Research Article |
Corresponding author: Stefano Mammola ( stefano.mammola@cnr.it ) Corresponding author: Marco Isaia ( marco.isaia@unito.it ) Academic editor: Oana Teodora Moldovan
© 2017 Stefano Mammola, Elena Piano, Pier Mauro Giachino, Marco Isaia.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Mammola S, Piano E, Giachino PМ, Isaia M (2017) An ecological survey of the invertebrate community at the epigean/hypogean interface. Subterranean Biology 24: 27-52. https://doi.org/10.3897/subtbiol.24.21585
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We studied the ecological continuum between caves and the associated network of fissures – Milieu Souterrain Superficiel (MSS) – in an hypogean site in the Graian Alps, Italy. Over one year, we surveyed the faunal assemblages by means of pitfall traps placed in the caves and specific subterranean sampling devices (SSD) buried in the MSS. We used generalized linear mixed models (GLMMs) and generalized additive mixed models (GAMMs) to compare the spatial and temporal dynamics of the subterranean invertebrates inhabiting the two environments. As expected, arthropod communities occurring near the surface were characterized by minor level of subterranean adaptations, and conversely, subterranean species were more abundant and diversified at higher depths, both in the caves and in the MSS. Diversity and abundance of external elements in the superficial layers were found to be highly seasonal dependent, with minor values in winter compared to the other seasons. We provided information about the faunal assemblages dwelling in the two hypogean compartments, and we characterized the microclimatic conditions therein. We discussed the existence of an ecological gradient of specialization extending from the surface to the deep hypogean layers, which can be interpreted in light of the microclimatic changes occurring at increasing depths and the parallel decrease in available organic matter.
Mesovoid Shallow Substratum, Cave fauna, Superficial Subterranean Habitats, Subterranean biology, Subterranean Sampling Device, Ecological gradient, Troglobionts
According to the modern view of subterranean biology, subterranean organisms do not exclusively inhabit underground vacuums of wide dimensions (i.e. caves), but also naturally occupy the network of fissures the size of which is not commensurable to the human scale (
Since MSS is not accessible to men unless by indirect means (see, e.g.,
We conducted a one-year ecological study in an alpine hypogean site, aiming at investigating simultaneously the cave environment and the surrounding MSS. Our aims were to 1) compare the faunal assemblages characteristic of the two subterranean compartments; 2) investigate whether a temporal (seasonal) and/or a spatial gradient of specialization exists in the MSS – i.e. higher richness and abundance of specialized elements at increasing depth; 3) investigate whether the same gradients exist in the cave – i.e. variation in richness and abundance related to season and/or to vertical distance from the surface (subjacency) or to distance from the cave entrance.
The study was set in the Pugnetto hypogean complex, in the nearby to the hamlet of Pugnetto, municipality of Mezzenile, Lanzo Valley, Graian Alps, Piedmont (NW-Italy). The site is protected under the European Habitat Directive 43/92 (S.C.I. IT 1110048) and hosts five natural caves classified as “Caves not open to the public” (H 8310), namely the Borna di Pugnetto (cadastre number Pi 1501, entrance at N45°16'19", E5°02'26"; altitude 820 m a.s.l.), the Tana del Lupo (Pi 1502, N45°16'19", E5°02'22"; 813 m a.s.l.), the Creusa d‘le Tampe (Pi 1503, N45°16'12", E5°02'33"; 870 m a.s.l.), the Tana della Volpe (Pi 1504, N45°16'13", E5°02'34", 885 m a.s.l.) and the Cavernetta (N45°16'17", E5°02'36", 895 m a.s.l.) which still lacks a cadastre number but was mentioned in the original description of the hypogean site by
Our study was conducted in the Borna di Pugnetto (hereinafter Borna) and in the Creusa d‘le Tampe (hereinafter Creusa) caves (Figs
a Main entrance of the Borna di Pugnetto (photo credit: Alberto Chiarle and Mauro Paschetta, 2014) b Main entrance of the Creusa d’le Tampe (photo credit: Elena Piano, 2013) c exposed soil/MSS profile in a fresh-cut along a slope in the vicinity of the Borna di Pugnetto (photo credit: Jacopo Orlandini, 2014) d the typical cave geo-morphology within the Borna di Pugnetto (photo credit: Alberto Chiarle and Mauro Paschetta, 2014) e detail of the MSS geo-morphological structure (photo credit: Jacopo Orlandini, 2014).
