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Monthly variation of soil (Ts) and subsoil (Tss) temperature and soil (Rhs) and subsoil (Rhss) relative humidity. The box-plots show the median, the upper and lower quartiles, the maximum and the minimum values and the outliers.
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The mesovoid shallow substratum (MSS) is a unique habitat that shelters and serves as a microrefuge for epigean, endogean and hypogean invertebrate species. Understanding the MSS community′s spatio-temporal structure and species diversity patterns in relation to the environmental parameters plays a crucial role in conservation. In this study we inv...
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Context 1
... and Rh were measured at the ground level (here- after soil temperature (Ts) and soil relative humidity (Rhs)) and at 0.5 m and 0.75 m depth (hereafter sub- soil temperature (Tss) and subsoil relative humidity (Rhss)). Figure 1 summarizes these measurements. ...
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... temperature and relative humidity in the MSS present decreased variation in amplitude in comparison with the external temperature and rel- ative humidity that show high fluctuations (Figs 1, 3). The CCA results for association of the ori- batid mite species abundance and environmental variables showed that Ts and Rhs are the strongest determinants of Oribatida community composition FIGURE 2: Biplots of the CCA model of the mite species abundance matrix in relation to habitat type; EDAF -edaphic, DMSS -deep MSS, SMSS -superficial MSS. ...
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... Figures 1, 3). Cepheus dentatus and Camisia biverru- cata (Koch, 1839) were the species correlated with Ts, while the presence of Oribatula tibialis (Nicolet, 1855), Neoribates aurantiacus (Oudemans, 1914) and H. initialis was influenced by Rhs ( Figure 3). ...
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... as pointed out by other studies, the two variables are interconnected and the higher the temperature, the greater the choice of organisms for higher humidity (Madge 1964). The low amplitude of temperature variation in MSS compared with high temperature variation at soil level (Figure 1) makes the MSS a micro-refuge for epigean species when conditions in the surface habitats become too harsh (e.g., too hot) ( Nitzu et al. 2014). ...
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The Carpathian Mts, showing the highest lichen biodiversity in Ukraine, i.e. 1,395 species of 2,106 of the total Ukrainian lichens, have been found to contain the highest proportion of regionally unique taxa (i.e. so far recorded only from the Carpathian Mts) (i.e. 551 species, or 39.5% of the Carpathian and 26.2% of the total Ukrainian lichen flor...
Citations
... MSSs are inhabited by species with different ecological requirements (epigean, edaphobionts, and troglobionts, among others) (e.g., Nitzu et al., 2014;Jiménez-Valverde et al., 2015;Mammola et al., 2016). Recently, new and rarely captured species have been frequently discovered in MSSs (e.g., Baquero et al., 2017;Gilgado et al., 2017;Ledesma et al., 2019;Jordana et al., 2020), allowing for a deeper understanding of the ecology of soil and subsoil habitats (Rendoš et al., 2012(Rendoš et al., , 2016Nitzu et al., 2014;Nae & Băncilă, 2017;Ledesma et al., 2020;Nae et al., 2021). MSS-research began in the Pyrenees (Juberthie et al., 1980(Juberthie et al., , 1981 and has been largely conducted in Europe (Mammola et al., 2016). ...
The species Aspilota umbrosa Belokobylskij, 2007 is recorded for the first time in Europe. The first photographsof this species, both of the holotype and of the new record, the second in the world, are provided. The holotype was foundnear the Klyuchevskaya Sopka volcano (Kamchatka peninsula, Russia). The new specimen of this species was collectedin a superficial subterranean habitat (the MSS – Milieu Souterrain Superficiel) in the rock glacier in Val Sassa (SwissNational Park, Grisons, Switzerland) by a subterranean trap in 2020. This is the first record of A. umbrosa in the MSS ofthe Swiss Alps and in a rock glacier. A key to the known Swiss species of Aspilota is provided
... Some, named troglobionts, are closely associated with caves, while others prefer caves but live in different subterranean habitats (troglophiles) and some temporarily inhabit caves (trogloxenes) [3,4]. Beetles are also found in shallow subterranean habitats, termed the Mesovoid Shallow Substratum (MSS) [5][6][7][8][9][10]. The MSS comprises small crevices and scree slopes in rock debris [8][9][10]. ...
