Does Size Matter? Two Subterranean Biodiversity Hotspots in the Lessini Mountains in the Veneto Prealps in Northern Italy

Abstract

In the Lessini Mountains, the southernmost prealpine area in the Veneto region, thousands of caves are found, many of which have been extensively studied from the biological point of view. Numerous studies have been carried out on taxonomic and biogeographic aspects over the last hundred years. Two caves, in particular, have been found to be extremely rich in species adapted to life in subterranean environments. These are the Arena Cave in the Monti Lessini Veronesi and the Buso della Rana cave system in the Monti Lessini Vicentini. The two caves have extremely different development: Arena Cave is about 100 meters in length, and the Buso della Rana-Pisatela cave system is more than 37 km in length. Despite this huge difference in size, they both have the highest number of subterranean dwelling species in northern Italy (16 troglobionts and 8 stygobionts in Arena Cave, and 7 troglobionts and 11 stygobionts in the Buso della Rana-Pisatela cave system).

Full text

Citation: Latella, L. Does Size
Matter? Two Subterranean
Biodiversity Hotspots in the Lessini
Mountains in the Veneto Prealps in
Northern Italy. Diversity 2024,16, 25.
https://doi.org/10.3390/
d16010025
Academic Editors: Tanja Pipan,
David C. Culver, Louis Deharveng
and Michael Wink
Received: 6 October 2023
Revised: 14 December 2023
Accepted: 26 December 2023
Published: 30 December 2023
Copyright: © 2023 by the author.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
diversity
Article
Does Size Matter? Two Subterranean Biodiversity Hotspots in
the Lessini Mountains in the Veneto Prealps in Northern Italy
Leonardo Latella
Museo di Storia Naturale of Verona, Lungadige Porta Vittoria, 9, 37129 Verona, Italy;
leonardo.latella@comune.verona.it
Abstract:
In the Lessini Mountains, the southernmost prealpine area in the Veneto region, thousands
of caves are found, many of which have been extensively studied from the biological point of view.
Numerous studies have been carried out on taxonomic and biogeographic aspects over the last
hundred years. Two caves, in particular, have been found to be extremely rich in species adapted to
life in subterranean environments. These are the Arena Cave in the Monti Lessini Veronesi and the
Buso della Rana cave system in the Monti Lessini Vicentini. The two caves have extremely different
development: Arena Cave is about 100 meters in length, and the Buso della Rana-Pisatela cave system
is more than 37 km in length. Despite this huge difference in size, they both have the highest number
of subterranean dwelling species in northern Italy (16 troglobionts and 8 stygobionts in Arena Cave,
and 7 troglobionts and 11 stygobionts in the Buso della Rana-Pisatela cave system).
Keywords:
caves; Arena Cave; Buso della Rana-Pisatella cave system; biospeleology; checklist;
contact karst
1. Introduction
The Lessini Mountains, located in the western part of the Veneto Prealps (southeastern
Italian Alps) [
1
], extend over a total area of 1403 km
2
with a maximum elevation of 1865 m
a.s.l. They are bordered by the Adige Valley to the west and the Leogra Valley to the east
and northeast. The Val dei Ronchi separates the range from the Pasubio-Carega Group to
the northwest. From the geological point of view, the Lessini Mountains are dominated
by limestones from the Mesozoic and Cenozoic ages, interspersed by Cenozoic volcanic
rocks and Eocenic limestone outcrops [
2
]. Lessinia includes the western Lessini Mountains
(between the Adige and Illasi Valleys) and eastern Lessinia Mountains (between the Illasi
and Leogra Valleys). The western Lessini Mountains (Dolomia Principale, Calcari grigi,
Rosso Ammonitico, Scaglia Rossa, and Maiolica) mainly consist of carbonate rocks; some
areas of the eastern Lessinia Mountains are primarily composed of volcanic rocks developed
during the Venetian Tertiary magmatism [2].
In this mountain range, two caves have a high number of obligate subterranean species:
Arena Cave (having 16 troglobionts and 8 stygobionts), which is in the Central-Western
Lessini Mountains, and the Buso della Rana-Pisatella cave system (having 7 troglobionts
and 11 stygobionts), in the Eastern Lessini Mountains [
3
,
4
]. The two caves are located
22 km apart as the crow flies (Figure 1).
Despite the short distance between the two caves, which are situated in the same
mountain range, they show enormous differences in development, rock formations, and
fauna composition, making the separate analysis and comparison of the two caves in-
teresting. Especially evident is the huge difference in the length of the two caves. This
difference in size would suggest that the larger and more diversified one has a greater
richness of cave-dwelling species. To verify this hypothesis, we compare here, the number
of subterranean species present in each of the two caves. However, the two caves have
one important characteristic in common: both can be considered cases of contact karst
caves, i.e., karst phenomena and forms influenced by the contact between two or more
Diversity 2024,16, 25. https://doi.org/10.3390/d16010025 https://www.mdpi.com/journal/diversity

Diversity 2024,16, 25 2 of 17
karstifiable rocks that differ in some characteristics, such as porosity, chemical composition
and fracture density, or a karstifiable rock and a non-karstifiable rock [5].
Diversity 2024, 16, x FOR PEER REVIEW 2 of 19
important characteristic in common: both can be considered cases of contact karst caves,
i.e., karst phenomena and forms influenced by the contact between two or more karstifia-
ble rocks that differ in some characteristics, such as porosity, chemical composition and
fracture density, or a karstifiable rock and a non-karstifiable rock [5].
Figure 1. Location of the two studied caves in the Lessinia Mountains (the red rectangle in the
small photo indicates the position in Italy).
1.1. Arena Cave (476 V/VR)
Arena Cave is registered as number 476 in the Cadastre of the Caves of Veneto Re-
gion. It is located in the Province of Verona, municipality of Bosco Chiesanuova, in the
Malga Bagorno area. Its location, at 11°06′02″ E 45°39′56″ N, has an altitude of 1512 m a.s.l.
(Figure 2).
Figure 1.
Location of the two studied caves in the Lessinia Mountains (the red rectangle in the small
photo indicates the position in Italy).
1.1. Arena Cave (476 V/VR)
Arena Cave is registered as number 476 in the Cadastre of the Caves of Veneto Region.
It is located in the Province of Verona, municipality of Bosco Chiesanuova, in the Malga
Bagorno area. Its location, at 11
◦
06
0
02
00
E 45
◦
39
0
56
00
N, has an altitude of 1512 m a.s.l.
(Figure 2).
The cave is 74 m long with a difference in elevation of
−
22 m from the entrance to the
bottom. It is formed by a large chamber, roughly elliptical in plane section, with a main
diameter of about 50 m. The roof coincides mostly with bedding planes. The southern part
of the floor is characterized by a large, asymmetrical, funnel-shaped depression, a type of
subterranean doline that developed in the collapse of debris [
6
] (Figure 3). The chamber is
connected to the surface through some narrow passages that start from an open collapse
depression located on a slope, which resembles a Roman theatre (i.e., an “Arena”, hence
the name of the cave).

Diversity 2024,16, 25 3 of 17
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Figure 2. Cave map of Arena Cave. Sezione = cross section; pianta = plan; ingresso = entrance.
Figure 2. Cave map of Arena Cave. Sezione = cross section; pianta = plan; ingresso = entrance.

