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Cave and rockshelter sites have long been the cornerstone of Western Stemmed Tradition (WST) research in the northwestern Great Basin; however, these sites likely offer a narrow view of Terminal Pl...

Semiarid regions experience conspicuous seasonal variations, especially related to precipitation. Caves in these areas can be exceptions since they are less affected by dry seasons. In the north of the Brazilian semiarid, there are structurally heterogeneous karst areas with significant speleological potential and several anthropogenic impacts, with a neglected subterranean fauna. Therefore, we aimed to evaluate the influence of caves and external environmental features on the species richness and composition of cave invertebrates of this region. We expected that the caves would have high species richness and endemism, high dissimilarity among spatially discontinuous regions, and caves with water or guano would have greater overall richness and troglobitic species richness than those without water or guano. We collected invertebrates in 40 caves and recorded 416 species from 145 families and 45 orders (38.28 ± 13.88 spp./cave). We identified 57 species with troglomorphic traits, most having narrow distributions, including phylogenetic and/or geographic (oceanic) relicts, suggesting a lengthy and complex evolutionary history. In addition to the faunal dissimilarity among hydrographic basins and caves with or without water, our data indicate the variety of resources, the native vegetation surrounding the caves, the area, and the number of entrances as relevant variables of species composition and richness variation. Caves with water or guano had the highest richness of troglobites. The study area is unique in South America for having such a high concentration of troglobites associated with the presence of karstic aquifers and paleoclimatic changes (including oceanic transgressions and regressions), thus deserving emergency conservation actions.

The island of Borneo is a global biodiversity hotspot. However, its limestone caves are one of its least-studied ecosystems. We report for the first time the fungal species richness, diversity and abundance from Madai cave, situated in north-eastern Borneo. Environmental samples from inside the cave environment were collected (guano, speleothem, and cavern water) via opportunistic sampling. The dilution method was performed for isolation of fungi. Morphological characterisation and molecular analysis of the ITS region were utilised for the identification of isolates. Fifty-five pure cultures of fungi were attained, comprising 32 species from 15 genera, eight orders, and two divisions, Ascomycota and Basidiomycota. Ascomycetes dominated the fungal composition, accounting for 53 (96%) out of 55 total isolates. Penicillium spp. accounted for more than 47.1% of fungal abundance in all sample types. However, Aspergillus spp. had the highest occurrence rate, being isolated from all environmental samples except one. Purpureocillium lilacinum was isolated most frequently, appearing in five separate samples across all three substrates. Annulohypoxylon nitens, Ganoderma australe, Pyrrhoderma noxium, and Xylaria feejeensis were discovered and reported for the first time from the cave environment. This study provides additional data for further research on the mycoflora of Sabah’s various ecosystems, especially limestone caves.

The discovery of two undisturbed caves in Jamaica with 14C and 210Pb dating indicating that the oldest layers of guano were ca. 200 years old in the first cave (81 cm long core), and as much as 4300 years old in the second cave (129 cm long core) provides exciting possibilities to examine past ecological communities. We analyzed genetic and pollen profiles in these ancient bat guano deposits and revealed DNA sequences most similar to arthropods, mainly insects such as caddisflies (Trichoptera), butterflies (Lepidoptera) and mayflies (Ephemeroptera), suggesting a potential source for past dietary data. Palynological analysis failed to recover well-preserved pollen grains and spores older than ca. 200 years, however in layers preserving pollen, angiosperms were the most abundant plants observed, with a regular occurrence of the potato (Solanaceae) and pepper families (Piperaceae), which are frequently bat-dispersed. In general, changes in frequency of particular plant taxa appear to reflect changes of vegetation and land use in the cave vicinity; however, some changes could be linked to hurricanes, leading to forest canopy damage and promoting the growth of light-demanding species such as guarumo (Cecropia). Higher amounts of mangrove (Rhizophora) pollen have also been recorded in the periods of heavy hurricane activities. Our data highlight the value of bat guano deposits in caves as a resource for the analysis of past ecological systems and stress the conservation values of these deposits.

There have been increasing reports on the presence of Histoplasma capsulatum in some European countries. The study investigated the presence of Histoplasma in bat guanos, speleologists with records of visiting Histoplasma-endemic regions and patients with histoplasmosis. A commercial ALPHA Histoplasma Antigen enzyme immunoassay was tested as an alternative methodology to detect Histoplasma in environment and compared with polymerase chain reaction (PCR) assays. The presence of Histoplasma antigen in bat guanos was not confirmed by PCR. Among 14 healthy speleologists, two were indicated as having the Histoplasma antigen in urine, but expressed negative PCR-specific results for the Histoplasma antigen. Five unequivocal cases of imported acute pulmonary histoplasmosis in Slovenia between years 2005 and 2016 were confirmed in patients returning from North and South America after visiting hazardous localities e.g., caves with guano, and places with dust. Currently there is no evidence of autochthonous histoplasmosis in Slovenia, or that bat guano is a source of H. capsulatum. Involvement of histoplasmosis in travellers’ and cavers’ morbidity might be underestimated in non-endemic areas. It is crucial to ensure the use of appropriate protective equipment in Histoplasma hazardous localities, to spread information about this hazardous microbe to vulnerable populations and to monitor the health of the environment. A differential diagnosis for a febrile respiratory disease outbreak in patients returning from endemic regions should trigger routine consideration of possible histoplasmosis.

