Elena Piano

Ecology and sampling techniques of an understudied subterranean habitat: the Milieu Souterrain Superficiel (MSS)

The term Milieu Souterrain Superficiel (MSS) has been used since the early 1980s in subterranean biology to categorize an array of different hypogean habitats. In general terms, a MSS habitat represents the underground network of empty air-filled voids and cracks developing within multiple layers of rock fragments. Its origins can be diverse and is generally covered by topsoil. The MSS habitat is often connected both with the deep hypogean domain—caves and deep rock cracks—and the superficial soil horizon. A MSS is usually characterized by peculiar microclimatic conditions, and it can harbor specialized hypogean, endogean, and surface-dwelling species. In light of the many interpretations given by different authors, we reviewed 235 papers regarding the MSS in order to provide a state-of-the-art description of these habitats and facilitate their study. We have briefly described the different types of MSS mentioned in the scientific literature (alluvial, bedrock, colluvial, volcanic, and other types) and synthesized the advances in the study of the physical and ecological factors affecting this habitat—i.e., microclimate, energy flows, animal communities, and trophic interactions. We finally described and reviewed the available sampling methods used to investigate MSS fauna.

Seasonal dynamics and micro-climatic preference of two Alpine endemic hypogean beetles

The climatic conditions occurring underground have a profound influence on hypogean fauna (Culver & Pipan, 2010). As far as hypogean beetles are concerned, it is well known that they generally dwell in environments which are relatively stable from an environmental point of view, characterized by constant temperature and an atmosphere saturated with water vapor (Moldovan, 2005). Indeed several hypogean beetles are adapted to narrow ranges of temperature and relative humidity, and thus classified as stenothermic and stenohygric (Howarth, 1980). Hence, it has been observed that they are able to react quickly when significant changes in the microclimatic conditions occurs (Juberthie, 1969), migrating inside the inner parts of the cave (and through the network Citation:

Environmental drivers of phototrophic biofilms in an Alpine show cave (SW-Italian Alps)

The proliferation of lampenflora is a major threat for the conservation of show caves, since phototrophic organisms cause physical, chemical and aesthetic damage to speleothems. In this paper we examine the environmental factors influencing the presence and the growth of the three main photosynthetic groups composing phototrophic biofilms in the Bossea show cave (SW-Italian Alps). The presence and the primary production of cyanobacteria, diatoms and green algae were detected with BenthoTorch®, an instrument for in situ measurement of chlorophyll a concentration that has never been used before in caves. By means of different techniques of regression analysis, we highlighted the response of the three photosynthetic groups to different environmental factors. Illuminance proved to be the main factor influencing positively both the probability of the presence and the productivity of the three groups. The presence of seeping water on the substrate and the distance from the cave entrance proved to play an important role in determining patterns of colonization. By means of GIS techniques, we provide thematic maps of the cave, providing a representation of pattern of the density of the three examined photosynthetic groups within different areas of the cave. The same approach may apply to other show caves, aiming at providing suggestions for the cave management (i.e. cleaning of the cave walls and positioning of artificial lights) and reduce impact caused by tourism.

Fine sedimentation affects CPOM availability and shredder abundance in Alpine streams

Fine sedimentation was quantified in the upper stretch of the Pellice river basin (Italian Alps), in an area partially disturbed by mining activities. Sediment traps were placed in 52 sites and benthic samples were collected to assess the amount of coarse particulate organic matter and the structure of macroinvertebrate community. The results indicated that the increase of fine sediment in the river reduced the amount of coarse particulate organic matter, affecting the abundance of invertebrate shredders. In small alpine streams, where allochthonous organic matter is the main energy source, fine sedimentation reduces the availability of trophic resources, affecting the abundance of benthic communities.

Body-size shifts in aquatic and terrestrial urban communities

Body size is intrinsically linked to metabolic rate and life-history traits, and is a crucial determinant of food webs and community dynamics1,2. The increased temperatures associated with the urban-heat-island effect result in increased metabolic costs and are expected to drive shifts to smaller body sizes3. Urban environments are, however, also characterized by substantial habitat fragmentation4, which favours mobile species. Here, using a replicated, spatially nested sampling design across ten animal taxonomic groups, we show that urban communities generally consist of smaller species. In addition, although we show urban warming for three habitat types and associated reduced community-weighted mean body sizes for four taxa, three taxa display a shift to larger species along the urbanization gradients. Our results show that the general trend towards smaller-sized species is overruled by filtering for larger species when there is positive covariation between size and dispersal, a process that can mitigate the low connectivity of ecological resources in urban settings5. We thus demonstrate that the urban-heat-island effect and urban habitat fragmentation are associated with contrasting community-level shifts in body size that critically depend on the association between body size and dispersal. Because body size determines the structure and dynamics of ecological networks1, such shifts may affect urban ecosystem function.The urban-heat-island effect drives community-level shifts towards smaller body sizes; however, habitat fragmentation caused by urbanization favours larger body sizes in species with positive size–dispersal links.

