The Biology of Caves and Other Subterranean Habitats
Abstract
Caves and other subterranean habitats with their often strange (even bizarre) inhabitants have long been objects of fascination, curiosity, and debate. The question of how such organisms have evolved, and the relative roles of natural selection and genetic drift, has engaged subterranean biologists for decades. Indeed, these studies continue to inform the general theory of adaptation and evolution. Subterranean ecosystems generally exhibit little or no primary productivity and, as extreme ecosystems, provide general insights into ecosystem function. The Biology of Caves and other Subterranean Habitats offers a concise but comprehensive introduction to cave ecology and evolution. Whilst there is an emphasis on biological processes occurring in these unique environments, conservation and management aspects are also considered. The monograph includes a global range of examples from more than 25 countries, and case studies from both caves and non-cave subterranean habitats; it also provides a clear explanation of specialized terms used by speleologists. This accessible text will appeal to researchers new to the field and to the many professional ecologists and conservation practitioners requiring a concise but authoritative overview. Its engaging style will also make it suitable for undergraduate and graduate students taking courses in cave and subterranean biology. Its more than 650 references, 150 of which are new since the first edition, provide many entry points to the research literature.
... Slovenia is home to a remarkable diversity of subterranean species known as troglobionts, making it a hotspot for subterranean biodiversity [1,2]. These subterranean species often have very restricted habitats, frequently existing in only small areas or even a single location [3]. ...
... Current research on terrestrial subterranean karst systems primarily focuses on several key areas: defining hydrogeological characteristics [6,12,16]; identifying drivers of species richness [1,2]; assessing groundwater quality by measuring nitrogen concentrations and other contaminants, as well as determining pollution sources [7,15,18,[20][21][22][23]; examining the toxicity of nitrogen to common organisms [9]; and using SWAT modelling to identify critical source areas and address non-point source pollution [19]. One of the most comprehensive works on agriculture and karst systems is the book Karst Management, published in 2011 [17], which discusses various agricultural activities that may impact soil and water in karst regions. ...
... Our analysis focused on 11 subbasins within the Dobličica River area (Figure 1). This focus was necessitated by the fact that only the hinterland of the selected subbasins (1,8,9,10,11,14,16,17,18,19,20) has a significant impact on the habitat of the black olm. ...
The black olm (Proteus anguinus parkelj Sket & Arntzen) is an endemic species found exclusively in the Dobličica River subterranean water systems of the Dinaric karst in southern Slovenia. These unique habitats are vulnerable to contamination due to rapid water flow, primarily from nitrates from agricultural fertilisers and untreated urban wastewater. The safe limit of nitrate concentration for olms is 9.2 mg NO3 − /L, yet measurements in karst springs have shown levels ranging from 3 mg to over 20 mg NO3 − /L. The SWAT modelling tool assessed agri-environmental and land use scenarios for their impact on nitrate leaching. Using the model, we identified hotspots with high nitrogen leaching potential that require immediate attention and implementation of better agricultural practices for fertiliser use. For these hotspots, the most effective approach combines scenarios of cover crops (R2), reduced fertilisation (R3), crop rotation (R4), and conversion of cropland to grassland (E2, E4, E5), potentially decreasing nitrate leaching by up to 60%. Implementing the best scenarios is expected to reduce nitrogen levels below the limit value of 9.2 mg NO3 − /L, essential for maintaining the black olm habitat.
... Despite the possibility of exploiting subterranean shelters, climate change remains one of the main threats to Speleomantes (Rondinini, Battistoni, and Teofili 2022). First, external climate is intimately connected with the inner microclimatic conditions of subterranean environments, where it mostly affects the areas close to the entrance (Badino 2004;Lunghi, Manenti, and Ficetola 2015;Culver and Pipan 2019). Thus, the increase of temperature and dryness at the surface caused by global warming will also be replicated in the shallowest areas of subterranean environments (Mammola et al. 2019), reducing the availability of suitable habitats and therefore representing a threat to many cave-dwelling species (Rizzo et al. 2015;Sánchez-Fernández et al. 2016;Mammola, Goodacre, and Isaia 2018). ...
... Thus, the increase of temperature and dryness at the surface caused by global warming will also be replicated in the shallowest areas of subterranean environments (Mammola et al. 2019), reducing the availability of suitable habitats and therefore representing a threat to many cave-dwelling species (Rizzo et al. 2015;Sánchez-Fernández et al. 2016;Mammola, Goodacre, and Isaia 2018). Secondly, as facultative cave-dwellers (so-called troglophiles; Howarth and Moldovan 2018), Speleomantes often forage on the surface, where prey is more abundant Culver and Pipan 2019). Climate change may reduce prey availability in both surface and subterranean habitats, confining salamanders to deep, resource-poor spaces (Salvidio et al. 1994;. ...
... Indeed, individuals probably will not disperse fast enough to reach new future suitable areas, thus remaining trapped in their subterranean refugia. Here, climate change will progressively confine them to the deepest oligotrophic areas where trophic supply is probably insufficient for the sustainment of the entire population Culver and Pipan 2019;Mammola et al. 2019). Additionally, natural barriers may play an important role in limiting Speleomantes dispersal. ...
Aim: Amphibians are particularly sensitive to rapid climatic shifts, due to their eco-physiology, life history traits and high frequency of narrowly distributed species. The genus Speleomantes encompasses the only extant Western Palearctic plethodontids, with three species occurring in peninsular Italy and the remaining five endemic to Sardinia Island. Given the restricted ranges of Speleomantes species and their vulnerability to environmental change, we implemented Ecological Niche Models (ENMs) to estimate the likely impacts of various global warming scenarios on the extent and geographical location of climatically suitable areas.
Time Period: Current, with ENMs projected to 2030, 2050 and 2070 under alternative Shared Socioeconomic Pathways.
Location: Italian Peninsula and Sardinia Island.
Major Taxa Studied: Speleomantes Dubois 1984 (Caudata: Plethodontidae).
Methods: Ensembles of ENMs were fitted for each Speleomantes species, using the 'biomod2' modelling platform in R environment. Then, post-modelling analyses were applied in GIS environment to highlight: (i) the primary geographic direction of predicted suitability shifts for each species and (ii) the proportion of stable, gained and lost suitable areas for each genetic lineage of the single species.
Results: We found a noticeable shrinking of suitable areas for all Speleomantes species, being particularly extensive under 'business-as-usual' scenarios for the Sardinian ones. Moreover, core suitable areas were predicted to shift for most species and suitability losses emerged to differently affect distinct genetic lineages, posing additional challenges for designing effective conservation measures.
Main Conclusions: The predicted shrinkage and shifting of climatically suitable areas for most Speleomantes species point to the urgency of evaluating in due time alternative conservation strategies for these endemic urodeles, to prevent losses of taxonomic and genetic diversity. Our modelling framework may be applied to other species with similar traits (e.g., low dispersal ability and narrow environmental niche breadth) to predict climate-induced range contractions or shifts, using the gained information to optimise conservation outcomes.
... Furthermore, the biodiversity within the epikarst is important, as studies of epikarst fauna have shown that small voids and fissures are highly diverse [6,7]. Even at a large scale, and where intensive sampling efforts were undertaken [8], the epikarst fauna was rich in species, but yet undersampled [9]. ...
... Even at a large scale, and where intensive sampling efforts were undertaken [8], the epikarst fauna was rich in species, but yet undersampled [9]. Based on sampling at different depths in caves, including deep sites and sites close to or in epikarst, Culver and Pipan [6] suggest that epikarst ...
... Six copepod species from the group Harpacticoida were collected (Speocyclops infernus, Bryocamptus balcanicus, Elaphoidella cvetkae, Moraria poppei, Morariopsis scotenophila, and Parastenocrais nolli alpina), as well as one amphipod species from the genus Niphargus; all but one were stygobionts. These are all species previously reported from the cave system [6]. While copepods were numerically common, amphipods were not and only three individuals were found (see Table 2). ...
Karst areas are characteristic landscapes formed by the dissolution of soluble rocks, whose hydrology is largely influenced by rapid infiltration through the karst massif. These areas are often hotspots of biodiversity, especially for epikarst and cave fauna. The epikarst, the uppermost layer of the unsaturated zone, plays a crucial role in regulating water flow in karst aquifers. The aim of this study was to investigate the extent of microplastic (MP) pollution, its relationship with precipitation and its correlation with copepod populations in karst areas. The study was conducted between April 2021 and October 2022 in the Postojna–Planina karst area in SW Slovenia at surface and underground sampling sites to determine the pathways of MP pollution from the surface to the depth of the karst massif. The results indicate that heavier rainfall flushes out more MP contaminants from the surface and epikarst environment. The transport dynamics of MP pollution are similar to the dynamics of copepods, which are the baseline organisms for the food chain in caves. One MP sample contained only polyamide particles, which could indicate clothing as a possible source of pollution, but the results are inconclusive. With this study, we provide the first insight into the transport of MP pollution from the surface environment to deeper karst massifs.
... The predominant focus of historical research has been on species descriptions, with only a limited number of recent publications delving into cave ecology, community, and conservation (Togouet et al. 2009;Kayo et al. 2012;Vandel et al. 2013;Deharveng and Bedos 2018;Du Preez et al. 2023). These remarkable species are uniquely adapted to thrive in challenging conditions, rendering them particularly intriguing subjects for researchers studying evolutionary biology and biogeography (Trontelj et al. 2012;Culver and Pipan 2019;Pipan et al. 2020;Sánchez-Fernández et al. 2021). ...
... Species restricted to subterranean environments, such as troglobionts and stygobionts, cannot complete their life cycles in aboveground habitats (Sket 2008). One way to identify these subterranean-adapted species is by recognizing specific morphological traits, known as troglomorphisms, commonly observed in troglobitic and stygobitic fauna (Sket 2008;Culver and Pipan 2019). These adaptations include reduced eyes, pigmentation, and hypertrophy of nonvisual sensory structures and locomotor appendages. ...
... Cave environments' extreme isolation and distinct conditions render many species rare and vulnerable. Numerous obligate cave-dwelling species are deemed threatened or endangered at regional or global scales, primarily due to their confinement to small geographical areas (Culver and Pipan 2019). ...
Aziza cave, which is also known as kef Aziza or Tazouguert cave, represents an important and large karstic system that consists of more than 3.5 km of surveyed galleries, standing as the fifth most extensive cave system in Morocco and one of the ten largest in North Africa. This study unveils Aziza cave as an important spot of subterranean diversity in Africa. Here, we provide the first checklist of subterranean fauna in this cave, with 26 taxa, comprising 22 troglobiotic and 4 stygobiotic species. Of this total, eight species still require further confirmation of their status. The richest taxa include Coleoptera (5 species), Araneae (4 species), Entomobryomorpha (3 species), and Isopoda (2 species). However, it is noteworthy that only around 34.6% of the cave-restricted species found in the cave have been formally described to date. Additionally, the biodiversity of large system areas remains to be discovered as these areas need to be further explored. Furthermore, this paper highlights the broader conservation challenges faced by subterranean habitats in Morocco, particularly considering human-induced impacts on these remarkable ecosystems. We aim to draw attention to the crucial ecological role of subterranean environments and their extraordinary biological diversity. By doing so, we aim to inspire increased research and conservation initiatives, not just in this area but across Africa.