Map of the study area. The shape and the topographic position of the four caves (Borna Maggiore di Pugnetto, Tana del Lupo, Creusa d’le Tampe, Tana della Volpe) was obtained from the original planimetric drawings of
We used twenty-four pitfall traps (diameter 9 cm, volume 40 ml) to collect invertebrates in caves. The pitfall traps were arranged in groups of three (hereinafter cave-triplets), at a distance of ~5 m to one another. Six cave-triplets (code: C1–C6) were placed in the Borna at 4, 30, 90, 150, 230 (“Ramo della Madonna”) and 350 (“Ramo della Fontana”) meters from the main entrance. Two cave-triplets (code: C7, C8) were placed in the Creusa at 5 and 25 meters from the main entrance (Fig.
For collecting invertebrates in the MSS, we utilized twenty-four Subterranean Sampling Devices (SSD; after
Three adjacent SSDs of three different length (40, 60 and 80 cm) were buried vertically in the ground (hereinafter MSS-triplet; Fig.
Six pitfall traps (with brine, not baited) were also placed in leaf litter habitat (epigean; code: L1–L6), as a reference to discriminate correctly between the epigean fauna and the specialized fauna – see paragraph “Specimens sorting and ecological classification” for details.
Figure
To characterize the subterranean microclimate, we placed one Hygrochron temperature and humidity datalogger in correspondence of each pitfall trap in the cave and at the lower-end of each SSD. Hygrochron were programmed to sample temperature (T) and relative humidity (RH) every three hours for the whole sampling period (accuracy of ± 0.5°C and ± 1%, respectively). We also derived the mean daily outside temperature to the same periods from the nearest thermo-hygro-pluviometric weather station (Fua, Lanzo Torinese, Cod. 111; N45°17'23", E7°29'38"; 550 m a.s.l.). The temperature values recorded by the weather station were corrected with the standard environmental lapse rate – the change of temperature with altitude for the stationary atmosphere. In all analyses relating the abundance and species richness of external and adapted elements with the explanatory parameters (see later sections), we used the pseudo-replicates of each trap as basic sample units.
Trapped individual were sorted, identified and classified either as epigean (category: “external”) or subterranean elements (category: “adapted”). According to our expertise (MI and SM: Araneae; PMG: Coleoptera and some other orders of Insecta) and the availability of specialists for additional taxa (see Acknowledgments), identification of the species levels was possible for the orders Araneae, Opiliones, Pseudoscorpiones, Chilopoda, Isopoda (one species), and for most orders of insects (especially Coleoptera; see Appendix
In subterranean biology, species are often classified into ecological categories (e.g. trogloxenes, troglophiles, troglobionts) according to their preferred habitat and general association with the subterranean domain (
We performed regression-type analysis following the general advices of
The first set of models was computed through mixed-design analysis of variance with Poisson distributed data (i.e. generalized linear mixed models, GLMMs; equation 1), whereas for the second (equation 2) and third (equation 3) sets, we primarily relied on Poisson generalized additive mixed models (GAMMs). The mixed part of both GLMMs and GAMMs was introduced in order to account for multiple observations from the same triplet over time, by specifying the triplet as random factor. The latter variable was included as random factor in order to account for the variation it introduced in our samples – and thus to correctly estimate the regression coefficients, – rather than to test for its direct effect on the dependent variables.
Prior to model fitting, we explored the three datasets following the standard protocol for data exploration proposed by
Comparing MSS and cave communities (aim 1)
Species richness of external elements and abundance of adapted and external elements were analysed in relation to the explanatory two-levels factor HABITAT (levels: “Cave” and “MSS”) using the glmer R command in the package ‘lme4’ (
The fixed structure of the model was:
(1) y ~ HABITAT
Where y is one of Next, Rext and Nad. The random part of the model allowed us to deal with repeated observations and measurements of the same triplet (temporal dependence) and the clumped distribution of the traps within the triplet (spatial dependence).
Spatial and temporal gradients in MSS (aim 2) and cave (3) communities
Species richness of external elements and abundance of adapted and external elements were analysed in relation to the explanatory variables using the gamm R command in the package ‘mgcv’ (
For both the cave and the MSS compartments, we also included the two-levels factor BAIT (level= “Fresh” and “Exhausted”) as covariates and the two-levels factor SITE (levels= “Borna” and “Creusa”). The first variable was included to evaluate the effect of the ageing of the bait on our dependent variables. The second variable was introduced to take into account for possible local effects, since we pooled together records from two different caves and associated MSS.