A total of 281 guano samples were collected from caves (N = 181) in eight European countries (Bulgaria, Czech Republic, France, Hungary, Italy, Romania, Slovakia and Slovenia) and attics in the Czech R. (N = 100). The correlation of detection of mycobacteria between Ziehl–Neelsen (ZN) microscopy and culture examination and qPCR was strong. ZN microscopy was positive in guano from caves (58.6%) more than double than positivity in guano from attics (21.0%, p &lt, 0.01). From 89 mycobacterial isolates (73 isolates from cave guano and 16 isolates from attics’ guano), 68 (76.4%) isolates of 19 sp., ssp. and complex were identified as members of three Groups (M. fortuitum, M.chelonae, and M. mucogenicum) and four complexes (M. avium, M. terrae, M.vaccae, and M.smegmatis). A total of 20 isolates (22.5%) belonged to risk group 1 (environmental saprophytes), 48 isolates (53.9%) belonged to risk group 2 (potential pathogens), and none of the isolates belonged to risk group 3 (obligatory pathogens). When comparing bat guano collected from caves and attics, differences (p &lt, 0.01, Mann–Whitney test) were observed for the electrical conductivity, total carbon, total organic, and total inorganic carbon. No difference (p &gt, 0.05, Mann–Whitney test) was found for pH and oxidation-reduction potential parameters.

Interpreting depositional settings of cave sites is generally problematic, especially in absence of palaeontological/archaeological evidence. This is the case of some deposits at San Teodoro Cave (Sicily), a key site for the Mediterranean Palaeolithic. In a stratigraphic level interrupted by a carbonatic concretion, phosphatic nodules are present only in the part enclosed between the concretion and the cave wall. The discovery of these nodules combined with the punctual lack of fossils had initially suggested an erosion phenomenon and subsequent formation of nodules at a vadose level. Here we show the usefulness of an integrated, geochemical-palaeoecological approach in defining stratigraphy and palaeoenvironmental reconstructions. XRD, ICP-OES, ATR-FTIR and EDS analyses allowed the formulation of a new hypothesis regarding the origin of the nodules, the depositional dynamics, and the role played by the guano produced by an extensive colony of bats. The role of barium and rubidium in detecting taphonomical processes has been highlighted.

Two cores of bat guano were recovered from Topolnița and Gura Ponicovei Caves in the south-west region of Romania and analyzed for δ15N, δ13C and pollen. Deposition of guano began in CE 1537 at Gura Ponicovei and CE 1694 at Topolnița, allowing for the development and interpretation of climate and vegetation records since the Little Ice Age. δ15N values were found to record phases of the North Atlantic Oscillation through the effect of winter precipitation on the local nitrogen cycle. Alternatively, δ13C values are interpreted to correspond with variation in summer precipitation. Both records suggest progressively drier conditions during the termination of the LIA in the region at CE 1870. The transition to landscape openness, with patchy forests of Tilia, Carpinus betulus, and Quercus during this time could be related to the lower water availability at both sites. Wetter conditions inferred from a 3‰ increase in δ15N values between CE 1879 and CE 1970 occurred contemporaneously with a similar trend in European winter precipitation and increased frequency of positive NAO phases during this interval. The δ13C values between CE 1885 and CE 1968 suggest a trend towards a wetter summer climate as well. Although there is at times evidence of forest dynamics being controlled by climate, vegetation appears to be more closely associated with human impact, particularly around Topolnița Cave. The contribution of ruderal pollen indicators from CE 1985 – present is likely related to agricultural practices.

White-nose syndrome (WNS) is a bat disease caused by the fungal pathogen Pseudogymnoascus destructans, which thrives in cold and very humid environments where bats frequently hibernate. Conidia of Pseudogymnoascus species are often documented on bats prior to the onset of WNS, but characterization of high-risk areas defined by microclimate cave conditions have been lacking. Investigating the occurrence of this fungal genus and appropriate environmental conditions to support P. destructans in southwestern U.S. caves is key to understanding the sites most likely to be impacted by WNS. Microclimate conditions in ten caves at El Malpais (ELMA) National Monument in New Mexico, USA were recorded using i-Button data loggers during the winters of 2011–2014 to assess appropriate environmental conditions (temperature and relative humidity) for P. destructans and other Pseudogymnoascus species. Optimal microclimate conditions for P. destructans and other psychrophilic fungi were found in all the caves with at least 50% of the caves identified as high-risk areas. Pseudogymnoascus species were detected in 70% of the caves using culturing methods and PCR, but no soil samples were positive for P. destructans using real-time PCR in soil and guano samples. Pseudogymnoascus destructans has a recognized range of appropriate temperatures and relative humidity for growth and cave microclimate can help define high-risk areas. This study offers resource managers guidance for establishing priority monitoring areas in their bat caves to determine which bat species are at higher risk.

SummaryA new species of a predator aphelenchoidid,Seinura cavernan. sp., is described and illustrated. The new species was isolated from bat guano collected from a calcareous cave in Japan. The new species is characterised by its three-lined lateral field, secretory-excretory pore at the level of the posterior two-thirds of the metacorpus, a long post-uterine sac, hermaphrodite tail shape elongate conoid with a filiform tip, and an androdioecious reproductive mode. The new species is typologically and biologically close toS. steineri, but is distinguished by its longer post-uterine sac, slightly longer stylet and slightly larger median bulb. The comparisons with other morphologically similar species,i.e.,S. chertkovi,S. tenuicaudataandS. steineri, are discussed. A molecular phylogenetic analysis based on two small and large subunit ribosomal RNA genes revealed that the new species is located at the basal position of clade 3 of the Aphelenchoididae, clearly separate fromS. demani, its congener with unclear rectum and anus, which is located at the derived position, suggesting that further generic revision is necessary for the genus.Aphelenchoides liin. comb. (=Seinura lii) is proposed.