... Beetles are also found in shallow subterranean habitats, termed the Mesovoid Shallow Substratum (MSS) [5][6][7][8][9][10]. The MSS comprises small crevices and scree slopes in rock debris [8][9][10]. The uniqueness of this environment relies on a connection between subterranean conditions-such as lack of light-with a direct inflow of organic matter from the surface [7,9,10]. ...
... The MSS comprises small crevices and scree slopes in rock debris [8][9][10]. The uniqueness of this environment relies on a connection between subterranean conditions-such as lack of light-with a direct inflow of organic matter from the surface [7,9,10]. ...
This paper presents the study's results on beetles (Insecta: Coleoptera) inhabiting caves in the Częstochowa Upland, southern Poland. During two years of research, 2084 specimens, representing 105 species from 19 beetle families, were collected. The obtained results indicate that many beetle species choose to inhabit caves despite lacking specific adaptations for living in such environments. The cave entrance zone is the most attractive place for surface species to inhabit because its climatic conditions are more stable than outside the cave, some sunlight is present, and the availability of organic matter is high. In the deeper parts of the studied caves, the number of occurring species rapidly decreased. Three species were recognised as troglobionts, four as troglophiles and 32 as subtroglophiles. Speonomus normandi hydrophilus, Choleva lederiana gracilenta and Quedius mesomelinus mesomelinus were identified as the most frequent in the studied caves. Speonomus normandi hydrophilus is endemic to the Arize massif in the Central Pyrenees (France) and was experimentally introduced into the Dzwonnica cave (Poland) in 1982, while Ch. lederiana gracilenta is one of two troglobitic beetles native to Polish caves and Q. imesomelinus mesomelinus is a widespread, very common troglophile.
... It has been reported from numerous localities in the Alps (Fischer & Schatz 2020, Schatz & Bruckner 2021, the Giant Mountains (Materna 2000, Starý 2006, Miko 2013, Carpathian Mts. (Nae & Băncilă 2017, Skubała & Maslak 2009, and is also known from Caucasus Mts. (Schatz & Bruckner 2021). ...
... It has been found in a peatland, 870 m a. s. l. (Starý 2006), at dry limestone plateau, 550 m a. s. l. (Lazarus & Krisper 2014), and in soils polluted with heavy metals in Romania, where it was associated with Cu, As and Mn (Manu et al. 2019). Most often it has been collected from soil or ground vegetation, but has been also reported from the unique microhabitat of the intermediate layer between the base of the soil and the bedrock, called "mesovoid shallow substratum" (Nae & Băncilă 2017) and from decaying wood (Skubała & Maslak 2009). Bog, decaying wood, some Sphagnum Unknown Mahunka (2006) Z. conjunctus Shaldybina, 1987 Russia (Stolby Nature Reserve, Krasnoyarsky Krai) Moss cushion at the base of a pine tree Unknown Shaldybina & Grishina (1987) Z. eoeryi (Mahunka, 1972) Z. lobatus (Hammer, 1977) Pakistan (Lowari Pass, 3400 m a. s. l.) Thin moss on the soil, grass, Potentilla, Epilobium , Fragaria , Geranium , pine cones Unknown Hammer (1977) Z. maritimus Shaldybina, 1973 Russia (Primorsky krai), USA (Alaska), Canada (Yukon, Vancouver Island, British Columbia) ...
