DOI: 10.3897/subtbiol.58.201189

Authors: William A. Shear, Shiloh McCollum, Jean Krejca, Gretchen M. Baker, Paul E. Marek

Abstract: We describe Bollmanella amorensis sp. nov. from Labor of Love Cave, White Pine County, Nevada, United States. From cave explorations in Great Basin National Park and Humboldt-Toiyabe National Forest in White Pine Co., Nevada, we provide new records of Idagona lehmanensis Shear, 2007, and Nevadesmus ophimontis Shear, 2009, and ecological notes on Labor of Love Cave, a list of species encountered, and remarks on the distinctiveness of caves in the Great Basin Desert.

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DOI: 10.3897/subtbiol.58.198345

Authors: Amazonas Chagas-Jr, Heloisa Barbosa Fernandes

Abstract: A new troglomorphic species of the genus Otostigmus Porat, 1876 has been discovered in two hypogenic caves in Serra da Bodoquena National Park, a hotspot for Brazil’s subterranean fauna. This represents the second troglobitic Otostigmus species in the world. The newly described species, Otostigmus (Parotostigmus) troglobius sp. nov., was based on molecular and morphological data. Although the specimens do not occur in the same cave, species delimitation analysis using a mitochondrial marker Cytochrome C Oxidase subunit I (COI) grouped the new species into a congruent MOTU. Morphologically, the new species closely resembles O. tidius Chamberlin, 1914 due to the median keels on the tergites and the short, incomplete sutures on the sternites. The new discovery underscores the importance of conserving the caves in Serra da Bodoquena.

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DOI: 10.3897/subtbiol.58.192708

Authors: Jozef Grego, Andrzej Falniowski, Sebastian Hofman, Ninon Lecoquierre, Yaser Fatemi, Jean-François Flot, Mohammad Javad Malek-Hosseini, Aleksandra Jaszczyńska

Abstract: The stygobiont gastropods of Iran remain poorly studied. In the present paper, two new species of Hydrobiidae are described from Tuveh seasonal spring, close to Tuveh village, Andimeshk, Khuzestan Province, Iran: one representing a new genus and the other a new species putatively assigned to the genus Pseudamnicola. The shell, soft part morphology, radula, female reproductive organs and penis are described. Phylogenetic analyses based on mitochondrial COI, nuclear histone H3, and ribosomal 28S genes place the new genus as a sister to a clade comprising the Balkan genus Montenegrospeum and Martensamnicola hissarica from Tajikistan. This clade is closely related to the Balkan genus Dalmatinella Radoman, 1973 and the European genus Belgrandia Bourguignat, 1869 within the subfamily Belgrandiinae De Stefani, 1877. Anatomical features also suggest affinities with the Turkish genus Turcorientalia Radoman, 1973, although molecular confirmation is not possible due to the lack of DNA data for the latter. A second new species from the same locality is described based on shell morphology and is provisionally assigned to Pseudamnicola Paulucci, 1878.

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DOI: 10.3897/subtbiol.57.196126

Authors: Saiyu Qiao, Yufeng Zhang, Huifeng Zhao

Abstract: A new earthworm species, Drawida xinjiangensis sp. nov., was described from the northwestern China using an integrative taxonomic approach that combined morphological examination and molecular data. The new species belongs to the Drawida japonica species complex (DjSC) and is distinguished from its congeners by its body length of 33–70 mm, clitellum in X–XIII (white laterally and dark brown medially), two to three gizzards, one pair of testis sacs suspended on septum 9/10 or 10/11, and cylindrical-shaped white prostate glands that are nearly as long as or longer than the testis sacs. Molecular species delimitation: Assemble Species by Automatic Partitioning (ASAP), Generalised Mixed Yule Coalescent (GMYC), and Bayesian Phylogenetics and Phylogeography (BPP) consistently supported its species status assignment. Phylogenetic analyses based on the nuclear 28S rRNA gene and mitogenomic data revealed that D. xinjiangensis sp. nov. forms an independent lineage within the DjSC. This discovery extends the west distribution of moniligastrid species to the arid and semi-arid regions in East Asia and highlights the hidden diversity of earthworms in this under-surveyed area.

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DOI: 10.3897/subtbiol.57.207619

Authors: Ulrich Higl, Dominic Wanke, Arnaud Faille

Abstract: The biodiversity of two caves in the Bavarian Alps, one at low altitude (Angerlloch) and the other at high altitude (Schrecksattel Ice Cave), was studied, with particular attention paid to the relative abundance of trogloxene, troglophile, and troglobitic species. Two excursions were organised for each cave to collect specimens and set pitfall traps for three months. A total of 32 species were identified in the Angerlloch and 16 in the Schrecksattel Ice Cave. Greater total biodiversity was observed in Angerlloch, but most of this fauna consisted of non-troglomorphic trogloxene species. In the Schrecksattel Ice Cave, only 16 species were found, but some of these were more highly adapted to life underground. Our results indicate that the low-altitude cave attracts a greater diversity of species with a low degree of cave adaptation, whereas the high-altitude cave harbours fewer species overall but has a more specialised troglobitic fauna. The results and species lists are presented herein. Furthermore, the occurrence of the springtail Protaphorura septempapillata (Palissa, 1986) was documented for the first time in the Schrecksattel Ice Cave, representing the second cave record for this species beyond its type locality.

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DOI: 10.3897/subtbiol.57.195204

Authors: Hannah R. Rigoni, Chase Corrington, Benjamin T. Hutchins, Benjamin F. Schwartz, Weston H. Nowlin, Annette S. Engel

Abstract: Karst groundwater ecosystems can rely on organic matter (OM) sourced from the surface, but dependance on this can be unreliable depending on how surface water recharges the subsurface. Instead, autochthonous, chemolithoautotrophically-produced OM can also support karst ecosystems. The reduced compounds fueling chemolithoautotrophy can be derived from both OM decomposition and mineral dissolution. The relative contributions of allochthonous versus autochthonous OM to stygofaunal food webs remain unassessed in most settings. Here, we examined OM contributions to metazoan communities in the Barton Springs segment of the Edwards Aquifer (Texas, USA). Geochemical gradients corresponded to variations in microbial community compositions, and biomass-weighted metabolic profiling provided evidence for chemolithoautotrophic OM production, particularly in sulfidic wells where heterotrophic contributions to fine particulate OM (FPOM) were the lowest. Chemolithoautotrophic contributions to OM were calculated using Bayesian isotope mixing models of bulk stable carbon and nitrogen isotope ratios of FPOM, with chemolithoautotrophic contributions as high as 44.8% in one groundwater well. However, bulk δ¹³C of metazoan assemblages were not strongly depleted (< -35‰), and Bayesian isotope mixing models indicated diets primarily supported by allochthonous OM instead of chemolithoautotrophic OM. The dampened isotopic evidence for chemolithoautotrophy may be due to the hydrologic dynamics of the Barton Springs segment, where rapid recharge and relatively short groundwater residence times likely promote allochthonous OM use by stygofauna. These findings highlight the importance of hydrologic context in determining energy pathways in subterranean food webs and have implications for understanding the vulnerability of stygobiontic communities to changes in land use, including urban development.

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DOI: 10.3897/subtbiol.57.198999

Authors: Qingyu Liu, Hao Zhang, Congsheng Li, Ye Chen, Yufeng Zhang, Huifeng Zhao

Abstract: Two new taxa of the earthworm genus Amynthas Kinberg, 1867 are described from northwestern China: Amynthas longnanensis sp. nov. from Longnan, Gansu Province, and Amynthas pectiniferus guanzhongensis ssp. nov. from Weinan, Shaanxi Province. Both belong to the A. sieboldi group, characterized by three pairs of spermathecal pores in 6/7–8/9. Species delimitation based on COI barcoding (ASAP and GMYC) and phylogenetic analysis using mitogenomic protein-coding genes (PCGs) support their recognition as distinct lineages. The mitogenomic phylogeny reveals that the A. sieboldi group as currently defined is not monophyletic. The new findings fill a distributional gap of Amynthas in northwestern China and provide new morphological and molecular data for understanding earthworm diversity.

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DOI: 10.3897/subtbiol.57.202620

Authors: Pablo Barranco, Francisco Javier Sánchez-Camacho, Jaime Mayoral

Abstract: The arachnid order Palpigradi remains poorly studied, particularly with respect to patterns of morphological adaptation to subterranean environments. Subterranean adaptation in palpigrades and comparison with endogean species has been documented scarcely. In this study, Eukoenenia serrana sp. nov. is described as a new species in the E. mirabilis-berlesei group from the cave Sima del Pla de Llomes in eastern Spain. The new species exhibits pronounced troglomorphic features, including large body size and strong elongation of appendicular elements, particularly leg I, while retaining diagnostic characters of the group like the single lateral prosomal organ. We explored the morphological differentiation between endogean and hypogean species in the E. mirabilis-berlesei group, a lineage traditionally regarded as endogean and only marginally associated with caves. An analysis of elongation patterns, with an emphasis on the palp and legs, showed clear differences between species from different habitats using a set of linear measurements and derived ratios related to body and their appendages size. Hypogean species consistently show marked elongation of tibia I, basitarsus IV, and combined tarsus–tibia lengths, whereas changes in overall body characters are more moderate. In contrast, the basitarsus IV/tibia IV ratio remains largely unchanged, indicating proportional elongation of these segments. Highly cave-adapted species in this group retained a single blade in the lateral organ, suggesting that sensory and appendicular traits may evolve independently. The results reveal a mosaic pattern of subterranean adaptation within the E. mirabilis-berlesei group and by extension in palpigrades.

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DOI: 10.3897/subtbiol.57.208540

Authors: Steven J. B. Cooper, Mattia Saccò, Michelle T. Guzik, Andrew D. Austin, Mark S. Harvey

Abstract: not applicable

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DOI: 10.3897/subtbiol.57.193389

Authors: Izabel Cristina Moreira, Laise de Holanda Cavalcanti, Roberth Lopes Marley-Santos, Lucas Leonardo-Silva, Luciene Araújo de Lima, Solange Xavier-Santos

Abstract: More than 26,000 caves occur in the extensive Brazilian territory, distributed across the country’s five geopolitical regions. Some of these caves have been studied for the biodiversity they harbor; however, many groups of organisms are neglected, as is the case with myxomycetes. Herein we present the second record of myxomycetes in Brazilian caves, based on specimens sampled in a cave in the Central-West region, in the state of Goiás. Leaf litter and decaying wood fragments from the photic, penumbra, and aphotic zones were incubated in humid chambers and monitored for 90 days. Of the 22 humid chambers prepared, 45% were positive. Three species were identified: Arcyria cinerea (Arcyriaceae), Cribraria elegans (Cribrariaceae), and Perichaena depressa (Trichiaceae), in addition to one unidentified specimen. This study, the first conducted in Cerrado caves, expands the known occurrence of myxomycetes in Brazilian subterranean environments, reporting Cribrariaceae and P. depressa for the first time in caves and A. cinerea for the first time in a Brazilian Cerrado cave.

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DOI: 10.3897/subtbiol.57.186957

Authors: Kenneth Sparks, Benjamin F. Schwartz, Safra Altman, Kathryn E. Perez, Zoey B. Chanin, Caroline Mierzejewski, Benjamin T. Hutchins

Abstract: Groundwater-obligate taxa (stygobionts) exhibit variable degrees of habitat specialization, ranging from those restricted to single micro-habitats to generalists occurring in a broad range of habitat types. The karstic Edwards Aquifer is the most biodiverse and well-studied groundwater habitat in the continental United States, but some Edwards Aquifer-associated species also occupy the hyporheic zone of rivers downstream from karst. The goals of this research were to 1) assess the potential of the off-karst hyporheic zone as habitat for ‘karst’ species and 2) to determine how stygobiont richness, diversity, and abundance related to distance from karst in the hyporheic zone of a spring-dominated river system. We enumerated stygobionts and quantified environmental characteristics for 55 samples from 11 sites in the San Marcos River between 3 and 140 river kilometers downstream from the rivers’ spring source. Linear mixed effects models and variance partitioning were used to investigate relationships between stygobiont assemblage metrics and environmental parameters (distance from karst, temperature, dissolved oxygen, and hydraulic conductivity). Stygobiont richness and diversity were significantly affected by distance from karst. Stygobiont richness and abundance were significantly affected by hydraulic conductivity. In richness and diversity models, predictive power was low, and additional environmental variables were significant in all models. This study demonstrates that stygobiont communities in the hyporheic zone can be structured by both local environmental conditions, via habitat filtering, and to a lesser extent, by spatial effects (i.e., proximity to source populations in adjacent aquifers). Furthermore, habitat specificity is poorly characterized for stygobiont taxa, inhibiting effective conservation planning and implementation.

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DOI: 10.3897/subtbiol.57.188080

Authors: Andrew G. Cannizzaro, Thomas R. Sawicki, Jacob Schaefer, Dominique Castrillo, Eric C. Maxwell, Elizabeth Burch, David C. Culver, Matthew L. Niemiller

Abstract: Shallow groundwater habitats, including hypotelminorheic seeps, support highly specialized and often narrowly endemic fauna, yet biodiversity in these systems remains poorly documented, particularly in urban landscapes. Here, we describe two new stygobiotic amphipod species, Stygobromus scottsruni sp. nov. and Stygobromus paveki sp. nov., from hypotelminorheic habitats in the Washington, D.C. metropolitan region, and report new locality records for S. anacostensis. Morphological and molecular analyses place both new species within the Stygobromus tenuis species group. Phylogenetic analyses based on mitochondrial (co1+16S) and nuclear (28S) loci, together with multiple species delimitation approaches, consistently support the recognition of both taxa as distinct evolutionary lineages. Despite occurring in close geographic proximity to other congeners, both species exhibit strong genetic differentiation and extremely restricted distributions, consistent with fine-scale isolation among shallow groundwater systems. With the addition of these taxa, described stygobiotic amphipod species richness from the Piedmont and Atlantic Coastal Plain of Maryland, Virginia, and the District of Columbia now totals 16 species, 10 of which are known from in and around the Washington, D.C. metropolitan area, based on currently described taxa. These results highlight hypotelminorheic habitats as important centers of micro-endemism and demonstrate that urban groundwater systems can harbor unexpectedly high levels of cryptic subterranean biodiversity, underscoring the need for targeted surveys and conservation attention to shallow subterranean ecosystems.