Speleothems are a recognized source of paleoclimatic information, but their value as a source of signals from human activities in caves with an archaeological record has rarely been explored. Previous studies of speleothems in the Sierra de Atapuerca karst system (Burgos, northern Spain) revealed an important human fossil record, provided information about human activities in and around these caves, and the impacts on their natural environment. The present study reports the results of molecular characterization of dark-colored laminae from the stalagmites Ilargi (Galería de las Estatuas) and GS1, GS2, and GS3 (Galería del Silo), by pyrolysis-GC-MS (Py-GC-MS) and thermally assisted hydrolysis and methylation (THM-GC-MS). The features of the organic matter demonstrate the presence of (1) a dominant aliphatic fraction probably from in situ bacterial and ex situ plant-derived lipids, (2) black carbon (from soot and/or charcoal), (3) polysaccharides and N-rich moieties (probably from bat guano and microbial sources), and (4) a signal of terpenoid derivatives that may originate from the overlying limestone (kerogen) and extant gymnosperm resin (soils) or cyanobacteria (cave). Some plant-derived lignin may be present as well but was not identified unambiguously. It is concluded that this approach allows identifying multiple general sources of organic matter which can help understand speleothem formation processes, and evidence of soot deposition could be clearly linked to human activities.

Bat populations are declining worldwide because of anthropogenic activities, including habitat destruction and hunting. Cambodia represents an important case study for studying human-bat interactions, as loss of karst caves and the destruction of forests threaten the stability of bat populations and the ecosystem services they provide. Cambodians rely on bats for tourism revenue, fertilizer from guano, and as a source of protein. However, there is a lack of information on people's attitudes towards and relationships with cave-roosting bats. In 2018, we interviewed 60 residents around three karst outcrops (Sampeau Hill, Banan Hill, and Reichiatra Hill) in Battambang Province, northwestern Cambodia, along with agricultural professionals in Battambang Town, the province's capital. The primary objectives of the interviews were to examine people's (1) attitudes towards bats, (2) experiences with bats, and (3) engagement in high-risk behaviors associated with transmission of bat-related diseases (e.g., guano mining, hunting, etc.). Most respondents (70%) held positive attitudes towards bats and listed guano production, pest control, and tourism as benefits bats provide. Additionally, all informants believed bats should be protected and stated that they would feel sad if bats were extirpated. Conversely, respondents noted that many people eat bats. We followed these semi-structured interviews with five key informant interviews involved with the conservation of bats, which provided information on the history of human-bat interactions within these communities. Respondents' positive attitudes towards bats and recognition of ecosystem services bats provide indicate they would support bat conservation policy and may be interested in developing community-based conservation programs around karst outcrops.

Primate taphonomy in cave deposits is complex, and the taphonomic signatures of different accumulation scenarios present some degree of equifinality, rendering their identification in the fossil record challenging. Here, we describe an assemblage of 30 modern chacma baboons, including nine partially mummified individuals, which died inside a cave chamber at Misgrot, South Africa. The mortality profile is attritional, favouring immature and older individuals. We describe skeletal part representation, body postures for the most complete mummies and preserved articulations. Bone surface modifications suggest limited impact by biotic agents besides small rodents and invertebrates. Several specimens were affected by heat-induced damage, most likely resulting from natural combustion of bat guano, abundant on the cave floor. Misgrot has a similar geology and similar dimensions to some Plio-Pleistocene primate-bearing cave deposits from the UNESCO World Heritage Cradle of Humankind area. Implications of the taphonomic features of the Misgrot baboon remains are discussed in the context of some of these fossil assemblages, including the recently discovered Homo naledi assemblages.

International audience; The decay of bat guano deposits in caves produces mineral accumulations, mainly phosphates and secondary sulfates. Chameau Cave, Eastern Morocco, is located in the semi-arid Bni Snassen Mountains. It is composed of semi-active and dry passages, and is featured by strong condensation-corrosion on the walls, presence of fluvial sediments, and old corroded flowstones. Due to forced and convective airflow, the cave is generally very dry, with some damp sites related to condensation. Samples collected on the surface of different passages and along two sediment profiles yielded minerals related to bat guano decay. On recent or fresh guano, precursor minerals correspond to sulfate (gypsum), phosphate-sulfate (ardealite) and phosphate (brushite). Phosphates (hydroxylapatite, fluorapatite) occur at the interface with host rock or carbonate speleothems. At the contact of phyllosilicates contained in allogenic fluvial deposits or shale partings, or with pyrite-rich sediments, various phosphates occur (Al-rich strengite, Fe-rich variscite, phosphosiderite, leucophosphite, spheniscidite, crandallite, minyulite, variscite, and strengite), the latter two minerals being the stable end-members. Black seams of oxyhydroxides (goethite, hematite, birnessite) line the contact between carbonate host rock and weathered fluvial deposits. After “digestion” by acidic guano leachates, fluvial deposits only display the most resistant minerals (quartz, muscovite, K-feldspars and Na-plagioclases) and weathering byproducts (kaolinite). We discuss the origin of a pure gypsum particle cone, possibly related to evaporation at the edge of a wet cupola and subsequent detachment of sulfate particles. Among environmental conditions, humidity is required for decay. In this dry cave, most of the damp originates from either permanent or seasonal condensation. Dust particle advection seems to be essential in providing compounds that are not present on fresh guano (quartz, clay minerals). Bat guano phosphatization has probably occurred since >100 ka. The Chameau Cave appears as an outstanding site for bat guano-related minerals (n = 12), including rare phosphates (spheniscidite and minyulite).

The Basura Cave (Toirano, Savona, NW Italy) hosts important cave bear bone assemblages and a numerous and varied, tracks and traces record left by humans and other producers. An outstanding element of the analysed material is represented by fossil bear fur fragments, which were found in the inner deposits of the cave, and that, to date, are virtually unknown in the cave global record. After analysing and discussing micromorphological features of the inedited material, we integrate and interpret new radiocarbon data, along with taphonomic, sedimentological, geochemical and mineralogical evidences, with the aim of improving our understanding about the nature and chronology of the bear fur-bearing deposit. The bear fur fragments are included in a stratigraphic succession corresponding to a secondary deposit, formed after the dismantling, reworking and redeposition of a former bear-bearing deposit, as a result of short but intensive flooding events that most probably took place at the end of the Last Glacial Maximum. After sediments redeposition, important diagenetic changes have occurred and probably driven by guano deposits, whose pre-existence, in absence of record, is inferred from corrosion features, nutrient concentrations, mineral species identified (REE bearing hydroxyapatite), and claw traces left by bats on the cave ceiling and walls. Diagenetic imprint derived by guano deposits caused mineralization of bear fur fragments by replacement with apatite, which faithfully copied the form and structure of hairs but also of vegetal tissues, phytoliths and pollen found within them. Our study demonstrates for the first time that the bear fur is one of the main vectors in introducing botanical microremains into the interior of the “Old World” caves.