... Caves are distinguished by their unique environmental conditions, notably the absence of light, which precludes the presence of photosynthetic organisms. This lack of primary production results in oligotrophic conditions, characterized by limited nutrient availability, compared with epigean habitats (Culver & Pipan, 2019). Caves typically have lower luminosity and more stable temperatures and humidity than surface environments. ...
... Caves typically have lower luminosity and more stable temperatures and humidity than surface environments. In response to these challenges, cave-dwelling organisms have evolved specialized traits to thrive in these unique habitats (Culver & Pipan, 2019;Mammola et al., 2020). ...
... These adaptations include changes in body structure, sensory systems, feeding strategies, locomotion, and more. Through these morphological, physiological, and behavioral adjustments, organisms effectively navigate and exploit specific cave microhabitats, thereby reducing competition and niche overlap with other species (Culver & Pipan, 2019). By occupying different microhabitats and using distinct resources, cave-dwelling organisms can coexist and compete for resources within subterranean environments (Mammola et al., 2020). ...
Stable temperature, humidity, and oligotrophy may be essential in microhabitat selection, shaping niche overlap and species coexistence within cave ecosystems. To explore this hypothesis, we assessed the distribution and microhabitat preferences of four non‐obligate cave‐dwelling spiders: Loxosceles anomala , Scytodes itapevi , Philoponella vittata , and the exotic Oecobius navus . Using 1‐m ² quadrats, we surveyed spider distribution and abundance in a limestone cave, considering environmental conditions and prey availability during wet and dry periods. Significant differences were observed between moist and dry periods regarding light intensity, wind speed, air temperature and moisture, potential prey, and the abundance of L. anomala . The exotic spider O. navus was more specialist, preferred brighter habitats, and showed lower niche overlap. Conversely, the three native species displayed greater niche overlap and preferred darker and more humid environments. Spatial disparities, rather than temporal variation, in species distribution underscored the influence of cave habitat conditions on species segregation. Furthermore, our findings highlighted the limited ability of the non‐native O. navus to occupy more illuminated cave areas, thereby potentially safeguarding these habitats against biological invasion by this species.
... Hydrological subterranean environments are primarily influenced by precipitation and surface watercourses, which infiltrate the underground system via gravity through small pores and fractures, forming complex pathways known as epikarst (Ford and Williams 2007;Bonacci et al. 2009;Audra and Palmer 2011;Culver and Pipan 2019). These flowing waters can penetrate the phreatic zone (i.e., saturated zone), either fully or partially replenishing subterranean water bodies. ...
... These flowing waters can penetrate the phreatic zone (i.e., saturated zone), either fully or partially replenishing subterranean water bodies. Such bodies may also originate from phreatic conduits regulated by elevation or pressure differentials (Gibert et al. 1994;Williams 2008;White 2019;Audra and Palmer 2011;Culver and Pipan 2019). Within caves, a variety of aquatic habitats, including those with lentic (e.g., lakes and pools) and lotic (e.g., rivers) features, are formed under the influence of both saturated and/or unsaturated zones (Bonacci et al. 2009;Souza-Silva et al. 2012;Pipan et al. 2020). ...
... They are considered to be a reliable indicator of water quality and environmental shifts, as both biotic and abiotic factors can influence the structure and composition of zooplankton communities (Deharveng and Bedos 2019;Shen and Liu 2021). Groundwater zooplankton have been extensively studied in Europe, Asia, and Oceania with research predominantly focused on the taxonomy and ecology of stygobiotic zooplankton, especially Copepoda (De Laurentiis et al. 1999;Pipan and Culver 2007;Galassi et al. 2009a, b;Karavonik et al. 2015, Hose et al. 2016, Cavite et al. 2017Culver and Pipan 2019;Bozkurt 2022;Guo et al. 2022). However, in Brazil, understanding of the diversity and ecology of subterranean zooplankton remains relatively limited, with only a handful of studies conducted thus far (e.g., Simões et al. 2013;Souza-Silva et al. 2017). ...
The subterranean environments of the Caatinga drylands (Brazilian biome) host a diverse array of aquatic systems and diversity, comprising both lentic and lotic ecosystems. Species diversity in these environmnts has been overlooked and remains poorly understood, especially regarding zooplankton. We studied the species richness and composition of zooplankton living in groundwater of the Caatinga drylands and explored how they vary in relation to habitat type (lentic and lotic) by testing two hypotheses: (1) species richness, gamma diversity, and zooplankton density are higher in lentic environments and (2) zooplankton species composition differs between habitat types. We sampled 12 lentic and lotic groundwater environments quarterly for 2 years, and identified 100 species of zooplankton in groundwater environments, including testate amoebae (50 species), Rotifera (25 species), Copepoda (16 species), and Cladocera (9 species). Rotifer species richness and copepod density differed between lentic and lotic habitats, as did zooplankton composition. Additionally, each habitat was found to harbor distinct indicator species based on their ecology and morphological characteristics. These findings contribute to current understanding of groundwater biodiversity and ecology, providing support for freshwater and speleological management and conservation programs.
... Cave-dwelling bats spend roughly half their life cycles roosting underground for resting, mating, and raising offspring [6,7], and the other half searching for and gathering resources during night foraging bouts in the external landscapes [8]. The energetic requirements of bats for their daily activities and the heterogeneity of the landscapes available for their foraging movements are among the main limiting factors shaping these nightly outside hunts driven by habitat selection [9]. ...
... Although caves have usually been the focus of conservation efforts for the cave-related biota [10], the surrounding landscapes can encompass most of the home range of a bat. These interplays between the underground and the surrounding landscapes shapes overall biodiversity patterns in subterranean environments worldwide [7] and are pivotal to maintain the diversity and abundance of cave bats over time [11]. The same holds true for subterranean invertebrate assemblages, which are largely sustained by the guano deposition made by bats [8,12]. ...
Individual movements of bats are triggered by their life requirements, limited by their recognition of the environment and risks of moving, and mediated by habitat selection. Mining adds fragmentation and heterogeneity to landscapes, with poorly understood consequences to the life activities of the bats. Cave dwelling bats spend most of their life cycles within caves, and as they constantly forage in external landscapes, their contribution in the input of organic matter to the caves is of paramount importance to the subterranean biodiversity. We investigated habitat selection by cave bats in a mosaic of Amazonian forests, iron-rich savannas and iron-caves, studying the movements of the aerial insect-catcher Furipterus horrens (Furipteridae) and of the foliage gleaning bat Lonchorhina aurita (Phyllostomidae), both cave-dependent in the area. We used radio telemetry to assess their use of habitats, under the prediction that these species prefer preserved habitats for their foraging bouts, avoiding human-modified habitats. We also anticipated larger areas and commuting distances for both species when foraging in disturbed landscapes. Thirty-one bats were monitored in conserved habitats, mining sites and pastures resulting in an average range of 415.1 ± 837.4 ha and average commuting distances of 2163 ± 2453 m (F. horrens) and 681.1 ± 1101 ha and 2781.5 ± 2333 m (L. aurita). Our results suggest that F. horrens are open space foragers, frequently recorded in cangas, and L. aurita are forest foragers that use all habitats proportionally to their availability in the landscape. We detected landscape-related effects mediated by seasonal variation in the maximum commuting distance performed by both species. These are the first radiotelemetry data available for F. horrens and L. aurita, delivering original baselines to understand their movement behaviors. This information sheds light into constraints related to the optimal and adjusted biological cycles of these two species and their range shifts under scenarios of disturbance and may subsidize programs for their conservation.
... Cave environments in the world support astonishing endemic biodiversity that have long been in a focus of scientific inquiry. Spatial isolation, together with absence of light, stable microclimates and food scarcity played important roles in cave fauna evolution [1]. The absence of photoautotrophic organisms such as plants, algae and cyanobacteria Ángel García-Bodelón and Najla Baković should be considered as joint first authors. ...
... These characteristics make them a perfectly suited model group to study microbial evolution in caves. Moreover, the stable year-long environmental conditions within the cave favour K-selected organisms [1] such as Arcellinida. But on the other hand, these organisms are microbial top-predators, and their presence depends on the presence of preys, which consist in smaller protists, metazoa, fungi, and organic debris [28]. ...
Karst caves, formed from the dissolution of soluble rocks, are characterized by the absence of photosynthetic activity and low levels of organic matter. Organisms evolve under these particular conditions, which causes high levels of endemic biodiversity in both macroorganism and microbes. Recent research has highlighted the presence of testate amoebae (Arcellinida) group in cave environments. This study investigates the diversity of Arcellinida in Dinaric karstic caves in Croatia, a global diversity hotspot, focusing on the influence of bat guano on community structure. Sediment samples were collected from two independent hydrosystems, and a metabarcoding approach was used to assess Arcellinida diversity at specific and intraspecific levels, using Arcellinid-specific primers to amplify the mitochondrial cytochrome oxidase subunit I (COI) region. Results reveal a significant impact of guano on both specific and intraspecific diversity of Arcellinida. Communities in guano-rich sites displayed higher diversity, abundance, and the presence of unique OTUs and genetic variants not observed in other habitats, highlighting the crucial role of bats as ecosystem engineers. In contrast, sites without guano hosted communities with low abundance and reduced biodiversity. These differences suggest the existence of guano-associated Arcellinida communities. This study provides new insights into the biodiversity of subterranean ecosystems and the ecological roles of Arcellinida in karstic environments.
Supplementary Information
The online version contains supplementary material available at 10.1007/s00248-024-02483-z.
... This subterranean biodiversity plays an indispensable role in ecological balance (Issartel, 2007), providing a unique habitat for species adapted to sometimes extreme conditions, thereby making a significant contribution to the biological diversity of our planet (Melun, 2012). Highly specialized groundwater fauna exhibits remarkable morphological, physiological, and behavioral adaptations reflecting their evolution to survive in specific hypogeal environments, such as eye reductions, elongated bodies, and reproductive strategies adapted to low-energy conditions (Gibert and Deharveng, 2002;Culver and Pipan, 2019). Various factors characterize groundwater and its habitats (Kallel et al., 2023), with primary production being strongly limited by the absence of photosynthesis (Stocker et al., 2019). ...
... The high diversity of stygofauna, especially the predominance of taxa such as crustaceans and Chironomidae, underscores the importance of these organisms as bioindicators of groundwater health. Stygofauna play a vital role in maintaining ecological balance, contributing to nutrient cycling and energy flow within subterranean environments (Culver, 1982;Culver and Pipan, 2019;Malard et al., 2023). Their sensitivity to environmental changes, such as variations in water quality, makes them invaluable for monitoring ecosystem integrity (Sket, 1999;El Adnani et al., 2007;Saccò et al., 2019a;Thanigaivel et al., 2023). ...
