knowledge for evolutionary and ecological research and consider ways of overcoming the seven shortfalls and dealing with the uncertainty they generate.
Article
Full-text available
The disoovery of the Ursilor (Bear's) Cave (Apuseni Mountains, Romania) in the 70th was followed by climatic and biological studies, before its opening as a show cave. Twenty years later these researches were undertaken again The tourism has impacted on the populations of two endemic cave beetles: Pholeuon leptoderum and Drirneotus n. sp. The first one became extremely rare in the tourist part of the cave. Drimeotus has adapted different dynamics during the year in the two parts of the cave; near the tourist path the populations evolves depending on the affluence of the tourists in the summer months, and in the protected part has kept the same pattern as in the early studies.
Article
Full-text available
Caves are considered buffered environments in terms of their ability to sustain near-constant microclimatic conditions. However, cave entrance environments are expected to respond rapidly to changing conditions on the surface. Our study documents an assemblage of endemic arthropods that have persisted in Rapa Nui caves, despite a catastrophic ecological shift, overgrazing, and surface ecosystems dominated by invasive species. We discovered eight previously unknown endemic species now restricted to caves—a large contribution to the island's natural history, given its severely depauperate native fauna. Two additional species, identified from a small number of South Pacific islands, probably arrived with early Polynesian colonizers. All of these animals are considered disturbance relicts—species whose distributions are now limited to areas that experienced minimal historical human disturbance. Extinction debts and the interaction of global climate change and invasive species are likely to present an uncertain future for these endemic cavernicoles.
Article
Full-text available
Despite Brazil’s role as a global environmental leader, most of its megadiverse and unique biomes are at risk (Ferreira et al. 2014). Currently, the Brazilian Congress is debating political proposals (e.g. PL 3682/2012) for the expansion of mining and hydropower generation activities across the country, including areas within the system of Brazil’s protected areas. Ferreira et al. (2014) showed a detrimental perspective for the integrity of Brazil’s biomes in a scenario where the ongoing proposals are approved. Areas of mining interest overlap 20 % of Brazil’s protected areas and indigenous lands, threatening the coverage integrity of the largest system of protected areas worldwide. Although there are compelling arguments, the authors argue that Brazil’s newly elected government should maintain a consistent stance with its influential role as a leader in the conservation of natural areas by expanding the protected areas system and reducing deforestation.Although we fully agree with the ...
Article
Full-text available
BackgroundA key question in evolutionary biology is the relationship between species traits and their habitats. Caves offer an ideal model to test the adjustment of species to their surrounding temperature, as they provide homogeneous and simple environments. We compared two species living under different thermal conditions within a lineage of Pyrenean beetles highly modified for the subterranean life since the Miocene. One, Troglocharinus fonti, is found in caves at 4-11°C in the ancestral Pyrenean range. The second, T. ferreri, inhabits the coastal area of Catalonia since the early Pliocene, and lives at 14-16°C.ResultsWe found no differences in their short term upper thermal limit (ca. 50°C), similar to that of most organisms, or their lower thermal limit (ca. -2.5°C), higher than for most temperate insects and suggesting the absence of cryoprotectants. In longer term tests (7 days) survival between 6-20°C was almost 100% for both species plus two outgroups of the same lineage, but all four died between 23-25°C, without significant differences between them.Conclusions Our results suggest that species in this lineage have lost some of the thermoregulatory mechanisms common in temperate insects, as their inferred default tolerance range is larger than the thermal variation experienced through their whole evolutionary history.
Article
Full-text available
Cave environments are characterized by possessing specialized fauna living in high environmental stability with limited food conditions. These fauna are highly vulnerable to impacts, because this condition can frequently be easily altered. Moreover, environmental determinants of the biodiversity patterns of caves remain poorly understood and protected. Therefore, the main goal of this work is to propose a cave conservation priority index (CCPi) for a rapid assessment for troglobiotic and troglophile protection. Furthermore, the troglobiotic diversity, distribution and threats have been mapped in the Brazilian Atlantic forest. To propose the CCPi, the human impacts and richness of troglobiotic and troglophile species of 100 caves were associated. Data related to troglomorphic/troglobiotic fauna from another 200 caves were used to map the troglobiotic diversity and distribution. The CCPi reveals extremely high conservation priority for 15 % of the caves, high for 36 % and average for 46 % of the caves. Fourteen caves with extremely high priorities should have urgent conservation and management actions. The geographical distribution of the 221 known troglobiotic/troglomorphic species allowed us to select 19 karst areas that need conservation actions. Seven areas were considered to have urgent priority for conservation actions. The two richest areas correspond to the "iron quadrangle" with iron ore caves (67 spp.) and the "Açungui limestone group" (56 spp.). Both areas have several caves and are important aquifers. The use of the CCPi can prevent future losses because it helps assessors to select caves with priorities for conservation which should receive emergency attention in relation to protection, management and conservation actions.
Article
Full-text available
We investigated the upper and lower vertical limits of the distribution of inhabitants of the most abundant freshwater habitat—groundwater. Distribution in photic habitats is limited by competition, predation, and risks of exposure to ultraviolet radiation. Nonetheless, a number of eyeless, depigmented subterranean species occur in twilight habitats, taking advantage of the higher food resources available and modulating their distribution by photophobic behavior. We argue that the upper boundary is an interesting system in the study of classic ecological and evolutionary questions. The lower boundary of the distribution of groundwater species (approximately 2000—4000 meters) is likely controlled by physicochemical parameters, including temperature, pressure, and oxygen. The lower boundary warrants further research, and it is one of the most poorly explored areas of the biosphere.
Article
Full-text available
Most organisms are able to survive shorter or longer exposure to sub-zero temperatures. Hypothetically, trogloxenes characterized as not adapted, and troglophiles as not completely adapted to thermally stable subterranean environment, have retained or partially retained their ability to withstand freezing, while most troglobionts have not. We tested this hypothesis experimentally on 37 species inhabiting caves in Slovenia, analyzing their lower lethal temperatures in summer and winter, or for one season, if the species was not present in caves during both seasons. Specimens were exposed for 12 hrs to 1°C-stepwise descending temperatures with 48 hr breaks. In general, the resistance to freezing was in agreement with the hypothesis, decreasing from trogloxenes over troglophiles to troglobionts. However, weak resistance was preserved in nearly all troglobionts, which responded in two ways. One group, withstanding freezing to a limited degree, and increasing freezing tolerance in winter, belong to the troglobionts inhabiting the superficial subterranean habitats. The other group, which equally withstand freezing in summer and winter, inhabit deep subterranean or other thermally buffered subterranean habitats. Data on cold resistance can thus serve as an efficient additional measure of adaptation to particular hypogean environments.