Escherichia coli, one of the primary intestinal commensal bacteria in humans and endothermic animals, is commonly considered an indicator of faecal pollution. E. coli strains were isolated from karst rivers under different hydrological conditions, from footpaths in tourist caves and from bat guano. Isolates were tested for phenotypic resistance to ampicillin, chloramphenicol, ciprofloxacin, nalidixic acid, tetracycline and trimethoprim. The highest percentage of antimicrobial resistant E. coli was found in karst waters, followed by those from surface swabs and from bat guano. Several isolates from rivers and swabs exhibited multidrug-resistant phenotype. Environmental conditions impact the populations of E. coli; a positive correlation between dissolved oxygen and E. coli counts, and a negative correlation between conductivity and E. coli concentrations have been observed for karst rivers. Malenščica (Slovenia), a drinking water resource with an extensive catchment area, contained a relative high percentage of antimicrobial-resistant E. coli strains. None of the isolates from bat guano was resistant to ampicillin, chloramphenicol, and tetracycline. Future monitoring of bats should consider a regular follow-up of indicative microbial disease indicators in fresh guano. Regular cleansing of tourist footpaths in caves and disinfection barriers at the cave entrances reduce the concentration and transmission of E. coli significantly. A future, more detailed, study on characterization of additional E. coli isolates is needed to reveal their pathogeneicity, mechanisms of antibiotic resistance, mobile genetic elements, and gene transfer frequencies to other members of the karst microbiome.

In this study, we examined the role of the eastern bent-winged bat (Miniopterus fuliginosus) in the dispersion of bat adenovirus and bat alphacoronavirus in east Asia, considering their gene flows and divergence times (based on deep-sequencing data), using bat fecal guano samples. Bats in China moved to Jeju Island and/or Taiwan in the last 20,000 years via the Korean Peninsula and/or Japan. The phylogenies of host mitochondrial D-loop DNA was not significantly congruent with those of bat adenovirus (m2XY = 0.07, p = 0.08), and bat alphacoronavirus (m2XY = 0.48, p = 0.20). We estimate that the first divergence time of bats carrying bat adenovirus in five caves studied (designated as K1, K2, JJ, N2, and F3) occurred approximately 3.17 million years ago. In contrast, the first divergence time of bat adenovirus among bats in the 5 caves was estimated to be approximately 224.32 years ago. The first divergence time of bats in caves CH, JJ, WY, N2, F1, F2, and F3 harboring bat alphacoronavirus was estimated to be 1.59 million years ago. The first divergence time of bat alphacoronavirus among the 7 caves was estimated to be approximately 2,596.92 years ago. The origin of bat adenovirus remains unclear, whereas our findings suggest that bat alphacoronavirus originated in Japan. Surprisingly, bat adenovirus and bat alphacoronavirus appeared to diverge substantially over the last 100 years, even though our gene-flow data indicate that the eastern bent-winged bat serves as an important natural reservoir of both viruses.

This study aimed at evaluating the effectiveness and the compatibility of two alternative treatments, in view of their possible use for conservation of prehistoric paintings in the Magura cave (Bulgaria). The paintings are made of bat guano applied over limestone, therefore, two sets of laboratory specimens were considered: stone specimens and stone specimens covered with a layer of sterilized bat guano. The two investigated treatments were a commercial product based on ethyl silicate (&ldquo, ES&rdquo, ) and a solution of diammonium hydrogen phosphate (&ldquo, DAP&rdquo, ), aimed at forming calcium phosphates. The results of the study indicated that both treatments were able to increase the mechanical properties of stone, the increase being higher for &ldquo, Both consolidants caused acceptable color changes, but the &ldquo, treatment significantly decreased stone wettability, water absorption, and water vapor permeability, while the &ldquo, treatment slightly affected those properties. In the stone + guano specimens, the presence of the guano layer affected the penetration of the consolidants, thus partly reducing their effectiveness. Compared to the stone samples, the guano layer experienced a more intense color change, alongside visible cracking. However, the adopted methodology to replicate the cave paintings was not completely successful, as the so-deposited guano layer was very prone to detachment when dry, unlike cave paintings. Future work will be dedicated to assessing the consolidant performance onto samples that resemble even more closely the conditions of the cave paintings, by improving the methodology for the guano layer deposition and by contaminating specimens with soluble salts before consolidant application.

Bat caves are very special roosts that harbour thousands of bats of one or more species. Such sites may hold an incredible “dark fungal diversity” which is still underestimated. We explored the culturable fungal richness in the air, on bats, and in the guano in a bat cave in Brazil’s Caatinga dry forest. Fungal abundance was 683 colony-forming units (CFU) in the guano, 673 CFU in the air, and 105 CFU on the bats. Based on morphological and phylogenetic analysis of ITS, LSU, and TUB2 sequences, fungal isolates of 59 taxa belonging to 37 genera in the phyla Ascomycota (28 genera, including Aspergillus, Penicillium, Cladosporium, and Talaromyces), Basidiomycota (eight genera, including Rhodotorula and Schizophyllum), and Mucoromycota (only Rhizopus) were identified. The fungal richness in the air was 23 taxa (especially Aspergillus taxa), mainly found at 15 m and 45 m from the cave entrance; on the bodies of bats it was 36 taxa (mainly Aspergillus taxa), especially on their wing membranes (21 taxa, nine of which were exclusively found in this microhabitat); and in guano 10 fungal taxa (especially Aspergillus and Penicillium) were found. The fungal richness associated with guano (fresh and non-fresh) was similar from bats with different eating habits (insectivorous, frugivorous, and haematophagous). Sampling effort was not sufficient to reveal the total fungal taxa richness estimated. Eight (21.6%) of the 37 genera and 17 (53.1%) of the 32 identified fungal species are reported for the first time in caves. Our results highlight bat caves in Brazil as hotspots of fungal diversity, emphasizing the need to protect such special roosts.