This study explored the biodiversity of organisms inhabiting subterranean waters and shed light on a complex diversity, bringing together species from both surface waters and those specifically adapted to subterranean environments of semiarid lands in North Africa. The focus was primarily on groundwater, mainly interstitial groundwater accessible through springs and wells. Each ecosystem was assessed using the most appropriate sampling method. Macroinvertebrate analysis in the watersheds of Garaat Tarf and Annk Djemel (Algeria), situated under a semi-arid climate, revealed significant diversity, encompassing 11 classes, 22 orders, and 36 families. The Annk Djemel watershed, with more extensive diversity (32 families) compared to the Garaat Tarf watershed, indicates that complex environments foster more diversified communities. Crustaceans, including copepods, ostracods, and amphipods, predominate in these subterranean habitats, confirming patterns observed globally. The prevalence of Copepoda, Cyclopoida, and Diaptomidae aligns with global studies. Species distribution shows a prevalence of accidental species, emphasizing their temporary nature. Only a few taxa (Diaptomidae, Cyclopoidae, and Chironomidae) were constant. The diverse feeding habits of these macroinvertebrates influenced their position in the trophic network. Environmental factors, such as water quality, habitat structure, and food availability, impact the composition of macroinvertebrate communities. The analysis reveals correlations between physicochemical water characteristics and the composition of assemblages. Pollution-resistant taxa show positive correlations with nitrates and orthophosphates. Innovative study methods and international collaboration are essential for a better understanding and preservation of these overlooked environments. Public awareness is crucial for the preservation and sustainable management of subterranean ecosystems on a global scale.
... The environmental variables of the hypogean environment are peculiar in that they are subject to regimes that are practically free of influences from the epigean environment. Variables linked to seasonality are hardly seen as mediators in the life cycles of troglobitic organisms (species restricted to the underground environment) and, in general, these species do not show ecological patterns linked to dry or rainy seasons [1]. detection of specializations related to the cave life and application for the conservation of R. krugi populations. ...
... When we compare it with the development of the gonadal stages, the same pattern is observed: the highest stomach volumes were recorded for the specimens in maturation or mature, mainly in the rainy season, which suggests a slight upward trend in food resources during this crucial breeding period. Besides food scarcity, many cave animals are faced with seasonal availability of resources or even infrequent input of food and can survive under starvation periods [1]. ...
Rhamdiopsis krugi is a highly specialized troglobitic (exclusively subterranean) catfish endemic to the phreatic water bodies of twelve caves located within two separated metasedimentary basins in the region of Chapada Diamantina, Bahia state, Brazil. This species is included in the List of Endangered Fauna of Brazil, under the Vulnerable category—VU. In general, troglobites have different strategies for searching for food and reproductive partners, as well as unique behaviors. Knowledge of the reproductive periods, as well as its feeding habits, provides fundamental data for effective protection and species conservation. Biological aspects related to feeding habits and reproduction of R. krugi were addressed across six annual cycles, considering both dry and rainy seasons. For this, stomach content analysis, using the frequency of occurrence and volumetric index methods, as well as observation of the sex ratio and stage of maturation of the gonads were carried out for 148 individuals of R. krugi sampled in eight caves in Chapada Diamantina. Stomach volumes correlated with reproduction aspects across the dry and rainy seasons. These populations showed opportunistic carnivorous feeding habits, consuming both autochthonous and allochthonous items, with a preference for foraging in submerged guano deposits, which demonstrates the catfish’s strong dependence on bats. Regarding sex ratios, there was no marked seasonality; however, in rainy seasons, there was a higher proportion of maturing females, showing a reproductive tendency. During these periods, there was also a significantly higher number of stomachs with contents, showing seasonality in the diet. Specialized diet and dependence on rainy periods, especially in diet, corroborate the fragility of R. krugi, especially considering the changes in rainfall regimes in Brazil, with dry seasons exceeding eight months per year in the last ten years.
... Typically, subterranean estuaries can be accessed via sinkholes that lead to complex networks of submerged cave systems [3,4]. Anchialine caves are typically subdivided into three sections: (i) the entrance, including the photic zone that is nutrient-rich and where photosynthesis dominates; (ii) cavern or twilight zone, the ecotone between the photic and aphotic regions; and (iii) the 'true' cave zone, which is aphotic and often considered oligotrophic [5,6]. Subterranean estuaries are inhabited by stygobionts (i.e. ...
... An exhaustive literature review was conducted to obtain records of published and unpublished COI and 16S sequence data. There is ample evidence that the entrance and cave zones are distinct environments [5,7,52]. Consequently, only sequences from species previously reported from the cave zone were considered in this study. ...
Subterranean estuaries are coastal ecosystems characterized by vertically stratified groundwater. The biota within these ecosystems is relatively understudied due to the inherent difficulty of accessing such extreme environments. The fauna inhabiting these ecosystems is considered vulnerable to extinction, and the presence of cryptic species has major implications for research and conservation efforts. Most species lack molecular data; however, the evaluation of genetic data for some taxa has revealed that undocumented species are common. This study employs molecular species delimitation methods and DNA barcoding through the analysis of publicly and newly generated sequences, including individuals from type localities and non-crustacean phyla; the latter are typically overlooked in biodiversity assessments of subterranean estuaries. We analysed 376 cytochrome c oxidase subunit I (COI) gene sequences and 154 16S rRNA gene sequences. The COI sequences represented 32% of previously described species and 50% of stygobiont species from the Yucatan Peninsula and Cozumel Island, while sequences of the 16S rRNA represented 14% of described species and 22% of stygobionts. Our results revealed cryptic genetic lineages and taxonomic misidentification of species. As several species from these ecosystems are recognized as endangered, the use of molecular approaches will improve biodiversity estimates and highlight overlooked cryptic lineages in need of evaluation of conservation status.
... Caves have internal characteristics that interact with the external environment, creating photic and aphotic zones. The deeper parts of the cave are aphotic and have a stable average temperature and near-saturation humidity (Culver 1982, Culver & Pipan 2009, Tobin et al. 2013. The absence of natural light leads to the total or partial lack of photosynthetic organisms, limiting primary production. ...
... Cave communities are classified based on species characteristics (morphological, physiological, evolutionary, and behavioral) and their affinity for subterranean habitats (Sket 2008, Culver & Pipan 2009, Díaz 2009). According to Sket (2008), based on the Schinner-Racovitza proposal, organisms are classified as follows: i) trogloxenes, animals regularly found in the underground environment but requiring the surface to complete their life cycle; ii) troglophiles, animals with an affinity for caves, possessing both underground and epigean populations, and potentially completing their life cycle entirely within cavities; iii) troglobites, obligate subterranean species exhibiting morphological, behavioral, and physiological adaptations to the hypogean environment, typically restricted to a specific cave; iv) "accidentals", animals lacking pre-adaptations, inadvertently entering the underground environment and failing to establish themselves (Barr 1968, Sket 2008, Juan et al. 2010. ...
The natural underground environment has unique characteristics when compared to surface environments. These environments feature a reduced complexity food web that includes organisms such as anuran amphibians. This study examines the occurrence of Neotropical anurans in Brazilian caves, utilizing taxonomic, geographic, geological, and environmental data extracted from the scientific literature. A total of 247 anuran records were found in caves across 18 scientific papers covering a 42-years period (from 1980 to 2022). Of these, 177 records (71.6%) displayed the anurans identified to the species level (54 species), with the families Leptodactylidae (13 species; 24.1%), Hylidae (12; 22.2%), and Bufonidae (10; 18.5%) being the most frequently recorded. The anuran records were predominant in the states of Minas Gerais (81 records; 34.8%) and São Paulo (35; 15.0%). The Atlantic Forest had the highest number of records (78; 33.5%), followed by ecotonal zones between the Atlantic Forest and Cerrado (43; 18.5%). Environmental information was scarce in the consulted records, with only 12 providing detail about the cave light zone where the anuran was found (nine in the entrance zone) and 63 indicating the presence/absence of water bodies. Carbonaceous (109; 46.8%) and ferruginous (76; 33.9%) lithology predominated among the caves considered. The low number of sampled caves (55 caves; 0.24%) compared to the total number of caves registered in Brazil (23,278 caves) underscores a knowledge gap regarding Neotropical anurans use of subterranean habitats.
... Caves represent ecosystems with unique conditions compared to the surface, characterized by climatic constancy, a tendency toward oligotrophy, and the absence of light-dark cycles. These environments have distinct zones marked by the presence or absence of light along their length (Culver & Pipan, 2009;Howarth, 1983;Moldovan et al., 2018;Poulson & White, 1969). Cave temperatures resemble the external annual average, while air humidity exceeds surface levels, potentially reaching saturation (H€ uppop, 2000). ...
... Cave temperatures resemble the external annual average, while air humidity exceeds surface levels, potentially reaching saturation (H€ uppop, 2000). These patterns underscore the singularity and dynamics of these subterranean environments, serving as biological filters that select species with varying degrees of adaptation and persistence in such conditions (Culver & Pipan, 2009;Mammola et al., 2019;Poulson & White, 1969). ...
The lack of understanding regarding how endogenous and behavioral factors affect the biological rhythms of amblypygid arachnids in cave environments underscores a gap in chronobiologic knowledge for this group. This study investigates the influence of specialization on subterranean habitats and the presence of biological rhythms on the locomotor activity patterns of the troglobitic and troglophilic species of the genus Charinus. Specimens collected from carbonate caves in Brazil were subjected to experimental treatments of constant light (LL), constant darkness (DD), and light–dark cycles (LD). The results revealed variations in the distribution of main periods among species, without a uniform pattern. Although some specimens showed greater variability in activity patterns in the DD and LL treatments, no significant differences were observed between troglobites and troglophiles. The lack of a clear distinction in rhythms between the two groups suggests the dynamic nature of circadian rhythms in these populations, where individual variations in activity patterns indicate this behavioral diversity. Additionally, intraspecific competition for food resources, probably intensified by the oligotrophic conditions of the cave environments, may play an important role in shaping these patterns and differences in activity phases. The presence of infradian rhythms and weak circadian rhythms in some individuals underscores the importance of considering non‐photic zeitgebers for a deeper understanding of these rhythms in cave organisms.
... INTRODUCTION Culver and Pipan (2009) define a cave as a big, easily accessible hole in the ground that is usually filled with dark. More than merely holes in the ground, caves have deeper meaning for people. ...
... Total darkness in cave areas interferes with circadian cycles, which aff ect many surf ace-level ecological processes (Langecker, 2000). Moreover, creatures inhabiting caves that lack any photosynthetic activity in their shadowy regions depend on outside carbon sources (Culver and Pipan, 2009).It is not possible for many fungi that are occasionally or frequently found in caves to develop there due to spores carried by water, air currents, or animals. It has been noted that conidia of Aspergillus, Penicillium, and Mucor species multiply swiftly in the organic debris present in caves. ...