Diversity 2024,16, 25 4 of 17
Diversity 2024, 16, x FOR PEER REVIEW 4 of 19
The cave is 74 m long with a difference in elevation of −22 m from the entrance to the
boom. It is formed by a large chamber, roughly elliptical in plane section, with a main
diameter of about 50 m. The roof coincides mostly with bedding planes. The southern part
of the floor is characterized by a large, asymmetrical, funnel-shaped depression, a type of
subterranean doline that developed in the collapse of debris [6] (Figure 3). The chamber
is connected to the surface through some narrow passages that start from an open collapse
depression located on a slope, which resembles a Roman theatre (i.e., an “Arena”, hence
the name of the cave).
Figure 3. The large chamber of the Arena Cave (Photo: L. Latella).
From the geological point of view, the cave is an expression of a contact karst, where
different limestone types are in contact both stratigraphically and along tectonic structures
[5,7,8]. The limestone formations present here are “Calcari del Gruppo di San Vigilio” of
lower-middle Jurassic, about 60 m in depth, both pure Oolitic and bio-sparitic/–ruditic, or
reef limestones, relatively densely fractured; “Rosso Ammonitico”, a condensed rock unit
of middle-upper Jurassic age, about 30 m in depth, made up of nodular micritic limestone
that is very resistant to erosion, crossed by widely spaced fractures; and “Biancone”, a
chalk-type unit, from the lower and middle Cretaceous, 100–200 m in depth, made up of
whitish marly limestone that is closely stratified and densely fractured, and very sensitive
to frost and atmospheric agents. Therefore, the cave develops at the stratigraphic contact
between the “Calcari del Gruppo di San Vigilio” and the “Rosso Ammonitico” and is close
to a fault plane, placing the two above formations in vertical contact with the “Biancone”.
Figure 3. The large chamber of the Arena Cave (Photo: L. Latella).
From the geological point of view, the cave is an expression of a contact karst, where
different limestone types are in contact both stratigraphically and along tectonic struc-
tures [
5
,
7
,
8
]. The limestone formations present here are “Calcari del Gruppo di San Vigilio”
of lower-middle Jurassic, about 60 m in depth, both pure Oolitic and bio-sparitic/–ruditic,
or reef limestones, relatively densely fractured; “Rosso Ammonitico”, a condensed rock unit
of middle-upper Jurassic age, about 30 m in depth, made up of nodular micritic limestone
that is very resistant to erosion, crossed by widely spaced fractures; and “Biancone”, a
chalk-type unit, from the lower and middle Cretaceous, 100–200 m in depth, made up of
whitish marly limestone that is closely stratified and densely fractured, and very sensitive
to frost and atmospheric agents. Therefore, the cave develops at the stratigraphic contact
between the “Calcari del Gruppo di San Vigilio” and the “Rosso Ammonitico” and is close
to a fault plane, placing the two above formations in vertical contact with the “Biancone”.
The overlying rocks of the cave are formed by the massive beds of lower Rosso Ammonitico,
whereas the inner cave is mostly developed inside the Calcari del Gruppo di San Vigilio.
At the topographical surface, the line of the normal fault runs along a small valley a few
meters to the east of the cave; the displacement of the fault is about 100 m.
From a hydrological point of view, the water circulates diffusely inside the dense
network of discontinuities of the Biancone unit; the preferential flow is sub-parallel to
the topographical surface and occurs mostly below the dry valley bottoms but is also
influenced by the structural setting; vertical losses occur along the fault and fracture zones.

Diversity 2024,16, 25 5 of 17
In contrast, the water circulation is more concentrated and mostly vertical in the Rosso
Ammonitico [6].
1.2. Buso della Rana-Pisatela Cave System (40 V–VI/1707 V–VI)
Buso della Rana cave (40 V–VI) opens at an altitude of 340 m a.s.l. in the province
of Vicenza, municipality of Monte di Malo. It has a length of 30,102 m and an altitudinal
range of 274 m.
In 2012, the cave, which had only one entrance, was connected to the Pisatela Cave
(1707 V–VI), a cavity with two entrances (Pater Noster and Pisatela), the highest of which
opens at 747 m a.s.l. at a development of 7510 m. The two cavities thus give rise to the
Rana-Pisatella cave system, with a development of 37,612 m and an altitude difference of
407 m between the upper (Pisatella) and lower (Rana) entrances (Figure 4).
Diversity 2024, 16, x FOR PEER REVIEW 5 of 19
The overlying rocks of the cave are formed by the massive beds of lower Rosso Ammonit-
ico, whereas the inner cave is mostly developed inside the Calcari del Gruppo di San
Vigilio. At the topographical surface, the line of the normal fault runs along a small valley
a few meters to the east of the cave; the displacement of the fault is about 100 m.
From a hydrological point of view, the water circulates diffusely inside the dense
network of discontinuities of the Biancone unit; the preferential flow is sub-parallel to the
topographical surface and occurs mostly below the dry valley booms but is also influ-
enced by the structural seing; vertical losses occur along the fault and fracture zones. In
contrast, the water circulation is more concentrated and mostly vertical in the Rosso Am-
monitico [6].
1.2. Buso della Rana-Pisatela Cave System (40 V–VI/1707 V–VI)
Buso della Rana cave (40 V–VI) opens at an altitude of 340 m a.s.l. in the province of
Vicenza, municipality of Monte di Malo. It has a length of 30,102 m and an altitudinal
range of 274 m.
In 2012, the cave, which had only one entrance, was connected to the Pisatela Cave
(1707 V–VI), a cavity with two entrances (Pater Noster and Pisatela), the highest of which
opens at 747 m a.s.l. at a development of 7510 m. The two cavities thus give rise to the
Rana-Pisatella cave system, with a development of 37,612 m and an altitude difference of
407 m between the upper (Pisatella) and lower (Rana) entrances (Figure 4).
Figure 4. Plan of the Rana-Pisatela cave system (scale bar: 100 m).
The cave system is located on the Faedo-Casaron Plateau, which occupies a geo-
graphical area of 15 square kilometers in the province of Vicenza.
The area lies between the latitude of 45°36′46″ (Cornedo) to the South and 45°39′35″
(Monte di Malo) to the North and a longitude of 11°19′46″ (Monte Faedo) to the West and
11°22′22″ (Priabona) to the East. Morphologically, it is made up of limestone that gives
rise to gentle and rounded surface forms typical of hills, with the valleys oriented accord-
ing to the main Lessinian tectonic lines.
The Buso della Rana-Pisatela cave system, with its 37,612 m of development, is one
of the longest caves in Italy. It developed in the Oligocene limestone through joint net-
works from the Faedo-Casaron plateau towards the less permeable basalt surface. Here,
several independent and differently sized brooks descend along the gradient of the trans-
gressive contact towards the entrance of the Buso della Rana cave, which is the main
spring of the karst system forming the Rana River. Pisatela Cave is an inactive sink located
Figure 4. Plan of the Rana-Pisatela cave system (scale bar: 100 m).
The cave system is located on the Faedo-Casaron Plateau, which occupies a geograph-
ical area of 15 square kilometers in the province of Vicenza.
The area lies between the latitude of 45◦3604600 (Cornedo) to the South and 45◦390350 0
(Monte di Malo) to the North and a longitude of 11
◦
19
0
46
00
(Monte Faedo) to the West and
11
◦
22
0
22
00
(Priabona) to the East. Morphologically, it is made up of limestone that gives rise
to gentle and rounded surface forms typical of hills, with the valleys oriented according to
the main Lessinian tectonic lines.
The Buso della Rana-Pisatela cave system, with its 37,612 m of development, is one of
the longest caves in Italy. It developed in the Oligocene limestone through joint networks
from the Faedo-Casaron plateau towards the less permeable basalt surface. Here, several
independent and differently sized brooks descend along the gradient of the transgressive
contact towards the entrance of the Buso della Rana cave, which is the main spring of the
karst system forming the Rana River. Pisatela Cave is an inactive sink located inside a
doline that reaches the main stream through a series of narrow meanders and shafts [
9
,
10
].
The drainage in the cave system is primarily controlled by contact with the less
permeable basalt surface, basal conglomerate, and terrigenous marls of the Priabona
Formation that rise above the contact in the eastern sector of the system. In some parts
of the cave, the conduits evolved entirely in the Boro conglomerate, which is almost 2 m
thick in the upstream part of Pisatela Cave. Here, the conglomerate covers the lower basalt
contact, while in Buso della Rana, it is often absent, and the calcarenites lie directly on the
basalts [9].