Trogloxenes particularly bats play an important role in subterranean habitat. They provide organic material and induce cave microclimate that influence cave-dwelling biota, including arthropods. This study aimed to learn how bats population influences cave-dwelling arthropods community. Data collections were performed in three caves which had different bats species in Tasikmalaya karst area namely Liang Boeh, Liang Seungit and Sarongge. We recorded bats population, guano production, physicochemical condition of caves passage, and arthropods community in each cave. All samplings were only conducted in the specific sites of the dark zone where bat populations were aggregated. Data indicated that Liang Boeh was inhibited by Hipposideros sp (±472 individuals), Liang Seungit by Pteripodidae and Miniopterus sp. (±188 individuals), and Sarongge by Rhinolophus sp. (±1194 individuals). Guano production was positively correlated with bats population. Chemical compositions of soil were varying among bats species. Bats population strongly induced caves physicochemical condition. A total 15986 individuals of cave-dwelling arthropods belonging to 5 Classes and 18 Orders were recorded. Our result revealed that bats population determined arthropods community. Caves with greater bats population size and dominated by insectivorous species would potentially have greater diversity and abundance of cave-dwelling arthropods.

Bats are emblematic hosts of caves. These small flying mammals deserve special attention because their presence has a great economic and ecological impact; they introduce organic matter, the guano, in the ecosystem they live in. Indeed, “guano” (a Quechua word meaning “fertilizer”) is the accumulation of their fecal matter (excreta); its deposition can reach several meters. The composition of guano is influenced by the bat’s food. In addition to its role within the caves, the bat guano is exploited for various purposes; it is used as an effective fertilizer for the cultivation of plants because of having exceptionally high content of nitrogen, phosphate, and potassium and also for the improvement of detergents and other products of great value for humans. The bat guano hosts various classes of microorganisms (viruses, bacteria, algae, fungi, and protists), which are adapted to the cave environment. Since guano is highly acidic, these microorganisms can be considered as extremophiles. They produce functional organic compounds in extreme conditions that could be of interest not only in the drug industry but also in different biotechnological areas. Here, we review already available information on the ecological and economic effects of bats and their guano. We report their food preferences, foraging behaviors, and environmental impacts. Information on these aspects may be useful in finding a solution about protection and preservation of bat populations.

Highlights • EMCV genome was widely found in fecal guanos in Taiwanese, Korean, and Japanese caves. • Miniopterus fuliginosus is the main source of the fecal guano. • It is possible that Miniopterus fuliginosus is one of the natural hosts of EMCV in East Asia., Bats are reservoir hosts of many zoonotic viruses and identification of viruses that they carry is important. This study aimed to use high throughput screening to identify the viruses in fecal guano of Taiwanese insectivorous bats caves in order to obtain more information on bat-derived pathogenic viruses in East Asia. Guano samples were collected from two caves in Taiwan, pooled, and then subjected to Multiplex PCR-based next generation sequencing for viral identification. Subsequently, encephalomyocarditis virus (EMCV) sequence was detected and confirmed by reverse transcription PCR. EMCV is considered as rodent virus and thus, animal species identification through cytochrome oxidase I (COI) barcoding was further done to identify the viral source. Finally, determination of distribution and verification of the presence of EMCV in guano obtained from Japanese and South Korean caves was also done. We concluded that the guano collected was not contaminated with the excrement of rodents which were reported and presumed to live in Taiwan. Also, EMCV genome fragments were found in guanos of Japanese and South Korean caves. It is possible that the eastern bent-wing bat (Miniopterus fuliginosus) is one of the natural hosts of EMCV in East Asia.

Recent environmental processes are studied in ʻA’rak Naʻasane Cave at the northern Judean Desert, Israel. The outer zone of the cave is heavily influenced by the outside environment through a large entrance, facilitating entry of air flow, fauna and humans, with minor cave-forming modifications. Conversely, the inner cave sustains humid and warm conditions, favoring modifications by condensation corrosion of convective air flow, associated with deposition of popcorn speleothems at the lower parts of dissolution pockets. The warm humid air of the inner cave may be associated with an underlying thermal water table. Active condensation corrosion is decreasing, possibly because of gradual change in the cave microclimate, associated with falling water table and ventilation. Increasing connection with the surface is indicated by high collapse domes, rare flood invasion, and a large Trident Leaf-nosed bat community which spends the winter within the innermost parts of the cave. Bat guano supports bedrock corrosion and a rich invertebrate fauna, but humans preferred the outer parts of the cave, particularly for refuge during the second Jewish revolt against the Romans. Rare occasions of ancient human entry into the inner cave support this scenario by the small number of artifacts compared with the outer cave. Enigmatic small cairns in the largest inner hall were probably erected during the Intermediate Bronze Age.

A 285‐cm core of bat guano was recovered from Magurici Cave in north‐west Romania and analyzed for δ13C, δ15N and pollen. Guano deposition occurred from AD 881 until 1240 and from AD 1651 to 2013, allowing for the interpretation of summer variations in precipitation and temperature during the Medieval Warm Period (MWP) and the Little Ice Age (LIA). A 2‰ increase in δ13C, 1.5‰ decrease in δ15N, and the presence of Ulmus, Quercus and Carpinus betulus indicate a warm and dry MWP occurred in the region. The lack of deposition during the beginning of the LIA suggests a possible climate‐induced change in prey availability resulting in bats vacating the cave. Variation of δ13C values between −25 and −23‰ at AD 1650 (LIA) indicates similar drier conditions as at the end of MWP. However, a 2‰ decrease in δ13C values that occurred between AD 1790 and 1900 suggests climate was trending towards wetter conditions at the end of the LIA. From AD 1938 to 2013, δ13C values appear to be more influenced by temperature, indicating that this parameter had a more significant effect on carbon discrimination than water availability.