The adults of the new species Zachvatkinibates svanhovdi A. Seniczak and S. Seniczak sp. nov. are described and illustrated from Norway, and this is the first finding of Zachvatkinibates Shaldybina, 1973 in Fennoscandia. This species is the most similar to Z. quadrivertex (Halbert, 1920), but differs from it mainly by the shape of notogastral setae, posterior tectum of notogaster and lack of postanal porose area Ap, which in Z. quadrivertex is present. In Z. svanhovdi, the prodorsal seta in is long, translamella is narrow, notogastral setae are short and distally pliable, notogastral porose areas are usually oval and of medium size, but Aa can be larger, especially in males. Dorsal crest on tarsus I is present. The cytochrome oxidase I (COI) barcodes (length: 658 bp) of five specimens of the new species are provided; the maximum variation within the species was 2.41% (p-dist). The morphology and ecology of the new species is compared with other Zachvatkinibates species. The knowledge on family Punctoribatidae in Fennoscandia is updated, and Mycobates carli (Schweizer, 1922) is first reported from Norway.
... In temperate zones the MSS plays a role as refuge for arthropod fauna, mainly at high altitudes where the cold weather can eliminate all the ectothermic fauna from the surface 10,11 . Similarly, seasonal migration movements are observed in the MSS for different taxa as response to hot dry summer 20,21 . When troglobitic fauna is concerned the MSS has a different role, cave restricted Collembola showed higher underground dispersal capacity than troglophiles 22 , therefore, the MSS can connect neighboring caves systems and extend their distribution range. ...
... nov. 21. Antennal chaetotaxy (no represented): Ant IV dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally with about two rod sens sub-apical on longitudinal row, ventrally with one subapical-organ and about three wrinkly sens on longitudinal row (Fig. 4A); Ant III dorsally and ventrally with several short ciliate mic and mac, and finger-shaped sens, dorsally without modified sens, ventrally with one apical psp, about three wrinkly sens and three smooth mic on external longitudinal row, apical organ with one finger-shaped sens, two coffee bean-like sens, and one rod sens (Fig. 4A); Ant II dorsally and ventrally with several short ciliate mic and mac, www.nature.com/scientificreports/ ...
A hotspot of subterranean Collembola in ferruginous rock caves and Mesovoid Shallow Substratum is revealed by the analysis of pseudocryptic diversity. The diversity is accessed by detailed description of chaetotaxy and slight variation in morphology of 11 new species of Trogolaphysa Mills, 1938 (Collembola, Paronellidae, Paronellinae) and the 50 previously recorded species of springtails from caves, using optical and electronic microscopy. When combined with recent subterranean surveys, our results show an important reservoir of cave diversity in the Mesovoid Shallow Substratum. Contrastingly the conservation policy for subterranean fauna in metallogenic areas in Brazil prioritizes the caves instead the cave species, which may be extremely detrimental to the fauna in the shallow subterranean habitats not accessible to humans.
... Similarly, seasonal migration movements are observed in the MSS for different taxa as response to hot dry summer [18,19]. When troglobitic fauna is concerned the MSS has a different role, cave restricted Collembola showed higher underground dispersal capacity than troglophiles [20], therefore, the MSS can connect neighboring caves systems and extend their distribution range. ...
A super diverse hot spot of subterranean Collembola in ferruginous rock caves and Mesovoid Shallow Substratum is revealed by the analysis of cryptic diversity. The diversity is accessed by detailed description of chaetotaxy and slight variation in morphology of 11 new species of Trogolaphysa Mills, 1938 (Collembola, Paronellidae, Paronellinae) and the 49 previously recorded species of springtails from caves, using optical and electronic microscopy. When combined with recent subterranean surveys, our results show an important reservoir of cave diversity in the Mesovoid Shallow Substratum, contrasting with the conservation policy for subterranean fauna in metallogenic areas in Brazil which prioritizes the caves instead the cave species, which may be extremely detrimental to the fauna in the shallow subterranean habitats not accessible to humans.