Diversity 2024,16, 25 6 of 17
Cave morphologies are mostly vadose, with deep canyons and meanders and large
collapse rooms at the intersection of different streams and fracture sets. Small paleo-
phreatic conduits, evolved entirely in the calcarenite, hanging about 10–15 m above the
basalt surface, are related to the phreatic primary drainage and locally ancient epiphreatic
conditions. Various vadose narrow shafts reach the contact galleries from the above
overlying plateau, showing condensation corrosion morphologies [9].
During periods of heavy rain, the level of the streams rises, often flooding some areas
of the cave. In the terminal parts (the last few thousand meters towards the entrance), the
river shows a well-structured hyporheic zone (Figures 5–9).
Diversity 2024, 16, x FOR PEER REVIEW 6 of 19
inside a doline that reaches the main stream through a series of narrow meanders and
shafts [9,10].
The drainage in the cave system is primarily controlled by contact with the less per-
meable basalt surface, basal conglomerate, and terrigenous marls of the Priabona For-
mation that rise above the contact in the eastern sector of the system. In some parts of the
cave, the conduits evolved entirely in the Boro conglomerate, which is almost 2 m thick in
the upstream part of Pisatela Cave. Here, the conglomerate covers the lower basalt contact,
while in Buso della Rana, it is often absent, and the calcarenites lie directly on the basalts
[9].
Cave morphologies are mostly vadose, with deep canyons and meanders and large
collapse rooms at the intersection of different streams and fracture sets. Small paleo-phre-
atic conduits, evolved entirely in the calcarenite, hanging about 10–15 m above the basalt
surface, are related to the phreatic primary drainage and locally ancient epiphreatic con-
ditions. Various vadose narrow shafts reach the contact galleries from the above overlying
plateau, showing condensation corrosion morphologies [9].
During periods of heavy rain, the level of the streams rises, often flooding some areas
of the cave. In the terminal parts (the last few thousand meters towards the entrance), the
river shows a well-structured hyporheic zone (Figures 5–9).
Since the Rana-Pisatela system has only recently been joined, much of the research
(especially on aquatics) was carried out in the Buso della Rana cave, which will often be
cited separately in the text.
Figure 5. “Ramo aivo di destra”, one of the hydrologically active branches in the Buso della Rana
(Photo: S. Sedran).
Figure 5.
“Ramo attivo di destra”, one of the hydrologically active branches in the Buso della Rana
(Photo: S. Sedran).
Diversity 2024, 16, x FOR PEER REVIEW 7 of 19
Figure 6. The beautiful coralloid concretions in the “Ramo franchignia, Salea broccoli” in the Buso
della Rana (Photo: S. Sedran).
Figure 7. The left main branch of the Buso della Rana (Photo: S. Sedran).
Figure 6.
The beautiful coralloid concretions in the “Ramo franchignia, Saletta broccoli” in the Buso
della Rana (Photo: S. Sedran).

Diversity 2024,16, 25 7 of 17
Diversity 2024, 16, x FOR PEER REVIEW 7 of 19
Figure 6. The beautiful coralloid concretions in the “Ramo franchignia, Salea broccoli” in the Buso
della Rana (Photo: S. Sedran).
Figure 7. The left main branch of the Buso della Rana (Photo: S. Sedran).
Figure 7. The left main branch of the Buso della Rana (Photo: S. Sedran).
Diversity 2024, 16, x FOR PEER REVIEW 8 of 19
Figure 8. The large dimensions of the “Sala dei Massi”. One of the many large halls that characterize
the Rana–Pisatella cave system (Photo: S. Sedran).
Figure 8.
The large dimensions of the “Sala dei Massi”. One of the many large halls that characterize
the Rana–Pisatella cave system (Photo: S. Sedran).

Diversity 2024,16, 25 8 of 17
Diversity 2024, 16, x FOR PEER REVIEW 9 of 19
Figure 9. The shaft of the “Ramo del Pantano”, a tributary of the Main Branch of the Buso della Rana
(Photo: S. Sedran).
2. Materials and Methods
2.1. Sampling and Museum Collections
Both caves have been known since the first half of the last century, and numerous
biospeleological investigations have been carried out within them. The results of these
surveys have largely been published, but many unstudied specimens are present in the
collections of the Museo di Storia Naturale of Verona (Italy).
In recent years, research campaigns aimed to increase the knowledge of the troglobi-
otic and stygobiotic fauna as a whole were carried out [11–13].
Sampling of terrestrial fauna was carried out by means of pitfall traps and direct cap-
ture. The pitfall traps consisted of a plastic or glass cup, usually with an opening diameter
of 10 cm and a depth of 15 cm. Each was filled with a preserving liquid (NaCl solution) in
which was placed a tube containing an aracting bait of blue cheese. The traps were used
on a few occasions to integrate hand-collecting and were left in place for about a month
each time. In some cases, when we were unsure whether to return to the cave within a
short time, the bait was placed without the trap, and fauna were collected at sight.
Aquatic fauna were sampled in both caves by direct capture, hand nets, or using a
syringe to collect the water filling the pools. In the Buso della Rana, a drip funnel was
placed in the main branch 600 m from the entrance [13]. A drip funnel consists of a funnel
supported by a bucket that allows it to direct the dripping water into a plastic container.
A 2 cm × 3 cm area on two sides of the square container was cut out and covered with a
net (mesh size 60 µm) to retain the animals in the container [14,15].
The specimens sampled in the caves in the province of Verona, both in historical and
recent times and not taxonomically identified, have been deposited in the ‘miscellanea
Figure 9.
The shaft of the “Ramo del Pantano”, a tributary of the Main Branch of the Buso della Rana
(Photo: S. Sedran).
Since the Rana-Pisatela system has only recently been joined, much of the research
(especially on aquatics) was carried out in the Buso della Rana cave, which will often be
cited separately in the text.
2. Materials and Methods
2.1. Sampling and Museum Collections
Both caves have been known since the first half of the last century, and numerous
biospeleological investigations have been carried out within them. The results of these
surveys have largely been published, but many unstudied specimens are present in the
collections of the Museo di Storia Naturale of Verona (Italy).
In recent years, research campaigns aimed to increase the knowledge of the troglobiotic
and stygobiotic fauna as a whole were carried out [11–13].
Sampling of terrestrial fauna was carried out by means of pitfall traps and direct
capture. The pitfall traps consisted of a plastic or glass cup, usually with an opening
diameter of 10 cm and a depth of 15 cm. Each was filled with a preserving liquid (NaCl
solution) in which was placed a tube containing an attracting bait of blue cheese. The traps
were used on a few occasions to integrate hand-collecting and were left in place for about
a month each time. In some cases, when we were unsure whether to return to the cave
within a short time, the bait was placed without the trap, and fauna were collected at sight.
Aquatic fauna were sampled in both caves by direct capture, hand nets, or using a
syringe to collect the water filling the pools. In the Buso della Rana, a drip funnel was
placed in the main branch 600 m from the entrance [
13
]. A drip funnel consists of a funnel
supported by a bucket that allows it to direct the dripping water into a plastic container. A
2 cm
×
3 cm area on two sides of the square container was cut out and covered with a net
(mesh size 60 µm) to retain the animals in the container [14,15].
The specimens sampled in the caves in the province of Verona, both in historical and
recent times and not taxonomically identified, have been deposited in the ‘miscellanea
Biospeleologica’ collection of the Museum of Verona. In the course of the study for this
paper, the material collected in the two caves and not yet identified (i.e., Isopoda, Opilionida,
and Collembola) was sent to specialists.