This correspondence aims to furnish details concerning the fungal diversity records of Pandav and Tiger Cave, located in the Chakrata forest sections of the Doon Valley in the Uttarakhand Himalayas. Samples from the entrance, dark areas, and dark sections of the Pandav and Tiger caves were taken to isolate soil fungi and examine their physicochemical characteristics.Malt Extract Agar and Czapek's Dox Agar were used for the maintenance and isolation of fungus. Soil dilution plate method was used to separate the samples of soil. The pH of the study varied from 6.60 to 7.61. There were variations in the moisture level between 6.23 and 11.23 percent, phosphorus between 10.42 and 192.64 kg/ha, sulfur between 11.02 and 35.2 ppm, zinc below 1.49 ppm, manganese between 0.36 and 5.46 ppm, copper between 0.36 and 5.46 ppm, and fossil carbon between 0.10 and 0.83%.Two hundred and nine colonies of soil fungi representing fifteen genera and twenty-four species were isolated. Of these, 12 genera and 21 species (EZ), 11 genera and 20 species (TW), 10 genera and 19 species (DZ), and 9 genera and 18 species (TW), were separated from Cave 1 and 11 genera and 20 species (DZ) from Cave 2.Four species of Aspergillus-the most common genera were identified from both the caves-Aspergillus wentii, A. niger, A. flavus, and Aspergillus fumigatus. Fusarium and Penicillium, with their species-F.dimerum, Fusarium solani, f. sp., P. raistrickii, P. steckii, and P. maximae-were next. Ten common species, ten moderate species, three frequent species, and one rare species are found in Pandav Cave. In contrast, 10 common species, 6 moderate species, 5 frequent species, and 3 uncommon species have been found in the tiger cave. The Pandav Cave's entrance zone and the Tiger Cave's Twilight zone showed the greatest resemblance, at 0.9047 (90.47%), while the Pandav Cave's Dark zone and the Tiger Cave's Dark zone showed the largest dissimilarity, at 0.3877 (38.77%). The EZ of Pandav Cave had the greatest Shannon-wiener Diversity Index (2.97), followed by the EZ of Tiger Cave (2.94). This study will provide important data regarding the distribution and diversity of cave ecology, which will also give researchers a platform to investigate the fundamental taxonomy of cave microfungi and the impact of fungal spores on the human population and vice versa.
... Despite this, they harbor a vast biodiversity, estimated to exceed 50,000 species globally (Culver & Pipan, 2009), turning them into natural laboratories for ecological, evolutionary, and biomedical research (Recknagel & Trontelj, 2021). Cave environments are characterized by minimal diurnal and annual temperature and humidity fluctuations, dim lighting, nutrient deficiencies, and limited plant presence, typically concentrated in illuminated zones (Monro et al., 2018;Romero, 2009;Souza-Silva et al., 2021). ...
... With decreasing light, resource scarcity further divides caves into twilight and dark zones, making them ideal for studying biological community adaptation and ecological relationships (Wilkens et al., 2000). Presently, cave biology primarily focuses on describing and interpreting species diversity, a vital aim of this study (Culver & Pipan, 2009). Environmental differences between internal and external cave environments and photic zones have resulted in varied animal adaptive characteristics. ...
Mites are among the most abundant invertebrates in subsurface ecosystems, and their community assemblages and distributions are often significantly influenced by the diversity of habitat resources. The cave ecosystem encompasses drastic changes in nonbiological factors, such as changes in lighting conditions from bright to extraordinarily dark and habitat gradients of surface plant resources from abundant to scarce or even disappearing, providing an ideal unique environment for evaluating the assembly mechanism of soil animal communities. Nevertheless, there still needs to be a sufficient understanding of the biodiversity patterns and drivers of mite communities across environmental gradients in karst caves. We conducted a comprehensive survey on the composition and diversity of soil mites in three photometric zones (dark, twilight, and light) of a typical karst cave and its adjoining extractive environments (forest scrub and farmland). Our research aimed to investigate the ecological relationships of mite communities between above‐ and below‐ground habitats and the effects of abiotic factors on mite communities. We collected 49 families, 86 genera, and 1284 mites. In the external cave environment, we captured 1052 mites from 72 genera and 45 families; in the internal cave environment, we captured 232 mites from 46 genera and 29 families. The abundance, richness of genera, and diversity parameters of the mite community decreased from the cave entrance to the cave interior with decreasing light intensity. Oribatid mites dominated the mite community. Protoribates and Scheloribates were the dominant genera, along with Tectocepheus and 11 other genera, which primarily distinguished the mite communities among different habitats. Forty endemic taxa were found in the external cave environment, compared to 14 endemic taxa in the internal cave environment. The mite community showed a strong preference for the cave ecosystem habitat. Temperature, humidity, and soil nitrogen content significantly influenced the distribution pattern of mite communities (VIP > 0.8, p < 0.05).
... The subsequent discovery of a chemoautotrophic sulfidic groundwater ecosystem in Movile Cave, Romania [2], and later in other sulfidic caves, showed that large amounts of organic material produced in situ by autotrophic sulfur-and methane-oxidizing microorganisms can often support abundant and diverse subterranean invertebrate communities including numerous endemic species [3]. All the sulfidic underground ecosystems explored to date emerged as hotspots of subterranean biodiversity [4][5][6] and proved able to support complex subterranean food webs. Chemolithoautotrophs, often deemed less efficient than phototrophs, were traditionally not considered significant primary producers, but they play a crucial role in sustaining many ecosystem-level processes in the absence of light and photosynthesis in subsurface environments [5]. ...
... All the sulfidic underground ecosystems explored to date emerged as hotspots of subterranean biodiversity [4][5][6] and proved able to support complex subterranean food webs. Chemolithoautotrophs, often deemed less efficient than phototrophs, were traditionally not considered significant primary producers, but they play a crucial role in sustaining many ecosystem-level processes in the absence of light and photosynthesis in subsurface environments [5]. Although they are very rare, ecosystems based on chemoautotrophy are of particular interest to researchers as they may represent analogs for Earth's earliest biological communities or for possible extraterrestrial life [7]. ...
Sulfidic caves support diverse and abundant subterranean communities, including numerous endemic species and complex food webs, though the full extent of species diversity and resource utilization in these ecosystems remains largely unexplored. This paper presents the results of biological surveys conducted from 2023 to 2024 in Sulfur Cave, located in the Vromoner Canyon on the Greek–Albanian border, focusing on microbial, vertebrate, and invertebrate communities and investigating the structure of the subterranean food web. The microbial communities from the different biofilms are dominated by chemosynthetic sulfur-oxidizing microorganisms, specifically filamentous bacteria such as Thiotrix and Beggiatoa. Two species of fish, an eel (Anguilla sp.) and a Cyprinid (Alburnoides sp.), and six bat species from three families (Rhinolophidae, Miniopteridae, and Vespertilionidae) were documented. The invertebrate fauna includes five aquatic species, 25 terrestrial species, and four amphibiotic species. Among these, eight species are endemic, and seven species exhibit troglomorphic traits. Stable isotope analysis showed light carbon and nitrogen values for the terrestrial and aquatic invertebrates, suggesting that subterranean communities rely on food produced in situ by chemoautotrophic microorganisms. Our results identified cave areas of significant biological relevance and provided reference data to inform conservation actions aimed at preserving the biodiversity of this sulfidic cave.
... For these reasons, subterranean ecosystems are more simplified than the energy-rich and seasonally variable surface ecosystems. Therefore, in subterranean environments, the effects of climatic parameters can be easily analyzed separately, allowing a better understanding of their relative influence on the ecology of biological communities (Culver and Pipan 2009;Mammola 2019). ...
... However, even if subterranean ecosystems are considered ideal models to better understand climate change effects, they are still among the most overlooked habitats at the global scale (e.g., Sánchez-Fernández et al. 2018Ficetola et al. 2019). This situation is surprising because subterranean environments are widespread all over the globe and are often highly threatened, not only by climate change but also by pollution, quarrying, water extraction, invasive alien species, and pathogens (e.g., Culver and Pipan 2009;McClure et al. 2020). ...
Long-term effects of current climate on animal populations living in subterranean habitats are still poorly understood. In this study, we analyzed the variability of the demographic structure and abundance of a population of terrestrial plethodontid Strinati's Cave Salamanders (Speleomantes strinatii) (Aellen, 1958) living inside a northwestern Italian artificial cave over 27 yr from 1996 to 2022. The study site is situated in the Northern Apennines, where average air temperatures measured at two local weather stations were steadily increasing, whereas precipitation was relatively constant over the past 60 yr. Our objective was to evaluate whether the demographic traits of the population of Strinati's Cave Salamanders were showing detectable signs of directional shift during the 27 yr. Each year in July, the population abundance was estimated by a three-occasion removal experiment, salamanders' snout-vent length (SVL) was measured, and the population polymodal body-size distribution was decomposed into estimated age classes. The annual population abundance, adult sex ratio, recruitment, SVL, and growth increments of first-and second-year immature salamanders were analyzed. The Strinati's Cave Salamander time series was modeled by autoregressive moving average (ARMA) analysis, and demographic parameters were tested for temporal trends. There were no directional trends observed in any of the demographic parameters of the focal cave salamander population over the study period. The best autoregressive model describing the population variation was an ARMA (2,1). Results indicated that the salamander population had a complex fluctuating pattern in which the interaction of lagged density dependence and an external autocorrelated factor were influencing the population long-term dynamics. Our findings suggest that the population of this long-lived salamander species was probably buffered from external conditions by the thermal inertia of its subterranean habitat that is known to delay and reduce the amplitude of aboveground climatic signals.
... For these reasons, subterranean ecosystems are more simplified than the energy-rich and seasonally variable surface ecosystems. Therefore, in subterranean environments, the effects of climatic parameters can be easily analyzed separately, allowing a better understanding of their relative influence on the ecology of biological communities (Culver and Pipan 2009;Mammola 2019). ...
... However, even if subterranean ecosystems are considered ideal models to better understand climate change effects, they are still among the most overlooked habitats at the global scale (e.g., Sánchez-Fernández et al. 2018Ficetola et al. 2019). This situation is surprising because subterranean environments are widespread all over the globe and are often highly threatened, not only by climate change but also by pollution, quarrying, water extraction, invasive alien species, and pathogens (e.g., Culver and Pipan 2009;McClure et al. 2020). ...
... The cave environment is a distinct element of the landscape, characterized by its stable abiotic conditions, permanent absence of light in deeper zones, high humidity near saturation, and mild temperature with little variation (Culver & Pipan 2009;Tobin et al. 2013). Biotic conditions include simplified ecological networks, minimal or no primary production within the system, and energy support from the oligotrophic system facilitated by physical and biological agents (Simon et al. 2007). ...