... Oribatid mite diversity varies among habitats (Schatz and Behan-Pelletier 2008), although the factors responsible for their variation in space are little understood (Maraun and Scheu 2000;Caruso et al. 2019). Oribatid mite communities of subterranean habitats such as mountain scree have been studied intensively (Skubała et al. 2013;Jiménez-Valverde et al. 2015;Nae and Băncilă 2017), but their trophic structure is little known. ...
Mountain scree habitats are intermediate habitats between the base of the soil and the bedrock. They are composed of a network of small cracks and voids, and are commonly situated at the lower levels of scree slopes. Their environment is defined by empty spaces inside the scree, the absence of light and photoperiod, low temperature, and resource poor conditions. Soil arthropod communities, their trophic structure as well as their use of basal resources in mountain scree are little studied despite the fact that they are important components of these systems. Here, we investigate stable isotope ratios (15 N/ 14 N, 13 C/ 12 C) of oribatid mites (Oribatida, Acari) to understand their trophic niches and their variation with depth (50 and 75 cm) at two mountain scree sites (Cerdacul Stanciului, Marele Grohotis) in the Romanian Carpathians. Further, we used existing data to investigate the reproductive mode of the species in that habitat, as this may be related to resource availability. We hypothesized that trophic niches of oribatid mites will not differ between the two mountain scree regions but will be affected by depth. We furthermore hypothesized that due to the resource poor conditions oribatid mite species will span a narrow range of trophic levels, and that species are sexual rather than parthenogenetic. Our results showed that (1) oribatid mite trophic structure only slightly differed between the two sites indicating that the trophic ecology of oribatid mites in scree habitats is consistent and predictable, (2) oribatid mite trophic structure did not differ between the two studied soil depths indicating that the structure and availability of resources that were used by oribatid mites in deeper scree habitats varies little with depth, (3) oribatid mite species spanned only three trophic levels indicating that the habitat is rather resource poor, and (4) that all studied oribatid mite species were sexual supporting the view that resource poor conditions favour sexual reproduction.
... The authors (Kovacs, et al., 2017) suggest that structural elements have a stronger influence on microclimate conditions than tree species composition of the overstorey. Thus, it was concluded that the structure of the saprophage community (Oribatid case) was influenced by temperature and humidity (Nae & Bancila, 2017). ...
The paper presents a study about the influence of two saprophage groups (Isopoda, Diplopoda) on leaf litter decomposition under different levels of humidification and chemical stress. Because of their worldwide distribution, we focused on the common pillbug Armadillidium vulgare (Latreille, 1804) (Isopoda, Armadillidiidae), and the common millipede species Rossiulus kessleri (Lohmander, 1927) (Julida, Julidae). The function of environment creation by the given saprophages, as destructors of dead plant matter, supporting such ecosystem services as soil fertility improvement and nutrients’ turnover, is highlighted. To conduct the experiment, the animals were collected manually and using pitfall trapping. In order to bring the experimental conditions closer to the natural, the individuals were not sexed. The maximum consumption of leaf litter by woodlice was recorded in the conditions with adequate moisture (0.5 mL of distilled water per box) and amounted to 2.52 mg/10 individuals per day, which exceeds its consumption with low and increased moisture, respectively, by 1.82 and 1.24 times. As for the effect of interaction, the consumption of maple litter with optimal moisture (4.77 mg/10 individuals per day) was the greatest. The largest absolute difference between broad-leaved tree species in the average weight of leaf litter consumed by woodlice was between maple leaf litter and oak leaf litter, the minimum – between robinia leaf litter and oak leaf litter. According to the results of the obtained data (Diplopoda), it can be stated that there is a statistically significant effect of chemical stress and discrepancy of the average zinc content in the object of study (in Diplopoda and their faecal pellets). We found that the diet provided did not affect the distribution of zinc in Diplopoda under conditions of chemical stress. According to the results of pairwise comparisons, we determined that the zinc content in the Diplopoda clearly differs for control and almost every concentration of zinc sulfate solution – 0.03 and 0.15 g/L, the samples of which do not form a homogeneous group. The species composition, abundance and distribution in space of soil invertebrates are informative indicators which reflect the ecological state of soils, intensity in development of soil horizons as well as intensity of processes occurring in them.