Diversity 2024,16, 25 9 of 17
2.2. Bibliographic Research
Many papers on speleological (especially the Buso della Rana cave), geological, and
biological aspects have been published from the first half of the last century. As far as
biological research is concerned, in addition to the many publications devoted to the
description of new species or studies and reviews of certain faunitic groups, there have
been few works on the fauna as a whole of the two caves [16–19].
Therefore, we collected all publications to date and checked and updated the scientific
names. This was also important so as to identify and list all the species for which the two
caves represent the type locality. More than 60 publications were found in which, in various
ways, the presence of animal species in one (or both) of the two (or both) caves under study
is mentioned. For reasons of space and usefulness, all of them are obviously not listed in
the bibliography.
3. Results
The subterranean fauna of the two caves, as a whole, consists of 46 cave-dwelling
species. A total of 35 species are troglobionts or stygobionts, while 12 can be considered
eutroglophiles (sensu Ruffo, 1957) [
20
,
21
]. In the Arena Cave, 24 obligate subterranean
species are known, of which 16 are troglobionts and 8 are stygobionts; for the Rana-Pisatella
cave system, 18 species are known, of which 7 are troglobionts and 11 are stygobionts
(Table 1).
Table 1.
The list of troglobiotic and stygobiotic species known in the two studied caves.
Tb—troglobiont; Stb—stygobiont; 1—present; 0—absent; Tl—Type locality.
Class Order Family Genus/Species/Subspecies Status Arena Rana-Pisatella
Gastropoda Ellobiida Ellobiidae Zospeum globosum Kušˇcer, 1928 Tb 1 1
Arachnida Opiliones Ischyropsalididae Ischyropsalis strandti Kratochvíl,
1936 Tb 1 1
Arachnida Pseudoscorpionida Neobisiidae Neobisium (Blothrus) torrei
(Simon, 1881) Tb 1 1
Arachnida Pseudoscorpionida Neobisiidae Balkanoroncus boldorii (Beier,
1931) Tb 1 0
Arachnida Pseudoscorpionida Chthoniidae Chthonius lessiniensis Schawaller,
1982 Tb 1 0
Malacostraca Isopoda Trichoniscidae Androniscus (Dentigeroniscus)
degener Brian, 1926 Tb 1 1
Diplopoda Chordeumatida
Craspedosomatidae
Lessinosoma paolettii Strasser,
1967 Tb 1 Tl 0
Diplopoda Chordeumatida Iulidae Trogloiulus boldorii Manfredi,
1940 Tb 1 0
Collembola Poduromorpha Onychiuridae Onychiurus hauseri Dallai, 1975 Tb 1 0
Collembola Entomobryomorpha Entomobryidae Pseudosinella concii Gisin, 1950 Tb 1 0
Collembola Entomobryomorpha Entomobryidae Pseudosinella sp. Tb 1 0
Insecta Coleoptera Carabidae Italaphaenops dimaioi Ghidini,
1964 Tb 1 0
Insecta Coleoptera Carabidae Lessynodytes pivai Vigna
Taglianti & Sciaky, 1988 Tb 1 Tl 0
Insecta Coleoptera Carabidae Orotrechus pominii Tamanini,
1953 Tb 1 1
Insecta Coleoptera Carabidae Orotrechus vicentinus juccii
Pomini, 1940 Tb 1 0

Diversity 2024,16, 25 10 of 17
Table 1. Cont.
Class Order Family Genus/Species/Subspecies Status Arena Rana-Pisatella
Insecta Coleoptera Leiodidae Halberria zorzii (Ruffo, 1950) Tb 1 Tl 0
Insecta Coleoptera Leiodidae Lessiniella trevisioli Pavan, 1941 Tb 0 1 Tl
Insecta Coleoptera Leiodidae Neobathyscia fabianii (Dodero,
1904) Tb 0 1
Copepoda Cyclopoida Cyclopidae Speocyclops infernus Kiefer, 1930 Stb 1 1
Copepoda Harpactoida
Camptocamptidae
Elaphoidella phreatica (Chappuis,
1925) Stb 0 1
Copepoda Harpactoida
Camptocamptidae Elaphoidella ruffoi Chappuis, 1953
Stb 0 1
Copepoda Harpactoida
Camptocamptidae
Elaphoidella sp. A1 Stb 1 0
Copepoda Harpactoida
Camptocamptidae
Elaphoidella sp. A Stb 0 1
Copepoda Harpactoida
Camptocamptidae
Ceuthonectes serbicus Chappuis,
1924 Stb 0 1
Copepoda Harpactoida
Camptocamptidae
Lessinocamptus insoletus
(Chappuis, 1928) Stb 0 1 Tl
Copepoda Harpactoida
Camptocamptidae
Lessinocamptus pivai Stoch, 1997 Stb 0 1 Tl
Copepoda Harpactoida
Camptocamptidae
Lessinocamptus caoduroi Stoch,
1997 Stb 1 Tl 0
Copepoda Harpactoida
Camptocamptidae
Bryocamptus (Limocamptus)
echinatus (Mrazek, 1893) Stb 1 0
Copepoda Harpactoida
Camptocamptidae
Moraria (M.) sp. A1 Stb 1 0
Copepoda Harpactoida
Parastenocaridiidae
Parastenocaris ranae Stoch, 2000 Stb 0 1 Tl
Copepoda Harpactoida Amaeridae
Nitocrella psammophila Chappuis,
1955 Stb 1 1
Malacostraca Amphipoda Niphargidae
Niphargus costozzae Schellenberg,
1935 Stb 0 1
Malacostraca Amphipoda Niphargidae Niphargus similis Karaman &
Ruffo, 1989 Stb 1 0
Malacostraca Isopoda Sphaeromatidae Monolistra (Typhlosphaeroma)
bericum bericum Fabiani, 1901 Stb 0 1
Malacostraca Bathynellacea Bathynellidae Bathynella sp. Stb 1 0
Tot. 24 18
3.1. Terrestrial Fauna
Mollusca Gastropoda is represented by Zospeum globosum Kušcer, 1928. It is a small
mollusc (rarely exceeding 2 mm in height) with a translucent or transparent shell, a body
diaphanous, and totally lacking in ocular spots. It is mainly encountered on walls that are
damp or wet from dripping water and often covered by silty materials [22].
Among the Opiliones, Ischyropsalis strandi Kratochvil, 1936 is present (Figure 10). This
species is endemic to the caves in the Verona Prealps (Monte Baldo and Lessini Mountains).
They are present in the two caves studied, and can be found in other caves in the Lessinia
Mountains, usually above 600 meters of altitude, being a rather cryophilous species [
23
].
Within the Rana-Pisatella system, they are more frequently found in the higher elevations
of the Grotta della Pisatela.
In regard to Pseudoscorpionida, three troglobiotic species are present in the caves:
Chthonius (Chthonius)lessiniensis Schawaller, 1982, with Balkan affinities being a subter-
ranean species. They show a high degree of troglomorphy, ranging from the western
Venetian Prealps to the eastern Venetian Prealps [
24
], and are quite easy to detect under the
collapsed stones at the bottom of the hall in the Arena Cave. Neobisium (Blothrus)torrei (E.
Simon, 1881) is present in many caves in the Prealps and Alps of Veneto and Friuli Venezia
Giulia regions [
3
]. This species is found in the Arena Cave and is the only troglobiotic
pseudoscorpion currently known in the Rana-Pisatella system. Balkanoronchus boldorii