Recent studies of ancient bat guano deposits in the caves of Gunung Mulu National Park and Niah National Park, Sarawak, Malaysia, have resulted in noteworthy records of phosphate minerals from these environments, including variscite, nano-particulate silica, fluorapatite, and niter.

Caves are important sites of fossil preservation, especially for Quaternary vertebrates. Taphonomic processes operating in caves are not well understood and have never been experimentally examined. This study focuses on the potential role of bat guano, which impacts environmental biogeochemistry and serves as the base of the food chain in cave ecosystems. The presence or absence of guano is expected to be a major control of preservation potential in caves. Bats first appear in the fossil record in the early Eocene so bat guano likely influenced cave preservation only during the Cenozoic. This is a probable megabias of the cave fossil record. Microcosm experiments were used to determine the impact of guano presence and composition, moisture, temperature, and time on preservation potential of small mammal bones, leaves, and crickets. Guano came from insectivorous and frugivorous bats. The guano of insectivorous bats has an acidic pH, while the guano of frugivorous bats is close to neutral. Lab stud...

International audience; The mineralogical and geochemical evolution of cave guano deposits in France has been investigated in detail. Two test pits were excavated in guano mounds from insectivorous bats and one in a guano mound from omnivorous choughs. Both bats and choughs are thought to be among the main accumulators of guano during the Pleistocene in southwest France. Thin section analysis, mineralogical identification and quantification, geochemical analysis, organic matter characterization through pyrolysis and thermochemolysis coupled to gas-chromatography, were conducted to better understand the evolution of guano in caves and to identify the underlying factors. Bat guano undergoes mineralization through loss of organic matter and precipitation of phosphate and sulfate minerals. The neoformed minerals include gypsum, ardealite, brushite, francoanellite, hydroxylapatite, monetite, newberyite, and taranakite, and vary according to the local availability of chemical elements released by the alteration of detrital minerals due to acidic solutions. Chough guano, located at higher altitude in a periglacial environment, does not show similar mineral formation. Organic geo-chemical analysis indicates strong differences between guano. Abundant hydrocarbons derived from insect cuticles were the dominant feature in bat guano, whereas a mostly vegetal origin typifies chough guano. Geochemical analysis points to an especially high content of copper and zinc in bat guanos, a few hundreds of µg/g and thousands of µg/g, respectively. Both organic matter analysis and geochemistry may help identification of bat guano in archeological contexts , where phosphate minerals can originate from multiple sources.

Authigenic phosphate mineralization was first studied on the territory of Russia on the basis of the Holocene and Pleistocene deposits of Denisova Cave. The formation of phosphates in the eastern gallery is related to biodegradation of the horizons of guano of insectivorous bats, which inhabited the cave in the absence man. The results confirmed the archaeological record of the Holocene and the upper part of Pleistocene sequences of the eastern gallery.

This study examines the pattern of preservation of pollen and non-pollen palynomorphs on bat guano samples in Eraaning Cave in relation to the current vegetation of the surrounding Garo hills of Meghalaya. It is observed that, the recorded pollen data does reflect the tropical mixed deciduous forest with an admixture of riparian and evergreen plant taxa from the local and regional vegetation. The presence of evergreen and riparian taxa namely, Mesua, Schima, Duabanga, and Syzygium in the pollen assemblage is strongly indicative of the high rainfall in the region. The recovery of fern and fungal spores in the pollen assemblage also reflects the warm and humid climatic condition in this region. The forestland and open-land samples from the vicinity of the cave were also studied to understand the nature of pollen preservation in relation to the bat guano samples. The presence of cereal, Areca, Psidium, and Citrus along with Melastoma pollen in both bat guano and open-land samples is indicative of the anthropogenic activities in and around the study area. Multivariate principal component analysis (PCA) and box plot were applied to the quantified data obtained from pollen frequency analyses which clearly revealed a significant variation and similarity in vegetation types among the surface samples collected in and around Eraaning Cave. The generated pollen data from the bat guano, forestland, and open-land samples reveals that, bat guano in cave sediments provides a reliable substrate to understand the modern pollen and vegetation relationship and can fruitfully be utilized as a baseline for the reconstruction of the palaeoecology in the Garo hills of Meghalaya. The study also suggested that the bat species utilized the mixed vegetation composition and different landforms as their preferred habitats as demonstrated by the pollen data from the bat guano samples.

International audience; The Prehistoric Cave at Azé (France) was divided in two parts by calcite flowstone. This isolated the innermost parts of the cave from the entrance, which remained connected to the surface. Since the closure, bats have not been able to access the cave beyond the flowstone blockage. They have been present only in the cave entrance. The byproducts released by bats had a considerable effect on this entrance part. Gaseous exhalations have changed the composition of the surrounding atmosphere, leading to the development of dome-shaped cupolas and other wall weathering features. Guano deposits built up causing the formation of aerosols and acidic leachates. The resulting corrosion of the limestone has led to the formation of phosphate deposits and biogenic karst. The rate of retreat of the cave walls has been estimated at 5 to 7 mm/ka. Corrosion has also affected archaeological artefacts, as well as any traces left on the walls by humans or animals. Bioglyphs linked to bears as well as anthropic graffiti have completely disappeared from the part of the cave used by bats.