For the analysis of the MSS compartment, in addition to the above mentioned variables we also included in the models the three-level categorical variable sampling depth (DEPTH; levels: “0.4m”, “0.6m” and “0.8m”). For the cave habitat, in addition to the aforementioned variables, we also included the distance from the cave entrance (Dst_i; continuous variable) and the three-levels categorical variable subjacency (SUBJ; levels: “0–20m”, “20–40m” and “60–80m”). We excluded from the regression analysis the microclimatic variables, given that due to malfunctioning, several dataloggers did not recorded reliable measurements of temperature and relative humidity during the sampling period (see results, further details in
The (fixed) structures of the initial models were (aims 2 and 3, respectively):
(2) y ~ DEPTH + s(Serie_i) + BAIT + SITE
(3) y ~ Dst_i + SUBJ + s(Serie_i) + BAIT + SITE
Where y is one of Next, Rext and Nad, and s(Serie_i) indicate the smoothing term. The random part of the model is equal to the previous model (1). For the 2nd and 3rd models, we adopted a statistical hypothesis testing framework, whereby model reduction was carried out on the full model by sequentially deleting non-significant terms and potential interactions according to AIC values (
Approximately 15,700 Arthropoda (20 Orders), 117 Mollusca (order Pulmonata), 868 Crustacea (order Isopoda), and 14 Anellida (Lumbricus sp.) were collected. We report the complete list of the taxa and number of specimen collected in each habitat in Appendix
Due to malfunction, several dataloggers did not sample either temperature or relative humidity during the sampling period. Specifically, nine out of eighteen in the Borna, two out of six in the Creusa and ten out of twenty-four in the MSS. Due to this significant loss of data, it was not possible to include climatic data in the regression analyses.
Relative humidity in the caves proved to be almost constantly close to saturation, with mean monthly values ranging from 85 to 100 %. However, in the vicinity of the entrance the relative humidity dropped down to 70–75 % in winter. In the MSS the mean monthly relative humidity was always above 90 %. With regard to temperature, changes and max–min ranges were attenuated with increasing distance from the entrance and delayed in respect to the outside values. The mean annual temperature values deep inside the two caves was comparable (Tmean ± SD: Borna = 9.0 ± 0.4 °C; Creusa = 8.9 ± 0.8 °C) and presented little variations over the year. In the outermost sections, temperature was nearly stable in summer, spring and autumn, with a max–min range around 4.5 °C in the vicinity of the entrance for both caves. However, the microclimate at the entrance zone drastically changed during winter, when we observed a drop in the mean temperature values (mean values always below 6 °C). The coldest temperature values were recorded in December and January (Tmin: Borna = –2.0 °C and Creusa = –0.9 °C).
The range of temperature variation, both daily and monthly, was lower in the MSS in respect to surface (Fig.
Initial data exploration revealed the presence of a few outlying values (mostly due to higher prevalence of Diptera and Hymenoptera in certain traps), which were removed from the dataset. Both abundance and species richness of external elements were lower in caves in respect to MSS (Next: t = –4.37, p < 0.001; Rext: z = –4.93, p < 0.001). Conversely, the abundance of subterranean elements was higher in cave (Nad: t = 5.39, p < 0.001; Table
Estimated regression parameters and approximate significance of smooth terms according to GLMMs and GAMMs, respectively, obtained from the 3 sets of models and the 3 dependent variables considered (Next, Rext, Nad, see text). The final model structures resulting from model selection are reported (only fixed terms are shown).
