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Abstract

We report data on the spatial structure and seasonal variation of the community of Valmarino cave, a medium sized sandstone cave, located a few kilometres from the coast line, in Central Italy. Due to both its habitat features and its relatively recent geological history, Valmarino cave is only inhabited by terrestrial, troglophilic elements, i.e facultative cave dwellers. By means of monthly censuses and density plot estimates we have investigated species abundance, diversity and their spatial organization, by considering separately samples from different cave sectors. Homogeneous sampling design allowed to compare series of samplings performed in 1974 and 1994. On the whole 21 arthropods and one snail species constitute the cave community. Ordination plots resulting from correspondence analyses of monthly samples outline a distinct spatial and temporal structure. Two main sub-communities can be identified: a inner subcommunity, mainly represented by eu-troglophilic species, showing a remarkable stability throughout the year and an outer sub-community, mainly represented by sub-troglophilic species, showing strong seasonal variation. Both spatial and temporal vectors show similar importance in shaping the community structure. An interesting result of this study is the long term stability of both spatial and seasonal components of the community structure which remained almost identical after 20 years, as shown by the comparison of ordination plots obtained from 1974 and 1994 sampling series. Therefore this study provides empirical evidence of a frequently hypothesised, albeit never demonstrated feature of the cave ecosystem.
... The biological studies trying to consider the ensemble of subterranean organisms, generally focus only on caves used by troglophile and more or less accidental species (Di Russo et al. 1997;Fenolio et al. 2005;Novak et al. 2010;Manenti et al. 2013;Lunghi et al. 2018), while ecological studies dealing with troglobionts often focus only on single species and rarely consider the whole community (Kozel et al. 2019). External seasonality and organic matter or prey abundance in the less deep sectors of caves seem to affect occurrence, abundance and interactions of troglophile species (Mammola et al. 2017;Ficetola et al. 2018). ...
... Climate and environmental variations affected the presence/absence of certain species (e.g. Carchini et al. 1982;Di Russo et al. 1997;Mammola et al. 2015;Bento et al. 2016;Lunghi et al. 2017;Mammola and Isaia 2018), therefore, sampling during different periods all year round was necessary to outline a more complete picture on species diversity and seasonal variations. An increase of fauna diversity was recorded during spring, while lower values were detected in colder months. ...
... This is related to the climate variations and probably to the presence of organic resources brought in from the surface. However, for some species it is probably linked also to their reproductive activity or to the availability of potential preys (Kane 1975;Di Russo et al. 1997;Mammola and Isaia 2018). Fauna variations can also be related to seasonal precipitation changes (infiltrating waters also bring nutrients into the cave) (Bento et al. 2016). ...
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Biological studies on factors shaping underground communities are poor, especially those considering simultaneously organisms with different degrees of adaptation to cave life. In this study, we assessed the annual dynamics and use of both horizontal and vertical microhabitats of a whole community with the aim of understanding whether cave-dwelling organisms have a similar distribution among vertical and ground-level microhabitats and to find out which microhabitat features influence such distribution. We monthly assessed from 2017 to 2018, by direct observation combined with quadrat sampling method on the ground and transects on the walls, richness and abundance of 62 cave-dwelling species in a cave of Northern Italy. Environmental factors such as light intensity, temperature, relative humidity and min-eralogical composition of the substrates were measured during each monitoring session, influencing the dynamics of the whole community and revealing significant differences between ground and wall micro-habitats. A gradient of variation of the species assemblages occurred from the entrance toward inner areas, however, evidence that the dynamics of the walls are very different from those occurring at the ground independent from the distance from the surface are shown. Biodiversity indices highlighted sampling area diversity and a discrete total cave fauna biodiversity with the highest values found near the entrance and the lowest in the inner part of the cave.