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DOI: 10.3897/subtbiol.57.189090

Authors: Canella Radea, Dimitrios Protopapas, Aristeidis Parmakelis, Efterpi Koskeridou

Abstract: A new stygobiont hydrobiid genus and species is described from the Stymphalia karst basin (Peloponnese, Greece). Cyllena hermes gen. et sp. nov. is distinguished from other valvatiform or valvatiform–planorboid stygobiont hydrobiids by a unique combination of shell and anatomical characters, supporting the establishment of a new monotypic genus. It is currently known from a single karstic spring connected to the subterranean hydrological network of the Stymphalia polje, indicating a highly localized distribution. Following a precautionary approach, and in accordance with IUCN criteria, the species is here considered to qualify as Vulnerable (VU D2). The discovery of the new genus contributes to the documentation of subterranean aquatic gastropod diversity in Greece and emphasizes the need for further targeted exploration of karst ecosystems in the region.

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DOI: 10.3897/subtbiol.56.194481

Authors: Naoyuki Nakahama, Takeshi Yamasaki

Abstract: Two new species of Galloisiana Caudell, 1924, G. harimensis sp. nov. and G. kawamurai sp. nov., are described. In general appearance, both new species resemble G. nipponensis (Caudell & King, 1924), the type species of the genus. Molecular phylogeny reconstructed in the present study shows that the new species belong to clades different from that of G. nipponensis. Morphologically, the three species can be distinguished based on the male genitalia, although the females resemble each other.

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DOI: 10.3897/subtbiol.56.183693

Authors: Emre Barlas, Derya Çetintürk, Nuri Yiğit

Abstract: The Cytochrome-b sequences of 41 Myotis capaccinii samples from eight localities in Türkiye were analyzed phylogenetically, taking into account European, Tunisian, and Iranian samples. Nine haplotypes were identified, including previous sequences from Türkiye, and three of these haplotypes were recorded for the first time from two Mediterranean locations. The genetic distance between the Turkish and European samples was 3.0%, ranging from 0.10% to 0.76% among the six Turkish population groups examined. Haplotype diversity (Hd) and nucleotide diversity (Pi) for the Turkish samples were 0.786 and 0.005, respectively, which were quite similar to those for the European samples. In the Bayesian Inference tree, the Bulgarian and Turkish samples were placed in a different clade from other European samples. In the constructed Median-joining network, although common haplotypes were found among the Tunisian, European, Turkish, and Iranian subpopulations, the haplotype differentiation in the Bulgarian and Turkish samples was evident. Accordingly, in previous studies, the European subpopulation was included in the nominal subspecies, while the Bulgarian and Turkish specimens were considered M. capaccinii bureschi. The low genetic diversity among subpopulations is thought to be related to the species’ dispersal ability, with European specimens dispersing into southeastern Mediterranean refugia during the Last Glacial Maximum, exchanging genes with Thracian and Anatolian specimens. Furthermore, the level of genetic diversity within the Bulgarian and Turkish specimens within the same haplogroup provides evidence that no other subpopulation within Türkiye has differentiated at the subspecies level.

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DOI: 10.3897/subtbiol.56.188497

Authors: Shiqi Zhang, Chunyan Xie, Shixiong Ma, Donghui Wu, Yufeng Zhang, Huifeng Zhao

Abstract: To elucidate the species composition and genetic diversity patterns of earthworms in urban green spaces in Northeast China, this study systematically investigated 42 urban parks across the region. Based on mitochondrial cytochrome c oxidase subunit I (COI) gene sequences, the genetic diversity, population structure, and phylogeographic patterns of the dominant species were analyzed. A total of 604 earthworm individuals were collected and identified, belonging to 12 species from seven genera and three families. Genetic analysis of the dominant species revealed that Drawida ghilarovi sensu lato Gates, 1969 exhibited the highest genetic diversity, with its haplotypes forming six independent clades in the phylogenetic tree. The haplotype network displayed a typical reticulate structure with no shared haplotypes, indicating significant population differentiation in this species. Drawida japonica sensu lato (Michaelsen, 1892) showed high haplotype diversity and nucleotide diversity, with a star-like radiation pattern in the haplotype network, suggesting that the population may have undergone a recent rapid expansion event. In contrast, Aporrectodea caliginosa (Eisen, 1874) and Metaphire tschiliensis (Michaelsen, 1928) exhibited relatively low genetic diversity; however, multiple shared haplotypes across geographically distinct populations were detected, which may reflect historical gene flow or shared ancestral origins. This study provides the first comprehensive elucidation of the species composition and genetic diversity patterns of earthworms in urban parks in Northeast China, offering an important foundation for understanding the biogeographic evolution of earthworm communities in urban green spaces in this region.

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DOI: 10.3897/subtbiol.56.188589

Authors: Giampaolo Rossetti, Andrei Ștefan, Rozalia Motoc, Serban M. Sarbu, Ilaria Mazzini

Abstract: Subterranean ecosystems host highly specialized and often endemic crustacean faunas, yet ostracods inhabiting chemically extreme sulfidic caves remain poorly documented. In this study, a new stygobiont species of the genus Pseudocandona is described from a sulfidic cave in Albania using an integrative taxonomic approach combining detailed morphological analyses and molecular data. Subterranean ostracods exhibit a suite of morphological, physiological, and life-history adaptations to the energetic constraints of groundwater habitats, and species thriving in sulfidic systems additionally tolerate high concentrations of hydrogen sulfide and hypoxia. The new species, Pseudocandona sulphurella sp. nov., belongs to the rostrata group, a clade otherwise dominated by surface-water taxa and with only one previously known stygobiont representative. Its valves and soft parts are frequently covered by sulfur-oxidizing bacterial epibionts, a feature shared with other crustaceans from sulfidic environments and potentially indicative of mutualistic interactions. Phylogenetic analyses based on COI and 28S sequences place the new species within a moderately supported clade of Western Palearctic Pseudocandona, while also highlighting the likely polyphyly of the genus. The discovery of P. sulphurella sp. nov. represents the first record of a stygobiont ostracod in Albania and expands current knowledge of biodiversity in chemoautotrophic subterranean ecosystems. This finding underscores the evolutionary significance of sulfidic caves as natural laboratories for studying adaptation, speciation, and the ecological dynamics of groundwater crustaceans.

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DOI: 10.3897/subtbiol.56.188075

Authors: Yue-Ming He, Hong-Quan Xiang, Jin-Mei Yang, Hui Zheng, Suo-Zhu Yang, Yu-Feng Zhou, Jian Zheng

Abstract: This article describes a new species of cavesnail, Erhaia dijiang Y.-M. He, H.-Q. Xiang, Y.-F. Zhou & H. Zheng, sp. nov., collected from a cave in Hongta District, Yuxi City, Yunnan Province, China. This species is a member of the family Erhaiidae, characterized by the absence of eyes. It represents the highest-altitude distribution among known cave-dwelling Erhaiidae species, and is the first freshwater cavesnail discovered in China.

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DOI: 10.3897/subtbiol.56.182808

Authors: Mahmood Mehrafrooz Mayvan, Roman Ozimec, Ľubomír Kováč

Abstract: We provide redescriptions of two subterranean species of the genus Oncopodura inhabiting caves of the Balkan Peninsula, namely O. cavernarum Stach, 1934 and O. jugoslavica Absolon & Kseneman, 1932, partly based on specimens from type localities. Oncopodura cavernarum differs from other species of the genus by a combination of the following characters: single lobe in PAO, unguis elongated, ungual basal lamella shorter than or equal to half the length of the unguis, mucro with 4 teeth and 2 basal scales, and pattern of hooks on dens: 1 dorsointernal hook on basal part, and 1 dorsoexternal and 1 dorsointernal hook on distal part. Oncopodura jugoslavica can be distinguished from other congeners by PAO with 4 equally short lobes, unguis elongated, ungual basal lamella longer than half the length of the unguis, mucro with 4 teeth and 2 basal scales, and pattern of hooks on dens: basal part with 1 dorsointernal hook, and distal part with 1 dorsoexternal and 2 dorsointernal hooks. In both species, the modification of the foot complex shows a moderate level of troglomorphy. Regarding the geographical distribution, these two Dinaric Oncopodura species occur in allopatry: O. cavernarum has north-western and central merodinaric distribution range, while O. jugoslavica is characterized by a south-eastern merodinaric distribution. For both redescribed species, tables with diagnostic characters of morphologically similar congeners are provided.

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DOI: 10.3897/subtbiol.56.190758

Authors: Michael Curran, Syngeon Rodman, Joel Huey, Stephanie Floeckner, Nihara Gunawardene, Morgan Lythe, Dean Main

Abstract: We report the first troglobitic members of the scorpion family Bothriuridae from Australia, discovered in the Pilbara region of Western Australia, and provide a brief review of Australian troglofaunal scorpions. Scorpions are rare in subterranean ecosystems, with only 28 species recorded globally. Two described and two undescribed subterranean scorpion species were previously known from Australia. The Pilbara specimen documented herein exhibit pronounced troglomorphic traits, including complete loss of ocelli and pigmentation and weakened/diminished appendages, consistent with obligate subterranean adaptation. Morphological and phylogenetic analyses place the specimen within Bothriuridae. Photographs, collection details, and molecular data are provided. Together with a previously collected undescribed bothriurid from the Pilbara, this discovery represents the first documented troglobitic Bothriuridae globally. Formal taxonomic treatment will follow in a subsequent dedicated study.

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DOI: 10.3897/subtbiol.56.190752

Authors: Zi-Wei Yin

Abstract: Subterranean ecosystems harbor a high diversity of narrow-range endemics, yet they remain largely invisible in conservation planning, representing a major bias that may trigger irreversible biodiversity loss under rapid climate change. This study applied an Ensemble of Small Models (ESMs) framework, coupled with a target-group background strategy, to assess the climate vulnerability and protection status of Zopherobatrus Yin & Li, 2015 (Coleoptera: Staphylinidae: Pselaphinae), an extremely rare troglobitic rove beetle genus endemic to the karst landscapes of southwest China. Habitat suitability was projected under current conditions and across the 2050s and 2070s (SSP1-2.6 and SSP5-8.5). Under current conditions, highly suitable habitat is strictly confined to specific karst terrains; future projections reveal that the retained habitat ratio declines to only 24.0–48.2%, while the conventional net area change metric (–36.4% to –63.1%) masks functional habitat loss because climatically “gained” areas remain ecologically inaccessible to this dispersal-limited group, and the projected range centroid shifts northwestward by up to 141 km. A systematic gap analysis, cross-referenced with national and provincial nature reserves, showed that only 2.6% of current suitable habitat is formally protected, all known localities fall entirely outside reserves, and 95.9% of identified climate refugia lack any legal protection. These findings quantified a “double jeopardy” for subterranean biodiversity: simultaneous habitat erosion driven by climate change combined with a near-total absence of formal protection. The establishment of targeted micro-reserves centered on climate refugia, the formal inclusion of Zopherobatrus in national protection lists, and the integration of subterranean fauna into regional conservation frameworks are recommended.

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DOI: 10.3897/subtbiol.56.189049

Authors: Najla Baković, Robert Baković, Ferry Siemensma, Tanja Pipan

Abstract: The Dinarides of Bosnia and Herzegovina are exceptionally rich in surface and subterranean geomorphological and hydrological karst features which make them an ideal setting for ecological studies of microorganisms. This work presents the results of a seven-year study of testate amoebae, polyphyletic eukaryotic microorganisms, across various karst habitats. The main objectives were to collect data on testate amoebae biodiversity and to deepen our understanding of the transformation of their assemblages across surface and subterranean habitats. More than 50 taxa of testate amoebae were identified, including the notable finding of Psammonobiotus dinarica – a species originally described from caves – in a pristine surface karst river. A Jaccard resemblance-based cluster analysis of overall testate amoebae biodiversity revealed a clear separation of lake assemblages from river, spring, soil and cave assemblages. These results were confirmed by site-based cluster analysis and NMDS ordination, which showed strong separation (SIMPROF test) of lake, soil and cave assemblages, but also distinct assemblages of hydrologically interconnected karst features (sinking river Šuica → sinkhole cave Ponor Kovači → the resurgence Vrilo Spring). This study confirms a high similarity between river, spring, cave, and soil assemblages but reveals a previously unknown distinction of lake assemblages. It also showed that spring assemblages have higher similarity to river assemblages than to those in caves, implying a possible impact of primary production due to diffuse light in springs during summer droughts. This study emphasizes the need for further research of microbial assemblages in relation to environmental factors in order to support science-based protection of both surface and subterranean karst ecosystems.

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DOI: 10.3897/subtbiol.56.180999

Authors: Lluc Garcia, Adrià Miralles-Núñez, Antonio Cruz-Suárez

Abstract: A new genus and two new species of terrestrial cave-dwelling isopods from the eastern Iberian Peninsula and the Balearic Islands, belonging to the family Philosciidae Kinahan, 1857, are described and fully illustrated. Iberoscia gen. nov. is erected to accommodate Iberoscia zaragozai sp. nov. from Catalonia, Murcia and the Valencian Community. Furthermore, the genus Paractenoscia Taiti & Rossano, 2015 is recorded for the first time in the Ibero-Balearic region with the description of a new species from Ibiza (Balearic Islands). Moreover, the male of Anaphiloscia simoni Racovitza, 1907, is illustrated for the first time based on topotypical specimens from Mallorca, and Anaphiloscia sicula Arcangeli, 1934, is proposed here as a subjective junior synonym of this species. Finally, the monotypic genus Parachaetophiloscia Cruz & Dalens, 1990, is revised and proposed as a subjective junior synonym of Chaetophiloscia Verhoeff, 1908.

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DOI: 10.3897/subtbiol.55.185185

Authors: Ivan N. Marin, Dmitry M. Palatov, Nguyễn Văn Thinh, Louis Deharveng, Lara Konecny-Dupré, Christophe J. Douady, Florian Malard

Abstract: Our current understanding of the molecular systematics of Stenasellidae, a species-rich family of obligate groundwater isopods known from Africa, Asia, Europe and North-America, is based primarily on specimens of the genus Stenasellus Dollfus, 1897 collected in Europe. Here, we provide an updated phylogeny of the family comprising specimens of Stenasellus collected from multiple countries in Asia, including Iran, Turkmenistan, India, Cambodia, and Vietnam as well as specimens of Metastenasellus Magniez, 1979 collected in Africa. First, we used COI-based molecular species delimitation methods to delimit putative species within the family Stenasellidae. Second, we built a phylogeny of the Stenasellidae comprising 48 putative species, for which two mitochondrial and two nuclear genes were available. Third, we describe Stenasellus angiangensis sp. nov. from the hyporheic zone of streams located in mountainous islands of the Mekong River Delta, the first representative of the family Stenasellidae described from Vietnam. We show that the number of putative species delimited using various COI-based molecular species delimitation methods largely surpass the number of species described based on morphological criteria. The four-gene phylogeny reveals, for the first time, a robustly supported and broadly distributed Asian clade comprising Stenasellus species from Iran, India, Cambodia, and Vietnam, including Stenasellus angiangensis sp. nov. as the sister taxon to Stenasellus cambodianus Boutin & Magniez, 1985 from southern Cambodia. Our phylogeny strongly questions the monophyly of the genus Stenasellus, even though the latter cannot be definitively rejected due to difficulties in resolving the phylogenetic relationships among the clades identified within stenasellids.