y | Final model | Model | Distribution | Variables | Parametric coefficients: | Smooth terms | |||||
---|---|---|---|---|---|---|---|---|---|---|---|
Estimate (α or β) | SE | p | edf | F | p | ||||||
MSS vs CAVE (Equation 1) |
Next | ~ HABITAT | GLMM | Negative binomial | Intercept (α) | 0.9722 | 0.4490 | – | – | – | – |
HABITAT (CAVE) | -2.9168 | 0.6669 | <0.001 *** | – | – | – | |||||
Rext | ~ HABITAT | GLMM | Poisson | Intercept (α) | 0.1418 | 0.2975 | – | – | – | – | |
HABITAT (CAVE) | -2.2727 | 0.4613 | <0.001 *** | – | – | – | |||||
Nad | ~ HABITAT | GLMM | Negative binomial | Intercept (α) | -1.4751 | 0.4491 | – | – | – | – | |
HABITAT (CAVE) | 3.2843 | 0.6087 | <0.001 *** | – | – | – | |||||
MSS (Equation 2) |
Next | ~ BAIT + DEPTH + s(Serie_i) | GAMM | Poisson | Intercept (α) | 0.5339 | 0.2475 | – | – | – | – |
BAIT (Fresh) | 0.9040 | 0.2014 | <0.001 *** | – | – | – | |||||
DEPTH (0.6m) | -0.1260 | 0.1939 | 0.516 | – | – | – | |||||
DEPTH (0.8m) | -0.7835 | 0.2436 | 0.001 ** | – | – | – | |||||
s(Serie_i) | – | – | – | 3.321 | 9.02 | <0.001 *** | |||||
Rext | ~ BAIT + DEPTH + s(Serie_i) | GAMM | Poisson | Intercept (α) | -0.440 | 0.1813 | – | – | – | – | |
BAIT (Fresh) | 0.4196 | 0.1396 | 0.002 ** | – | – | – | |||||
DEPTH (0.6m) | 0.0497 | 0.1410 | 0.724 | – | – | – | |||||
DEPTH (0.8m) | -0.5601 | 0.1706 | 0.001 ** | – | – | – | |||||
s(Serie_i) | – | – | – | 4.478 | 10.68 | <0.001 *** | |||||
Nad | ~ BAIT + DEPTH | GLMM | Poisson | Intercept (α) | -3.4961 | 0.6420 | – | – | – | – | |
BAIT (Fresh) | 1.3337 | 0.2422 | <0.001 *** | – | – | – | |||||
DEPTH (0.6m) | 0.8145 | 0.3283 | 0.013 * | – | – | – | |||||
DEPTH (0.8m) | 1.3131 | 0.3030 | <0.001 *** | – | – | – | |||||
CAVE (Equation 3) |
Next | ~ SITE + s(Serie_i) x SUBJ | GAMM | Poisson | Intercept (α) | -2.7564 | 0.5078 | – | – | – | – |
SITE (Creusa) | 2.9922 | 0.8183 | <0.001 *** | – | – | – | |||||
s(Serie_i) x SUBj (0–20m) | – | – | – | 5383 | 5.544 | <0.001 *** | |||||
s(Serie_i) x SUBj (20–40m) | – | – | – | 1.000 | 0.193 | 0.661 | |||||
s(Serie_i) x SUBj (60–80m) | – | – | – | 1.000 | 0.194 | 0.660 | |||||
Rext | ~ SUBJ + Dst_i | GLMM | Poisson | Intercept (α) | –0.7185 | 0.2413 | – | – | – | – | |
SUBj (20–40m) | –1.0541 | 0.6363 | 0.097 | – | – | – | |||||
SUBJ (60–80m) | –2.5317 | 1.1017 | 0.021 * | – | – | – | |||||
Dst_i | –0.0077 | 0.0029 | 0.010 * | – | – | – | |||||
Nad | ~ SITE + BAIT + SUBJ + Dst_i | GLMM | Negative binomial | Intercept (α) | 1.3087 | 0.2343 | – | – | – | – | |
SITE (Creusa) | 0.8874 | 0.2993 | 0.003 ** | – | – | – | |||||
SUBj (20–40m) | 1.3205 | 0.2918 | <0.001 *** | – | – | – | |||||
SUBJ (60–80m) | 0.6406 | 0.2853 | 0.024 * | – | – | – | |||||
Dst_i | –0.0017 | 0.0009 | 0.072 | – | – | – |
The best model structures resulting from model selection concerning the analysis of MSS and caves (equations 2 and 3) are reported in Table
Abundance and richness of external elements showed a significant non-linear U-shaped trend in respect to the sampling series (Next: F = 9.02, p < 0.001; Rext: F = 10.68, p < 0.001), with higher values in summer and early autumn, followed by a drastic decline in winter and an uprising in spring (Fig.
Predicted values (black line) and 95% confidence intervals (grey surface) of the effect of the sampling series (Serie_i) on the abundance of external elements in the MSS (a), on the species richness of external elements in the MSS (b) and on the abundance of external elements in the cave at subjacency of 0–20m (c) derived from GAMM analyses. Only fixed effects are shown.