... Such studies showed that the cave community tends to change over time, in contrast to what was previously assumed (Poulson and White, 1969). Such changes are usually caused by accidental or transient species that use the caves in part of the year to hibernate, reproduce, as a refuge, among other uses (Di Russo et al., 1999). On the other hand, a long-term study conducted inside an artificial cave in Italy, during seven successive summers, indicated that the entire species composition remained constant (Salvidio et al., 2020). ...
... The species composition was significantly different among the sampling events and changed gradually from the entrance to the inner regions of the cave. Di Russo et al. (1999), studying the spatial structure and periodic variation of the troglophilic community of Valmarino cave (a medium-sized sandstone cave, located in Central Italy), reported similar results on the distribution of fauna in space, identifying communities of entrance and inner regions. However, such authors recorded long-term community stability which remained almost identical after 20 years, a result quite distinct from to the observed in this study. ...
Article
Community stability and carrying capacity are essential core concepts in ecology because they reflect the interactions between organisms and their environment and provide clues to predictability. Since caves are considered stable habitats, we used long-term invertebrate monitoring to understand temporal and spatial carrying capacity in a neotropical cave based on species richness and beta diversity. Invertebrates were monitored from the entrance to the inner parts of the cave in plots (25m long each) in five sampling events over 10 years. Overall, 181 species were recorded. Although the turnover in species composition was considered high, the number of species remained almost constant along time, obviously considering the species oscillations between dry and rainy periods. The partitioning of beta diversity showed that in the rainy period and in the inner regions, the species replacement (turnover) was responsible for the differences in species composition over time. The richness stability and the turnover along time seem to indicate a maximum number of species that may coexist in the cave (∼64 species), probably a proxy of environmental carrying capacity. Then, despite the environmental stability of the caves, the permanent absence of light and scarcity of food resources makes these habitats restrictive to colonization, printing an intense temporal turnover in faunal composition, although with constant richness. Hence, at least for the studied cave, it seems that there is a maximum number of species it can shelter, regardless of which taxa.
... NMDS and Adonis analysis showed that the microbial communities of black stains and unstained surfaces differed significantly from each other and were stable. This parallels the stability of cave environmental conditions (Bourges et al., 2014;Di Russo et al., 1997;Engel and Northup, 2008) and especially the stability of black stains (once formed) in this part of the cave. Black stains in the Absidiole displayed counter-selection of Pseudomonas bacteria (from 73.9% of bacterial sequences to 0.03%) and selection of Exophiala fungi (from 0.4% to 30.4% of fungal sequences), as in Lascaux's Passage (Alonso et al., 2018), but surprisingly the relative abundance of Ochroconis fungi was of the same level in unmarked and stained surfaces (an average of 2%). ...
Article
Anthropization of Paleolithic karstic caves can cause an imbalance of cave microbiota and may trigger formation of wall alterations including black stains. In Lascaux Cave, a previous attempt to mechanically remove black stains was followed by reformation of the stain in months, suggesting that microbial recolonization had taken place. On this basis, we hypothesized that mechanical cleaning (a routine cleaning method for conservation of heritage sites) leaves a residual microbial community that can also serve as pioneer community, i.e. a community of early microbial residents that triggers subsequent microbial successions involved in the reformation of black stains. We monitored post-cleaning microbial recolonization over 19 months in the Apse of Lascaux Cave (France), after using two methods of mechanical cleaning (scalpel alone, or scalpel + sponge). Illumina MiSeq metabarcoding evidenced various taxa i.e. the bacteria Pseudomonas, Pedomicrobium and black-melanized fungi Ochroconis (=Scolecobasidium) during early recolonization of cleaned surfaces, and at later stages the establishment of several other taxa including the bacteria Luteimonas, Chitinophaga and the black fungus Exophiala. Surfaces at 19 months after cleaning were visually and microbiologically different from stained surfaces immediately after cleaned and from unstained surfaces, but also from non-cleaned stained surfaces, probably because of a particular microbial succession, distinct from the original succession during stain formation. Variations in relative abundance of Bacteroidota and Eurotiomycetes classes and Exophiala genus were higher when the sponge was used in addition to the scalpel. The bacteria Filomicrobium and the fungi Isaria and Cephalotrichum were identified on sponge-cleaned surfaces and on the sponge itself, pointing to a contaminant status due to the cleaning method. Overall, it suggests that post-cleaning pioneer communities may play an important role in orienting stain reformation in caves. Sponges routinely used by restorers to curb microbial stains may bring microbial contaminants, which questions current cleaning practices in show caves.