Cave environments present stable abiotic conditions, including permanent darkness, high humidity, and mild temperatures, while biotic factors reflect simplified ecological networks and energy constraints. Cave invertebrates, primarily detritivores and generalists, demonstrate specific adaptations to these conditions. Predation and necrophagy are critical behaviors shaped by the cave's scarcity of food resources. In this study, we report a rare necrophagy event involving Discocyrtanus canjinjim Carvalho & Kury, 2017 (Arachnida: Opiliones) feeding on a deceased cricket (Eidmanacris sp.) (Orthoptera: Phalangopsidae) in the Ponte de Pedra I Cave, Brazil. This limestone cave features large entrances and supports an oligotrophic system. The event occurred in an aphotic zone, where the harvestman displayed no flee behavior despite external stimuli. The presence of other predatory arachnids suggests potential intra-guild competition. Opiliones in the Cerrado and Caatinga exhibit opportunistic carnivory, consuming various prey taxa, including insects and vertebrates. Necrophagy, intra-guild predation, and cannibalism are behaviors reported in subterranean populations due to limited food availability. Climate change and anthropogenic pressures, such as deforestation and mining, threaten the stability of cave environments. Observations like this contribute to understanding the ecological dynamics within caves, highlighting the importance of preserving these fragile ecosystems.
... Cave animals are commonly classified into three categories based on their degree of adaptation to the cave environment: troglobites, troglophiles, and trogloxenes (Sket 2008;Howarth and Moldovan 2018;Culver and Pipan 2019). The present study describes 12 new cave-dwelling species and classifies their cave adaptability mainly based on the troglomorphic features exhibited in their external morphology, such as the degeneration of eyes, unpigmented bodies and the development of legs (Liu et al. 2017). ...
A taxonomic study of the genus Skleroprotopus Attems, 1901 from nine provinces in China was conducted utilizing morphological comparisons and molecular phylogenetic analyses. The results reveal thirteen new species, i.e. Skleroprotopus yutiantianae sp. nov. , S. tiankeng sp. nov. , S. megistus sp. nov. , S. penglai sp. nov. , S. longissimus sp. nov. , S. genjudi sp. nov. , S. laiyuanensis sp. nov. , S. longiflagellatus sp. nov. , S. change sp. nov. , S. ampullaceus sp. nov. , S. incisodentatus sp. nov. , S. multistriatus sp. nov. , and S. conicus sp. nov. This significantly enriches the diversity of Skleroprotopus in China, bringing it to a total of 18 species. With the exception of S. yutiantianae sp. nov. , all these species were collected in caves. In terms of their degree of adaptation to the cave environment, the latter six species are presumed troglophiles, while the others are likely troglobites. DNA-barcoding based on the COI mitochondrial gene is documented for the first time in this genus. The specific p -distances between Skleroprotopus species range from 6.6–17.0%, while intraspecific p -distances are only 0.2–1.4%. Additionally, the morphological features of male leg-pair 1, the penis and leg-pair 7 are also discussed.
... Infested intermediate hosts are less photophobic and darker pigmented (Lyndon, 1996) than uninfested ones, which presumably increases the likelihood that they will be preyed upon, thus facilitating transmission of the parasite to the final host. Cave habitats are lightless, nutrient-limited, with buffered daily and seasonal environmental fluctuations and simpler communities with lower population densities (Culver & Pipan, 2019). Among abiotic conditions, constant darkness represents the most significant disruption to existing predator-prey interactions, rendering manipulations of photophobia and pigmentation ineffective. ...
Acanthocephalans are obligatory endoparasites that often alter the phenotype of their invertebrate intermediate host to facilitate trophic transmission to their final vertebrate host. Acanthocephalus anguillae , a widespread parasite of European freshwater fishes and isopod Asellus aquaticus , was recently discovered also in Postojna-Planina Cave System (Slovenia) parasitising olms ( Proteus anguinus ) and cave populations of A. aquaticus. This setting offers a unique opportunity to investigate potential fine-tuning of parasitic manipulations to the specifics of the highly divergent subterranean environment where some common phenotypic alterations lose functionality, but others might gain it. We measured three behavioural traits: movement activity, shelter-seeking, and response to light of infested and uninfested isopods from surface and cave populations. All behaviours were quantified from 1-h video-recordings via video-tracking isopod’s movement in empty or custom modified (half-sheltered/half-illuminated) Petri dishes. Infested isopods of both populations spent significantly less time sheltering and were significantly less photophobic than uninfested ones, whereas the activity of isopods was not altered. However, we observed almost no cave-specific responses upon infestation in the two altered behaviours. It seems phenotypic alterations are not particularly fine-tuned to the subterranean environment and its hosts, and likely still reflect the parasite’s surface origin.
... Hypogean species, which live in underground ecosystems, exhibit common morphological characteristics despite inhabiting different geographical areas and belonging to different groups: no eyes and lack of pigmentation. This is an example of convergent evolution, as eyes and pigmentation are useless in total darkness, so they gradually disappeared over time, while other structures, such as appendages, have become more prominent (Culver and Pipan 2019). ...
Despite the various criteria presented in the literature, most authors engaged in the debate about emergence agree on a fundamental distinction between strong/ontologically robust cases of emergence and weak/metaphysically innocent ones. The former typically involve entities that exhibit new causal capacities, while the latter are primarily associated with deductive unpredictability, conceptual novelty, and other qualities that highlight our epistemic limitations in understanding them. In this paper, I initially examine a paradigmatic example of weak emergence, namely the higher-level patterns generated by virtual cellular automata (CAs) as analyzed by Mark Bedau. Then, I demonstrate that the same mechanism can be observed in real biological systems, such as the dynamics governing the pigmentation ontogeny of the ocellated lizard (Timon lepidus). Unlike virtual CAs, however, real CAs produce patterns that seem to perform non-reducible functions. Therefore, I propose that despite the similarities between virtual and real CAs, the pigmentation pattern of the ocellated lizard should be regarded as a case of strong emergence. Moreover, I suggest that this analysis may shed light on the nature of biological emergent entities in general. Finally, the paper includes an addendum introducing an issue that, while not exhaustively addressed here, is highly relevant: how to metaphysically conceptualize the causal efficacy exhibited by the pigmentation patterns.
... Communications Biology | (2024) 7:1500 palaeovalleys 52,53 without groundwater connectivity between them, supporting the scenario of multiple colonizations from the surface. The transition to life in groundwater ecosystems is associated with significant morphological changes (e.g., reduction of eyes and pigments, but gain of long tactile setae and antennae) 54,55 , and the reduction/loss of wings in subterranean beetles 56 . The precise origin of certain phenotypes associated with an underground lifestyle (i.e., reduction of eyes and pigments) remains insufficiently studied, leaving the question open as to whether these changes result from degeneration (or pseudogenization) of associated genes following relaxation of selective pressures 57 . ...
Understanding the profound influence of climatic and tectonic histories on adaptation and speciation is a crucial focus in biology research. While voyages like Humboldt’s expedition shaped our understanding of adaptation, the origin of current biodiversity remains unclear – whether it arose in situ or through dispersal from analogous habitats. Situated in the geologically complex Australopacific region, our study focuses on Limbodessus diving beetles (Dytiscidae), a diverse genus distributed from underground aquifers in Western Australia to alpine meadows in New Guinea. Using low-coverage whole-genome sequencing, we established a time-calibrated phylogenetic tree, elucidating Limbodessus’ origin in the mid-late Miocene, most likely in the Sahul continent (i.e., Australia and New Guinea) and western Pacific archipelagos. Our results provide evidence for parallel colonization and speciation at extreme altitudinal ends, driven by aridification in Australia, influencing subterranean colonization, and in situ diversification of alpine taxa by passive-uplifting of local biota in New Guinea. Furthermore, our findings highlight instances of subterranean speciation in isolated underground aquifers, marked by recurrent independent colonizations of this habitat.
... In addition to preserving the cave's natural beauty, or speleothems, protection of the cave and its surroundings is also necessary to minimize disruption to the occasionally exceedingly delicate subterranean ecology. In actuality, the subterranean spaces, which include both caverns and small fissures, serve as a home to a remarkably diverse and relatively specialized cave fauna (Culver and Tanja, 2009). ...
Karst refers to any topography or landform that forms over soluble rocks. The classic karst landforms, which
include enclosed depressions, sinking streams, and caves, are primarily generated by surface and subsurface
waters dissolving rocks; mechanical erosion plays a secondary role. This review paper attempted to evaluate the
Karst topography from an international standpoint, taking into account its creation, features, resultant landforms, denudation, and restoration. The systematic review of the literature, which has been shown to be a
trustworthy approach, served as the foundation for this review article. The review shower that iypically, karst
topography forms on evaporites, dolostones, limestones, and rock salt. Caves can contain significant paleontological, paleoenvironmental, and archeological remnants because they can serve as traps for surface material,
shielding it from surface erosion. Karst cannot be produced solely by a significant amount of rock solubility. The
best karst is developed by dense, massive, pure, and coarsely fractured rocks. Although karst terrain is most
common in humid areas with carbonate rock, it can also be found in temperate, tropical, alpine, and polar
climates. According to the Koppen-Geiger ¨ classification, arid climates account for approximately 34.2% of all
carbonate rock areas worldwide. Cold climates account for 28.2%, moderate temperatures for 15.9%, tropical
climates for 13.1%, and polar climates for 8.6% of the total. Ten percent of the Earth’s surface is made up of karst
topography, and up to 25% of the world’s population depends on karst areas for their water supply, making an
understanding of karst essential. The peculiarities of karst environments make them highly vulnerable to geohazards such as collapse and dropout dolines (sinkholes), slope movements, and floods. Karst areas are also
extremely specialized habitats, with important and often very vulnerable subterranean ecosystems. The
vulnerability of karst requires unique management and protection strategies to be devised to preserve this
natural landscape for its future sustainability
... Further, about 33% of bat species are fruit or nectar feeding and are thus pollinators and dispersers of numerous plants. In caves, bats are umbrella species that act as surrogates for cave biodiversity and conservation value and provide organic nutrients to cave ecosystems in the form of guano [15][16][17]. Despite the numerous ecosystem services bats provide, roughly 25% of bat species are threatened due to anthropogenic disturbance, lending urgency to bat community conservation and management [18,19]. ...