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DOI: 10.3897/subtbiol.55.188007

Authors: Adrienne Jochum, Jeannette Kneubühler, Eike Neubert, Jacques Chauvin, Sophie Gansoinat, Quentin Wackenheim, Peter Michalik, Bernhard Ruthensteiner

Abstract: Finding live individuals of the minute, subterranean snail genus, Iberozospeum Jochum, Kneubühler, Prieto & Neubert, 2022 is a serendipitous, challenging endeavor. Its diversity has largely been described from caves explored in the Spanish, Cantabrian-Pyrenean part of its range, but only rarely from the French side of the western Pyrenees. Recent exploration has thus targeted cave systems in the Départment Pyrénées-Atlantiques to reveal iberozospeid diversity on the southwest, French Pyrenean side. In this study, four live populations were discovered in four different French caves and subjected to integrative taxonomic study, incorporating new genetic data and 3D X-ray microscopic imagery. We show that the new individuals are genetically and morphologically most similar to I. bellesi (Gittenberger, 1973) and consequently, assign them to this species. 3D X-ray image data of type material of I. bellesi and that of individuals within this study are presented. The new genetic data is presented within the latest phylogenetic reconstruction of Iberozospeum while the prevalence of thriving, live populations of I. bellesi is confirmed in multiple caves for the first time in France.

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DOI: 10.3897/subtbiol.55.184889

Authors: Jure Jugovic, Mohammad-Javad Malek Hosseini, Yaser Fatemi, Jean-François Flot, Matjaž Kuntner, Valerija Zakšek

Abstract: A new cave species of atyid shrimp of the genus Caridina H. Milne Edwards, 1837 was discovered during a survey of a subterranean habitat in southwestern Iran (Khuzestan Province, Tashan Cave). This is the first record of a cave atyid shrimp in Iran and the first documented occurrence of a Caridina species inhabiting sulphidic groundwater. Phylogenetic analyses and genetic distances support the status of Caridina tashanica sp. nov. as a distinct species, closely related to C. shahrazadae. Morphologically, the new species is characterised by reduced eyes and pigmentation, and by an extremely elongated rostrum with numerous teeth. Males have specific, triangular endopodite of pleopod I, and a rod-like appendix masculina of pleopod II with numerous long setae. Ovigerous females lay relatively large, oval, white eggs, which clearly indicate a landlocked life cycle. This study contributes to the knowledge of atyid shrimps in the Middle East and to the subterranean biodiversity of the sulphidic Tashan Cave ecosystem in Iran.

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DOI: 10.3897/subtbiol.55.177524

Authors: Pablo Alberto Hernández-Solis, Víctor Eduardo Gómez-Bretón, Dalia Angélica de la Cruz-Fernández, Nuno Simões

Abstract: For over five decades, the biodiversity of freshwater sponges from Yucatan cenotes has remained largely unexplored. Scientific documentation is limited to studies from 1936 and 1968, which highlights a significant knowledge gap. It is here reported the presence of the freshwater sponge Ephydatia fluviatilis in the Sabak-Ha and Suhem cenotes of Yucatan, Mexico. It is provided detailed descriptions of the spicules to accurately characterize morphology and ease comparison with other Ephydatia species. Additionally, we conducted molecular analyses using DNA sequences from the ITS1, 5.8S, and ITS2 regions, as well as partial 18S and 28S sequences, to determine phylogenetic relationships within the Spongillidae family. DNA sequences from both cenotes were nearly identical to E. fluviatilis GenBank sequences (<3% divergence), confirming species identification. Spicule analysis revealed gemmuloscleres birotules with shafts densely covered by spines as long as the rotular rays as recorded throughout the entire geographic range of this species in both lentic and lotic water bodies. This study expands our knowledge of the diversity and distribution of freshwater sponges in Yucatan cenotes, where only Spongilla cenota and Radiospongilla crateriformis had been previously reported. Our results lay the groundwork for future research on cenote sponge biodiversity in this unique karst ecosystem and underscore the importance of integrating molecular and morphological data for accurate species identification.

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DOI: 10.3897/subtbiol.55.186321

Authors: Cene Fišer, Ester Premate, Valerija Zakšek

Abstract: Epizoic-host species interactions in subterranean environments are rare and poorly understood. A little-known case of epizoic-host interaction involves flatworms (Temnocephalidae) and large subterranean crustaceans (cave shrimps of the family Atyidae and amphipods of the genus Niphargus) living along the Dinaric Karst. The taxonomy and distribution of Dinaric temnocephalids were thoroughly studied in the previous century. There are 15 subterranean temnocephalid species living on different crustacean species in the area. However, analyses of associations between epizoic and host species, as well as co-occurrence of epizoic species, are lacking. In this study, we extracted literature data and analysed epizoic-host interactions using network analysis, complemented by co-occurrence patterns among temnocephalid species. The network analysis indicates low host specificity between temnocephalids and crustaceans; one species is specialized on Niphargus, whereas the others attach to different cave shrimp species. Of these, only one species seems to be specialized to a specific host body part, while the rest can attach to different body parts of cave shrimps. The analysis of co-occurrence patterns implies that many temnocephalid species pairs generally compete for their hosts. We conclude that epizoic-host specificity in subterranean temnocephalids, detectable at the order level, evolved long ago, and that most species cannot be highly specialized due to oligotrophic conditions in caves. As a result, there is substantial competition among them for a host. We discuss potential caveats of the study based solely on the literature and provide a roadmap for future research.

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DOI: 10.3897/subtbiol.55.181079

Authors: Dhanusha Kawalkar, Pooja Patil, Hemant V. Ghate, Shirish Manchi

Abstract: Thread-legged assassin bugs (Emesinae: Reduviidae) are cryptic, slender-bodied predators recognized for their distinctive morphology and varied hunting strategies. Despite their widespread distribution, especially in tropical regions, many species remain poorly studied, particularly those inhabiting subterranean environments. Myiophanes kempi China, 1924, a species previously known only from its type locality in the Siju Cave, Assam (now Meghalaya), had not been encountered for a century. During a systematic survey of limestone caves on the Andaman Islands, a tropical archipelago in the Bay of Bengal, two male specimens of M. kempi were collected in 2019. The specimens were preserved in ethanol, examined morphologically, and identified using established keys and the original species description. This discovery marks the first record of this bug from the Andaman Islands and confirms existence of M. kempi population after 100 years; this extends its known distribution by over 1,000 km southeast of its original locality. The present report provides a redescription of the species, including the first detailed illustrations of the male genitalia. The rediscovery raises intriguing biogeographical questions and emphasizes the ecological importance of caves in the Andaman Islands as a home for this lesser-known assassin bug. This study enhances existing knowledge of cave-dwelling Reduviidae and underscores the significance of cave ecosystems.

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DOI: 10.3897/subtbiol.55.176926

Authors: Maria M. Gomes, Arnaud Faille, Ana Sofia P. S. Reboleira

Abstract: Understanding the composition and functioning of subterranean habitats is particularly critical, as these ecosystems host a rich array of specialized, often endemic species that remain among the least explored and protected globally. We investigated the evolutionary history of endemic troglobiont beetles of the genus Trechus Clairville, 1806, from continental Portugal, that have been recently red listed as Critically Endangered and Endangered of extinction. We sequenced the mitochondrial cox1 gene of three species with restricted distributions: T. gamae Reboleira & Serrano, 2009, T. lunai Reboleira & Serrano, 2009, and T. tatai Reboleira & Ortuño, 2010. Our results show low genetic divergence among them and their hypothesized epigean common ancestor, T. fulvus Dejean, 1831. Despite this, consistent morphological differences, including reduced eyes and distinctive aedeagus shape, suggest divergent evolutionary trajectories and adaptations towards subterranean lifestyle. The unexpected lack of molecular differentiation points to a complex evolutionary history for this group of species. Future research should integrate additional molecular markers and expand taxonomic and geographic sampling, as this approach has the potential to enhance the identification of precise conservation units and, if necessary, improve existing strategies.

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DOI: 10.3897/subtbiol.55.164068

Authors: Brenda Durán, Fernando Álvarez

Abstract: With the aim of describing the isotopic niche of the anchialine shrimps of the genus Typhlatya from the Yucatan Peninsula (YP) we analyze a dataset that includes three of the four species present in the region using published and new δ13C and δ15N values. Typhlatya shrimps are recognized as ecologically important in the anchialine food web due to their ample distribution, relatively high abundance and feeding characteristics. The variation in δ13C values shows that Typhlatya shrimps cover almost the complete interval of the whole anchialine community reflecting the different nutrient sources they can feed on; the three species separately have δ13C ranges that partially overlap, becoming more negative starting with T. mitchelli, followed by T. dzilamensis and T. pearsei. The δ13C ranges for T. mitchelli and T. pearsei from four cenotes did not change between the rainy and dry seasons. By region, the δ13C ranges for the whole anchialine community in the Ring of Cenotes are narrower and less negative than those from the Caribbean Cave Area, reflecting the different geomorphology and hydrology of the two regions. The isotopic niches of T. mitchelli and T. pearsei, the two dominant species in the freshwater lens, overlap around 20% leaving an ample margin for coexistence. Typhlatya pearsei has the most negative δ13C values which are consistent with feeding on microbial biomass, including methanotrophic bacteria. The isotopic niches of the three Typhlatya species studied although somewhat variable, are well defined and compose a well structured community.

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DOI: 10.3897/subtbiol.55.175751

Authors: Thalia Rodovanski, Maria Elina Bichuette, Mauricio Cetra, George Mendes Taliaferro Mattox

Abstract: Brazilian cavefishes are among the richest troglobitic ichthyofauna worldwide including 13 genera. Most cavefish in the Neotropical region belong to Siluriformes. Trichomycteridae are highly diverse siluriforms, and the subfamily Copionodontinae are notably primitive in body shape, showing unique specializations to cave life. We studied the sensorial morphology referring to perception of the environment and processing of this information in four species Copionodon pecten, C. lianae, Glaphyropoma spinosum, and G. rodriguesi, of which G. spinosum is the only hypogean species, involved in perception of presenting some classical troglomorphisms such as reduction of eyes and body pigmentation, in addition to brain morphometry. Previous studies have demonstrated that troglobitic copionodontines are highly agonistic; therefore we measured the barbels, eyes and counted pores of the lateral canal system on the head. A PERMANOVA was performed to test for statistically significant differences in these morphological traits among the species groups. These similarities were significant among both genera and epigean and hypogean populations in terms of external morphometric/meristic and morphometry of the brain. G. spinosum has a longer telencephalon than the other studied species, a region related to olfaction, spatial memory and species-specific behavior such as social interactions and reproductive behavior, a region typically wider in the studied hypogean fishes. We detected troglomorphisms (character states related to isolation in subterranean habitats) in the laterosensory cephalic system of the whole group, such as the reduction of anterior canal segments in relation to other trichomyterids, indicating that all four species have exaptations to inhabit hypogean habitats. Although we could not confirm the existence of morphological differences between the two genera, the hypogean and epigean populations of these endemic Chapada Diamantina cave catfishes may exhibit morphological similarities.

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DOI: 10.3897/subtbiol.55.175661

Authors: Júlio C. C. V. Santos, Manfred Asche, Hannelore Hoch, Rodrigo L. Ferreira

Abstract: A new obligately cavernicolous planthopper species belonging to the Delphacidae was discovered in a cave in Bahia State, Brazil. As no epigean relatives could be identified so far, the species is isolated among the Asiracinae, and accordingly, a new genus Spelaeodelphax gen. nov. is erected to accommodate it. Worldwide, this discovery represents only the fourth record of subterranean Delphacidae and the second confirmed case of a cave-adapted delphacid, and the first such record in the Americas. The Spelaeodelphax nexus sp. nov. is the seventh troglobiont Fulgoromorpha recorded in Brazil. The discovery of this spectacularly troglomorphic species is yet another example of the unique diversity and ecological complexity of biological communities in Brazilian cave environments, and underlines the imperative need for the conservation and in-depth study of subterranean ecosystems.

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DOI: 10.3897/subtbiol.54.168846

Authors: Andrew G. Cannizzaro, Corey J. Lange, David J. Berg

Abstract: In North America, the genus Hyalella (Crustacea: Amphipoda) is primarily epigean, with only two described species found in groundwater. In contrast, Stygobromus is an amphipod genus occupying subterranean habitats throughout most of the Nearctic. Using both molecular and morphological traits, we describe two new species of stygobitic amphipods, Hyalella plutonia sp. nov. and Hyalella keepuikantun sp. nov. from groundwater habitats in Nye County, Nevada, USA. These species form a monophyletic group that also contains the similar H. cretae from Ash Meadows National Wildlife Refuge. Members of this “H. cretae” complex are similar in terms of overall morphology but can be distinguished through examination of taxonomically relevant characters within the genus such as the third uropod, gnathopods, and pereopod 7. The description of these species highlights the unexpected diversity of stygobitic Hyalella within the northern Mojave Desert, where four such species comprise two independent lineages. Stygobromus is absent from these locations. This diversity of Hyalella is hypothesized to be due in part to the evolutionary history the genus and the ecological/climatic conditions of the region itself, which may exclude other more wide-ranging Nearctic stygobites like Stygobromus.

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DOI: 10.3897/subtbiol.54.174031

Authors: Raquel Colado, Alberto Sendra, Susana Pallarés, Jorge Plaza-Buendía, Josefa Velasco, David Sánchez-Fernández

Abstract: Although the effects of climate change on biodiversity are increasingly well documented, subterranean ecosystems remain largely understudied. These environments, characterized by stable temperatures and permanent darkness, offer a unique opportunity to assess the vulnerability of low-dispersal species to warming. While recent studies have shown that subterranean invertebrates are highly sensitive to rising temperatures, most research has focused exclusively on adult stages, overlooking the thermal sensitivity of other life stages. Here, we compared the upper thermal limits (LT50) of larvae and adults of Anillochlamys tropica, a cave-dwelling beetle endemic to the southeastern Iberian Peninsula. Individuals were exposed to constant temperatures ranging from 17 °C to 27.5 °C for seven days in controlled laboratory experiments. Adults tolerated temperatures up to 25 °C but died within 24 h at 27.5 °C. In contrast, larvae were significantly more sensitive, showing reduced survival at 23 °C and 25 °C, and exhibiting a lower LT50 (21.47 ± 0.53 °C) than adults (26.1 ± 0.73 °C). Notably, the current cave temperatures approach the larvae’s thermal limit, meaning that the species could have a narrow thermal safety margin. These findings demonstrate that the early life stages of subterranean beetles are particularly vulnerable to climate warming and underscore the importance of considering all developmental stages when evaluating the impacts of climate change on subterranean biodiversity.