Concerning the cave habitat, we detected an higher abundance of external and adapted elements in the Creusa cave (test relative to “Borna”; Next: t = 3.693, p < 0.001; Nad: t = 2.965, p = 0.003), whereas the richness of external elements was not significantly influenced by this parameter (Fig.
We observed a decrease in the richness of external elements at increasing distance from the cave entrance (Rext; z = –2.575, p = 0.010). Moreover, the richness of external elements was lower at 60–80m subjacency (test relative to “0–20 m”; Rext: z = -2.298, p = 0.021). No significant effect was detected in respect to the 20–40m subjacency (Fig.
Despite subterranean animal communities being relatively simple, their precise characterization still represents an interesting topic in subterranean ecology. This is mostly because spatial boundaries and species composition of the communities are difficult to define, especially when considering MSS and its interconnections with the deep hypogean domain. Relying on the classical definitions (
In this contribution, we aimed to investigate this ecological continuum both spatially and temporally, by comparing the arthropod communities inhabiting caves and the adjacent MSS compartments (Appendix
Although several species sampled at the Pugnetto hypogean complex are unique for obvious biogeographical reasons, the composition of the animal community was in general terms quite similar to that reported from other MSS sites in the Canary Islands (
We found that the abundance of specialized organisms was higher in the cave compartment, whilst we documented a higher diversity and abundance of epigean species in the MSS. The dominance of epigean species in the superficial layers of the MSS has been reported by several authors (
Concerning the cave habitat, we detected a higher diversity and abundance in the Creusa rather than in the Borna (Fig.
When analysing the cave and MSS habitats separately, we detected a gradient of subterranean specialization of the biological community in both compartments. In the cave, regression analyses suggests that there was a gradient of subterranean specialization of the biological community from the entrance zone toward the deepest sectors (see, e.g.,
At the same time, we observed how the presence of external elements was seasonally dependent, their abundance being highly fluctuating during the year. We observed this trend both in the MSS (Fig.
As a side question, with this study we were able to test the effect of the quality of the bait – at least up to two months – on the probability of capturing invertebrates in the two subterranean environments. This was possible since we renewed the bait in the trap every two sampling sessions, so that at each sampling session the condition of the bait changed from fresh to exhausted. According to the regression analysis, we demonstrated that a fresh bait is more effective in capturing individuals in the MSS (Fig.
We would like to thank all the taxonomists that helped us to identify specific taxa, namely Axel L. Schonhofer (Opiliones), Giulio Gardini (Pseudoscorpiones), Marzio Zapparoli (Chilopoda) and Massimo Meregalli (Coleoptera, Curculionidae). We are greatful to Paolo Debernardi (Natural Park of La Mandria) for allowing us to access the Borna di Pugnetto during winter, and to all friends and colleagues which helped in the samplings: Michele and Luigi Motta, Massimo Meregalli, Jacopo Orlandini, Mauro Paschetta, Fabio Ferrero, Enrico Lana, Elena Piano Sr., Davide Giuliano and Martina Dalle. We are indebted to Alberto Sendra, Vlastmil Růžička and Oana Moldovan for their help in improving the quality of the manuscript through their useful comments during the review process.
This study was set within the CaveLab project “From microclimate to climate change: caves as laboratories for the study of the effects of temperature on ecosystems and biodiversity”, funded by University of Torino and Compagnia di San Paolo [grant number ORTO11T92F].
Taxonomic groups sampled in this study according to the habitat. The total number of sampled individuals in each habitat is reported. The column “adaptation” provides the attribution to the category “external” or “adapted”. Classification was based on the criteria defined in the relative paragraph.