In a rapidly changing world, where species conservation needs vary by local habitat, concentrated conservation efforts at small spatial scales can be critical. Bats provide an array of value to the ecosystems they inhabit; many bat species are also of conservation concern. San Diego County, California, contains 22 of the 41 bat species that occur in the United States, 16 of which are on conservation watchlists. Thus, management of bat communities in San Diego County is a pressing need. Because bats exploit vast areas of the landscape and historical sampling strategies have shifted over time, a standardized way of prioritizing areas of the landscape for management would provide an integral asset to bat conservation. We leveraged long-term bat community survey data from sampling areas across San Diego County to prioritize areas with the most management need. We calculated two types of scores: species scores and threat scores. Species scores incorporated richness and conservation status, and threat scores included landscape level threats that bats could encounter. We found that urbanization, the presence of artificial lights, and areas sampled on unconserved land were all significantly associated with decreases in species richness. Further, using species and threat scores, each sampling area was placed into one of four conservation categories, in order from greatest to least conservation need, ranging from highest priority (high species score, high threat score) to lowest (low species score, low threat score). Additionally, we focused on sampling areas in which Townsend’s big-eared bat (Corynorhinus townsendii) and/or pallid bat (Antrozous pallidus) occurred. These two species are of exceptional conservation concern in San Diego County and across the western United States. We identified urbanization, the presence of artificial lights, and areas sampled on unconserved land as threats that were all significantly associated with the absence of Townsend’s big-eared bat, but not pallid bat. The strategy, methodology, and solutions proposed in our study should assist bat conservation and management efforts wherever bats occur, and can be extended to other species that require conservation attention.
... Despite being the largest freshwater reservoir on earth and a keystone ecosystem (Ferguson et al. 2021;Saccò et al. 2024), it is one of the least studied ecosystems, mainly due to difficulties in accessibility (Mammola et al. 2019;). Yet, it harbors a unique range of organisms that are adapted to the dark and energy deprived conditions (Culver and Pipan 2019). These obligate groundwater dwellers (stygobites) exhibit a high rate of endemism, and some of the species originate from ancient adaptive radiations (Trontelj et al. 2009;Borko et al. 2021). ...
Implementing and optimizing biodiversity monitoring is crucial given the current, worldwide biodiversity decline. Compared to other ecosystems, monitoring of biodiversity is lagging behind in groundwater ecosystems, both because of sparse taxonomic knowledge and methodological constraints. We here assessed temporal variation in the occurrence and abundance of macroinvertebrates collected systematically from shallow groundwater aquifers of Switzerland to establish general principles on seasonality and repeatability of assessment outcomes. We found no seasonal abundance pattern for obligate groundwater amphipods and isopods, indicating temporal consistency. In contrast, other macroinvertebrates (predominantly stygophiles and stygoxenes) showed pronounced seasonality in their detection rate. However, we found variability in detection rates across groundwater amphipod species and especially across sampling sites. For groundwater communities, characterized by narrowly-distributed and rare species, our results highlight the need for tailored and extensive sampling strategies. When setting up monitoring programs on groundwater fauna, detection probability, temporal autocorrelation, and standardization of sampling effort should be carefully considered. Applying novel, systematic approaches, can offer promising methodologies for understanding and conserving groundwater ecosystems.
... Species richness and abundance of the invertebrate fauna are expected to decrease with increasing depth in the MSS [56][57][58] , as the result of the decrease in available organic matter with increasing depth 59 and change in conditions in the rock (e.g., water retention, lower temperature compared to top layers) and the surface (e.g., more cold/hot or rainy/dry conditions) 60,61 . This can cause a migration of epigean species into MSS to escape Fig. 6. ...
Iron Formations (IF) are threatened by mining, particularly the Mesovoid Shallow Substratum (MSS), an understudied subterranean environment. We evaluate the spatiotemporal patterns of subterranean fauna in MSS of iron duricrust (canga) in the Iron Quadrangle and Southern Espinhaço Range, southeastern Brazil. Samplings took place between July 2014 and June 2022 using five trap types. We sampled 108,005 individuals, 1,054 morphospecies, and seven phyla, globally the largest dataset on MSS in IF. Arthropoda represented 97% of all invertebrates sampled. We identified 31 troglomorphic organisms, primarily Arthropoda and Platyhelminthes. MSS traps were the most efficient method, capturing 80% of all invertebrates. Morphospecies were more prevalent in each locality than shared among localities. Species replacement was the main processes to spatial differences. Over time, we found a decrease of total dissimilarity and importance of species replacement for troglomorphic organisms. A positive correlation between spatial distance and compositional dissimilarity of invertebrates was found. Iron Quadrangle and Southern Espinhaço Range showed marked differences in the spatiotemporal patterns of subterranean fauna. Brazilian IF are threatened, with their biological significance not fully understood but highly endangered due their limited distribution. Conservation efforts require a comprehensive understanding of both biotic and abiotic factors shaping the entire IF ecosystem.
... Though we know little about the ecology of the respective caves, cave systems in the region of Loka pri Mengšu are densely represented, with many being contiguous and thus, sharing subterranean drainage systems. Zospeum snails get washed through these systems during seasonal water flows and displacement of fauna within a concentrated system of caves is not unusual (Culver and Pipan 2009). Considering that Z. s. schmidtii was reported from many caves in the region and in several cases sympatrically with Z. costatum (Bole, 1974), the potentiality of mixing of populations is high. ...
The syntypes of Carychium reticulatum Hauffen, 1856 and Carychium bidentatum Hauffen, 1856, thought to be lost to science, have been rediscovered in a recent inventory of the shell collection of Domink Bilimek (1813–1884) at the University of Natural Resources and Life Sciences in Vienna, Austria (BOKU). In this work, we present (1) the historical context of the Dominik Bilimek and Heinrich (also: Henrik) Hauffen collections and (2) provide a contemporary image dataset of Hauffen’s original material while considering today’s taxonomic understanding of the genus Zospeum. We clarify the taxonomic status of the syntypes by using light microscopy, Scanning Electron Microcopy (SEM) and 3D X-ray Micro-CT data in conjunction with Hauffen’s original illustrations and compare them to contemporary investigations of their closest congeners. Analysis of Zospeum reticulatum (Hauffen, 1856) is supported by historical documentation from Heinrich Hauffen’s original annotations and Bilimek’s diary entries, as well as from individual collection labels in Hauffen’s own script (verified by archive material from the Archive of the National Museum of Slovenia, NMS). Hauffen’s descriptions are based on apertural morphology and shell microstructure. Our re-investigation of Zospeum reticulatum provides corroborating evidence supporting its assignment to Zospeum spelaeum (Rossmaessler, 1839) while that of Carychium bidentatum corroborates its status as junior synonym of Zospeum costatum (Freyer, 1855).
... Flooded cave entrances represent ideal natural protection from predators and human disturbances. Important traits of caves are also the small variations of temperatures that are very close to the annual temperatures of the area (Culver & Pipan, 2009). This means that in winter months temperatures in caves are higher than outside the cave which could be an advantage in winter periods when food availability is limited and energy saving represents an advantage (Nolet & Rosell, 1994). ...
After its reintroduction in northern Croatia, the Eurasian beaver (Castor fiber L.) successfully spread to new areas along streams. Although the beaver has spread in a part of Croatia covered with karst, the information published to date about beavers using caves has been sparse. Since the first observation in the Matešićeva Cave – Popovačka Cave System near Slunj in 2021 and two caves in Plitvice Lakes National Park in 2022, the beaver and traces of its activity was recorded several times in Jopićeva Cave near Krnjak in the period from March to June 2022. In the same year, beaver activity was recorded near Krnjak in the surface stream Bent connected with cave streams. The spatial and field analysis of this locality showed it is likely that the entry of the beaver into the cave, diving through karst spring Vrelce, was induced by limited access to compatible surface habitats and additional pressures induced by humans. More research is needed for a better understanding of the importance of caves for beavers and the species limitations determined by karst areas.
... The fauna in caves is classified according to how long the organisms stay in these environments. Therefore, fauna that occurs facultatively or occasionally in these environments is known as trogloxene, the fauna that uses these sites seasonally is known as troglophile, and the fauna that completely develops inside caves is known as troglobiont [10][11][12]. ...
The mountain region of central Veracruz, Mexico hosts a large system of karst and volcanic caves that are unexplored. In particular, the vertebrates that inhabit these subterranean ecosystems are unknown. This study evaluated the diversity of mammals, birds, reptiles, amphibians, and fish in three environments (euphotic, disphotic, and aphotic) of 16 caves of different geological origin (12 karst caves and 4 volcanic caves) distributed along an altitudinal gradient (300–2400 m a.s.l.). We found a richness of 242 vertebrate species (184 birds, 30 mammals, 15 reptiles, 12 amphibians, and 1 fish) and an abundance of a total of 11,323 individuals (4,969 mammals, 6,483 birds, 36 reptiles, 27 amphibians, and 5 fish). The richness of all vertebrate classes was higher in karst than in volcanic caves. Vertebrate diversity was also higher at mid-altitudes between 600–899 m a.s.l. Diversity varied between environments, where bird and reptile richness was higher in the euphotic environment, while mammal and amphibian diversity was higher in the aphotic environment. The similarity in the composition of vertebrate species does not depend on the distance between karstic and volcanic caves. Volcanic and karst caves shared on average up to 70% and 55% of vertebrate species, which indicates that only 30% and 45% of species, respectively, is different in each cave type. Given the vulnerability and fragility of these subterranean ecosystems, as well as the important diversity that they contain, we recommend including the caves of the central region of Veracruz in the conservation agenda of local governments and communities. Community-based conservation can help ensure the presence of vertebrate species in the caves of this region.
... Geometrical features of karst landforms can decode paleoenvironments, paleoclimates, and help visualize the geological processes that formed them. Caves are "windows" to the past, providing answers and information to several field of studies, such as geomorphology (e.g., Ford, 2006), structural geology (e.g., Forbes and Nance, 1997), ore deposit geology (e.g., Piló et al., 2015), hydrology (e.g., Moreno et al., 2017), climatology (e.g., Luetscher et al., 2005), bioclimatology (e.g., Finlayson et al., 2008), paleontology (e.g., Anderson et al., 2010), biology (e.g., Culver and Pipan, 2019), archeology (e.g., Straus, 1990), anthropology (e.g., Healy, 2007), and many others. Webster and Sheets (2010) proposed in their landmark-based geometric morphometrics study that morphometrics entails the quantitative examination of biological shape, its variation, and the correlation of shape with other biotic or abiotic variables. ...
Morphometric analysis is the quantification of shapes, which makes irregular shapes found in nature analyzable and comparable. Cave morphometry has been used for the genetic classification of caves, the digital reconstruction of their conduits, the decoding of their paleoenvironment, and other research purposes. Ratios and indices that have been derived from Euclidean geometry and application of fractal geometry onto karst features and topological parameters are the basic methodologies that have been used for shape quantification. This paper reviews the literature that focuses on methodologies used for morphometric analyses and the applications that these methodologies have found.
... Surface habitats differ markedly from subterranean ones in many aspects, such as the presence of light, higher degree of daily and seasonal environmental fluctuations (Lauritzen, 2018), increased food availability (Culver & Pipan, 2019), more diverse communities (Gibert & Deharveng, 2002), which can result in higher risk of predation and increased interspecific competition (Manenti et al., 2023;Sket, 2008). ...