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DOI: 10.3897/subtbiol.54.157228

Authors: Ľubomír Kováč, Marko Karaman, Mahmood Mehrafrooz Mayvan

Abstract: Two new subterranean springtail species of the genus Schaefferia Absolon, 1900 are described: S. greensladeae sp. nov. from Danial Cave in the Alborz Mountains, Iran, and S. montenegrina sp. nov. from Babatuša Cave in Dinaric Karst, Montenegro. Both species are characterized by the absence of eyes and postantennal organ (PAO) having more than 5 lobes in the contrary to other eyeless congeners that have less than 5 lobes in PAO. Morphological traits of both species indicate high level of association with the subterranean environment. The phylogenetic relationships and morphological adaptations of two new species to subterranean life are discussed. Schaefferia greensladeae sp. nov. and S. montenegrina sp. nov. serve as a good example of convergent evolution that is characteristic of invertebrates highly adapted to deep subterranean realm. An updated key to the world species of the genus and a table summarizing diagnostic characters of the world species of Schaefferia are provided.

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DOI: 10.3897/subtbiol.53.157901

Authors: Jorge Plaza-Buendía, David Sánchez-Fernández, Raquel Colado, Enrique Beruete Azpilikueta, Susana Pallarés

Abstract: Climate change is altering the environmental conditions of subterranean ecosystems, leading to warmer and drier caves across many regions. Despite growing evidence of the high heat sensitivity of cave-dwelling arthropods, the role of humidity as a limiting factor remains largely unexplored. To address this knowledge gap, we experimentally assessed the combined effects of temperature and humidity on the survival of four Pyrenean cave beetles (Euryspeonomus eloseguii, Troglocharinus impellitieri, T. hustachei, and Stygiophyes ribagorzanus). Adult beetles were exposed to eight combinations of temperature (11, 20, 23, and 25 °C) and relative humidity (100% and 75% RH) for seven days under controlled laboratory conditions. Survival was monitored daily, and Cox proportional-hazards models were used to analyze treatment effects. Our results showed a significant interaction between temperature and humidity across all species. By the end of the experiment, no individuals survived under 75% relative humidity at any temperature, nor at 23 °C or 25 °C in any humidity condition. Particularly, E. eloseguii and T. impellitieri were highly sensitive to humidity reduction, with no survival beyond 48 hours at 75% RH regardless of temperature. E. eloseguii also exhibited the lowest heat tolerance, with complete mortality at 20 °C even under saturated humidity. Our findings highlight the extreme vulnerability of subterranean beetles to desiccation and heat. Even moderate humidity reductions can drastically reduce survival, underscoring the need to consider combined climatic stressors when assessing the conservation status of subterranean species under climate change.

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DOI: 10.3897/subtbiol.53.162344

Authors: István Urák, Blerina Vrenozi, Zofia Głąbiak, Ninon Lecoquierre, Cord Eiberger, Mark Maraun, Andrei Ştefan, Jean-François Flot, Traian Brad, Luisa Dainelli, Serban M. Sarbu, Raluca I. Băncilă

Abstract: We report the discovery and detailed analysis of an extraordinary colonial spider assemblage in Sulfur Cave, a chemoautotrophic sulfidic ecosystem located on the Albania-Greece border. The colony, comprising an estimated 69,000 individuals of Tegenaria domestica (Agelenidae) and more than 42,000 of Prinerigone vagans (Linyphiidae), spans a surface area of over 100 m²—representing the first documented case of colonial web formation in these species. Stable isotope analyses (δ¹³C and δ¹⁵N) revealed that the trophic web sustaining this assemblage is fueled by in situ primary production from sulfur-oxidizing microbial biofilms then transferred through chironomid larvae and adults to higher trophic levels. Morphological and molecular data confirmed the identity of the two spider species and revealed that their populations in Sulfur Cave are genetically distinct from other populations. Regarding T. domestica, we found a seasonal pattern in fecundity, with significantly larger egg clutches in early summer. Microbiome analysis of this species also revealed a lower Shannon diversity in the cave population compared with a surface individual captured nearby. Our findings unveil a unique case of facultative coloniality in this cosmopolitan spider, likely driven by resource abundance in a chemoautotrophic cave, and provide new insights into the adaptation and trophic integration of surface species in sulfidic subterranean habitats.

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DOI: 10.3897/subtbiol.53.163632

Authors: Rita P. Eusébio, Stefano Taiti, Ana Sofia P. S. Reboleira

Abstract: The Mesovoid Shallow Substratum (MSS) is a largely unknown subterranean habitat formed by the accumulation of rock fragments in steep slopes. In this study, we investigated the diversity, abundance and seasonal dynamics of terrestrial isopods in MSS habitats across Portugal. We identified 12 species belonging to five families, with Porcellionidae and Armadillidiidae being the most represented. The new species, Eluma cristata, endemic to the MSS of Alcobertas, is described. The terrestrial isopod communities included both surface-dwelling and subterranean-adapted species, pointing out the connectivity between these habitats and highlighting the adaptability of terrestrial isopods to diverse environments. Widespread species occurring in multiple locations, and species confined to single sites are present. Seasonality does not seem to affect the community composition. Our findings highlight the rich diversity of MSS habitats, thus their importance for conservation.

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DOI: 10.3897/subtbiol.53.141956

Authors: Vu Dang Hoang Nguyen, Thinh Gia Tran, Truong Minh Nhat Nguyen, Huy Duc Hoang, Thao Thi Phuong Nguyen

Abstract: During explorations conducted in lava caves within Dak Nong UNESCO Global Geopark, Dak Nong Province, Vietnam, from 2020 to 2023, we collected five specimens belonging to the genus Cyrtodactylus. Morphological and molecular analyses identified the population as C. ziegleri. This finding represents an important step in documenting the herpetofauna of Vietnam’s lava caves.

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DOI: 10.3897/subtbiol.53.157293

Authors: Luis Espinasa, Jordi Espinasa, Olivia Castañeda

Abstract: The Allegheny Cave Amphipod (Stygobromus allegheniensis Holsinger, 1967) is a troglobiotic crustacean native to caves in the northeastern United States, including the unique tectonic ice caves of New York’s Shawangunk Ridge. These caves act as natural cold air traps, retaining ice even through summer, creating an extreme environment for aquatic cave-adapted stygobionts. A key question is how these amphipods survive winters when the caves freeze, encasing floors and walls in ice. While past lab studies suggested they can survive freezing, our research shows limitations: individuals fully encased in ice do not survive beyond four hours. However, when surrounded by ice but buffered by a thin layer of liquid water at 0.5 °C, amphipods can endure prolonged exposure to near-freezing temperatures. Field observations confirmed that live specimens persist in the liquid layer beneath the ice and remain active during winter, displaying movement and foraging for food in baited traps. Additionally, specimens varied widely in size, suggesting the presence of multiple reproductive cohorts rather than a single annual reproductive cycle triggered by spring thaw. These findings indicate that reproduction and mortality occur year-round and that adults are capable of surviving the ice cave’s extreme winter conditions.

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DOI: 10.3897/subtbiol.53.152803

Authors: Rostislav R. Borisov, Elena S. Chertoprud, Dmitry M. Palatov, Anna A. Novichkova, Lada V. Vorobjeva, Elizaveta M. Chertoprud

Abstract: A study was conducted on the macrozoobenthos fauna of 15 watercourses flowing through the caves of the Duripshski, Gumistinsko-Panavskiy and Dzhalski speleo districts in Abkhazia (Western Caucasus). Assemblages inhabiting the inner part of caves, entrances and surface biotopes are described. A total of 84 species of aquatic invertebrates were identified in the surveyed streams, with 23 species belonging to the ecological group of stygobionts, 17 classified as stygophiles, and 44 stygoxenes. The species richness and abundance of stygoxenes exhibited a decline from surface stations to stations situated deeper within the caves. Stygobionts were found to be most diverse and abundant within subterranean cavities. The number of stygophile species was found to be highest in the ecotone zone. The similarity of the species structure of macrozoobenthos assemblages was the highest in watercourses flowing from neighboring caves and was mainly determined by the distance factor between sites. At the taxonomic level of families, on the contrary, a large proportion of structural variation in assemblages was attributed to the light factor. Ecotone assemblages, although having their own specificity, occupied a dependent position in relation to surface and subterranean. It is substantiated that illuminance is the primary regulatory factor in the inhibition of the penetration of surface and subterranean faunas into habitats that are not typical for them. The active and passive ways of the invasion of stygobionts into the surface biotopes and of stygoxenes into the subterranean cavities are discussed.

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DOI: 10.3897/subtbiol.53.139979

Authors: Matthew D. Shaw

Abstract: Nullarbor caves were sampled during the Yalata Bush Blitz 2021. Collecting methods commonly used above ground were employed. A large diversity of cave mites and other arthropods were uncovered, including two genera of Neotrombidiidae showing remarkable extremes in larval body form. The first of these genera, Cruralonga gen. nov. (Gracile Velvet mites) has unique features, such as elongate appendages and unusually large numbers of leg sensory setae. Morphotype adults matching these larvae were found widespread in the Nullarbor karst and these also possess putative troglomorphic traits, with longer appendages than previously recorded for the family, and loss of the posterior pair of eyes. Two other species from eastern Australian caves, formerly Neotrombidium, are also transferred to this genus. The second cave-dwelling taxon, has a compact larval body form and a separate set of unique characters such as an expanded anterior region of the scutum and setae on coxa II modified as spurs. It was collected from the folded wings of a trogophilic beetle. Integrative systematics of Australian cave-dwelling Neotrombidiidae could offer a previously unexplored insight into troglomorphic adaptation.

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DOI: 10.3897/subtbiol.53.165710

Authors: Mattia Saccò, Abdul Elmasri, Mahima Tawal, Roman Alther, Florian Altermatt, Alejandro Martínez, Stefano Mammola, Maja Zagmajster, Pablo Saenz-Agudelo, Miwa Takahashi, Jake Thornhill, Aude Andouche, Mariella Baratti, Muhammad Bilal, Helena Bilandžija, Rossano Bolpagni, Sarah Boulamail, Fulvio Celico, Claire A. Chauveau, Efrain Chavez, Olivier Collard, Steven J. B. Cooper, Marjorie Couton, Francesco Cozzoli, Kat Dawkins, Tiziana Di Lorenzo, Ximena Dubinsky, Hans Jürgen Hahn, Jean-François Flot, Fabien Glatting, Christian Griebler, Simone Guareschi, Tamar Guy-Haim, Shane Herbert, Grant Hose, William F. Humphreys, Benjamin Hutchins, Sanda Iepure, Tim Johns, Clemens Karwautz, Dominik Kirschner, Oren Kolodny, Marjeta Konec, Raoul Manenti, Tarik Meziane, Nataša Mori, Elena Pazhenkova, Maxine P. Piggott, Alejandra Prieto-Davó, Zoé Raphalen, Hannah Rau, Ana Sofia Reboleira, Hans Recknagel, Anne Robertson, Coline Royaux, Benjamin Schwartz, Klaus Schwenk, Nuno Simoes, David Stanković, Fabio Stoch, Joe D. Taylor, Mieke Van Der Heyde, Živa Vehovar, Valerija Zakšek, Nicole E. White, Morten E. Allentoft, Paul Nevill, Kathryn L. Korbel, Michelle T. Guzik

Abstract: NONE

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DOI: 10.3897/subtbiol.53.161072

Authors: Ngoc-Son Tran, Khanh Chuyen Phung, Santi Watiroyram

Abstract: A new species of the genus Elaphoidella Chappuis, 1929 has been discovered in the hyporheic zone of a stream in Quang Nam Province, central Vietnam. This represents the second endemic species of Elaphoidella recorded in Vietnam, following E. vietnamica Borutzky, 1967. The newly described species, E. quangnamensis sp. nov., is morphologically most similar to E. brancelji Sanoamuang & Watiroyram, 2021 from Thailand, a member of Group VIII (the sewelli group) sensu Lang (1948). Both species share several key morphological features, including a serrated free margin on urosomites 2–4 dorsally, anal operculum with more than 25 spinules along free distal margin, caudal rami lacking spinules on the inner margin, and similar armature on P1 and P3–P6. The new species differs markedly from E. brancelji in some characters: I) caudal rami parallel in the new species but slightly divergent in E. brancelji; II) caudal ramus length/width ratio in the new species about 1.2 times but 2.0 times in E. brancelji; III) base of caudal ramus with two spinules ventrally but four spinules in E. brancelji; IV) P1 Enp-1 about 1/2 length of Exp-2 but 2/3 in E. brancelji; V) P2 Enp-2 of the new species with three setae but four setae in E. brancelji; VI) P5 baseoendopod of the new species with prominence at insertion of seta II but absent in E. brancelji. The new species represents the sixth known species of Elaphoidella recorded from Vietnam.

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DOI: 10.3897/subtbiol.53.161945

Authors: Gökhan Yürümez

Abstract: The genus Ellobius Fischer, 1814, adapted to subterranean life, consists of five species distributed in the Palaearctic region and only Ellobius lutescens Thomas, 1897 occurs in Turkey. Although karyological, morphological, and biological studies have been conducted on populations of this species in Turkey, there is no study focused on nucleolar organizer regions (NORs) or C-heterochromatin characteristics. Nine specimens (five females and four males) were collected from four different localities in Van and Hakkâri provinces in Eastern Anatolia in this study. Cytogenetic analyses on these specimens confirmed a conserved diploid number (2n = 17) and the unusual sex chromosome system (XO in both sexes). An interstitial NOR on chromosome 1 and numerous telomeric NORs on the other chromosomes were detected by silver nitrate staining. In addition, C-banding analyses detected no C-band heterochromatin in any of the chromosomes, including the X. These cytogenetic results, along with current morphometric and genetic data, indicate subspecies-level variation between Turkish and Iranian populations, emphasizing the influence of physical barriers on micro-evolutionary divergence. This study is the first investigation of NORs and C-heterochromatin in Turkish populations of the subterranean rodent E. lutescens to date. In addition, our study expands the southernmost distribution limit of this species up to Karabey village (Yüksekova-Hakkâri) in Turkey.