... However, even if a high spatial heterogeneity was demonstrated in the composition of this fauna, this, by itself, does not mean that there would or would not be evidence of gene flow between populations of the same species, since the data tested belong exclusively to the superficial layer of the MSS. Furthermore, although some studies have discussed that IF caves with more troglobiotic species have lower species replacement values, considering them more ecologically "stable" (Di Russo et al. 1997;Mammola et al. 2019), contrary to our expectations, the values of total β-diversity in the MSS were high for both troglobiotic and non-troglobiotic animals. This fact reinforces the idea of moderate-to-low ecological connectivity between the sampling sites in the MSS, even if the environment presents favorable conditions for the evolution and permanence of the fauna. ...
Article
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Iron Formations (IF) are among the most threatened environments due to the extensive mining activities. Mesovoid Shallow Substratum (MSS) in IF represents a poorly known subterranean environment and evaluating its fauna has the potential for expanding knowledge about the distribution of troglobiotic populations. We evaluated the spatiotemporal distribution of the subterranean fauna in the MSS of IF in Brazil. We sampled the MSS invertebrate fauna and described the community patterns of troglobiotic and non-troglobiotic species. A total of 22,821 individuals and 276 morphospecies belonging to two phyla were found: Annelida and Arthropoda. Acariformes, Diptera, Hymenoptera, Blattodea, and Collembola represented 92.2% of the individuals sampled. Nine troglobiotic morphospecies belonging to four groups were sampled: Araneae (1), Entomobryomorpha (6), Poduromorpha (1), and Pseudoscorpiones (1). We found a high compositional dissimilarity of troglobiotic and non-troglobiotic species in terms of spatial β-diversity (among MSS sites) and temporal β-diversity (among months). The observed spatial β-diversity of troglobiotic species sampled in the MSS is greater than that of non-troglobiotic species. The temporal variation is similar for both groups. The richness difference component contributed more to spatial and temporal β-diversity for troglobiotic species, while higher replacement values for non-troglobiotic species were observed. Average values of temporal β-diversity and the replacement component were greater for non-troglobiotic than for troglobiotic species, while the richness difference component had an opposite pattern. The spatiotemporal β-diversity patterns suggest a medium-to-low connectivity of invertebrate populations that colonize the MSS, favoring the adoption of strategies for conserving broader areas in the context of IF.
... Estudos que buscam compreender variações composicionais em ambientes subterrâneos costumam comparar comunidades em diferentes períodos do ano, como o período chuvoso e o período seco (LUNGHI et al., 2015;BENTO et al., 2016;MAM-MOLA et al., 2020), ou são realizados em curto período (TOBIN et al., 2013), evidenciando variações consistentes em relação à composição faunística em virtude da sazonalidade. Nesse sentido, nossos resultados corroboram com estudos que investigaram essa variação nas comunidades entre sucessivos períodos de seca (DI RUSSO et al., 1997;SALVIDIO et al., 2020;SIMÕES et al., 2022). ...
Conference Paper
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In this study, we used a multimodal statistical approach to assess the influence of physical, trophic, and shelter components on the spatial and temporal structuring of terrestrial invertebrate communities in limestone caves located in Iuiú and Malhada, southwestern Bahia. Seven caves were sampled during two distinct events, in 2016 and 2021, both during the dry season. 1 x 1 m quadrants were used for invertebrate collection and to measure organic and inorganic substrate components. Species richness and composition remained constant between sampling events, indicating that caves may exhibit a stable ecological carrying capacity over time. The increase in species richness was influenced by the amount of boulders in both sampling events, highlighting the importance of environmental heterogeneity even in the long term. These results emphasize the importance of maintaining key elements for the conservation of these communities.