Subterranean and surface habitats are in stark contrast in several environmental factors. Therefore, adaptation to the subterranean environment typically impedes the (re)colonisation of surface habitats. The genus Niphargus includes amphipod crustaceans that primarily occupy subterranean habitats. All its species show typical adaptations to the subterranean environment. However, some Niphargus species occur in surface‐subterranean ecotones. To understand whether (i) habitat‐based phenotypic divergence is present between the cave and the ecotone species and (ii) similar phenotypes emerge independently in each ecotone, we studied morphological divergence between four cave and four ecotone Niphargus species based on 13 functional morphological traits. To account for different selection acting on the sexes, we included both males (N = 244) and females (N = 222). Nine out of 13 traits showed habitat‐divergence. Traits related to feeding and crawling were shorter, while traits related to oxygenation were larger in ecotone species. Eleven out of 13 traits were sexually dimorphic. Traits related to oxygenation and crawling were larger in females, while the trait related to swimming was larger in males. We found that the extent of sexual dimorphism differs between the habitats in eight traits related to sensing, feeding, oxygenation and crawling. Additionally, we found that in certain traits related to sensing and oxygenation, habitat‐related differences are only present in one sex, but not the other. We conclude that the detected differences between the cave and the ecotone species indicate divergent evolution, where similarities among the different species within habitat type indicate convergent evolution. The high degree of sexual dimorphism paired with differences in sexual dimorphism between the habitats in certain traits suggest that sexual and fecundity selections have comparable effects to environmental selection. Thus, studies of habitat‐dependent adaptations investigating one sex only, or not considering sexual dimorphism, can lead to erroneous conclusions.
... Flooded cave entrances represent ideal natural protection from predators and human disturbances. Important traits of caves are also the small variations of temperatures that are very close to the annual temperatures of the area (Culver & Pipan, 2009). This means that in winter months temperatures in caves are higher than outside the cave which could be an advantage in winter periods when food availability is limited and energy saving represents an advantage (Nolet & Rosell, 1994). ...
After its reintroduction in northern Croatia, the Eurasian beaver (Castor fiber L.) successfully spread to new areas along streams. Although the beaver has spread in a part of Croatia covered with karst, the information published to date about beavers using caves has been sparse. Since the first observation in the Matešićeva Cave-Popovačka Cave System near Slunj in 2021 and two caves in Plitvice Lakes National Park in 2022, the beaver and traces of its activity was recorded several times in Jopićeva Cave near Krnjak in the period from March to June 2022. In the same year, beaver activity was recorded near Krnjak in the surface stream Bent connected with cave streams. The spatial and field analysis of this locality showed it is likely that the entry of the beaver into the cave, diving through karst spring Vrelce, was induced by limited access to compatible surface habitats and additional pressures induced by humans. More research is needed for a better understanding of the importance of caves for beavers and the species limitations determined by karst areas. Europski dabar (Castor fiber L.) nakon reintrodukcije u sjevernu Hrvatsku uspješno se proširio na nova područja uz vodotoke. Iako se dabar proširio u dijelu Hrvatske prekrivenom kršem, dosad je malo objavljenih podataka o dabrovima koji koriste špilje. Od prvog opažanja u Sustavu Matešićeva špilja-Popovačka špilja kod Slunja 2021. godine i dvije špilje u Nacionalnom parku Plitvička jezera 2022. godine, dabar i tragovi njegove aktivnosti više su puta zabilježeni u Jopićevoj špilji kod Krnjaka u razdoblju od ožujka do lipnja 2022. godine. Iste godine zabilježena je aktivnost dabra kod Krnjaka u površinskom potoku Bent koji je povezan sa špiljskim potocima. Prostorna i terenska analiza ovog lokaliteta pokazala je da je vjerojatno da je ulazak dabra u špilju, roneći kroz krški izvor Vrelce, bio potaknut ograničenim pristupom kompatibilnim površinskim staništima i dodatnim pritiscima izaz-vanima od strane čovjeka. Potrebna su dodatna istraživanja za bolje razumijevanje važnosti špilja za dabra i ograničenja ove vrste koja uvjetuju krška područja.
... Caves are common in limestone formations and are karstic structures carved by water. In general, the cave environment is dark, humid, and with minor temperature fluctuations, and, although offering a scarce supply of nutrients to the cave biota, provides habitats with stable conditions for many unique and endemic species [3][4][5][6][7]. The preservation and research on the fauna and flora of this uncommon environment is nowadays a priority in many countries (e.g., USA, Turkey, Slovenia). ...
Dimba Cave is a large array of natural galleries in limestone mountains of the Democratic Republic of the Congo that contains highly valued pre-historic archaeological artifacts. The cave attracts a high number of tourists every year and is used by local populations as a water supply source. The main objective of the research undertaken in Dimba Cave consisted of assessing the quality of water and sediments from Dimba Cave ponds through evaluating contamination by heavy metals (15 elements analyzed, including As, Cd, Pb, and Hg) and by microbial populations (including Escherichia coli and total coliforms) in order to estimate the ecotoxicological risk to humans and to non-human biota. All water samples collected in the cave ponds showed very high metal concentrations exceeding the internationally recommended limits for drinking water, particularly for Cr, Mn, As, Pb, and Hg. Most sediment samples from cave ponds also displayed high heavy metal concentrations. The calculated pollution parameters, such as the enrichment factor (EF), and ecological risk parameters, such as the ecological risk index (Eri), indicated that the sediment may be toxic to aquatic biota. Furthermore, the microbiological analysis of pond waters indicated a widespread contamination with bacteria such as Escherichia coli, Enterococcus spp., total coliforms, and Pseudomonas spp., probably from anthropogenic and/or animal sources. Therefore, the consumption of Dimba Cave water as a drinking water represents a threat to public health. Urgent management measures should be enforced to protect public health and the cave ecosystem.
... Not only do they provide water from large springs, widely utilized as a source of drinking water supply (Stevanovic, 2010), but they also constitute surface and underground habitats that act as ecological niches (Culver and Pipan, 2019). Additionally, karst regions hold cultural and historical value: many artifacts associated with early human development have been found in these areas (Sherwood and Goldberg, 2001). ...
Urbanization and agricultural activities are increasingly threatening karstic systems and the water resources they provide, which are crucial for the livelihood of rural communities. Over the past decade, the Río Grande de Comitán-Lagos de Montebello (RGC-LM) watershed in Chiapas, Mexico, has experienced significant deterioration in water quality. The objective of this study was to analyze the water quality dynamics within the RGC-LM watershed and their potential interaction with land use change. We conducted assessments of the physicochemical and microbiological parameters across the lotic and lentic systems within the watershed over eight monitoring campaigns (from 2013 to 2020). Cluster Analysis (CA) and Principal Component Analysis (PCA) statistical approaches were performed to identify the main drivers of surface water quality variability. This approach aims to enhance water quality assessments within environmental management strategies, with the goal of promoting sustainable land-use practices aimed at protecting and improving water resources within the watershed. Our findings indicate that there is a substantial difference between the upper and lower watershed water quality. The good water quality conditions observed in the higher altitude lakes (mountain lakes) can be attributed to favorable topographical features, specific land use patterns, and effective conservation measures. In contrast, the lotic system in the middle watershed and the lentic water bodies located at lower altitudes (plain lakes) exhibit signs of water quality degradation due to inefficient wastewater treatment and agricultural runoff. This study highlights the importance of implementing adequate environmental management strategies to address these water quality challenges effectively. Given the pressing issue of water quality degradation in karstic environments, we emphasize on the need for increased monitoring frequency of both physicochemical and microbiological parameters to capture seasonal variations and to further understand the vulnerability of water resources in karstic environments. Future assessment of nutrient concentrations, pesticide levels, and blue-green algae populations will also be crucial for evaluating the trophic state at a watershed scale.
This dissertation addresses knowledge gaps in cave-dwelling bat ecology in two distinct countries, namely Costa Rica and Bulgaria, with a strong conservation focus. Originally centered on Costa Rica's Brunca region, the study expanded to encompass the entire country As the pandemic restricted fieldwork and travel to Costa Rica, the original methodology was performed in Bulgaria, further expanding the geographical scope of the research. Not all aspects of the research follow a parallel trajectory. While Bulgarian cave fauna and cave-dwelling bats have been subject to numerous studies, the Costa Rican underground roosts needed more thorough investigation. This dissertation accordingly dedicates three chapters to the latter. A consistent methodology is applied in both locations, with the final chapter integrating data from both countries to present a comprehensive view. Utilizing insights from both nations, the dissertation proposes conservation measures suitable for both contexts. The first chapter offers a comprehensive literature review, combining the existing data on Costa Rican cave-dwelling fauna with original field research. Chapter two centers on crucial bat roosts in Bulgaria, emphasizing urgent conservation measures. Chapter three is dedicated to exploring the diversity of cave-dwelling bats in Costa Rica's Brunca region, identifying vulnerable roosts and assessing bat abundance. Chapter four examines the influence of various environmental and anthropogenic factors on cave-dwelling bats, while the last chapter delves into the impact of bat guano nutrients on surface ecosystems, particularly nitrogen and phosphorus transportation in underground streams. This dissertation takes a pragmatic and conservation-oriented approach, yielding scientific research as well as strategic documents, capacity building, and outreach materials. It represents a sustainable, long-term effort aimed at enhancing knowledge and preserving underground ecosystems and cave bat colonies on two continents, fully achieving its objectives and generating additional material for future studies.
Most cave-obligate species (troglobionts) have small ranges due to limited dispersal ability and the isolated nature of cave habitats. The troglobiontic linyphiid spider Phanetta subterranea (Emerton, 1875), the only member of its genus, is a notable exception to this pattern; it has been reported from more counties and caves than any other troglobiont in North America. As many troglobionts exhibit significant genetic differentiation between populations over even small geographic distances, it has been hypothesized that Phanetta may comprise multiple, genetically distinct lineages. To test this hypothesis, we examined genetic diversity in Phanetta across its range at the mitochondrial cytochrome c oxidase subunit I gene for 47 individuals from 40 caves, distributed across seven states and 37 counties. We found limited genetic differentiation across the species’ range with haplotypes shared by individuals collected up to 600 km apart. Intraspecific nucleotide diversity was 0.006 +/- 0.005 (mean +/- SD), and the maximum genetic p-distance observed between any two individuals was 0.022. These values are within the typical range observed for other spider species. Thus, we found no evidence of cryptic genetic diversity in Phanetta. Our observation of low genetic diversity across such a broad distribution raises the question of how these troglobiontic spiders have managed to disperse so widely.