International audience; The Towakkalak System located in the Maros karst of South Sulawesi is currently the richest of Southeast Asia in obligate subterranean species. It comprises several caves and shafts that give access to the subterranean Towakkalak river as well as many unconnected fossil caves, stream sinks, and springs located within its footprint. The total length of the caves linked to the active system is 24,319 m and comprises two of the longest caves of Indonesia, Gua Salukkan Kallang and Gua Tanette. Studies of its fauna began in 1985. There are 10 stygobionts and 26 troglobionts that are known from the system. The smaller adjacent system of Saripa has 6 stygobionts and 18 troglobionts, of which 1 and 3, respectively, are absent from Towakkalak. Like all tropical cave inventories, our dataset has limits due to identification uncertainties, gaps in habitat (waters, guano) and taxonomic coverage (micro-crustaceans, mites), sampling methods (pitfall trapping, Karaman–Chappuis), and problems of ecological assignment. A number of additional species are therefore expected to be found in the future. The Towakkalak and Saripa cave systems are included in the Bantimurung-Bulusaraung National Park and are under efficient protection, but parts of the Maros karst outside the park are under serious threat, mainly from quarrying.

The lava tubes at Undara became internationally recognised in the late 1980s, when 24 species of terrestrial cave-adapted invertebrates (troglobionts) were recorded from Bayliss Cave, making it one of the 20 richest known cave communities in the world at the time. Over the last decades, several of the Undara species have been taxonomically described and a great deal of research has been undertaken in other parts of Australia, which has revealed additional subterranean hotspots. It is therefore timely to update the list of Undara cave fauna, and to evaluate the Undara cave system in relation to other subterranean hotspots in Australia. The updated species list was compiled from the published literature and museum databases. Minimally, 78 species of arthropods have been recorded from 17 lava tube caves in the Undara Basalt. Sixteen species have been taxonomically described, 30 identified to genus and/or morpho-species, and 32 remain unidentified to species or genus level. Thirty troglobionts and one stygobiont species were recorded. Seven caves harboured obligate subterranean species, Bayliss Cave harboured the most obligate subterranean species: 23 troglobionts and one stygobiont. All these caves contained deep zone environments with high humidity, of which three also contained ‘bad air’ (CO2). The unique combination of geomorphic structure and environmental parameters (high humidity) and multiple energy sources (tree roots, bats and guano, organic material wash-in) are the main factors responsible for Bayliss Cave’s extraordinary local richness. Further research is needed to investigate CO2 as a factor influencing troglobiont richness and distribution in ‘bad air’ caves. Undara remains the richest subterranean hotspot in humid tropical Australia, however, significantly richer subterranean assemblages are found in arid and semi-arid calcrete aquifers, karst and iron-ore terrains, mostly in Western Australia.

The purpose of this study was to isolate and identify phosphate solubilizing Fungi from phosphate guano in Pawon Cave, Karst area, Citatah, West Java. The research design was descriptive. The data obtained were presented descriptively based on macroscopic and microscopic morphological characteristics and the result of qualitative phosphate capability screening on pikovskaya medium. The results showed that there were five isolates in the phosphate guano in Pawon Cave. Karst area, which were Penicillium sp. PF1, Aspergillus sp. PF2, Aspergillus sp. PF3, Mycelia sterilia, and Mucor sp. Based on the screening test, there were two isolates which could solubilize the phosphate, Aspergillus sp. PF3 and Mucor sp.

Based on samples collected during surveys of Brazilian cave fauna, seven earwig species are reported: Cylindrogaster cavernicola Kamimura, sp. n., Cylindrogaster sp. 1, Cylindrogaster sp. 2, Euborellia janeirensis, Euborellia brasiliensis, Paralabellula dorsalis, and Doru luteipes, as well as four species identified to the (sub)family level. To date, C. cavernicola Kamimura, sp. n. has been recorded only from cave habitats (but near entrances), whereas the other four organisms identified at the species level have also been recorded from non-cave habitats. Wings and female genital structures of Cylindrogaster spp. (Cylindrogastrinae) are examined for the first time. The genital traits, including the gonapophyses of the 8th abdominal segment shorter than those of the 9th segement, and venation of the hind wings of Cylindrogastrinae correspond to those of the members of Diplatyidae and not to Pygidicranidae. This is the first synopsis of cave-dwelling earwigs of Brazil, one of the most species-rich areas of Dermaptera in the world.

The presence of mycobacteria was studied in Bull Rock Cave (“Byci skala”) and the water catch - ment area of Jedovnice Brook (“Jedovnicky potok”) using direct microscopy after Ziehl-Neelsen (ZN) staining, culture examination and molecular techniques. Mycobacteria were detected in 47.1% of a total of 68 samples. The mycobacterial genes hsp65 and dnaA were detected and sequenced in 37 (74.0%) out of the 50 cave environmental samples and in 10 (55.6%) out of the 18 samples of water catchment sediments. Nine species of slowly growing mycobacteria ( M. terrae , M. arupense , M. gordonae , M. lentiflavum , M. parascrofulaceum , M. parmense , M. sas - katchewanense , M. simiae and M. xenopi ) and two subsp. ( M. avium subsp. avium and M. avium subsp. hominis - suis ) were detected. Fourteen species of rapidly growing mycobacteria ( M. chelonae , M. chubuense , M. poriferae , M. flavescens , M. fortuitum , M. porcinum , M. rhodesiae , M. gilvum , M. goodii , M. peregrinum , M. mageritense , M. vanbaalenii , M. gadium and M. insubricum ) were detected. The highest mycobacterial presence was docu- mented by ZN staining and/or culture examinations in earthworm castings and bat guano (73.3% positivity out of the 15 samples) in the cave environment and in the water sediments collected under the outflow from the wastewater treatment plants (77.8% positivity out of nine samples). The highest total organic carbon (TOC) was detected in wooden material and earthworm castings with pH values between 5.0 and 7.7 in the cave environment and in water sediments collected under the outflow from the wastewater treatment plants with pH between 5.8 and 7.0. It could be concluded that the karst cave environment with its running surface water contaminated with different microorganisms or chemical substances creates favourable conditions not only for animals (especially earthworms) but also for mycobacteria. This fact is also demonstrated by the presence of these mycobacteria in the cave environment mainly in earthworm castings and bat guano.