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DOI: 10.3897/subtbiol.52.155412

Authors: Tuba Yağcı Gurbanov, Rafig Gurbanov, Uygar Kabaoğlu, Burcu Tekin, Nursel Aşan Baydemir

Abstract: The blind mole rats which have adapted excellently to subterranean life, exhibit remarkable navigation abilities despite structural regression in their visual systems and decline in auditory functions. This study investigates the sensory capabilities enabling blind mole rat possible adaptation to dark environments through gene expression analyses, evolutionary sequence comparisons, and cranial morphometric examinations. Comparative analyses reveal a significant reduction in the expression of genes related to vision (Pax6, Impg) and hearing (Coch, Necap1) in subterranean blind mole rat Nannospalax xanthodon compared to surface dwelling Rattus rattus. In contrast, genes associated with sound perception (Foxp2, Bmp7, Kcnq4, Tmc1) show a significant increase in expression. These findings indicate that sensory losses are aligned with the classical model observed in species that compensate for sensory deficits through echolocation. Remarkably, the gene expression profiles of N. xanthodon show similarities to those of mammals with highly developed echolocation abilities, revealing common molecular patterns between seismic and ultrasonic echolocation. Phylogenetic analyses of echolocation related genes show that blind mole rats share specific amino acid motifs with certain echolocating bat and whale species. Notably, positive selection signal observed in the Kcnq4 gene suggest that this gene may play a critical role in seismic vibration detection. Cranial morphometry studies support the notion that N. xanthodon possess not only putative morphological adaptations specific to their digging lifestyle but also specialized anatomical structures for detecting seismic vibrations. This study provides contributions to understanding the sensory adaptation of subterranean mammals and evaluates blind mole rats in this context for the first time.

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DOI: 10.3897/subtbiol.52.154293

Authors: Michel Perreau, Iva Njunjić, Michiel D. de Groot, Warre Van Caenegem, Danny Haelewaters

Abstract: The only known extinct species of the subfamily Catopocerinae (Leiodidae), Archaeocerus uenoi Perreau, 2019, is redescribed. The female is described for the first time and the internal morphology (genitalia) of both sexes is described and illustrated using X-ray synchrotron micro-tomography and nano-tomography. These new observations allow a detailed description of the aedeagus, ensuring a complementary comparison with extant genera and confirming the placement of the genus Archaeocerus in the subfamily Catopocerinae.

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DOI: 10.3897/subtbiol.52.142334

Authors: Darrell C. Acuña, Lorenz Rhuel P. Ragasa, Myla R. Santiago-Bautista, Volker von Wirth, Leonardo A. Guevarra Jr

Abstract: In 1892, French naturalist Eugène Simon described samples of cave-dwelling tarantula species collected from Culapnitan Caves in the Philippines, a cave system that is now part of the present-time Libmanan Caves Natural Park (LCNP). One of these is the new monotypic tarantula genus and species of that time, Orphnaecus pellitus Simon, 1892. Based on Simon’s notes and quick observation of the eyes of the syntypes, this species is highly suspected to be troglobitic. In the present study, we rediscovered O. pellitus from its reported site and investigated its troglobitic characteristics. Morphological analysis of O. pellitus shows troglomorphic characteristics which include reduced eye size, attenuated limbs, shortened tactile setae, and diminishing pigmentation. Tolerance to hypoxia and heightened sensitivity to ground movements are the other subterranean adaptations that were observed. Phylogenetic analysis revealed a cladistic relationship among tarantula morphologically identified from the genus Orphnaecus. Our findings provide evidence that O. pellitus is a true troglobitic tarantula reported worldwide, and currently the only one known from Asia. We also report two new species of Orphnaecus, Orphnaecus libmanan sp. nov. and the dwarf Orphnaecus tangcongvaca sp. nov., collected from the forest grounds of the LCNP. The ecological niche differentiation of theraphosid species in the LCNP is also provided. Our findings are supported by morphology, molecular phylogeny, and ecology.

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DOI: 10.3897/subtbiol.52.158440

Authors: José D. Gilgado, Hans-Peter Rusterholz, Bruno Baur

Abstract: Invertebrate communities living in the superficial subterranean environment (MSS, Milieu Souterrain Superficiel) have rarely been studied. One possible reason for this is the labor-intensive sampling method that must be used. The most suitable method to collect these arthropods is by using subterranean sampling devices (SSDs, vertically buried tubes with side holes and a pitfall trap at the bottom). However, there is a possibility that surface-active species that encounter the tube from the outside will follow it into the depths and eventually fall through a hole into the trap at a greater depth, which may influence the recorded species composition within a particular depth range. We examined for the first time the depth specificity of SSDs in the scree layer of a rock glacier in the Swiss National Park. We compared the catches from 1 m long standard SSDs over a period of one year with catches from short sections of standard tubes buried at different depths (15–35, 40–60, 70–90 cm) in the scree layer so that they only captured arthropods at their particular depth range. In total, we found individuals of 11 arthropod species and one gastropod species in the different SSDs. Catches from standard SSDs did not differ in species composition from catches from short tubes buried at a depth range of 40–90 cm, confirming the depth specificity of standard SSDs.

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DOI: 10.3897/subtbiol.52.142113

Authors: Giampaolo Rossetti, Serban M. Sarbu, Andrei Ștefan, Rozalia Motoc, Ilaria Mazzini

Abstract: The genus Mixtacandona Klie, 1938 (Crustacea, Ostracoda, Candonidae) includes 21 living non-marine species, all subterranean, with Palearctic distribution. Here we report on Mixtacandona thessalica sp. nov., collected in a sulfidic cave in central Greece. It can be considered an extremophile species because of its ability to thrive in an environment with high concentrations of sulfide and reduced chemical compounds. Mixtacandona thessalica sp. nov. belongs to the species group laisi–chappuisi, one of the five groups in which species of the genus are conventionally placed. A detailed morphological description of the valves and the soft parts of the new species is offered. In addition, COI and 28S genetic markers were sequenced. Mixtacandona thessalica sp. nov. is easily distinguishable from the other species in the genus by its peculiar carapace outline and the marked sexual dimorphism of the posterior margin of valves, as well as by details of chaetotactic characters. The discovery of Mixtacandona thessalica sp. nov. increases the number of non-marine ostracod species known from Greece, which is still rather scarce compared to other Mediterranean countries due to the scarcity of studies. Ten hypogeal species of living ostracods have already been reported from Greece, six of which are considered endemic, and among them three belong to the genus Mixtacandona. It is stressed that a comprehensive review of this genus by combining a thorough morphological approach and molecular techniques, is most needed to assess its phylogenetic relationships within the family Candonidae.

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DOI: 10.3897/subtbiol.52.158075

Authors: David C. Culver

Abstract: none

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DOI: 10.3897/subtbiol.52.154132

Authors: Ștefan Cătălin Baba, Andrei Giurginca, Rodica Plăiașu, Robert Opran, Andreea Mîrleneanu, Octavian Pacioglu

Abstract: The current study presents the first confirmed record of Lithobius peregrinus Latzel, 1880, in Romania, based on specimens collected from Gaura cu Muscă Cave (Banat Mountains). Over a three–year period (2022–2024), 39 specimens were sampled, revealing significant morphological variations, asymmetries, and teratological features. Morphological assessments, including variations in ocelli count, coxosternal teeth, plectrotaxy, and gonopod structures, were conducted. Directional asymmetry was analyzed, showing significant right–side bias in antennal segment counts but no clear asymmetry in other measured traits. Behavioural observations highlighted the close water affinity of this centipede, with individuals frequently found near water bodies, possibly contributing to its dispersal ability. The study also discussed the broader cave affinities of L. peregrinus, comparing its current geographic distribution to related species and highlighting changes in the cave microclimate that may have impacted its subterranean adaptations. These findings contribute to the understanding of the ecological preferences and morphological diversity of the species, enhancing the knowledge of centipede biodiversity in Romania.

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DOI: 10.3897/subtbiol.52.140425

Authors: Violeta Zhelyazkova, Jure Tičar, Stanimira Deleva, Angel Ivanov, Nia Toshkova, Sebastien Puechmaille

Abstract: White-nose disease (WND) is a deadly disease in hibernating bats. Its causal agent is the fungus Pseudogymnoascus destructans (Pd) that was only recently introduced to North America, where it causes pervasive mortality. In contrast, it is native to Eurasia, where bats do not experience mass mortality. The pathogen successfully survives for long periods in cave substrates (e.g., mud, soil) as well as on contaminated caving clothes and equipment. This emphasizes the importance of limiting its human-mediated spread, particularly over long distances and from areas where it is native and genetically highly diverse. To assess associated conservation risks more precisely, we conducted a global online survey on cavers’ WND knowledge, travel frequencies, and hygiene habits regarding caving equipment, and we analysed the relationship between those variables. We interpreted our results based on respondents’ continents of origin, separating them into 3 categories: continents where Pd was recently introduced and continues to cause significant bat mortality (North America); continents where Pd is currently not present but is hypothesized to cause bat mortality if introduced (Australia and South and Central America); and continents where Pd is native and currently not associated with bat mortality (Europe and Asia). We found significant differences between categories, showing that further awareness of WND is most needed in Europe and Asia where specific guidelines for limiting Pd spread are still to be developed. Interestingly, however, WND knowledge alone explained very little variance in respondents’ hygiene habits, showing that other factors such as habits, emotions, or personal values have a stronger influence on cavers’ behaviour.

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DOI: 10.3897/subtbiol.52.140693

Authors: Sarah Boulamail, Salvatore Inguscio, Sara Ventruti, Benedetta Barzaghi, Damiano Brognoli, Raffaele De Giorgi, Joachim Langeneck, Elia Lo Parrino, Emanuele Mancini, Raoul Manenti, Alejandro Martínez, Martina Pulieri, Emanuela Rossi, Francesco Cozzoli

Abstract: The study of hypogean fauna is critical for preserving subterranean biodiversity and ecosystem functions despite climate change. However, underground habitats, often hosting unique and endemic species, present significant challenges for traditional biodiversity assessments owing to their inaccessibility and the specialised nature of their inhabitants, resulting in a paucity of academic studies. Furthermore, even when these studies exist, data are often held in personal notes and databases that are not interoperable according to current standards, making them less usable for research purposes. Reawakening this dormant data, standardising, and sharing it according to modern criteria offers enormous opportunities to expand existing knowledge and provide support for future studies. In the Apulian region of Southern Italy, a biodiversity hotspot for subterranean life, the paucity of recent systematic surveys and reliance on expert knowledge poses both opportunities and challenges for ecological research. By integrating data from various sources, this study provides an overview of the subterranean faunal assemblages documented in this region. Overall, the dataset is comprehensive, comprising 109 species (29 of which are considered endemic to Apulia) and 224 sites, totalling 622 presence-only records. As our records have expanded over 93 years, this dataset represents a unique resource for elucidating the characteristics of Apulian subterranean ecosystems and potentially their changes over time. Integrating observations already published in previous studies with unpublished records, this is the first complete data collection on Apulian subterranean fauna organised according to modern standards of data sharing.

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DOI: 10.3897/subtbiol.52.150357

Authors: Osvaldo Villarreal, Leonardo Delgado-Santa, Carlos A. Lasso

Abstract: Subterranean amblypygids, or whip spiders, are important yet understudied components of cave ecosystems, playing key roles as apex predators in these fragile, nutrient-limited environments. Their restricted distributions make them particularly valuable for understanding cave ecosystem dynamics and conservation needs. However, the diversity and distribution of these arachnids in Colombian caves remain poorly documented. This study examines 53 specimens of amblypygids from 12 Colombian caves, and compiles all available literature records to provide an updated list of cave-associated species in Colombia. A new species of Charinus is described, based on females collected in a cave in Santander. The genus Heterophrynus is recorded from six Colombian caves; H. yarigui is newly recorded from Tolima, and for the first time to hypogean microhabitats from Cueva de Los Guácharos, Cunday, Tolima); H. cervinus is recorded from Boyacá, Las Cacas Cave, while H. batesii is newly recorded from Guaviare Caves in Serranía La Macarena/La Lindosa, Caquetá, and for the first time in hypogean microhabitats. The male gonopodes of H. guacharo are described from topotipic specimens from Natural National Park Los Guácharos in the Suaza River basin. Phrynus araya is recorded for two additional caves in Santander, and finally Paraphrynus laevifrons, is recorded for the first time from Providencia Island and for insular hypogean environments in Colombia from the cave (Cueva Bat Hole, Chay Hill, and Represa Freshwater Bay). These findings expand the distribution of amblypygids in Colombian caves, improve our understanding of their diversity, and provide important data for developing conservation strategies for subterranean ecosystems.

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DOI: 10.3897/subtbiol.51.152663

Authors: Jerry H. Carpenter

Abstract: Amphicutis stygobita Pomory, Carpenter & Winter, 2011 was the world’s first known cave brittle star. It has been found only in two anchialine caves: Bernier Cave (type locality and current study area) and Lighthouse Cave on San Salvador Island, The Bahamas. Bernier Cave’s low salinity (14–28 ppt) reduces ionic precipitation in A. stygobita’s endoskeleton to produce fewer and lighter ossicles. Scanning electron microscopy (SEM) revealed details of internal skeletal structures including elongated arm segment ossicles with greatly reduced density and increased fenestration. The large ceiling entrance of Bernier Cave is directly above the water allowing abundant growth of algae and accumulation of detritus. Small (disk diameter = 3-4 mm) microphagous deposit-feeding brittle stars survived and grew in captivity by consuming energy-rich detritus containing algae, bacteria, invertebrates, and a sticky biofilm containing extracellular polymeric substances (EPS). Reproductive structures are described for this hermaphroditic brooding species, as are morphology and growth rates for adults and three babies born in captivity. Comparisons are made to three recently described cave species that appear to be cave endemics and to several epigean brittle stars including the brackish-water species Ophiophragmus filograneus and two deep-sea species: Amphilepis patens and Amphilepis platytata herein removed from synonymy. Several of these species show paedomorphy, including reduced mouth structures and arm ossicles. Paedomorphy conserves energy by not producing, maintaining, and transporting adult structures not needed for survival. Paedomorphic traits that are adaptive and occur in cave organisms are considered troglomorphic traits, as in A. stygobita. Correlations are made between specific paedomorphisms and environmental features.