Diversity 2024,16, 25 11 of 17
(Beier, 1931) is present in some caves of the Prealps of Brescia, in the Monte Baldo and the
Lessini Mountains. These specimens were collected both with traps and hand-collecting in
the Arena Cave. This species frequents the same habitats as C. lessiniensis.
Diversity 2024, 16, x FOR PEER REVIEW 12 of 19
Figure 10. A specimen of Ischyropsalis strandi: this species is present in both investigated caves
(Photo: F. Rosseo).
The Terrestrial Isopoda species is represented by Androniscus (Dentigeroniscus) de-
gener Brian, 1926, and is troglobiont and endemic to the Lessini Mountains in the Verona
and Vicenza Provinces. It is quite common in both caves under stones in weer areas.
To date, troglobiotic millipedes have only been found within Arena Cave; these are
the Julidae Troglojulus boldorii Manfredi, 1940n, a species endemic to the caves of the
Figure 10.
A specimen of Ischyropsalis strandi: this species is present in both investigated caves (Photo:
F. Rossetto).

Diversity 2024,16, 25 12 of 17
The Terrestrial Isopoda species is represented by Androniscus (Dentigeroniscus)degener
Brian, 1926, and is troglobiont and endemic to the Lessini Mountains in the Verona and
Vicenza Provinces. It is quite common in both caves under stones in wetter areas.
To date, troglobiotic millipedes have only been found within Arena Cave; these are the
Julidae Troglojulus boldorii Manfredi, 1940n, a species endemic to the caves of the Prealps of
Lombardia, Veneto, and Trentino, and the Craspedosomatidae Lessinosoma paoletti Strasser,
1977, which is endemic to the Arena Cave [17].
The Troglobiotic Collembola are also known to date only from Arena Cave. The
Onychiuridae species are represented by Onychiurus hauseri Dallai, 1975 and are endemic
to the caves in the Veneto and Trentino regions, and the Entomobryidae Pseudosinella concii
Gisin, 1950 is a species distributed in different caves in Italy and Switzerland and is quite
common in the Arena Cave [
3
]. Some specimens belonging to the genus Pseudosinella
Schaeffer, 1897 are not identifiable on a species level since they are very damaged; these
were also sampled in the Arena Cave. However, it is possible to assert that they do not
belong to Pseudosinella concii on the basis of the different number of labium setae.
Coleoptera are the most interesting among the terrestrial animals found in the caves
considered in this study. Carabidae Trechinae, in particular, shows particularly robust
adaptations for life in subterranean environments, and among them is Italaphaenops dimaioi
Ghidini, 1964. Endemic to the Lessini Mountains in the Verona area, this species is one of the
largest subterranean Trechinae in the world. The colonization of caves by this troglobiont
can be traced back to an epoch preceding the Quaternary glaciations [
25
]. I.dimaioi is
known from some caves of the Veronese Lessini, which develop at the contact between
different types of karst formations like the Arena Cave. It is not present in the Vicenza
Province, so it is not present in the Rana-Pisatella cave system.
Another extremely specialized genus of ancient pre-Quaternary origin of Trechinae is
Lessinodytes Vigna Taglianti, 1982. This genus is distributed in the Lombardy and Veneto
Prealps with three species, which are all endemic to one or a few caves and is present in the
Arena Cave with the rather rare species Lessinodytes pivai Vigna Taglianti & Sciaky, 1988,
which is endemic to that cave [26,27].
Orotrechus vicentinus juccii Pomini, 1940, is endemic to the Lessini Mountains in Verona
Province and belongs to a group of species distributed in the Venetian Prealps and which,
in Arena Cave, co-occurs with Orotrechus pominii Tamanini, 1953. O.pomini is the only
known troglobiont Trechinae from the Rana-Pisatella system [28].
With regard to the Leiodidae Cholevinae, Halberria zorzii (Ruffo, 1950) is present in
Arena Cave, while in the Buso della Rana, we find Lessiniella trevisioli Pavan, 1941 and
Neobathyscia fabianii (Dodero, 1904) (Figure 11). The genus Halberria Conci & Tamanini,
1951 is present, with nine species in the caves in Eastern Veneto and Southern Trentino. In
the Western Lessini Mountains, H.zorzi is present only in the caves that open at higher
altitudes (above 1400 m a.s.l.), while in the caves at lower altitudes, the species of the genus
Neobathyscia show vicariant distributions [17]. It is quite common in the Arena Cave [18].
The genus Lessiniella Pavan, 1941 is phylogenetically close to Halberria [
29
,
30
] and
consists of two species: Lessiniella trevisoli, of which Buso della Rana is the typical locality
where it is found rare in the innermost areas, and Lessiniella berica Piva, 1993, from the
nearby Berici Mountains, which are also in the Vicenza Province [29].
To the genus Neobathyscia belongs nine species endemic to the Venetian Prealps,
distributed between the Adige and Piave Rivers [
30
]. N. fabiani is known from several
caves in the province of Vicenza that open in localities not far from the Rana-Pisatella cave
system [
29
]. Within the system, it is rather common, especially in the branches of the Buso
della Rana cave, while it seems rarer in the Pisatela cave.

Diversity 2024,16, 25 13 of 17
Diversity 2024, 16, x FOR PEER REVIEW 14 of 19
Figure 11. A specimen of Neobathyscia fabianii from the Rana-Pisatela cave system (Photo: L. Latella).
The genus Lessiniella Pavan, 1941 is phylogenetically close to Halberria [29,30] and
consists of two species: Lessiniella trevisoli, of which Buso della Rana is the typical locality
where it is found rare in the innermost areas, and Lessiniella berica Piva, 1993, from the
nearby Berici Mountains, which are also in the Vicenza Province [29].
To the genus Neobathyscia belongs nine species endemic to the Venetian Prealps, dis-
tributed between the Adige and Piave Rivers [30]. N. fabiani is known from several caves
in the province of Vicenza that open in localities not far from the Rana-Pisatella cave sys-
tem [29]. Within the system, it is rather common, especially in the branches of the Buso
della Rana cave, while it seems rarer in the Pisatela cave.
3.2. Aquatic Fauna
In Arena Cave, six copepod species were found in a small pool fed by percolating
water and a small drain, all stygobiotic. In Buso della Rana, 19 species of copepods are
known, and 9 of the copepods are stygobiotic.
The Cyclopidae are represented by Speocyclops infernus (Kiefer, 1930), a stygobiotic
species that is widespread over a broader geographical area in the epikarst and vadose
zones in the eastern Alpine region and is present in both the caves under study [14,31,32].
First collected in Buso della Rana by Chappuis in the first half of the last century [33–35],
S. infernus has since been found in many parts of the cave, in both small lakes and pools.
It is also found in puddles inside Arena Cave, although the species aribution is not yet
certain and is therefore currently reported as S. cf. infernus [3].
Among the Harpacticoida, those of the family Canthocamptidae are the most abun-
dant stygobiotic copepods. The genus Elaphoidella is present in the Arena Cave and Rana-
Pisatela system. Elaphoidella is widespread in almost all groundwater habitats in Italy, in
both karstic and porous aquifers, as well as the hyporheic zones of rivers, springs, the
epikarst, and the saturated karst. Elaphoidella phreatica (Chappuis, 1925) is widely
Figure 11.
A specimen of Neobathyscia fabianii from the Rana-Pisatela cave system (Photo: L. Latella).
3.2. Aquatic Fauna
In Arena Cave, six copepod species were found in a small pool fed by percolating
water and a small drain, all stygobiotic. In Buso della Rana, 19 species of copepods are
known, and 9 of the copepods are stygobiotic.
The Cyclopidae are represented by Speocyclops infernus (Kiefer, 1930), a stygobiotic
species that is widespread over a broader geographical area in the epikarst and vadose
zones in the eastern Alpine region and is present in both the caves under study [
14
,
31
,
32
].
First collected in Buso della Rana by Chappuis in the first half of the last century [
33
–
35
], S.
infernus has since been found in many parts of the cave, in both small lakes and pools. It is
also found in puddles inside Arena Cave, although the species attribution is not yet certain
and is therefore currently reported as S. cf. infernus [3].
Among the Harpacticoida, those of the family Canthocamptidae are the most abundant
stygobiotic copepods. The genus Elaphoidella is present in the Arena Cave and Rana-Pisatela
system. Elaphoidella is widespread in almost all groundwater habitats in Italy, in both karstic
and porous aquifers, as well as the hyporheic zones of rivers, springs, the epikarst, and the
saturated karst. Elaphoidella phreatica (Chappuis, 1925) is widely distributed in Italy and
across Europe [
36
]. Elaphoidella ruffoi Chappuis, 1953 is endemic to the Buso della Rana and
is rather rare. It was first found during research in 1952 and was not found in the epikarst.
Only one species of this genus was found in Arena Cave: Elaphoidella sp. A1 [
37
]. An
Elaphoidella, different from the others, was also found in the Buso della Rana, but the scarce
material made it impossible to identify at the species level (Bruno et al. 2018). Ceuthonectes
serbicus Chappuis, 1924 was detected in recent research on epikarst fauna in the Buso della
Rana cave [13].
The genus Elaphoidella is closely related to the genus Lessinocamptus. Known until a
few years ago only from the vadose zone of the Lessinian caves, Lessinocamptus is now also