Class | Order | Family | Species | Adaptation | Litter | MSS | Cave |
---|---|---|---|---|---|---|---|
Arachnida | Acarina | Fam. | Morphospecies I | Adapted | 87 | 24 | 137 |
Arachnida | Acarina | Fam. | Morphospecies II | External | 75 | – | – |
Arachnida | Acarina | Fam. | Morphospecies III | External | 18 | 26 | – |
Arachnida | Acarina | Fam. | Morphospecies IV | Adapted | – | 79 | 63 |
Arachnida | Acarina | Fam. | Morphospecies V | Adapted | – | 122 | – |
Arachnida | Araneae | Agelenidae | Histopona leonardoi Pantini & Isaia, 2013 | External | 17 | 14 | – |
Arachnida | Araneae | Agelenidae | Tegenaria silvestris L. Koch, 1872 | Adapted | 2 | 8 | 4 |
Arachnida | Araneae | Amaurobiidae | Amaurobius sp. | – | 2 | 1 | – |
Arachnida | Araneae | Dysderidae | Harpactocrates drassoides (Simon, 1882) | External | 37 | 34 | 1 |
Arachnida | Araneae | Fam. | Immatures indet. | – | 2 | 3 | – |
Arachnida | Araneae | Gnaphosidae | Drassodes sp. | – | – | 1 | – |
Arachnida | Araneae | Linyphiidae | Gen. sp. | Adapted | – | 6 | 2 |
Arachnida | Araneae | Linyphiidae | Mansuphantes aridus (Thorell, 1875) | Adapted | – | 1 | 1 |
Arachnida | Araneae | Linyphiidae | Micrargus alpinus Relys & Weiss, 1997 | Adapted | – | 1 | – |
Arachnida | Araneae | Linyphiidae | Porrhomma convexum (Westring, 1851) | Adapted | – | 2 | – |
Arachnida | Araneae | Linyphiidae | Troglohyphantes bornensis Isaia & Pantini, 2008 | Adapted | – | 4 | 15 |
Arachnida | Araneae | Linyphiidae | Troglohyphantes lucifer Isaia, Mammola & Pantini, 2016 | Adapted | – | 6 | 1 |
Arachnida | Araneae | Linyphiidae | Troglohyphantes n. sp. | Adapted | 2 | 6 | – |
Arachnida | Araneae | Lycosidae | Trochosa hispanica Simon, 1870 | External | 1 | – | – |
Arachnida | Araneae | Nesticidae | Kryptoesticus eremita (Simon, 1879) | Adapted | – | 3 | 2 |
Arachnida | Araneae | Philodromidae | Philodromus sp. | External | – | 1 | – |
Arachnida | Araneae | Phrurolithidae | Phrurolithus festivus (C. L. Koch, 1835) | External | – | 1 | – |
Arachnida | Araneae | Pimoidae | Pimoa graphitica Mammola, Hormiga & Isaia, 2016 | Adapted | – | – | 2 |
Arachnida | Araneae | Salticidae | Saitis barbipes (Simon, 1868) | External | 1 | – | – |
Arachnida | Araneae | Tetragnathidae | Meta menardi (Latreille, 1804) | Adapted | – | 1 | 2 |
Arachnida | Araneae | Tetragnathidae | Metellina merianae (Scopoli 1763) | Adapted | – | 1 | – |
Arachnida | Araneae | Theridiidae | Gen. sp. | – | – | 1 | – |
Arachnida | Araneae | Theridiidae | Pholcomma gibbum (Westring, 1851) | External | 3 | – | – |
Arachnida | Opiliones | Dicranolasmatidae | Dicranolasma soerensenii Thorell, 1876 | External | 35 | 3 | – |
Arachnida | Opiliones | Ischyropsalidae | Ischyropsalis dentipalpis Canestrini, 1872 | Adapted | – | – | 1 |
Arachnida | Opiliones | Nemastomatidae | Nemastoma dentigerum Canestrini, 1873 | External | 29 | 1 | – |
Arachnida | Opiliones | Nemastomatidae | Paranemastoma quadripunctatum (Perty, 1833) | External | 19 | 4 | – |
Arachnida | Opiliones | Phalangiidae | Gen. sp. | – | 1 | – | – |
Arachnida | Opiliones | Phalangiidae | Astrobunus bernardinus Simon, 1879 | External | 22 | 1 | – |
Arachnida | Opiliones | Phalangiidae | Leiobunum religiosum (Simon, 1879) | – | – | – | 1 |
Arachnida | Opiliones | Phalangiidae | Odiellus coronatus (Roewer, 1911) | External | 4 | – | – |
Arachnida | Opiliones | Trogulidae | Anelasmocephalus rufitarsis Simon, 1879 | External | 17 | – | – |