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DOI: 10.3897/subtbiol.51.141192

Authors: Miriam Steinitz-Kannan, Jerry H. Carpenter, Mark A. Nienaber

Abstract: The rare endemic brittle star Amphicutis stygobita (Echinodermata: Ophiuroidea: Amphilepididae) is found in Bernier Cave on San Salvador Island in The Bahamas. We report here on the diatom-rich detritus on which the brittle star feeds. The detritus contains a microbial biofilm dominated by a large diatom Campylodiscus neofastuosus Ruck & Nakov that thrives in the cave’s brackish water. The biofilm diatoms and bacteria produce sticky, carbohydrate-rich, extracellular polymeric substances (EPS) that add to the detritus’ nutritional value and help give the detritus a consistency for A. stygobita to pull it into its mouth. Of particular interest is that Campylodiscus neofastuosus has distinct cell wall features that make efficient use of the limited light in the cave. It appears these morphological adaptations might focus light onto the chloroplasts increasing light capture. Furthermore, large, dark, and highly lobed chloroplasts serve as an additional low-light adaptation. This is a rare instance where efficient photosynthetic activity by diatoms produces a complex biofilm that feeds an endemic cave species population.

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DOI: 10.3897/subtbiol.51.139380

Authors: Julio Cifuentes, Enrique Peña Pérez, Álvaro Luna

Abstract: We use a morphological approach to describe a new species of isopod in the genus Baeticoniscus, found so far only in an underground gallery system created during the Roman period, approximately two thousand years ago, located beneath the modern town of Carmona (Seville, Spain). Specimens have been observed inhabiting rotten wood in the aphotic zone. The new species, Baeticoniscus carmonaensis sp. nov. differs from related species in the presence of the eyes as well as the number and arrangement of the tubercles and ribs on the cephalon and pereion. The description of this new species of Baeticoniscus represents one of the few cases worldwide in which the description of a new taxon has been described in a subterranean archaeological site.

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DOI: 10.3897/subtbiol.51.146465

Authors: Jianzhou Sun, Xiangbo Guo, Feng Zhang

Abstract: Three new Tyrannochthonius species are described, including detailed diagnosis and illustrations: T. rudongyanensis sp. nov., T. tiani sp. nov., and T. yanwuensis sp. nov. All samples were collected from the hypogean habitats in Guangxi, China. A distribution map of all Tyrannochthonius species in Guangxi is provided.

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DOI: 10.3897/subtbiol.51.141075

Authors: Luca Coppari, Enrico Lunghi

Abstract: The Monte Albo cave salamander, Speleomantes flavus, is the species endemic to the namesake massif located in the northeastern part of Sardegna, Italy. Speleomantes are the only plethodontid species in Europe, a genus composed of eight troglophilic species living in epigean and subterranean environments. Most ecological studies on these species deal with species occupancy (i.e., presence/absence), while studies aiming to identify drivers for their abundance are lacking. Here, we present the first study aiming to determine which ecological factors influence the abundance of S. flavus. We identified three main hypotheses: (1) salamanders are more abundant where microclimatic conditions are the most suitable for their physiological requirements; (2) Speleomantes are more abundant where prey richness is the highest; and (3) salamanders tend to avoid potential predators. Our results suggested that cave air temperature, humidity, and illuminance are strongly affected by season, and individuals of S. flavus tended to aggregate in relatively cold and humid areas not far from the cave entrance. For most Speleomantes, there was a significant correlation between their abundance and the presence of the considered invertebrate species. This study produced the first information on which ecological features affect the abundance of S. flavus individuals. Additional studies extending to further Speleomantes populations and species may support our hypotheses, including factors not considered here.

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DOI: 10.3897/subtbiol.51.144111

Authors: Hannelore Hoch, Heriberto López, Manuel Naranjo, Dora Aguín-Pombo, Pedro Oromí

Abstract: The Canary Islands harbour a rich and diverse fauna of obligate subterranean arthropods (i.e. troglobionts). Among the insect taxa which have repeatedly undergone the evolutionary switch from life on the surface to underground environments are the Fulgoromorpha, or planthoppers: Cixiidae and Meenoplidae. Previously, a total of 13 troglobitic planthopper species have been described from El Hierro, La Palma, Tenerife and Gran Canaria. Here we describe three new troglobitic cixiid species: Cixius palmirandus sp. nov. from La Palma, Cixius theseus sp. nov. from El Hierro and Tachycixius gomerobscurus sp. nov. from La Gomera, and one new meenoplid species: Meenoplus skotinophilus sp. nov. from El Hierro. Tachycixius gomerobscurus sp. nov. is the first record of a subterranean adapted Fulgoromorpha on La Gomera. With now 17 documented species of strictly hypogean planthoppers, the Canary Islands hold the highest number of subterranean planthoppers of any region worldwide, representing ca. ¼ of all known species. We provide a key to all subterranean planthopper species known from the Canary Islands as well as information on their habitat, distribution, ecological classification and conservation status. As all highly specialized, narrow range troglobitic planthopper species must be regarded as vulnerable, if not endangered, climate change poses a major risk of extinction. We hypothesize on island colonization and subterranean speciation underlying taxonomic diversity and high endemicity. We conclude that the currently observed zoogeographic patterns imply the existence of an ancient fauna which is now extinct.

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DOI: 10.3897/subtbiol.51.151034

Authors: Gonzalo Giribet, Pooja A. Anilkumar, Abbio Goodwin, Ronald S. Stewart, Claud A. Watkis, Damion Whyte, Gustavo Hormiga

Abstract: The velvet worm Speleoperipatus spelaeus Peck, 1975 is one of the rarest velvet worm species reported, as it is only known from its type locality, Pedro Great Cave, Clarendon Parish, Jamaica. The type material of the species, the only four specimens available in known scientific collections, was obtained in the early 1970’s, and since then, no additional specimens have been available for research. More recently, observations of three probably conspecific specimens by the Jamaican Caves Organisation, not collected, have been made in a different location, Swansea Cave, Saint Catherine Parish. Here we report and document five specimens of this rare species from the type locality, Pedro Great Cave, as well as some observations about their behavior. Placing this species in a phylogenetic context should be attempted in the future, to better understand the significance of Speleoperipatus spelaeus and its evolutionary origins, its relationship to the Swansea Cave specimens, and to determine what are its closest relatives and whether those are other Jamaican species or velvet worms from other geographical areas.

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DOI: 10.3897/subtbiol.51.140856

Authors: Jorge Hernández-Lozano, Fernando Córdova-Tapia, Ramses Miranda-Gamboa, María de Lourdes Vázquez-Cruz, Carlos Garita-Alvarado, Norman Mercado-Silva, Claudia Patricia Ornelas-García

Abstract: Stable isotope analysis allows the study of element cycles in ecosystems and trophic ecology. δ13C reflects the diversity of primary productivity, while δ15N is a good indicator of trophic levels of organisms. Caves have limited resources due to the absence of light, reducing the trophic chains in these ecosystems. These extreme conditions impose strong selection pressures on cave-dwelling organisms, known as troglobites, which exhibit specific adaptations such as vision and pigment loss, and metabolic and physiological differences with their surface counterparts. The species Astyanax mexicanus corresponds to a model organism in the study of regressive evolution, which presents two different ecotypes, a widely distributed surface morph, and a cave-dwelling morph present in at least 34 caves in three karst regions of San Luis Potosí and Tamaulipas, Mexico. In the present study, we characterized the δ13C and δ15N of seven cave populations of A. mexicanus, corresponding to two karst regions: Sierra de El Abra and Sierra de Guatemala, representing distinct genetic cavefish lineages. We also developed a Nutrient Input Index (NI), to assess whether cave geomorphology influences resource availability. We found isotopic differences between caves and regions analyzed. Caves in the Sierra de El Abra showed higher δ15N values and wider trophic niche ranges compared with those in the Sierra de Guatemala, reflecting a more complex trophic network tentatively associated also with its geological history. In addition, a relationship was observed between the proximity of pools to the surface and the δ13C values, which could suggest differences in NI directly associated with cave geomorphology, impacting selective forces across the different cave systems.

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DOI: 10.3897/subtbiol.51.147950

Authors: Rosario Ruggieri, Houssein Elbaraasi, Mohammed H. Al Riaydh, Abdelsalam Elshaafi, Awad Bilal, Fathi Salloum, Mohamed Abdelmalik

Abstract: This study confirms the existence of the blind cave shrimp Typhlocaris lethaea Parisi, 1920, in Lethe Cave, Benghazi, Libya, nearly a century after its initial discovery, and documents its new distribution in the Al-Coeffiah caves. Field surveys conducted in 2023 and 2024 revealed its presence in the El-Khadim and Al-Jebah caves, extending its known range by 9 km. Specimens were found in subterranean lakes characterized by complete darkness, with water parameters including an average temperature of 22 °C, pH of 7.67, and salinity of 4.72 ppt. Two specimens were used for further analysis. These findings suggest that the species is more widespread than previously thought and highlight the potential hydrological connections within the karstic system. Additionally, the discovery of a depigmented isopod in El-Khadim cave suggests further hidden biodiversity. Given its IUCN data-deficient status, our findings emphasize the need for conservation efforts to protect these fragile ecosystems from human impacts, ensuring the preservation of Libya’s unique subterranean biodiversity.

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DOI: 10.3897/subtbiol.51.139954

Authors: Stefano Mammola, Jagoba Malumbres-Olarte, Ilaria Vaccarelli, Veronica Nanni, Adrià Bellvert, Ivan Jarić

Abstract: Caves, with their unique geologic and biological features, have inspired human imagination throughout history. From photography to movies, through comics, painting, and poetry, subterranean ecosystems feature prominently in various forms of artistic expression, often incorporating scientific ideas or concepts. Integrating art with science offers a powerful way to convey the uniqueness and importance of the organisms that inhabit subterranean ecosystems, emphasizing their importance as providers of key ecological and cultural services. This, in turn, would help promote their conservation. We discuss realized and unrealized connections between subterranean biologists and artists, aiming to achieve broader protection for subterranean ecosystems. We showcase the different art forms that depict subterranean environments, explain how each aligns with conservation science, and highlight the mutual benefits artists and researchers in subterranean biology can derive from collaboration. In doing so, we pose two important questions: How can we effectively bridge the cultural divide between subterranean scientists and artists? And how can we assess the effectiveness of art in enhancing science communication about subterranean ecosystems and their conservation? By addressing these questions, we envision a future where art and science intersect to safeguard the rich and diverse subterranean biological and cultural heritage.

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DOI: 10.3897/subtbiol.50.138174

Authors: Kathryn E. Perez, Vanessa Saenz, Yamileth Guerrero, Lisa Gonzalez, Evan Guerrero, Pete Diaz, Benjamin T. Hutchins, Benjamin F. Schwartz

Abstract: The freshwater gastropod family Cochliopidae is a diverse component of the subterranean fauna in the karst and hyporheic habitats of western Texas and northern Mexico, with 11 genera and 27 described species found in the region. Building on previous phylogenetic studies of the family in the region we incorporate additional sampling, DNA, and morphological study for systematic revision of species found in the western Edwards-Trinity Aquifer System and associated karst and hyporheic habitats. We describe three new species, Stygopyrgus variabilis sp. nov., Stygopyrgus gracilis sp. nov., and Phreatodrobia embossa sp. nov. and revise the generic placement of two others, Texapyrgus diaboli comb. nov. and Balconorbis coronae comb. nov.

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DOI: 10.3897/subtbiol.50.139914

Authors: Carlos Mario López-Orozco, Ricardo Borja-Arrieta, Yesenia Margarita Carpio-Díaz, Jonas Eduardo Gallão, Ivanklin Soares Campos-Filho, Maria Elina Bichuette

Abstract: Three new troglobitic amphibious species of Xangoniscus (Styloniscidae) have been described from limestone caves of the Bambui Group semiarid ecosystems (Chacoan subregion) in the state of Bahia: Xangoniscus antiquus sp. nov., X. chaimowiczi sp. nov., and X. jonasi sp. nov. Natural history information is provided for X. chaimowiczi sp. nov. and X. jonasi sp. nov. Considering the differences between the known species of Xangoniscus and those described in this study, we redefined the diagnostic characteristics of the genus. Moreover, the distribution of Xangoniscus is discussed. The species described in this study, along with those previously reported, serve as fundamental tools for decision-making processes aimed at conserving Brazil’s speleological natural heritage.

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DOI: 10.3897/subtbiol.50.135200

Authors: Kathryn A. Kennedy, Kirk S. Zigler, Brendan Cramphorn, Curt W. Harden, Kurt Helf, Julian J. Lewis, Thomas E. Malabad, Marc A. Milne, Matthew L. Niemiller, Charles D. R. Stephen

Abstract: 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.

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DOI: 10.3897/subtbiol.50.134613

Authors: Ioseb Natradze, Alexander Bukhnikashvili, Suren Gazaryan, Giorgi Sheklashvili, Levan Mumladze

Abstract: In Georgia, investigations of bat species using caves have been conducted since the beginning of the 20th century. However, robust knowledge is still largely unavailable, as only about 15% of the roughly 2,000 underground sites have been surveyed for bats. The primary objective of this article was to document the use of underground habitats by various bat species. To achieve this, we consolidated field observations with existing data from literature and assessed potential threats to these underground habitats, specifically focusing on caves that harbor significant bat colonies. In this study, we considered underground sites across Georgia where at least one bat species has been documented. In total, twenty out of the thirty bat species recorded in Georgia were found in these underground sites. We characterize the spatial distribution, elevational range, usage of underground habitat, and colony size for these species. These twenty species belong to three families and ten genera. Three of these species have conservation status on the Georgian Red List, and one species is on the Red List of Threatened Species, according to the International Union for Conservation of Nature (IUCN). We also report the spatial and elevational distribution of bat species richness and frequency of records in order to rectify conservation priorities in this important biodiversity hotspot.

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DOI: 10.3897/subtbiol.50.136402

Authors: Ryan M. Jones, Zachary C. Adcock, Andrew R. MacLaren, Kemble White IV

Abstract: In this paper, we describe modifications to a sampling technique for surface, stream-dwelling salamanders for use in subterranean settings. Leaf litter bags are an effective and commonly used trap for salamanders, and their construction purposefully allows animals to move freely in and out of the trap. However, this presents a problem in subterranean deployment because retrieving the trap over long vertical distances, such as well sampling, allows time and space for the animals to escape. To overcome this challenge, we enclosed a leaf litter bag in a suspended net system contained by a lanyard to sample a 3-meter deep well. Our trap modifications resulted in the live capture of adult and immature federally threatened Salado Salamanders (Eurycea chisholmensis) from the well in addition to aquatic invertebrates. This represents a novel trapping technique within a habitat system for which stygofauna sampling options are limited.