... a. Hervant 2011). Di Russo et al. (1997) zeigten allerdings in einer küstennahen Höhle in Mittelitalien, dass 21 Arthropoden-und eine Schneckenart die Höhle bewohnten, darunter eu-troglophile Spezies wie auch fakultativ Höhlen nutzende Arten. Interessant hierbei ist jedoch der Fakt, dass diese Lebensgemeinschaft an Tieren in Anzahl und Artenzusammensetzung für 20 Jahre nahezu stabil, sprich unverändert blieb. ...
Article
Grottenolme Proteus anguinus (Laurenti, 1768) zeigen eine sehr geringe Fortpflanzungsrate, werden sehr alt und haben eine im Vergleich zum Menschen verzögerte Generationenfolge. Damit sind Olme Lebewesen für spannende evolutionsbiologische Fragestellungen, die sich um Populationswachstum drehen. Die vorliegende Arbeit zeigt auf, dass eine stabile Umwelt, wenig Krankheiten, das Fehlen von Fressfeinden und ausreichende Nahrungsressourcen evolutiv einen starken Einfluss auf das fehlende Populationswachstum beim Olm haben. Ein Vergleich mit dem Menschen eröffnet, neben der Bildung und dem Aufbrechen von Bräuchen, auch Wege naturgegebener Mechanismen, wie das Wachstum der Weltbevölkerung sich stabilisieren ließe. Der verzögerte Eintritt in die Reproduktion zeigt beim Grottenolm zudem einen positiven Effekt auf die Langlebigkeit. Human fish and human beings – European Cave Salamander Proteus anguinus (Laurenti, 1768) explains longevity and population growth Abstract: European Cave Salamanders, more specifically olms, show a low reproduction rate, reach methuselah ages and exhibit extended generation frequencies in comparison to humans. Olms are therefore considered an interesting study object for evolutionary questions about population growth. The paper discusses stable environment, good health, top predator position and continuous basic food supply as the key factors for stagnation of population growth in olms as well, if combined with education, as in humans. A late start of reproduction triggers in turn a positive impact on longevity.
... glaphyroptera in Luxembourg caves, representing at the same time the first record of Cs. glaphyroptera in Luxembourg ( Table 2). Studies of subterranean mosquito populations in Europe are available from Austria [30,33,34], Germany [23][24][25][26][27][28][29]33], Croatia [35], Czech Republic [34,36,37], France [38], Hungary [18,34], Italy [39], Luxembourg [16], Poland [40], Norway [17], Slovakia [41][42][43], and Sweden [44] (Table 2). Several mosquito taxa are reported from subterranean habitats at larger geographical scales (i.e., spanning more than three European countries), including Anopheles maculipennis s. l., Cs. alaskaensis, Cs. annulata, Cs. glaphyroptera, and members of the Cx. ...
Article
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The common house mosquito, Culex pipiens s. l. is part of the morphologically hardly or non-distinguishable Culex pipiens complex. Upcoming molecular methods allowed us to identify members of mosquito populations that are characterized by differences in behavior, physiology, host and habitat preferences and thereof resulting in varying pathogen load and vector potential to deal with. In the last years, urban and surrounding periurban areas were of special interest due to the higher transmission risk of pathogens of medical and veterinary importance. Recently, surveys of underground habitats were performed to fully evaluate the spatial distribution of rare members of the Cx. pipiens complex in Europe. Subterranean environments and their contribution to mosquito-borne pathogen transmission are virtually unknown. Herein, we review the underground community structures of this species complex in Europe, add new data to Germany and provide the first reports of the Cx. pipiens complex and usually rarely found mosquito taxa in underground areas of Luxembourg. Furthermore, we report the first finding of Culiseta glaphyroptera in Luxembourg. Our results highlight the need for molecular specimen identifications to correctly and most comprehensively characterize subterranean mosquito community structures.