Arachnida | Opiliones | Trogulidae | Trogulus nepaeformis (Scopoli, 1763) | External | 13 | 1 | – |
Arachnida | Pseudoscorpiones | Chthoniidae | Chthonius (C.) tenuis L. Koch, 1873 | External | 10 | 7 | 2 |
Arachnida | Pseudoscorpiones | Chthoniidae | Chthonius (Globochthonius) globifer Simon, 1879 | External | 6 | 3 | 2 |
Arachnida | Pseudoscorpiones | Chthoniidae | Chthonius sp. | – | – | 3 | – |
Arachnida | Pseudoscorpiones | Neobisiidae | Roncus sp. | External | 11 | – | – |
Chilopoda | Ord. | Fam. | Gen. sp. | External | 161 | – | – |
Chilopoda | Geophilomorpha | Linotaeniidae | Strigamia acuminata (Leach, 1814) | – | – | 2 | – |
Chilopoda | Lithobiomorpha | Lithobiidae | Eupolybothrus tridentinus (Fanzago, 1874) | – | – | 1 | – |
Chilopoda | Lithobiomorpha | Lithobiidae | Lithobius pellicensis Verhoeff, 1935 | – | – | – | 3 |
Chilopoda | Lithobiomorpha | Lithobiidae | Lithobius pilicornis Newport, 1844 | – | – | 1 | – |
Chilopoda | Scolopendromorpha | Cryptopidae | Cryptops parisi Brolemann, 1920 | – | – | 1 | – |
Clitellata | Haplotaxida | Lumbricidae | Lumbricus sp. | External | 12 | 2 | – |
Diplopoda | Chordeumatida | Craspedosomatidae | Gen. sp. | External | 511 | 33 | 4 |
Diplopoda | Glomerida | Glomeridae | Gen. sp. | External | 26 | 6 | |
Entognatha | Collembola | Fam. | Morphospecies I | External | 45 | 79 | 285 |
Entognatha | Collembola | Fam. | Morphospecies II | Adapted | – | – | 323 |
Entognatha | Collembola | Fam. | Morphospecies III | Adapted | – | 321 | 477 |
Entognatha | Collembola | Fam. | Morphospecies IV | Adapted | 1 | 127 | – |
Entognatha | Collembola | Fam. | Morphospecies V | External | 22 | – | – |
Entognatha | Collembola | Fam. | Morphospecies VI | External | 75 | – | – |
Entognatha | Diplura | Fam. | Gen. sp. | External | 76 | – | – |
Gastropoda | Pulmonata | Helicidae | Helix sp. | External | 35 | – | – |
Gastropoda | Pulmonata | Limacidae | Limax sp. | External | 22 | 36 | – |
Gastropoda | Pulmonata | Oxychilidae | Oxychilus sp. | Adapted | – | 22 | 2 |
Insecta | Blattodea | Fam. | Larvae indet. | External | – | 4 | – |
Insecta | Blattodea | Blattellidae | Blattella sp. | External | 10 | 2 | – |
Insecta | Coleoptera | Fam. | Larvae indet. | – | 18 | 42 | 44 |
Insecta | Coleoptera | Carabidae | Abax continuus Ganglbauer, 1891 | External | 163 | 3 | – |
Insecta | Coleoptera | Carabidae | Bembidion (Peryphanes) deletum Audinet Serville, 1821 | – | – | 1 | – |
Insecta | Coleoptera | Carabidae | Binaghites affinis ovalipennis (Ganglbauer, 1900) | External | 1 | – | – |
Insecta | Coleoptera | Carabidae | Carabus intricatus Linne, 1761 | External | 3 | – | – |
Insecta | Coleoptera | Carabidae | Carabus monticola Dejean, 1826 | External | 26 | – | – |
Insecta | Coleoptera | Carabidae | Cychrus italicus Bonelli, 1810 | External | 28 | 1 | – |
Insecta | Coleoptera | Carabidae | Limodromus assimilis (Paykull, 1790) | External | 1 | – | – |
Insecta | Coleoptera | Carabidae | Platynus complanatus Dejean, 1828 | External | 4 | 2 | 1 |
Insecta | Coleoptera | Carabidae | Pterostichus (Oreophilus) externepunctatus (Dejean, 1828) | External | 106 | 38 | – |
Insecta | Coleoptera | Carabidae | Pterostichus (Pterostichus) rutilans (Dejean, 1828) | Adapted | – | 5 | – |
Insecta | Coleoptera | Carabidae | Sphodropsis ghilianii ghilianii (Schaum, 1858) | Adapted | 2 | 189 | 381 |