Chemical weathering of archaeological material is well known; however, while there have been a number of experimental studies demonstrating different types and degrees of chemical alteration on faunal remains, little attention has been paid to stone tools, and the few studies that exist relate mainly to siliceous materials and water-induced chemical alteration. Azokh Cave, located in the South Caucasus, contains a Middle Pleistocene to Holocene infill, and detailed macro- and microscopic examination of the lithic assemblages recovered there indicates potential chemical weathering of the stone artefacts. The cave is also currently home to one of the largest bat colonies in the region, and their guano forms a significant component of the infill of the inner galleries. Based on these two factors, an experimental pilot study was set up to artificially chemically alter a range of stone flakes produced specifically for the task, in order to determine the nature and likely cause of weathering. The experimental flakes, produced from different raw materials, were buried in fresh bat guano for up to two years. The results reported herein demonstrate that in a relatively short time, the highly acidic composition of bat guano strongly affects calcium-bearing rocks (e.g., limestone, basalt) altering their entire surface. Similar comparisons may be made with chemical alteration evident on archaeological lithics from Azokh Cave, suggesting that bat guano has played a significant role in diagenetic alteration.

Bat guano cores have been used as a source of palaeoenvironmental information to aid in the reconstruction of past climates and vegetation. We collected a 104-cm-long (43 cm compacted) guano core from Fern Cave, Alabama, USA, that provided a c. 6000-year record of guano accumulation. Pollen, nutrients (C, N, P) and stable isotopes (δ13C, δ15N) were measured on the guano core with the objective of reconstructing the environmental history of the area from the mid-Holocene to present. Our data indicate that bats have utilized Fern Cave for at least 6000 years and that Woodland Indians also utilized the cave for a short period. A 3-cm charcoal layer was dated to 2720±30 cal. a BP and inferred to be Woodland Indian in origin from microscopic inspection and thickness. Pollen and geochemical data showed that bat diets changed in the late Holocene possibly linked to food supply and climate changes. These results demonstrate that guano cores are a useful tool of palaeoenvironmental reconstruction when other forms of palaeorecords do not exist and can add to local archaeological information.

In neotropical regions, fossil bat guano accumulated over time as laminated layers in caves, hence providing a high-resolution temporal record of terrestrial environmental changes. Additionally, cave settings have the property to preserve such organic sediments from processes triggered by winds (deflation, abrasion and sandblasting) and intense rainfall (leaching away). This study reports both stable carbon and nitrogen isotope compositions of frugivorous bat guano deposited in a well-preserved stratigraphic succession of Blanchard Cave on Marie-Galante, Guadeloupe. These isotopic data are discussed with regard to climate changes and its specific impact on Eastern Caribbean vegetation during the Late Pleistocene from 40 to 10 ka cal. BP. Guano delta C-13 values are higher than modern ones, suggesting noticeable vegetation changes. This provides also evidence for overall drier environmental conditions during the Pleistocene compared to today. Meanwhile, within this generally drier climate, shifts between wetter and drier conditions can be observed. Large temporal amplitudes in both delta C-13 and delta N-15 variations reaching up to 5.9 parts per thousand and 16.8 parts per thousand), respectively, also indicate these oceanic tropical environments have been highly sensitive to regional or global climatic forcing. Stable isotope compositions of bat guano deposited from 40 to 35 ka BP, the Last Glacial Maximum and the Younger-Dryas reveal relatively wet environmental conditions whereas, at least from the end of the Heinrich event 1 and the Bolling period the region experienced drier environmental conditions. Nevertheless, when considering uncertainties in the model age, the isotopic record of Blanchard Cave show relatively similar variations with known proxy records from the northern South America and Central America, suggesting thus that the Blanchard Cave record is a robust proxy of past ITCZ migration. Teleconnections through global atmospheric pattern suggest that islands of the eastern Caribbean Basin could be also under the influence of a bipolar temperature gradients that impact the mean location of the ITCZ, with a Southern Hemisphere imprint during the glacial period and a more significant role of Northern Hemisphere during the last deglaciation.

Magura Cave, north-western Bulgaria, possesses valuable rock-art paintings made with bat guano and dated from the period between the Eneolithic and Bronze Ages. Since 2008, the Art Gallery is closed to the general public in order to protect the paintings from vandalism, microclimatic changes caused by visitors and artificial illumination, and the consequent growth of fungi and phototrophs. Nevertheless, some tourist visits are allowed under the supervision of cave managers. This study provides the first scientific report on cultivable fungal assemblages dwelling different substrata in the Art Gallery. A total of 78 strains, belonging to 37 OTUs (Ascomycota 81%, Zygomycota 13%, Basidiomycota 5%), were isolated in the study. This fungal diversity was clearly dominated by Penicillium (50% of strains) and Aspergillus (13%). The most relevant visible fungal colonies were detected in sediments rich in bat guano, where, besides Penicillium, other guanophilic fungi such as Mucor, Mortierella, Trichosporon and Trichoderma were dominant. Conversely, scarce fungi were detected on rock surface of painted walls. Based on the biocide susceptibility assay, octylisothiazolinone (OIT) and benzalkonium chloride (BAC) were effective inhibiting the in vitro growth of dominant fungal species in Magura Cave, when applied at concentrations ranged from 100 to 1,000 mg/L. These data provide a valuable knowledge about Magura fungi, and exemplify a type of preliminary test that may be conducted before planning any biocide treatment. However, considering the irreversible effects of biocides on the ecological balance in caves, and the low fungal contamination in painted walls of Magura Cave, there is no reason to use conventional biocides in this cave. Further studies, monitoring microbial communities and microclimatic parameters, should be conducted to improve the knowledge on microbial ecology in Magura Cave and possible human impacts, as well as to allow the early detection of potential microbial outbreaks.