The Kotumsar cave is situated in the Eastern Ghats and has been reported by the existence of 14 different organisms morphologically. To reassess the living taxa and hitherto unreported organisms, intervention of molecular tool is required to corroborate the exact faunal diversity. In the present study, we dealt with the environmental samples and opportunistically encountered living specimens from both deep and transition zones of the Kotumsar cave. The morphological and integrated approach confirmed the existence of Rhinolophus rouxii (Medellin et al., 2017) (bat), Kempiola shankari Sinha and Agarwal, 1977 (cricket), Heteropoda leprosa Simon, 1884 (spider). Further, the collected environmental DNA (eDNA) samples were successfully identified as Fejervarya pierrei (Dubois, 1975) (frog), Indoreonectes evezardi (Day, 1872) (fish), Metrocoris sp. (true bug), Barytelphusa cunicularis (Westwood, 1836) (crab), Trigoniulidae sp. (millipede), and Megascolecidae sp. (worm). Hence, the present investigation through combined approaches by both morphological and molecular data helps to add six more organisms to the faunal checklist of Kotumsar cave. The study also contributed the genetic information of cavernicoles in the global database from India. This genetic information would further help to pursuing other biological studies and adopt better conservation strategies of cave-dwelling organisms and restoration of the colligated ecosystem.
El uso de métodos complementarios de muestreo para murciélagos es recomendable para mejorar los inventarios de especies. En este estudio implementamos muestreo directo e indirecto en cuevas, en el río principal y en distintas coberturas de vegetación forestal en un paisaje kárstico del centro–norte de Guatemala, y determinamos la eficiencia y complementariedad entre métodos. Con redes de niebla y un esfuerzo de 2.670 m²h, capturamos un total de 296 individuos de 19 especies; con micrófonos ultrasónicos y un esfuerzo de 71 horas de grabación, obtuvimos 1.598 archivos efectivos de 16 especies/sonotipos; y con una trampa de arpa y un esfuerzo de 79 horas de operación, capturamos 939 individuos de 12 especies. En total, reportamos 34 especies/sonotipos. Con respecto al porcentaje de la riqueza de especies muestreada, según una extrapolación al doble del esfuerzo de muestreo, este varió entre 69,8 % y 91,5 %, lo cual consideramos satisfactorio. Los porcentajes de complementariedad entre métodos resultaron superiores al 70 %, y fue notablemente alto entre redes de niebla y micrófonos ultrasónicos (91 %). En el país, los inventarios han sido particularmente incompletos, porque están sesgados al uso de redes de niebla y, por ende, a la captura de murciélagos de la familia Phyllostomidae. Es necesaria la capacitación e implementación de métodos que no son utilizados extensivamente, como la detección acústica, para mejorar el conocimiento sobre este grupo taxonómico y brindar información fundamental para las estrategias de conservación.
Little is known about the role of guano from quartzite caves in forming biodiverse plant communities in the surrounding areas due to direct or indirect nutrient and energy inputs. This study investigates four caves in the Quartzite Endokarst of Ibitipoca State Park (PEI), Brazil, by analyzing surface soils from the cave interior to surrounding forests. We evaluated the following hypotheses: (i) guano deposits modify the cave floor and influence external plant communities; (ii) there is a nutrient gradient from the cave interior to its surroundings, linked to guano deposition and redistribution; (iii) forest communities near the caves rely on nutrient cycling from guano to achieve greater biomass and structural complexity. Results confirmed a significant incremental effect of macro and micronutrients in the studied caves, with notable variability between interior and exterior environments and between forested areas and more distant soils. This nutrient input profoundly alters soil morphology and physico-chemical properties through long-term cumulative processes. Many trees and shrubs in these forested areas are larger than their counterparts in the adjacent Rupestrian Quartzite Complex. Although both birds and bats are abundant in these caves, the insectivorous diet of these animals complicates the differentiation of their roles in guano deposition. Further studies using precise biogeochemical markers are needed to clarify the distinct contributions of birds and bats and the temporal dynamics of this phenomenon. Ultimately, this biogenic input creates nutrient-rich oases in an otherwise oligotrophic environment.
As a key life-history trait, growth rates are often used to measure individual performance and to inform parameters in demographic models. Furthermore, intra-specific trait variation generates diversity in nature. Therefore, partitioning out and understanding drivers of spatiotemporal variation in growth rate is of fundamental interest in ecology and evolution. However, this has rarely been attempted due to the amount of individual-level data required through both time and space, and issues with missing data in important covariates. Here we implemented a Bayesian state-space model using individual-level data from 20 populations of Arctic charr (\emph{Salvelinus alpinus}) across 15 capture occasions, which allowed us to: (1) integrate over the uncertainty of missing recapture records; (2) robustly estimate size-dependence; and (3) include a covariate (water temperature) that contained missing data. Interestingly, although there was substantial spatial, temporal, and spatiotemporal variation in growth rate, this was only weakly associated with variation in water temperature and almost entirely independent of size, suggesting that spatiotemporal variation in other environmental conditions affected individuals across sizes similarly. This fine-scale spatiotemporal variation emphasises the importance of local conditions and highlights the potential for spatiotemporal variation in a size-dependent life-history trait, even when environmental conditions are apparently very similar.
Background
We report the first finding of Bathynellacea, discovered in Mongolia. We also report a new species of the genus Altainella Camacho, 2020, which was previously recorded only in the western edge of Russia.
New information
Altainella mongoliensis sp. nov. was found in the interstitial hyporheic region of the Onon River Basin, Mongolia, by inserting a core approximately 1.2 m deep and extracting the underground interstitial water. The new species exhibited sexual dimorphism in thoracopod VII, uniquely observed within the genus Altainella. We provide a morphological description and remarks on the new species with molecular information based on the 18S rDNA and partial CO1 gene sequences. We emphasise the need for continued research on the subterranean biodiversity in previously neglected regions by reporting the first discovery of macro-stygobionts in Mongolia.
Stygobionts, namely animals with strong adaptations to subterranean environments that are unable to complete their life cycles outside groundwater, can be observed in spring ecotones, but their occurrence is generally considered accidental. The aim of this paper is to assess if stygobiont occurrence in springs is linked to specific environmental conditions or if it is random, irrespective of their features. For three years, we surveyed 59 spring sites recording the occurrence of vertebrate and invertebrate stygobiont species and assessing if spring features were related to their distribution. Moreover, we recorded the escape reactions of two easily identifiable stygobiont species. We detected six taxa usually considered as strictly stygobiont based on their troglomorphic features. Two of them were quite widespread: the salamander Proteus anguinus and the shrimp Troglocaris planinensis. Environmental characteristics were significantly related to the distribution of stygobionts. Hydroperiod and occurrence of flooding were the factors that played the strongest role in affecting occurrence. Our study suggests that the occurrence of stygobionts in springs is linked to specific habitat features rather than being a random mechanism and that the exploitation of ecotones can be important for the lifecycle of some species usually assumed to be strictly associated to caves.
Species diversity in caves, can vary according to the ecological
requests of each cavernicolous species and bats are one of the significant
components of the cave ecosystem. Numerous karstic caves suitable for
the cave inventory have been explored and many have been mapped in
Turkey. Various authors have noted that most of the 40 bat species
distributed in Türkiye, preferred karstic caves for nursery,
maternity roosts and hibernaculum. In the present study, one
rhinolophid, one vespertilionid and one miniopterid bat species were
detected in lava tunnels, providing ease of flight and appropriate
temperature, relative humidity and other abiotic factors in the
Mediterranen region of Turkiye for the first time. In addition, this
study also presents the records of porcupine, striped hyena, glasssnail, tarantula and tailles whip scorpion species in the lava tunnel
Keywords Yazıların tüm teknik ve hukuki 37 sorumluluğu yazarlarına aittir. İleri sürülen fikir ve iddialar Doğa ve Sürdürülebilirlik Derneğinin görüşünü yansıtmayabilir. ABSTRACT Species diversity in caves, can vary according to the ecological requests of each cavernicolous species and bats are one of the significant components of the cave ecosystem. Numerous karstic caves suitable for the cave inventory have been explored and many have been mapped in Turkey. Various authors have noted that most of the 40 bat species distributed in Türkiye, preferred karstic caves for nursery, maternity roosts and hibernaculum. In the present study, one rhinolophid, one vespertilionid and one miniopterid bat species were detected in lava tunnels, providing ease of flight and appropriate temperature, relative humidity and other abiotic factors in the Mediterranen region of Turkiye for the first time. In addition, this study also presents the records of porcupine, striped hyena, glass-snail, tarantula and tailles whip scorpion species in the lava tunnels. M AŞAN BAYDEMİR N. (2024). "Hatay ili, Hassa ilçesinde bulunan lav tünelleri ve mağaralarının faunası" Doğa ve Sürdürülebilirlik Derneği, Doğanın Sesi, 7 (13): 37-48 Doğanın Sesi,
Sleep and locomotor rhythms are nearly universal behaviors that are critical for the function of diverse biological processes. While the timing and duration of sleep vary dramatically throughout the animal kingdom, little is known about the genetic or evolutionary basis underlying species-specific differences in these behaviors. Growing evidence suggests differences in sleep duration and timing are influenced by the ecological environment and evolutionary history. Cave environments insulated from the effects of the day-night cycle provide a unique opportunity to investigate the function and evolution of sleep and circadian clocks. Over 200 species of cavefish have been identified throughout the world, allowing for the investigation of trait evolution in a relatively constant environment. The Mexican cavefish and Somalian cavefish, in particular, have emerged as highly useful laboratory models to investigate the mechanistic and ecological basis for the evolution of circadian rhythms. Furthermore, the Mexican cavefish has provided new insight into the evolution and functional significance of sleep. These species display loss of both circadian processes and light detection under very different environmental conditions and timescales. Here, we review recent findings that identify roles for sensory processing, neuromodulation, and DNA repair pathways in the evolution of sleep and circadian rhythms. Furthermore, we highlight the implementation of gene-editing and transgenic tools that allow for mechanistic studies of the neural basis for the evolution of sleep and circadian rhythms. These studies have potential to shed light on the fundamental functions of sleep-wake regulation and the genetic basis for interspecies trait diversity throughout the animal kingdom.
Astyanax mexicanus offers a unique model for studying parallel evolution, with cave-dwelling forms arising from at least two distinct surface lineages. Here, we examined repeated evolution in the two cave-adapted lineages across four phenotypic datasets: (i) meristic data; (ii) ecomorphological data; and geometric morphometric data for (iii) head and (iv) premaxillary bone shape. Additionally, we assessed the association of these phenotypes with their phylogenetic history. Certain traits exhibited evidence of parallel evolution for the cave ecotype across lineages, such as increased maxillary tooth count, reduced lateral line scale count, and shorter ascending process of the premaxillary bone. Geometric morphometric data showed a phenotypic divergence gradient in head shape between cave and surface populations, with the strongest divergence between Sierra de El Abra and surface populations and with less differentiation between surface populations and caves from Sierra de Guatemala and Micos. Cave populations that hybridize with surface populations (e.g. Micos) displayed the greatest overlap with the surface ecotype. There was a similar divergence gradient for premaxillary bone shape, but with weaker differentiation between cave and surface ecotypes. This evidence of repeated evolution in several cave traits emphasizes the significance of selective pressures exerted by the environment on the evolution of cave ecotypes.
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