Diversity 2024,16, 25 14 of 17
known from a site in Northern Slovenia. In fact, Lessinocamptus pivai Stoch 1997, which was
considered endemic to the Buso della Rana, was also found in the Lipnik spring complex
in the Julian Alps (NW Slovenia) [
38
]. Lessinocamptus caoduroi Stoch, 1997, present in the
pool in the Arena Cave, was found only in the percolating waters of the vadose zone of
caves with an elevation of more than 1000 m a.s.l. in the Lessini Mountains. Lessinocamptus
insoletus (Chappuis, 1928) was collected by Chappuis for the first time in the hypogean
brook inside Buso della Rana [
34
]; however, further intensive sampling in the brook did
not yield any specimen of L. insoletus and is therefore probable that the vadose zone is the
main habitat of the species, from which it can be transported into the brook by percolating
water [
39
]. The Ameiridae are represented by Nitocrella psammophila Chappuis, 1955 and
is a stygobiotic species endemic to Italy. It is a widely distributed harpacticoid in the
interstitial zone of subterranean streams in caves and the hyporheic in the Po Valley and
has been reported in Apennine wells in Central Italy and caves in southern Italy. It is
commonly found in the two caves under study. The Parastenocarididae are represented
by Parastenocaris ranae Stoch, 2000, which was collected in the Buso della Rana in the large
residual pools of the subterranean brook in a dry period [40].
Aquatic isopods have only been found in the Buso della Rana, where Monolistra
(Typhlosphaeroma)bericum bericum (Fabiani, 1901) is present. It is a stygobiotic isopod
endemic to the Lessini Mountains and Berici Hills (Vicenza Province).
Bathynellacea were found in the Arena Cave. Not yet identified at a species level,
Bathynella sp. from Arena Cave was collected in a pond fed by a small water flow.
Amphipoda are present with two stygobiotic species: Niphargus similis G. Karaman &
Ruffo, 1989 (Figure 12) in Arena Cave, and Niphargus costozzae Schellenberg, 1935 in the
Buso della Rana cave [3].
Diversity 2024, 16, x FOR PEER REVIEW 16 of 19
Figure 12. Niphargus similis from Arena Cave (Photo: L. Latella).
4. Discussion
The obligate subterranean fauna of Arena Cave and the Rana-Pisatella cave system
is exceptionally rich. It comprises 35 troglobionts and stygobionts, representing 74% of
the obligate subterranean fauna of the whole caves in the Lessini Mountains (more than
200 caves surveyed).
Despite its small size, Arena Cave is the richest one, with 15 troglobionts and eight
stygobionts. The Rana-Pisatella system has a higher number of stygobionts (seven tro-
globionts and 11 stygobionts).
The geographical proximity between the two caves (22 km) would lead one to suspect
a high taxonomic similarity in the fauna inhabiting them. These, on the contrary, have a
very different obligate subterranean fauna in terms of terrestrial but especially aquatic
species. Only seven species (five troglobionts and two stygobionts) out of thirty-five are
in common for the two caves—applying the Jaccard similarity index (and expressed as a
percentage similarity), the results in the similarity between the two caves is 21%.
The high richness and differences in faunal composition of the two caves can be ex-
plained by the paleogeographical events that occurred in the study area.
During the last glacial period, the Italian Alps were covered by glaciers, except at the
top of the highest mountains [41]. In contrast to the Alps, Prealpine areas were only par-
tially covered by glaciers [2,42], and glacial tongues occupied only a few deeper valleys
[43]. This favored the colonization of ice-free zones by invertebrates from moist and cold
habitats, like forest lier and soil, alpine grasslands, and talus areas. During interglacial
periods, as glaciers retreated, populations became isolated in the highest parts of the Pre-
alpine mountains or took refuge in cold, moist interiors of caves [5,6]. Surface populations
became extinct or isolated, and there was, therefore, lile or no gene flow between cave
communities, boosting the evolution of the troglobiont [11]. This is known as the climatic
relict hypothesis [44–47]. The effects of Quaternary glaciations also shaped the stygobiotic
species distribution, as the Massif was only marginally covered by ice, and the extensive
networks of fractures of the karstic system represented a refuge for stygobionts, boosting
isolation and speciation [13]. In fact, the vadose zone and the epikarst of the Lessinian
Figure 12. Niphargus similis from Arena Cave (Photo: L. Latella).
4. Discussion
The obligate subterranean fauna of Arena Cave and the Rana-Pisatella cave system
is exceptionally rich. It comprises 35 troglobionts and stygobionts, representing 74% of
the obligate subterranean fauna of the whole caves in the Lessini Mountains (more than
200 caves surveyed).