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DOI: 10.3897/subtbiol.50.134387

Authors: Jairo Arroyave, Arturo Angulo, Sonia Gabriela Hernández-Ávila, Martín Alonso Buenavad-González, Pablo Rojas-Rodríguez, Stanimira Deleva, Andrés Ulloa, Sophie Picq, Caleb D. McMahan

Abstract: Dedicated ichthyological surveys in two karstic caves in Costa Rica resulted in the discovery of hypogean populations from three epigean species of catfishes of the genus Rhamdia. The taxonomic identity of these populations was initially determined based on morphological traits and subsequently corroborated with comparative DNA sequence data in a phylogenetic framework. Individuals from all hypogean populations documented herein exhibit only partial troglomorphism, characterized by only moderate (vs. complete) integumentary depigmentation without extreme eye reduction/loss. A similar pattern of incomplete troglomorphism at the individual level has been observed in other cave-dwelling species/populations of Middle American Rhamdia, and tentatively attributed to gene flow with and/or incipient speciation from epigean lineages. Since most hypogean forms of Rhamdia derive from/are part of a larger clade of primarily R. laticauda, our discovery of cave-dwelling populations assignable to R. nicaraguensis and R. guatemalensis is noteworthy, particularly in the case of the former, which represents the first taxonomically verified record of a cave-dwelling population of this epigean species. Among our findings is the remarkable discovery of hypogean populations from two different species of Rhamdia (R. laticauda and R. nicaraguensis) inhabiting the same cave (Gabinarraca). This finding is particularly significant because it represents the first time that cave-dwelling populations from different species of Rhamdia are reported to be living in syntopy. Continued discovery of cave-dwelling populations during targeted ichthyological surveys reinforces the notion that our understanding of the diversity of hypogean Rhamdia is incomplete and that sustained exploration and taxonomically sound documentation work are paramount to advancing knowledge about the diversity and evolution of these group of Neotropical catfishes.

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DOI: 10.3897/subtbiol.50.113919

Authors: Soumia Moutaouakil, Marconi Souza-Silva, Lais Furtado Oliveira, Mohamed Ghamizi, Rodrigo Lopes Ferreira

Abstract: 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.

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DOI: 10.3897/subtbiol.49.135303

Authors: Santi Watiroyram, Kamonwan Koompoot

Abstract: Siamcyclops isanus sp. nov. has been found from a cave in northeastern Thailand. It differs from the other member of its genus by having: 1) smooth free margin of anal operculum, 2) a different armature on the second endopod of the first and fourth legs, 3) the armature of the second endopod of the male third leg, and 4) the shape of spermatophore. The discovery of the second species from the genus Siamcyclops thus leads to the proposal for the generic diagnosis. The morphological differences between the genus Siamcyclops and other closely related genera is added.

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DOI: 10.3897/subtbiol.49.120956

Authors: Enrique Baquero, Rafael Jordana, Floren Fadrique

Abstract: The Collembola fauna of the Cova Urbana located under the city of Tarragona (Spain) is studied, and the climatic characteristics of the cave, its partial geomorphology and its planimetry are presented. For the first time in this cave, some specimens of Collembola have been captured and studied, and two new species for science belonging to the genera Coecobrya and Pygmarrhopalites are described and illustrated. In addition, a specimen of the genus Arrhopalites probably belonging to another new species has been studied, but it could not be described because only one specimen was available. The presence of so few specimens and only four species is attributed to the absence of organic matter from percolation and detritus (oligotrophic cave).

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DOI: 10.3897/subtbiol.49.132515

Authors: Mara Knüsel, Roman Alther, Marjorie Couton, Florian Altermatt

Abstract: 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.

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DOI: 10.3897/subtbiol.49.132266

Authors: Johannes Mattes

Abstract: The Moravian Cistercian monk and collector Dominik Bilimek (1813–1884) is considered one of the earliest European naturalists to conduct zoological research on subterranean fauna in Latin America. During the second French invasion of Mexico, from 1861 to 1867, he accompanied Maximilian of Habsburg, the newly enthroned emperor of Mexico, to the region. There, he explored the Grutas de Cacahuamilpa near Taxco de Alarcón, comparing its fauna to his earlier discoveries in Postojnska jama in Carniola (Slovenia). After the victory of the Mexican republican forces and Maximilian’s execution, Bilimek retained his role as curator of the emperor’s collections, which he took back with him to Europe and exhibited at Miramare Castle near Trieste. Prior to his Mexican venture, Bilimek had embraced the imperial goals and intellectual agendas of the Viennese central administration while teaching at the monarchy’s military institutes. Following the revolutions of 1848/49, extensive research undertakings—including geoscientific, biological, and archaeological surveys—were initiated to preserve the Habsburg monarchy as a supranational entity amidst internal and external crises, and to legitimize its territorial framework as both a natural and cultural unit. These developments coincided with the initial surge of zoological interest in Carniolan caves, which spurred efforts to gather similar findings in other karst regions of the monarchy. Drawing on historical sources, this article explores the intersection of political concepts and fieldwork practices in the early study of cave biology. Special attention is given to the sites, networks, and modes of collecting during this transformative period.

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DOI: 10.3897/subtbiol.49.130692

Authors: Petra Lukeneder, Franz Ottner, Mathias Harzhauser, Viola Winkler, Brian Metscher, Bernhard Ruthensteiner, Adrienne Jochum

Abstract: 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).

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DOI: 10.3897/subtbiol.49.125131

Authors: Tiziana Di Lorenzo, Nataša Mori, Tatjana Simčič

Abstract: Research on the metabolic physiology of groundwater species, particularly regarding oxygen consumption rates (OCR), has made significant advancement, revealing valuable insights into the adaptations of exclusively groundwater-dwelling (stygobitic) species. However, a comprehensive understanding of how these metabolic rates scale with body mass and respond to temperature changes remains elusive. This study aims to bridge this gap by reviewing published data on OCR across a variety of groundwater organisms to elucidate patterns of metabolic rates in relation to body size and temperature. We employed a combination of literature review and quantitative analyses, focusing on the allometric scaling of OCR with body weight and the effect of temperature on metabolic rates. Our findings indicate that OCR scales with body weight in an allometric pattern, with an inter-species slope of 0.80, suggesting non-isometric scaling. Furthermore, our analysis showed that stygobitic species’ metabolic rates are less responsive to warming than those of non-stygobitic species at low to moderate temperatures. However, at higher temperatures, metabolic rates in stygobitic species decline faster than in non-stygobitic taxa, highlighting a potential vulnerability to global climate change. This study contributes to our understanding of the metabolic strategies of groundwater species, underscoring the need for further research to fully grasp the eco-evolutionary implications of these findings for groundwater conservation.

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DOI: 10.3897/subtbiol.49.128521

Authors: G. T. Harrison, Howard P. Dunleavy, Luisa F. Vasquez-Valverde, Alejandro I. Del Pozo-Valdivia, Kaloyan Ivanov, Paul E. Marek

Abstract: Subterranean arthropods are important components of soils and contribute essential food-web functions and other ecosystem services, however, their diversity and community composition has scarcely been assessed. Subterranean pitfall traps are a commonly used method for sampling soil habitats in Europe but have never been widely implemented in the Americas. We used subterranean pitfall traps to sample previously unsurveyed arthropod communities in southwestern Virginia, U.S. Traps were placed in shallow subterranean habitats (SSHs), underground habitats close to the surface where light does not penetrate, and more specifically at the interface between the soil and underlying “milieu souterrain superficiel”—a microhabitat consisting of the air-filled interstitial spaces between rocks (abbreviated MSS). In total, 2,260 arthropod specimens were collected constituting 345 morphospecies from 8 classes, 33 orders, and 94 families. A region of the mitochondrial cytochrome c oxidase subunit I (COI) gene was amplified and sequenced, and objective sequence clustering of 3% was used to establish molecular operational taxonomic units (mOTUs) to infer observed species richness. In all, 272 COI barcodes representing 256 mOTUs were documented for rare soil-dwelling arthropod taxa and are published to build a molecular library for future research in this system. This work is the first taxonomically extensive survey of North American soil-dwelling arthropods greater than 10 cm below the soil surface.

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DOI: 10.3897/subtbiol.49.118293

Authors: Mahmood Mehrafrooz Mayvan, Andrea Parimuchová, Ľubomír Kováč

Abstract: A new species of Oncopodura (Collembola, Entomobryomorpha, Oncopoduridae) from the Moghan cave in northeastern Iran, built in a carbonate complex of Kopet Dag mountain range, is described. Oncopodura moghanensis sp. nov. can be distinguished from other congeners by (1) 6 long subequal lobes in PAO, each subdivided into 3–6 fingers, (2) dens with 7 dorsal feather-like macrosetae; at basal part with 1 dorsoexternal and 2 dorsointernal hooks, at the distal part with 1 dorsoexternal and 3 dorsointernal hooks, (3) distal part of manubrium with long feather like macrosetae reaching middle part of dens, and (4) mucro with 4 teeth, apical tooth very sharp, and 2 scales at its basal half. A table with diagnostic characters of species related to the Oncopodura moghanensis sp. nov. and an updated key to the world species of Oncopodura are provided.

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DOI: 10.3897/subtbiol.49.119986

Authors: Luis Espinasa, Abrianna Gutierrez, Amata Hinkle, Matthew L. Niemiller

Abstract: A new genus and species of troglobiotic nicoletiid (Insecta, Zygentoma, Nicoletiidae) is described from northern Alabama, USA. The type species was collected from three caves in the Highland Rim section of the Interior Low Plateau physiographic province on the northern side of the Tennessee River Valley. Morphological and genetic analysis using the mitochondrial 16S rRNA locus show that Spinanycta alabamensis sp. nov. is quite distinct from related nicoletiids in North America. The species differs from members of other genera by its urosternum I, which in males is modified with a central pointy extension. The new species significantly extends the distribution of cave-dwelling members of the family into the southeastern United States and suggests that additional nicoletiid diversity remains to be discovered from karst regions of the eastern United States.

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DOI: 10.3897/subtbiol.49.121747

Authors: Jordi Espinasa, Luis Espinasa

Abstract: The Astyanax fish exhibits two morphs: an eyed, pigmented surface morph and an eyeless, depigmented cave morph. Previous studies have shown that blind morphs swim nearly parallel to the wall and can sense detailed information about objects by gliding alongside them and sensing changes in the flow field around their body using their lateral line sensory system. Hence, cavefish can build hydrodynamic images of their surroundings. Field observations showed that one of their presumptive prey, mysid shrimp, is predominately found not on the floor, but crawling on the walls. In our study, the angle of the body axis with respect to a vertical wall was measured while fish swam in a tank. Results show that when swimming by a wall, cavefish incline the vertical axis of their body away from the wall. But most significantly, this angle is different when the right side or the left side of their body is oriented towards the wall. Intriguingly, cavefish have a leftward-biased dorso-cranial bend, where the convex side of the head is towards their right side. Other studies have shown behavioral “handedness”. When exhibiting Vibration Attraction Behavior (VAB), cavefish in the field show laterality on the preponderant side they circle to explore a vibrating stimulus. Likewise in larval prey capture (LPC) behavior, larvae strike towards prey preferentially located on one side. Our results support that cavefish also express behavioral lateralization during passive swimming by walls and/or when searching for food that is perched on the walls, such as mysid shrimp.

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DOI: 10.3897/subtbiol.48.122645

Authors: Cene Fišer, Gregor Bračko, Teo Delić, Žiga Fišer, Jure Jugovic, Ajda Moškrič, Simona Prevorčnik, Rudi Verovnik, Maja Zagmajster, Valerija Zakšek, Peter Trontelj

Abstract: NA

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DOI: 10.3897/subtbiol.48.117086

Authors: Maxime Le Cesne, Hannelore Hoch, Yalin Zhang, Thierry Bourgoin

Abstract: Planthoppers are an interesting and contrasting model among insects for studying the subterranean environments. Their morphological and ethological adaptations to the underground conditions (complete darkness, lower temperatures, high hygrometry, stability of environmental constants, rarefied food sources, etc.), and their worldwide distribution in both temperate and tropical areas make them an interesting model among invertebrates. In this review, we highlight why cave planthoppers study matters, with particular emphasis on the Cixiidae. The two hypotheses proposed, the ‘climatic relict hypothesis’ and the ‘adaptive shift’, are not sufficient enough to clearly understand and explain the drivers to cavernicoly. Phylogenetic analyses approaches might help to better document and increase our knowledge on such peculiar environments. The singularity of the distribution pattern of the adaptation to cavernicoly in planthoppers raises also interesting questions to investigate and suggest contrasting scenarios to explore further, particularly should the Cixiidae be defined as a subtroglophile lineage?

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DOI: 10.3897/subtbiol.48.112984

Authors: Matthew L. Niemiller, Andrew G. Cannizzaro, Thomas R. Sawicki, David C. Culver

Abstract: We describe a new species of subterranean amphipod (Amphipoda: Crangonyctidae) in the genus Stygobromus from a hypotelminorheic seepage spring at Shepherd Parkway, part of National Capital East Parks, Washington, D.C., USA, part of the National Park System, using both morphological and genetic approaches. The Anacostia Groundwater Amphipod, S. anacostensis sp. nov. is a member of the S. tenuis species group but differs from related congeners based on body size, serrate blade-like edge of both palms of gnathopods 1 and 2, presence of rastellate setae on the posterodistal margin of the carpus of gnathopod 2, and aspects of the second antennae, mandibular palp, pereopods 5–7, uropods 1 and 2, and telson. Moreover, S. anacostensis sp. nov. is genetically distinct from S. tenuis in the Washington D.C. metropolitan area. The description of S. anacostensis sp. nov. increases the number of described Stygobromus species to eight in the Washington D.C. area and highlights the need for continued biodiversity studies, even in regions that have received considerable attention.

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DOI: 10.3897/subtbiol.48.113383

Authors: Danae Karakasi, Kaloust Paragamian, Moissis Mylonas, Katerina Vardinoyannis

Abstract: Greece is covered by a high percentage of carbonate rocks, resulting in a significant number of over 10,000 caves within its territory. Their fauna is not well known. Concerning gastropods, 68 terrestrial species have been reported from 70 caves. In this study, we contribute to the knowledge of the snails inhabiting Greek caves. We analyzed the literature and studied all the material deposited in the Natural History Museum of Crete, to create a first comprehensive list of gastropods of Greek caves. The number of caves from which gastropod species were reported increased to 182, while the number of known terrestrial species increased to 113. Fourteen of these species live only in caves, twelve of which are endemic of Greece. However, the fauna of Greek caves is still understudied. With the increasing exploration of Greek caves, the number of the known cave-dwelling gastropod species will increase and new species will be discovered.