... In general, it has been suggested that extreme environmental conditions acting at a variety of temporal and spatial scales, such as those occurring during periods with harsh climates or along altitudinal gradients, generate pressures that drive the fauna to seek refuge in subterranean environments (Bellés, 1991;Culver and Pipan, 2010). Temperature has been suggested to be an important factor conditioning the migration of fauna from the surface and upper soil layers into the MSS to avoid severe temperature values, conditioning the richness, abundance, and composition of the assemblages (e.g., Crouau-Roy et al., 1992;Růžička, 1999;Nitzu et al., 2006;Růžička and Zacharda, 2010;Pipan et al., 2011;Polak, 2012;Nitzu, 2016;Nae and Băncilă, 2017). The availability of nutrients is another crucial limiting factor in the MSS (Gers, 1992(Gers, , 1998. ...
... Thus, the MSS arises as a potentially accessible climatic refuge for the arthropod fauna, allowing many species to survive and thrive by avoiding the unfavorable conditions of the surface. In fact, seasonal migrations in different taxa have been observed in the MSS, generally as a response to hot and dry summer conditions (e.g., Hernando et al., 1999;Nitzu et al., 2010Nitzu et al., , 2011Nitzu et al., , 2014Pipan et al., 2011;Rendoš et al., 2012;Nae and Băncilă, 2017;among others). ...
The determinants of biodiversity patterns in the subterranean habitat called Mesovoid Shallow Substratum (MSS) are not well-understood. In this study, thirty-three scree slopes at high altitudes were selected across the Sierra de Guadarrama National Park in central Spain to investigate the effect of ten environmental variables on the abundance and species diversity of the spider and springtail assemblages from the colluvial MSS. In each locality, a multiperforated PVC tube with a pitfall trap inside was buried up to 1 m deep, and generalized linear models and Mantel tests were used to analyze the effect of mainly climate- and habitat-related variables on the diversity patterns of both taxa. A total of 1143 individual spiders belonging to 54 species and 40,811 springtail individuals belonging to 62 species were collected. The analyses indicated that cold temperatures and the presence of forest cover on the surface significantly enhance richness and abundance in the two taxa. Environmental similarity also had a small positive effect on faunistic similarity. However, the effects of temperature and habitat detected on spider richness and abundance were stronger than on springtails, whereas the reverse was found regarding faunistic similarity. These results indicate that subterranean dwellers respond differently to the same environmental factors, which in turn, points to a different degree of affinity for the MSS. The MSS plays an important role in the survival of high mountain arthropod species, acting as a climate refuge, so the protection of this habitat should be prioritized.
... Some previous studies were focused on ecology, revealing the special characteristics of this subterranean environment (Juberthie et al. 1980(Juberthie et al. , 1981Gers 1992Gers , 1998R u zi cka 1999;Pipan et al. 2011;R u zi cka et al. 2012;Ortuño et al. 2013;Nitzu et al. 2014). Other studies were conducted to explore faunal aspects and therefore were focused on characterising the invertebrate communities that inhabit the MSS (R u zi cka 1990;R u zi cka et al. 1995;Nitzu et al. 2010;Deltshev et al. 2011;Rendo s et al. 2012;Langourov et al. 2014;Jim enez-Valverde et al. 2015;Mammola et al. 2017;Nae & B ancil a 2017). Although the all the previous studies cited above have made a substantial contribution to the understanding of the MSS, taxonomic studies have been predominant, which has led to the discovery of new species in a range of taxonomic groups. ...