Diversity 2024,16, 25 15 of 17
Despite its small size, Arena Cave is the richest one, with 15 troglobionts and eight sty-
gobionts. The Rana-Pisatella system has a higher number of stygobionts (seven troglobionts
and 11 stygobionts).
The geographical proximity between the two caves (22 km) would lead one to suspect
a high taxonomic similarity in the fauna inhabiting them. These, on the contrary, have
a very different obligate subterranean fauna in terms of terrestrial but especially aquatic
species. Only seven species (five troglobionts and two stygobionts) out of thirty-five are
in common for the two caves—applying the Jaccard similarity index (and expressed as a
percentage similarity), the results in the similarity between the two caves is 21%.
The high richness and differences in faunal composition of the two caves can be
explained by the paleogeographical events that occurred in the study area.
During the last glacial period, the Italian Alps were covered by glaciers, except at the
top of the highest mountains [
41
]. In contrast to the Alps, Prealpine areas were only partially
covered by glaciers [
2
,
42
], and glacial tongues occupied only a few deeper valleys [
43
]. This
favored the colonization of ice-free zones by invertebrates from moist and cold habitats,
like forest litter and soil, alpine grasslands, and talus areas. During interglacial periods,
as glaciers retreated, populations became isolated in the highest parts of the Prealpine
mountains or took refuge in cold, moist interiors of caves [
5
,
6
]. Surface populations became
extinct or isolated, and there was, therefore, little or no gene flow between cave commu-
nities, boosting the evolution of the troglobiont [
11
]. This is known as the climatic relict
hypothesis [
44
–
47
]. The effects of Quaternary glaciations also shaped the stygobiotic species
distribution, as the Massif was only marginally covered by ice, and the extensive networks
of fractures of the karstic system represented a refuge for stygobionts, boosting isolation
and speciation [
13
]. In fact, the vadose zone and the epikarst of the Lessinian Massif are
known to harbor a high diversity of microcrustaceans, including many endemic species
due to the ancient geological age of the aquifers, high habitat fragmentation, and isolation
of microhabitats, factors of which concurred to promote speciation by vicariance [13,48].
The exceptional subterranean diversity of Arena Cave and the Rana-Pisatella cave
system, so different from each other in shape and development (less than 100 m for Arena
Cave and about 38 km for the Rana-Pisatella system), can be explained only by their
geology, where the two caves developed in different typologies of rocks, namely “contact
caves” [5,6].
Contact karst is considered, in a strict sense, a karst phenomenon, where forms are
influenced by the contact between a karstifiable rock and a non-karstifiable rock. In a
wide sense, the karst phenomena and forms that are influenced by the contact between
two karstifiable rocks differ in some of their characteristics, such as chemical composition,
porosity, and fracture density [5].
These different rock characteristics create a number of different microhabitats that also
influence the life and dispersion of the subterranean animals. Depending on the amount
of water retained, humidity, and other factors that are not yet fully known, specimens of
different species may prefer one microhabitat over another. This explains the rarity of the
findings in Arena Cave of the trechine Lessynodites pivai, a species that most probably do
not frequently inhabit the cave proper but rather lives in the wetter interstices of the Rosso
Ammonitico formations. The same can be said for Italaphaenops dimaioi; ongoing studies by
the lab of the Museum of Verona show that I. dimaioi was sampled almost exclusively in
the caves in contact with Rosso Ammonitico rock in the Lessini Mountains.
The same microhabitat characteristics are probably the reason for the abundance
of copepods and the presence of Bathynella sp., present in waters that flow from the
environments in the small pool inside the cave. It is in similar conditions that, collecting
water from the epikarst, we found Ceuthonectes serbicus for the first time in Buso della
Rana [13].
The smaller cave is, therefore, the richest in diversity. As is often said, size does not
always matter.

Diversity 2024,16, 25 16 of 17
Funding: This research received no external funding.
Institutional Review Board Statement: Not applicable.
Data Availability Statement: Data are contained within the article.
Acknowledgments:
I would like to thank Dave Culver (Washington, DC, USA), Louis Deharveng
(Paris, France), Valeria Lencioni (Trento, Italy), and Francesco Sauro (Verona, Italy) for their helpful
suggestions in the elaboration of the text; Ivan Petri (Trento, Italy), Barbara Valle (Milan, Italy), Fabio
Stoch (Brussels, Belgium) for their taxonomic identification or suggestions; Arianna Spada (Verona,
Italy) for the graphic elaboration of some figures; Sandro Sedran (Dolo, Italy) and Filippo Rossetto
(Verona, Italy) for allowing the use of photographs of the Buso della Rana and Ischyropsalis strandii.
I am also grateful to all the speleologists who shared the cave surveys with me, especially Giorgio
Annichini (Verona, Italy), Tarcisio Battagini (Verona, Italy), and Andrea Pasotto (Verona, Italy)—and
a special thanks to Dave Culver for revising the English text and a thanks to the four anonymous
reviewers.
Conflicts of Interest: The author declares no conflict of interest.
References
1. Marazzi, S. Atlante Orografico Delle Alpi—SOIUSA; Priuli and Verlucca: Scarmagno, Italy, 2005.
2.
Sauro, U. Il Paesaggio Degli Alti Lessini: Studio Geomorfologico; Museo Civico di Storia Naturale di Verona: Verona, Italy, 1973;
Volume 6, 161p.
3.
Caoduro, G.; Ruffo, S. La Grotta dell’Arena, un biotopo di eccezionale interesse negli alti Lessini. In La Lessinia Ieri Oggi Domani:
Quaderno Culturale; Editrice La Grafica: Lavagno, Italy, 1998; pp. 39–44.
4.
Culver, D.C.; Deharveng, L.; Bedos, A.; Lewis, J.J.; Madden, M.; Reddell, J.R.; Sket, B.; Trontelj, P.; White, D. The mid-latitude
biodiversity ridge in terrestrial cave fauna. Ecography 2006,29, 120–128. [CrossRef]
5. Sauro, U. Aspects of contact karst in the Venetian Fore-Alps. Acta Carsologica 2001,30, 89–102.
6.
Latella, L.; Sauro, U. Aspects of the Evolution of an Important Geo-Ecosystem in the Lessinian Mountain (Venetian Prealps, Italy).
Acta Carsologica 2007,36, 69–75. [CrossRef]
7.
Pasa, A. Carsismo ed idrografia carsica del Gruppo del Monte Baldo e dei Lessini Veronesi. CNR Cent. Studi Geogr. Fis. Ric. Sulla
Morfol. Idrogr. Carsismo 1954,5, 1–150.
8.
Sauro, U. Aspetti dell’evoluzione carsica legata a particolari condizioni litologiche e tettoniche negli Alti Lessini. Boll. Soc. Geol.
Ital. 1974,93, 945–969.
9.
Tisato, N.; Sauro, F.; Bernasconi, S.M.; Bruijn, R.H.; De Waele, J. Hypogenic contribution to speleogenesis in a predominant
epigenic karst system: A case study from the Venetian Alps, Italy. Geomorphology 2012,151, 156–163. [CrossRef]
10. Sistema carsico Rana-Pisatella. Available online: www.busodellarana.it (accessed on 1 August 2023).
11.
Latella, L.; Verdari, N.; Gobbi, M. Distribution of terrestrial cave-dwelling arthropods in two adjacent Prealpine Italian areas with
different glacial histories. Zool. Stud. 2012,51, 1113–1121.
12.
Avesani, D.; Latella, L. Spatio-temporal distribution of the genus Chionea (Diptera, Limoniidae) in the Buso del Valon ice cave
and other caves in the Lessini Mountains (Northern Italy). Boll. Mus. Civ. St. Nat. Verona 2016,40, 11–16.
13.
Bruno, M.C.; Cottarelli, V.; Grasso, R.; Latella, L.; Zaupa, S.; Spena, M.T. Epikarst crustaceans from some Italian caves: Endemisms
and spatial scales. Biogeographia 2018,33, 1–18.
14. Pipan, T. Epikarst—A Promising Habitat; ZRC Publishing: Ljubljana, Slovenia, 2005; pp. 1–101.
15.
Pipan, T.; Christman, M.C.; Culver, D.C. Dynamics of epikarst communities: Microgeographic pattern and environmental
determinants of epikarst copepods in Organ Cave, West Virginia. Am. Midl. Nat. 2006,156, 75–87. [CrossRef]
16.
Allegranzi, A.; Bartolomei, G.; Broglio, A.; Pasa, A.; Rigobello, A.; Ruffo, S. Il Buso della Rana (40 V-VI). Ras. Speleol. Ital.
1960
,12,
99–163.
17.
Caoduro, G.; Osella, G.; Ruffo, S. La fauna cavernicola della regione veronese. Memorie del Museo Civico di Storia Naturale di Verona.
Sez. Biol. 1994,6, 1–144.
18.
Latella, L.; Sauro, U. Note di Storia Naturale del sottosuolo dei Monti Lessini e del suo popolamento. Quaderno Culturale Lessinia
Ieri Oggi Domani 2006,34, 57–64.
19.
Latella, L. Il contributo del Museo Civico di Storia Naturale di Verona allo sviluppo della biospeleologia. Studi Trentini Di Sci. Nat.
Acta Biol. 2004,81, 15–22.
20. Ruffo, S. Le attuali conoscenze sulla fauna cavernicola della Regione Pugliese. Mem. Biogeogr. Adriat. 1957,3, 1–143.
21. Sket, B. Can we agree on an ecological classification of subterranean animals? J. Nat. Hist. 2008,42, 1549–1563. [CrossRef]
22.
Pezzoli, E. Il genere Zospeum Bourguignat, 1856 in Italia (Gastropoda Polmonata Basommatophora). Censimento delle stazioni
ad oggi segnalate. Nat. Brescia. 1992,27, 123–169.
23.
Petri, I.; Ballarin, F.; Latella, L. Seasonal abundance and spatio-temporal distribution of the troglophylic harvestman Ischyropsalis
ravasinii (Arachnida, Opiliones, Ischyropsalididae) in the Buso del Valon ice cave, Eastern Italian Prealps. Subterr. Biol.
2022
,42,
151–164. [CrossRef]

