Subterranean Habitats - EnviMod

Transcription

Faculty of the Environment
Jan Evangelista Purkyně University in Ústí nad Labem
Subterranean Habitats
M. Holec, R. Pokorný
Ústí nad Labem, 2012
Faculty of the Environment
Jan Evangelista Purkyně University in Ústí nad Labem
Název:
Autoři:
Mgr.
Michal Holec, Ph.D.
Ing. Richard Pokorný, DiS.
Vědecký redaktor: Prof. Ing. Jaroslav Boháč, CSc.
Recenzenti:
©
RNDr.
Miloslav Zacharda, CSc,
RNDr. Karel Tajovský, CSc.
Fakulta životního prostředí Univerzita J. E. Purkyně 2012
ISBN: 978-80-7414-416-5 (brož.)
ISBN: 978-80-7414-897-2 (online: pdf)
Tato publikace vznikla v rámci projektu OPVK EnviMod - Modernizace výuky technických
a přírodovědných oborů na UJEP se zaměřením na problematiku ochrany životního prostředí.
Reg. č.: CZ.1.07/2.2.00/28.0205
Obsah
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Emergence of Selected Underground Spaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Pseudokarst Caves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Definition of Types of Natural Underground Spaces . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Cave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Cavelet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Chasm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Rock overhang . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Superficial subterranean habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Stony accumulations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Most Important Cave Regions in the Czech Republic . . . . . . . . . . . . . . . . . . . . . . . . . 16
Bohemian Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Moravian Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Branná Belt Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Hranice Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Javoříčko Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Mladeč Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Pálava Hills Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Chýnov Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Karst of the Kamenice and Železný Brod catchment basin . . . . . . . . . . . . . . . . . . . . . . . . . 21
Železné hory Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Tišnov Karst . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Main pseudokarst areas in the Czech Republic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Subterranean habitats from the viewpoint of protection and administration . . . . . . 23
Cave utilization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Cave management and care . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Cave research and evidence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Environmental Conditions and Their Measuring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Subterranean Microclimates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Heat (temperature) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Humidity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Organic matter in subterranean habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Atmosphere in subterranean habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Methods of animals’ collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Direct searching for animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Pitfall trapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Core sampling of soil, sediments or litter and fauna extraction . . . . . . . . . . . . . . . . . . . . . . 49
Ecological and Evolutionary Classifications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
The origin and emergence of subterranean terrestrial fauna . . . . . . . . . . . . . . . . . . . . 55
Survey of selected groups of organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Nematodes (Nematoda) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Annelids (Annelida) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Molluscs (Mollusca) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Spiders and other related groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Crustacea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Millipedes and centipedes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Hexapods (Hexapoda) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Bats (Chiroptera) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Other vertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Future of underground research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Acknowledgements
With regard to the fact that this material is mainly based on the book of POKORNÝ and HOLEC
(2009), we would like to thank here all who already participated on the creation of the cited book.
Here we would like to thank especially to RNDr. Vlado Papáč for providing photos of springtails. We
also thank to language agency Skřivánek s.r.o. for the English translantion of the main parts of the
text and to Ing. Lenka Zoubková for the final language improvement. We also thank to both opponents
for their valuable comments on manuscript. We would also like to thank our workplace, which allows
us to explore this issue, especially within teaching activities. We also thanks to a donation from the
Ministry of Environment R&D (MŽP VaV) No. SP/2d3/4/07 for the purpose of research in the years
2007–2009, and later approved by University Development Fund (UDF, FRVŠ in Czech) project No.
305/2010/F4/a – grant of the Ministry of Education, Youth and Sports (MEYS, MŠMT in Czech) and
project of the Internal Foundation of UJEP – ”Paleontological research of the pseudokarst in north
Bohemia”.
Introduction
The authors have been paying considerable attention to the caves in the North Bohemian neovolcanites for several years. The issue includes not only mapping caves in the field, but also the acquisition
of other more detailed characteristics (part of data was published in POKORNÝ and HOLEC 2009).
The colonization of caves by invertebrates was studied, althought data have not been still published.
In the region, however, data on the occurrence of vertebrates, esp. bats, are available as well, e.g.
BENDA and CHVÁTAL (2011).
This was the background for the idea to create a compulsory school course, especially for foreign
students of Jan Evangelista Purkyně University – Faculty of the Environment in Ústí nad Labem,
which would call attention to the general scientific value of less known subterrannean environment.
In this text we generalised our former unpublished text for students oriented mainly on regional localities. The presented material brings the basic general information about the terrestrial caves and
other natural terrestrial subterranean habitats. Water habitats, including interstitial habitats, were not
mentioned because this text will serve as theoretical background for terrain excursion with students
of Faculty of the Environment, Jan Evangelista Purkyně University in Ústí nad Labem. Our field
trips with students will be oriented to the localities with terrestrial caves only and description of water habitats and water biota would stay without practical usage. For the same reason we prefer, for
example, description of epigenetic pseudokarst. Description of other cave types, rarely surrounded
our workplace or dangerous for excursions with students, was neglected.