The material for this study was obtained after intensive sampling in the colluvial milieu souterrain superficiel (mesovoid shallow substratum, or MSS) of the Sierra de Guadarrama National Park using 33 subterranean sampling devices (SSD). The data were obtained from the first extraction of the traps between May and October of 2015. This paper presents the results for the Poduromorpha taxon, which was part of the total Collembola captured. Of the 17 species captured in this study, 11 had previously been cited in Guadarrama, four are new references, and two new species. Friesea ortunoi Jordana and Baquero sp. nov. is characterised as having seven eyes, tibiotarsus as having one clavate chaetae and only two spiniform chaetae on Abd VI. Schaefferia sendrai Jordana and Baquero sp. nov. is characterised as having six eyes, seven sensilla on Ant IV, a bi- or tri-lobed apical vesicle on the antennal tip, claws with internal tooth and lateral (posterior) tooth, dens with six chaetae and retinaculum with 5 + 4 teeth (asymmetrically). Based on the results of the study of the group of species of Xenylla Tullberg, 1869 that appeared, Xenylla xavieri Gama, 1959 is separated from Xenylla lotharingiae Thibaud, 1963 and a brief description of the latter is provided. The comparative study of the Schaefferia Absolon, 1900 species related to the new species found in the Sierra de Guadarrama National Park led to the description of two new species: Schaefferia fjellbergi Jordana and Baquero sp. nov. and Schaefferia babenkoi Jordana and Baquero sp. nov., accepting the previous descriptions. The mesovoid shallow substratum (MSS) was revealed as an important habitat for the richness of the Collembola species. Furthermore, the findings showed the value of MSS as an indicator of the richness of the Collembola species. As many as 3000–4000 specimens were gathered in one sample device SSD. Among all specimens, the most abundant was Hypogastrura meridionalis Steiner, 1955 (77.7 %) and the least abundant was Xenylla schillei Börner, 1903 (11.5 %).
... Oppiidae), were rarely found. Recently, Nae and Băncilă (2017) described oribatid communities of an MSS site in the Romanian Carpathians, where common forest soil or tree-bark dwellers predominated. Similarly, Jiménez-Valverde et al. (2015) recorded three epigeic species of Oribatida when performing a study in Spain of subterranean fauna of bare scree slopes sparsely covered by the soil. ...
... Worldwide, there are only a few studies on Acari occupying shallow subterranean environments (Borges 1993, Arillo et al. 1994Růžička and Zacharda 1994;Schlick-Steiner and Steiner 2000;Zacharda et al. 2005). Within Europe, previously published studies on Oribatida in MSS have been carried out in France, Romania and Spain (Cassagne et al. 2008;Sendra et al. 2014;Jiménez-Valverde et al. 2015;Nae and Ivan 2015;Nae and Băncilă 2017). However, they used a different type of pitfall traps, which were monitored for shorter period than one year. ...
Mesovoid shallow substratum (MSS) of scree slopes constitutes a transition habitat between the soil and the network of voids in the vadose zone of a bedrock massif. In the present study, the vertical distribution of oribatid communities along a depth of 95 cm was studied at five forested MSS sites in the Western Carpathians, Slovakia. The sites differed in type of bedrock, topography and gradient of the microclimate and nutrients content. In all, 909 specimens were captured in subterranean traps exposed for one year. Most Oribatida represented edaphic forms, and their presence in the depth profile of the screes was accidental. Pantelozetes cavatica (Kunst, 1962) was the only species closely linked to deep subterranean environments found in the deeper part of the single limestone site studied. Species richness and the activity of oribatids along the scree profile at the sites clearly reflected the content of organic carbon in the soil substratum. The communities had very low numbers of individuals and low species richness at three sites with soil pH < 7 and organic carbon content in the upper soil layer ≤ 10%. However, they differed markedly in internal temperature dynamics. The other two sites, with a slightly alkaline soil pH and a higher carbon content, showed distinctly higher activity and a relatively uniform pattern of oribatid distribution across the depth profile. The soil pH and organic carbon content in the topsoil layer were substantial factors that determined the Oribatida diversity and vertical distribution in the forested screes.