Diversity 2024,16, 25 17 of 17
24.
Gardini, G. The species of the Chthonius heterodactylus group (Arachnida, Pseudoscorpiones, Chthoniidae) from the eastern
Alps and the Carpathians. Zootaxa 2014,3887, 101–137. [CrossRef]
25.
Casale, A.; Vigna Taglianti, A. Note su Italaphaenops dimaioi Ghidini (Coleoptera, Carabidae). Boll. del Mus. Civ. Stor. Nat. di
Verona 1976,2, 293–314.
26. Sciaky, R.; Vigna Taglianti, A. The genus Lessinodytes (Coleoptera, Carabidae, Trechinae). Mém. Biospéol. 1990,17, 169–173.
27.
Monguzzi, R. Nuovi Dati Per La Conoscenza Del Genere Lessinodytes Vigna Taglianti, 1982 (Coleoptera Carabidae Trechinae).
Nat. Brescia. 1993,29, 179–183.
28.
Faille, A.; Casale, A.; Balke, M.; Ribera, I. A molecular phylogeny of Alpine subterranean Trechini (Coleoptera: Carabidae). BMC
Evol. Biol. 2013,13, 248. [CrossRef] [PubMed]
29.
Latella, L. The Subterranean Cholevinae of Italy. In Cave Biodiversity: Speciation and Diversity of Subterranean Fauna; Johns Hopkins
University Press: Baltimore, MD, USA, 2022; p. 164.
30.
Giachino, P.M.; Vailati, D. I Cholevidae delle Alpi e Prealpi italiane: Inventario, analisi faunistica e origine del popolamento del
settore compreso fra i corsi dei fiumi Ticino e Tagliamento (Coleoptera). Biogeogr. J. Integr. Biogeogr.
2005
,26, 229–378. [CrossRef]
31.
Deharveng, L.; Stoch, F.; Gibert, J.; Bedos, A.; Galassi, D.; Zagmajster, M.; Brancelj, A.; Camacho, A.; Fiers, F.; Martin, P.; et al.
Groundwater biodiversity in Europe. Freshw. Biol. 2009,54, 709–726. [CrossRef]
32.
Papi, F.; Pipan, T. Ecological studies of an epikarst community in Snežna jama na planini Arto an ice cave in north central Slovenia.
Acta Carsologica 2011,40, 3. [CrossRef]
33.
Chappuis, P.A. Nouveaux Copépodes cavernicoles des genres Cyclops et Canthocamptus (Note préliminaire). Bul. Soc. De Stiint.
Din Cluj 1923,1, 584–590.
34. Chappuis, P.A. Nouveaux Copépodes cavernicoles. Bul. Soc. De Stiint. Din Cluj 1928,2, 20–34.
35.
Chappuis, P.A. Biospeologica LIX. Copepodes (premiere serie), avec l’énumeration de tous les copepodes cavernicoles connus en
1931. Arch. De Zool. Expérimentale Et Générale 1933,76, 1–56.
36.
Galassi, D.M.P. Groundwater copepods: Diversity patterns over ecological and evolutionary scales. Hydrobiologia
2001
,453,
227–253. [CrossRef]
37.
Ruffo, S.; Stoch, F. Checklist and Distribution of the Italian Fauna. Memorie del Museo Civico di Storia Naturale di Verona, II Serie, Sezione
Scienze Della Vita 17; Ministero dell’Ambiente e della Tutela del Territorio e del Mare: Roma, Italy, 2006; with CD-ROM.
38.
Mori, N.; Brancelj, A. Differences in aquatic microcrustacean assemblages between temporary and perennial springs of an alpine
karstic aquifer. Int. J. Speleol. 2013,42, 3–9. [CrossRef]
39.
Stoch, F. A new genus and two new species of Canthocamptidae (Copepoda, Harpacticoida) from caves in northern Italy.
Hydrobiologia 1997,350, 49–61. [CrossRef]
40.
Stoch, F. New and little known Parastenocaris (Copepoda, Harpacticoida, Parastenocarididae) from cave waters in Northeastern
Italy. Boll. Del Mus. Civ. Di Stor. Nat. Di Verona 2000,24, 195–206.
41. Hughes, P.D. Quaternary glacial history of the Mediterranean mountains. Progr. Phys. Geogr. 2006,30, 334–364. [CrossRef]
42.
Bassetti, M.; Borsato, A. Evoluzione geomorfologica e vegetazionale della bassa valle dell’Adige dall’ultimo massimo glaciale:
Sintesi delle conoscenze e riferimenti ad aree limitrofe. Studi Trentini Sci. Nat. Acta Geol. 2007,82, 31–42.
43.
Sommaruga, M.; Zorzin, R. Evidences of Morphologies and Glacial Deposits into High Valle del Chiampo (Northern Italy, Vicenza
Province): First Results. Boll. Del Mus. Civ. Di Stor. Nat. Di Verona Geol. Paleontol. Preist. 2018,42, 43–71.
44.
Sbordoni, V. Advances in speciation of cave animals. In Mechanisms of Speciation; Barigozzi, C., Ed.; Liss: New York, NY, USA,
1982; pp. 219–224.
45.
Humphreys, W.F. Relict fauna and their derivation. In Ecosystems of the World; Wilkens, H., Culver, D.C., Humphreys, W.F., Eds.;
Subterranean Ecosystems; Elsevier: Amsterdam, The Netherlands, 2000; Volume 30, pp. 417–432.
46.
Assmann, T.; Casale, A.; Drees, C.; Habel, J.C.; Matern, A.; Schuldt, A. Review: The dark side of relict species biology: Cave
animals as ancient lineages. In Relict Species: Phylogeography and Conservation Biology; Assmann, T., Habel, J.C., Eds.; Springer:
Berlin/Heidelberg, Germany, 2010; pp. 91–103.
47. Hampe, A.; Jump, A.S. Climate relicts: Past, present, future. Annu. Rev. Ecol. Evol. Syst. 2011,42, 313–333. [CrossRef]
48.
Galassi, D.M.; Stoch, F.; Fiasca, B.; Di Lorenzo, T.; Gattone, E. Groundwater biodiversity patterns in the Lessinian Massif of
northern Italy. Freshw. Biol. 2009,54, 830–847. [CrossRef]
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