... It is likely that D. lustriae is a subtroglophile species that uses caves and cave-like habitats as a shelter, but requires external ventures to complete its life-cycle. This contrasts to most of its Greek congeners, that are eutroglophiles, i.e. can sustain permanent populations in caves (Di Russo et al. 1999). ...
Article
We record Dolichopoda lustriae Rampini & Di Russo, 2008 (Orthoptera: Ensifera: Rhaphidophoridae) from the Pindus mountain range in Epirus, Greece, more than 200 km to the north of the type locality in Aetolia-Acarnania. We show that this species has an unusually broad distribution for a Balkan cave cricket, and can also live independently of caves, instead inhabiting the crevices of large rock formations. We found D. lustriae to be sympatric with Troglophilus zorae Karaman & Pavićević, 2011, whose presence in Greece is recorded for the first time in this study. We provide high-quality photographs of the habitus and genitalia of D. lustriae and re-describe its morphology. Detailed information about the habitat, behaviour of the D. lustriae is also provided, and possible measures for its conservation are suggested.
Article
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Iron Formations (IF) are threatened by mining, particularly the Mesovoid Shallow Substratum (MSS), an understudied subterranean environment. We evaluate the spatiotemporal patterns of subterranean fauna in MSS of iron duricrust (canga) in the Iron Quadrangle and Southern Espinhaço Range, southeastern Brazil. Samplings took place between July 2014 and June 2022 using five trap types. We sampled 108,005 individuals, 1,054 morphospecies, and seven phyla, globally the largest dataset on MSS in IF. Arthropoda represented 97% of all invertebrates sampled. We identified 31 troglomorphic organisms, primarily Arthropoda and Platyhelminthes. MSS traps were the most efficient method, capturing 80% of all invertebrates. Morphospecies were more prevalent in each locality than shared among localities. Species replacement was the main processes to spatial differences. Over time, we found a decrease of total dissimilarity and importance of species replacement for troglomorphic organisms. A positive correlation between spatial distance and compositional dissimilarity of invertebrates was found. Iron Quadrangle and Southern Espinhaço Range showed marked differences in the spatiotemporal patterns of subterranean fauna. Brazilian IF are threatened, with their biological significance not fully understood but highly endangered due their limited distribution. Conservation efforts require a comprehensive understanding of both biotic and abiotic factors shaping the entire IF ecosystem.
Article
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Community assembly processes are complex and understanding them represents a challenge in microbial ecology. Here, we used Lascaux Cave as a stable, confined environment to quantify the importance of stochastic vs deterministic processes during microbial community dynamics across the three domains of life in relation to an anthropogenic disturbance that had resulted in the side-by-side occurrence of a resistant community (unstained limestone), an impacted community (present in black stains), and a resilient community (attenuated stains). Metabarcoding data showed that the microbial communities of attenuated stains, black stains, and unstained surfaces differed, with attenuated stains being in an intermediate position. We found four scenarios to explain community response to disturbance in stable conditions for the three domains of life. Specifically, we proposed the existence of a fourth, not-documented yet scenario that concerns the always-rare microbial taxa, where stochastic processes predominate even after disturbance but are replaced by deterministic processes during post-disturbance recovery. This suggests a major role of always-rare taxa in resilience, perhaps because they might provide key functions required for ecosystem recovery. IMPORTANCE The importance of stochastic vs deterministic processes in cave microbial ecology has been a neglected topic so far, and this work provided an opportunity to do so in a context related to the dynamics of black-stain alterations in Lascaux, a UNESCO Paleolithic cave. Of particular significance was the discovery of a novel scenario for always-rare microbial taxa in relation to disturbance, in which stochastic processes are replaced later by deterministic processes during post-disturbance recovery, i.e., during attenuation of black stains.
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