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DOI: 10.3897/subtbiol.48.113022

Authors: Moïssou Lagnika, Raoul Polycarpe Tuekam Kayo, Gontran Sonet, Jean-François Flot, Patrick Martin

Abstract: This study aimed to explore the species diversity within the isopod genus Metastenasellus in Benin and Cameroon. Compared to other parts of the world, the described diversity of stygobiotic crustaceans in Africa is low due to a dearth of studies and taxonomic expertise. However, recent research activities in Benin and Cameroon suggest higher groundwater diversity than previously envisioned. Recent sampling campaigns in these countries have shown that Metastenasellus is a major group in the underground aquatic environment. The accumulation of biological material provided an opportunity to explore species diversity within the genus using a DNA taxonomy approach based on the cytochrome c oxidase subunit I (COI) gene fragment. Despite the limitations of using a single-locus approach for species delimitation, an overview of the diversity within the genus Metastenasellus was obtained, revealing the presence of 23 distinct lineages. Several elements suggest that most, if not all, of these lineages represent valid species. These include high genetic distances between lineages, morphologically distinct species separated by genetic distances of the same order of magnitude as between other described lineages, and the coexistence of different lineages at the same stations. Despite a limited sampling effort, these first results indicate a high level of species diversity and endemism within Metastenasellus in the studied regions. The narrow geographic distribution of the lineages suggests strong isolation and limited dispersal abilities. This study highlights the potential for discovering a significant number of new species within this genus and emphasizes the need for further research to uncover the extent of diversity in African stygobiotic isopods.

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DOI: 10.3897/subtbiol.48.114404

Authors: Curt W. Harden, Robert L. Davidson, Thomas E. Malabad, Michael S. Caterino, David R. Maddison

Abstract: Horologion Valentine, one of the rarest and most enigmatic carabid beetle genera in the world, was until now known only from the holotype of Horologion speokoites Valentine, discovered in 1931 in a small cave in West Virginia. A single specimen of a new species from Virginia was collected in 1991, but overlooked until 2018. DNA sequence data from specimens of this new species, Horologion hubbardi sp. nov., collected in 2022 and 2023, as well as a critical examination of the external morphology of both species, allow us to confidently place Horologion in the supertribe Trechitae, within a clade containing Bembidarenini and Trechini. A more specific placement as sister to the Gondwanan Bembidarenini is supported by DNA sequence data. Previous hypotheses placing Horologion in or near the tribes Anillini, Tachyini, Trechini, Patrobini, and Psydrini are rejected. The existence of two species of Horologion on opposite sides of the high mountains of the middle Appalachians suggests that these mountains are where the ancestral Horologion populations dispersed from, and predicts the discovery of additional populations and species. All specimens of H. hubbardi were collected in or near drip pools, and most were found dead, suggesting that the terrestrial epikarst, rather than caves, is the true habitat of Horologion, which explains their extreme rarity since epikarst has not been directly sampled. We recognize the tribe Horologionini, a relict lineage without any close relatives known in the Northern Hemisphere, and an important part of Appalachian biodiversity.

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DOI: 10.3897/subtbiol.47.113219

Authors: Stanimira Deleva, Andres Ulloa, Hernani F. M. Oliveira, Nikolay Simov, Ferdinando Didonna, Gloriana Chaverri

Abstract: This study focused on the cave fauna of Costa Rica, which has remained relatively understudied despite the presence of more than 435 recorded natural caves and artificial subterranean sites. We collected and reviewed all available literature data on cave fauna in Costa Rica and created the first comprehensive review of the existing information. In addition, we report new records from field surveys conducted between 2015 and 2018. This study reported approximately 123 animal species, whereas the remaining records (n = 82) represented taxa that could not be identified at the species level. Data were collected from 127 locations throughout the country, with new cave fauna records from 41 sites. Notably, we reported the first occurrence of the true bug Amnestus subferrugineus (Westwood 1837) within Costa Rican caves, which represents an addition to the country’s faunal inventory. As this study highlights the knowledge gaps in the subterranean fauna, it will serve as an important stepping stone for future research and conservation efforts related to caves in Costa Rica.

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DOI: 10.3897/subtbiol.47.113186

Authors: Kathryn E. Perez, Yamileth Guerrero, Roel Castañeda, Peter H. Diaz, Randy Gibson, Benjamin Schwartz, Benjamin T. Hutchins

Abstract: The Edwards and Edwards-Trinity Aquifers of Texas have diverse stygofauna, including fifteen species of snails found in phreatic and hyporheic habitats. These species have the hallmarks of adaptation to subterranean environments including extremely small body size and the loss of pigmentation and eyes. Here we use an integrative taxonomic approach, using shell, radula, and anatomical features as well as mitochondrial and nuclear DNA data, to circumscribe a new genus and two new cavesnail species from Central Texas. Vitropyrgus lillianae gen. et sp. nov. is described from Comal Springs (Comal County) and Fessenden Springs (Kerr County) and distinguished by a glassy, highly sculptured shell and distinctively simple, unornamented penial morphology. We also describe Phreatodrobia bulla sp. nov. from Hidden Springs (Bell County), and several other springs in Bell & Williamson Counties, Texas. This species has a smooth, unsculptured teleoconch, a reflected and flared lip, and deeply concave operculum.

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DOI: 10.3897/subtbiol.46.114190

Authors: Aldemaro Romero Jr., Max Moseley

Abstract: Carl Leavitt Hubbs (1894–1979) was a prominent and internationally renowned American ichthyologist whose publications include taxonomic descriptions of several North American blindfishes including the Mexican Cave Characin. His archived personal papers reveal a wide-ranging interest in the biology and evolutionary origins of cave and blindfishes, and his discussions and disputes with colleagues about their taxonomy. He also took opportunities to collect other fauna from American caves during the inter-war decades. Drawing upon his unpublished archive and other relevant sources his biospeleological work is chronicled in detail and discussed in the context of his other work.

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DOI: 10.3897/subtbiol.46.112569

Authors: Ana Sofia Schneider, Mara Knüsel, Florian Altermatt

Abstract: Groundwater is a vast ecosystem harboring a high diversity of specialized taxa. Despite its diversity, groundwater is a still relatively unexplored and threatened ecosystem. Especially the linkage of groundwater with other ecosystems remains largely unknown from the perspective of groundwater fauna. Here, we used citizen science data to get a first baseline knowledge of the occurrence, diversity, and biomass of major macroinvertebrate groups found in shallow groundwater systems of Switzerland. We investigated all organisms collected from the groundwater in 346 spring catchment boxes of municipal drinking water providers. We morphologically identified the organisms on a broad taxonomic level and estimated their biomass and pigmentation using automated image processing. Crustaceans, particularly Niphargus and groundwater isopods, were the most common taxa of obligate groundwater organisms. We also found a surprisingly high number of macroinvertebrates associated with surface and subsurface ecosystems. These taxa might be accidentally entering the groundwater or use it as temporary habitat. In both cases they possibly contribute essential allochthonous energy imports from the surface. We found a positive relationship between the estimated biomass of macroinvertebrates in the samples and the occurrence and abundance of Niphargus. Owing to the widespread occurrences of surface and subsurface macroinvertebrates in our groundwater samples, our study provides evidence for common interactions between groundwater, soil, and surface ecosystems.

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DOI: 10.3897/subtbiol.46.108082

Authors: Job Alí Díaz-Hernández, Paul Ugalde-Silva, Christian Berriozabal-Islas, Alejandro Novelo, Jaqueline Hernández-Uc, Abigail Arana-May, Sheila Denisse Pech-Patrón, Iris Aurora Nava-Jiménez, Jessica Borbolla-Vázquez

Abstract: The microdiversity of cenotes in the Yucatan Peninsula, Mexico has been little studied, with the phytoplankton and protists being the most representative species. However, all previous studies have been focused on cenotes associated with touristic activities, leaving a gap in the understanding of cenotes located within urban areas. The present study is dedicated to the identification of phytoplankton and protists in the cenotes of Cancun, Quintana Roo, Mexico. We conducted our research in four urban cenotes, collecting samples using a 150 µm plankton net, filtering them with a 45 µm membrane, and examining them under optical microscopy. Subsequently, we calculated the abundance, richness, and completeness of the samples. Our findings revealed a total of 6 phyla, 4 subphyla, 10 classes, 8 subclasses, 15 orders, 15 families, 18 genus, and 17 species and 4 species indeterminata in the cenotes of Cancun, Quintana Roo, Mexico. Among these, there were 8 species of phytoplankton and 1 species indeterminata, while 9 species of protists and 3 species indeterminata. These results highlight the remarkable species richness and the complex structure and composition of urban cenotes, suggesting that some species may be unique to this particular ecosystem. Currently, there is limited knowledge regarding the behavior of these aquifers (urban cenotes), and a comprehensive inventory or characterization of their microdiversity is lacking. Such information could be instrumental in the management, conservation, and sustainable use of these valuable aquifers.

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DOI: 10.3897/subtbiol.46.108396

Authors: Mohammad Javad Malek-Hosseini, Jean-François Flot, Yaser Fatemi, Hamid Babolimoakher, Matjaž Kuntner, Oleg A. Diripasko, Dušan Jelić, Nina G. Bogutskaya

Abstract: We report the first finding of the stygobiotic form of the cyprinid fish Garra rufa (Heckel, 1843), discovered in a single locality in southwestern Iran, while the epigean form of the species is widely distributed in western Asia (Iran, Jordan, Lebanon, Turkey, and Syria). We also report a new locality for its hypogean congener, Garra tashanensis, about 5 km east of its type locality. The two species occur in syntopy in outflows of the Tang-e-Ban, a seasonal karstic spring that only has flowing water during winter and spring, when fish individuals are washed from the cave to the surface. Identification of the investigated samples was confirmed by morphological analyses, COI distances, and a phylogenetic tree. These findings suggest the existence of a large karst aquifer in the Tashan area that harbours several cave species of fish, crustaceans, and gastropods and may have considerable conservation implications.

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DOI: 10.3897/subtbiol.46.113328

Authors: Jean-Jacques Geoffroy

Abstract:

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DOI: 10.3897/subtbiol.46.102043

Authors: Luis Espinasa, Aidan Pech

Abstract: Pachón cave in the Sierra de El Abra, in Northeast Mexico, stands out as hosting the world’s most widely studied cavefish population – with over 500 scholarly articles published about the population. Refugio Cave was recently discovered in the El Abra region. This cave hosts the mysid cave shrimp Spelaeomysis quinterensis and the blind cave tetra fish, Astyanax mexicanus. This study aims to understand how the aquatic community of Refugio Cave es related to other cave populations in the area. For this purpose, the Histone H3 gene of mysid shrimps and the OCA2 gene that confers albinism in Astyanax fish was sequenced. Results support that the Refugio and Pachón aquatic communities, which are only 4.5 km away apart, are closely related. Thus, the Refugio Cave population may contribute to better understand the evolutionary history of such an important population and, perhaps, help with Pachon’s cavefish conservation.

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DOI: 10.3897/subtbiol.46.110269

Authors: Martín Alonso Buenavad-González, Jesús M. López-Vila, David Torres-Vázquez, Sonia Gabriela Hernández-Ávila, Kaleb Zárate-Gálvez, Jairo Arroyave

Abstract: Dedicated ichthyological surveys in four active karstic caves in the Mexican state of Chiapas (Grijalva River drainage basin) resulted in the discovery of the same number of hypogean populations of Rhamdia catfishes assignable to two different species: R. laticauda and R. guatemalensis. The taxonomic identity of these populations was initially determined based on morphological traits and subsequently corroborated with molecular data in a phylogenetic framework. For the most part, these newly discovered populations exhibit partial and variable troglomorphism (vs. fixed), a pattern that has been observed in most other cave-dwelling species/populations of Mexican Rhamdia, and possibly caused by gene flow with and/or incipient speciation from epigean lineages. Since most hypogean forms of Mexican Rhamdia derive from/are part of a larger R. laticauda clade, our discovery of cave-dwelling populations assignable to R. guatemalensis is noteworthy and includes the very first record of a R. guatemalensis population with pronounced and widespread troglomorphism. Our discovery of hitherto unrecorded populations of hypogean Rhamdia highlights the continued importance of exploration in the process of documenting subterranean biodiversity, particularly in regions of the world rich with cave systems. Our findings corroborate the notion that, among Neotropical fishes, the catfish genus Rhamdia is one of the most prone and effective at colonizing subterranean habitats and establishing viable hypogean populations.

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DOI: 10.3897/subtbiol.46.105707

Authors: Luis Espinasa, Kayla-Ann Lewis

Abstract: In zebrafish larvae, the first response when detecting prey is an oculomotor behavior; eye convergence. Eye convergence increases the overlap between the visual fields of the left and right eyes to prepare for tracking prey. A high vergence angle is maintained throughout the prey-tracking and capture swim phases, enhancing binocular depth. Since the discovery of eye convergence, hundreds of articles reporting on this behavior in zebrafish have been published. In this study, we found that the larvae of blind tetra cavefish, Astyanax mexicanus, despite being adapted to the absence of visual stimuli due to the lack of light in the cave, have retained the oculomotor behavior of eye convergence in their vestigial eyes. In Astyanax, eye convergence responses can be triggered singlehandedly by vibrations elicited with a glass rod at frequencies similar to those generated by its prey (10–35 Hz). The blind cave tetra offers an intriguing combination of regression of the eye structure, while retaining several of the physiological functions and actions performed in the eye, including light-entrained retinomotor rhythms and eye convergence.

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DOI: 10.3897/subtbiol.46.104548

Authors: Srećko Ćurčić, Nikola Vesović, Maja Vrbica, Slađana Popović, Željko Radovanović, Nina B. Ćurčić, Anatoliy A. Yamashkin, Dejan Radović, Stanislav A. Yamashkin, Sofija Vranić, Tonći Rađa

Abstract: Two new subterranean leiodid taxa of the genus Spelaeobates Müller, 1901 from three pits in northern Dalmatia (Croatia), S. coriniensis sp. nov. and S. coriniensis nonveilleri ssp. nov., are described. The morphological traits of the new taxa are enumerated and illustrated. These two taxa are placed in the subgenus Spelaeobates Müller, 1901. The relationships of these two taxa and their close relatives are clarified. Data on the sexual dimorphism of the two new taxa and on the intrasubspecific variability of S. coriniensis nonveilleri ssp. nov. are presented. We also redescribed S. novaki, the type species of both the genus Spelaeobates and the subgenus Spelaeobates and the closest relative of the newly described species. A key for identification of the taxa of the genus Spelaeobates is included. The new taxa are endemic to the Dinaric Alps of Croatia. Spelaeobates (Pretneriella) kraussi Müller, 1903 and S. (P.) pharensis langhofferi Müller, 1931 were found for the first time outside their type locality.

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