Insecta | Coleoptera | Carabidae | Stomis (Stomis) elegans Chaudoir, 1861 | External | 6 | – | – |
Insecta | Coleoptera | Carabidae | Trechus modestus Putzeys, 1874 | – | – | 1 | – |
Insecta | Coleoptera | Cholevidae | Apocatops monguzzii Giachino & Vailati, 1987 | Adapted | – | 4 | 1 |
Insecta | Coleoptera | Cholevidae | Bathysciola pumilio (Reitter, 1884) | Adapted | 49 | 150 | 34 |
Insecta | Coleoptera | Cholevidae | Catops fuliginosus Erichson, 1837 | – | – | 1 | – |
Insecta | Coleoptera | Cholevidae | Catops subfuscus Kellner, 1846 | Adapted | – | 195 | 175 |
Insecta | Coleoptera | Cholevidae | Catops tristis (Panzer, 1794) | – | 1 | 2 | – |
Insecta | Coleoptera | Cholevidae | Catops ventricosus rotundatus Szymczakowski, 1963 | External | 161 | 113 | 2 |
Insecta | Coleoptera | Cholevidae | Choleva angustata (Fabricius, 1781) | Adapted | – | – | 4 |
Insecta | Coleoptera | Cholevidae | Dellabeffaella roccae (Capra, 1924) | Adapted | – | 1 | 991 |
Insecta | Coleoptera | Cholevidae | Sciodrepoides watsoni (Spence, 1815) | Adapted | – | 38 | – |
Insecta | Coleoptera | Colonidae | Colon sp. | – | – | 4 | – |
Insecta | Coleoptera | Curculionidae | Otiorhynchus salicicola Heyden, 1908 | External | 11 | 3 | – |
Insecta | Coleoptera | Elateridae | Gen. sp. | – | – | 1 | – |
Insecta | Coleoptera | Latridiidae | Gen. sp. | External | 4 | – | – |
Insecta | Coleoptera | Leiodidae | Gen. sp. | – | 2 | 1 | – |
Insecta | Coleoptera | Ptinidae | Gen. sp. | – | 2 | 1 | – |
Insecta | Coleoptera | Scaphydidae | Gen. sp. | – | 3 | – | – |
Insecta | Coleoptera | Scarabeidae | Gen. sp. | External | 111 | – | 1 |
Insecta | Coleoptera | Scydmaenidae | Gen. sp. | External | – | – | 1 |
Insecta | Coleoptera | Silphidae | Nicrophorus sp. | External | 137 | 14 | – |
Insecta | Coleoptera | Silphidae | Silpha sp. | External | 6 | – | – |
Insecta | Coleoptera | Staphylinidae | Bryaxis brachati Besuchet, 1980 | – | – | 2 | – |
Insecta | Coleoptera | Staphylinidae | Gen. sp. | External | 111 | 133 | – |
Insecta | Coleoptera | Staphylinidae | Gen. sp. (Pselaphinae) | Adapted | – | 4 | 2 |
Insecta | Dermaptera | Forficulidae | Forficula sp. | External | 2 | 2 | – |
Insecta | Diptera | Fam. | Larvae indet. | – | 289 | 421 | 787 |
Insecta | Diptera | Limoniidae | Chionea sp. | Adapted | – | 37 | 1 |
Insecta | Diptera | Limoniidae | Gen. sp. | Adapted | 7 | 570 | 2085 |
Insecta | Diptera | Muscidae | Gen. sp. | External | 100 | 79 | – |
Insecta | Diptera | Phoridae | Gen. sp. | External | 1071 | 1220 | 802 |
Insecta | Hymenoptera | Formicidae | Gen. sp. | External | 360 | 678 | 3 |
Insecta | Hymenoptera | Vespidae | Gen. sp. | External | 4 | 1 | – |
Insecta | Orthoptera | Acrididae | Gen. sp. | External | 9 | – | – |
Insecta | Orthoptera | Gryllidae | Gryllus sp. | – | 2 | – | – |
Insecta | Orthoptera | Rhaphidophoridae | Dolichopoda azami septentrionalis Baccetti & Capra, 1958 | Adapted | – | 1 | 14 |
Insecta | Rhyncota | Fam. | Gen. sp. | – | – | 3 | – |
Insecta | Rhyncota | Pentatomidae | Pentatoma sp. | External | 7 | – | – |
Insecta | Rhyncota | Pyrrhocoridae | Pyrrhocoris apterus (Linnaeus, 1758) | External | 3 | – | – |
Malacostraca | Isopoda | Armadillidiidae | Gen. sp. | External | 569 | 14 | – |
Malacostraca | Isopoda | Trichoniscidae | Alpioniscus feneriensis caprae Parona, 1880 | Adapted | – | 6 | 279 |
Zygentoma | Thysanura | Lepismatidae | Lepisma sp. | External | 11 | 1 | – |