Trophic availability buffers the detrimental effects of clogging in an alpine stream

Clogging, the streambed colmation by fine sediments, is an important widespread source of impact affecting freshwaters. Alterations in stream morphology and hydrology, added to the effects of global climate change, are responsible for this phenomenon, that is particularly pernicious in mountainous lotic systems naturally characterized by coarse substrates. Among the studies investigating this issue some were descriptive, while others used artificial substrates to compare ongoing fine sediment accumulation and macroinvertebrate assemblage recruitment. Other studies used from the outset artificial substrates arranged with different levels of clogging. Our study fits into this line, but adding an innovative element simulating different availability of coarse particulate organic matter, i.e. the main trophic input in low-order, mountainous stream. To investigate how clogging and CPOM can influence macroinvertebrate communities, we placed 135 artificial substrates in the upper Po river (NW Italy). We set up a three way factorial design with three different levels of sedimentation and terrestrial leaf material. Artificial substrates were removed on three different dates. Benthic invertebrates were identified and classified according to their bio-ecological traits. We also measured macroinvertebrate dry mass and CPOM degradation in the different trap types. Our findings show that clogging acts as a selective filter influencing taxa richness, density, functional composition and biomass of benthic assemblage. Moreover, fine sediments affect the energetic dynamics in the river ecosystem, decreasing the mass loss rate of terrestrial leaves. Interestingly, our results clearly demonstrate that high availability of CPOM can buffer the negative effect of clogging, suggesting that an adequate input of allochthonous organic matter may lessen the impact of fine sediment deposition. Because land use transformation and removal of wooded riparian areas increase clogging and simultaneously reduces the input of CPOM, our findings stress the importance to include the management of river basins in the conservation strategies of mountainous streams.

Ecological speciation in darkness? Spatial niche partitioning in sibling subterranean spiders (Araneae : Linyphiidae : Troglohyphantes)

Abstract. Speciation in subterranean habitats is commonly explained as the result of divergent selection in geographically isolated populations; conversely, the contribution of niche partitioning in driving subterranean species diversification has been rarely quantified. The present study integrated molecular and morphological data with a hypervolume analysis based on functional traits to investigate a potential case of parapatric speciation by means of niche differentiation in two sibling spiders inhabiting contiguous subterranean habitats within a small alpine hypogean site. Troglohyphantes giachinoi, sp. nov. and T. bornensis are diagnosed by small details of the genitalia, which are likely to be involved in a reproductive barrier. Molecular analysis recovered the two species as sister, and revealed a deep genetic divergence that may trace back to the Messinian (∼6 million years ago). The hypervolume analysis highlighted a marginal overlap in their ecological niches, coupled with morphological character displacement. Specifically, T. giachinoi, sp. nov. exhibits morphological traits suitable for thriving in the smaller pores of the superficial network of underground fissures (Milieu Souterrain Superficiel, MSS), whereas T. bornensis shows a greater adaptation to the deep subterranean habitat. Our results suggest that different selective regimes within the subterranean environment, i.e. deep caves v. MSS, may either drive local speciation or facilitate contiguous distributions of independently subterranean adapted species.

On climate change and subterranean spiders

Subterranean ecosystems offer intriguing opportunities to study mechanisms underlying responses to changes in climate because species within them are often adapted to largely constant temperatures. However, responses of specialized subterranean species to anthropogenic climate warming are still largely undiscussed. We combined physiological tests, species distribution models and genetic data to investigate the potential effect of raising temperatures on subterranean coenosis. We used spiders of the genus Troglohyphantes Joseph, 1881 (Araneae: Linyphiidae) as model organisms, focusing on a coherent biogeographic area of the Western Alps in which the distribution of these spiders has been well documented. Thermal tolerance experiments in climatic chambers pointed at a reduced physiological tolerance to temperature fluctuations at increasing levels of troglomorphism. This result suggests that, during their subterranean evolution, spiders have progressively fine-tuned thermal tolerance to the constant and narrow temperature ranges of their habitats. Further evidence of the sensitivity of our model species to temperature increase derives from species distribution models projected onto different climate change scenarios. Model projections point toward a future decline in habitat suitability for subterranean spiders. Moreover, genetic data at the population/species interface are suggestive of limited gene flow between subterranean populations, testifying reduced dispersal capacity and habitat connectivity. In light of these results, we predict the potential extinction of the most restricted endemic species. Our findings therefore emphasize the importance of considering subterranean organisms as model species for ecological studies dealing with climatic changes, and to extend such investigations to other subterranean systems worldwide. ‡ ‡ ‡ ‡ ‡