6
Emergence of Selected Underground Spaces
Processes leading to the emergence of spaces below the surface of the earth can be divided into two
basic groups – chemical and physical (= mechanical). By chemical processes we mean those which
change the essence of the chemical substance of the primary rock. Generally these processes involve
dissolution or oxidation. In the case of dissolution, the rock is dissolved by the action of a suitable solvent, such as water or a weak naturally occurring acid. This creates underground cavities of varying
size and layout. As the dissolution equation generally works both ways, the reverse can happen, with
dissolved substances accreting to create typical cave formations, known as speleothems.
The most common chemical process leading to the creation of underground spaces is karstification,
the result of which is karst, a characteristic type of landscape typified by the presence of surface and
underground phenomena. The condition for this is the reaction of rainwater and other CO2-enriched
surface waters with the rock (generally carbonate in composition – limestone, marble, dolomites) on
the surface of the earth. Water travels down deep through fissures, where it dissolves the rock to form
chimneys, passages and caves. In contrast to established opinion, karst phenomena do not only occur
in limestone, but may be present in all rocks that have been at least partially dissolved. In the case of
carbonates, the basic matter is dissolved according to the following schema (equations I, II) (PŘIBYL
et al. 1992).
(I) (II)
CO2 + H2O ↔ H+ + (HCO3) – ↔ H+ + CO32–
(HCO3)- + CaCO3 ↔ Ca2+ + 2(HCO3)2-
The work of CIGNA (1978) states that when describing the creation of the natural underground, we
must base our arguments on the so-called three-component equilibrium:
- CaCO3 (calcium carbonate, calcite) – the basic building block of calcareous rocks; in the
broader sense any dissolved parent rock is considered to be a reaction component
- water – a medium with mechanical and chemical action
- CO2 (carbon dioxide) – in the form of ions (HCO3)- in a water solution acts as a kind of solvent
CIGNA divides up the different types of formation depending on the number of components involved in the creation of the cave.
7
Non-karst
Karst
Tab. I. Division of karst and non-karst categories in accordance with POKORNÝ and POKORNÁ
(2007), MAVLYUDOV (2006), FIELD (2002) and CIGNA (1978).
Group
Number
of components
Subgroup
Examples
hyperkarst
>3
---
hydrothermal, acidic springs
karst
3
holokarst
pure limestone
merokarst
dolomites, marl limestone
parakarst
2
tachykarst
evaporites – gypsum, halites
bradykarst
quartzites, tufas, vulcanites?
hypokarst
1
glaciokarst
glaciers
vulcanokarst
lava tunnels
syngenetic
gas inclusions in vulcanites
epigenetic
tectonic, erosion caves (sandstone,
etc.)
pseudokarst
0
The development of the karst in Bohemia can be divided up into several phases separated by orogenic processes or sea flooding. The oldest provable period of karstification is linked to the sedimentation of old Palaeozoic limestone and is broken up into a series of secondary phases. The next
karstification period is linked to the final phase of the Variscan orogenic process and lasted until the
Upper Cretaceous floods during the Cenomanian. This period resulted in a number of fossil karsts
below Upper Cretaceous deposits in the Bohemian and Moravian Karst. The most recent karstification period has lasted since the end of the Upper Cretaceous floods to the present day. It is broken up
into a series of secondary phases linked to the Alpine buckling in the forefield of the Bohemian Massif
and separated by either brief periods of sea flooding (the eastern edge of the Bohemian Massif) or by
marked phases involving the accumulation of continental sediments (neogenic coal basins, tertiary
and quaternary terraces). The Tertiary period saw the creation of the most important underground
cave systems in the Bohemian, Moravian and North Moravian Karst, as well as in other isolated karst
islands. These karstification phases produced typologically different forms of karst, particularly as the
result of tectonic, climatic and hydrological factors. Also, from a regional viewpoint, the development
of karst areas was differentiated.
Within the Czech Republic the most widespread karst subgroups are holokarst and merokarst. If a
karst region develops in chemically pure limestone and it is possible to identify all the typical surface
and subsurface karst forms of relief, it is referred to as holokarst. The closest thing to a holokarst
region in this country is the Moravian and parts of the Bohemian Karst, while in Europe they can be
found in the Balkans, in Italy, and in France. Merokarst occurs in marl limestone and dolomites with
a lower percentage of calcite, where the karstification process is incomplete. Examples of merokarst
include Chýnov Cave and the Zbrašov Aragonite Caves among others (HROMAS et al. 2009).
If natural underground spaces are the result of nothing more than mechanical processes, they are
classed as non-karst, or pseudokarst. These may occur in any type of rock, i.e. including those which
could have been subject to chemical dissolution (although in this case it played no part in the genesis
of the underground spaces).
PŘIBYL et al. (1992) distinguish pseudokarst as being either syngenetic or epigenetic. The first
category represents underground spaces created around the same time as the surrounding rock. One
8
ideal example is the “bubble caves”, formed at the site of gas inclusions sealed within the surrounding igneous rock. Similar structures reaching a length and depth of approximately 25 m have been
described by BELLA (1998) at Štiavnické vrchy and Ragáč in Slovakia. PŘIBYL et al. (1992) also
include features created during the formation of metamorphic rock in this category of syngenetic
pseudokarst. A unique cave of this type was uncovered at the end of the 19th century in a quarry to
the west of Džbán. The cave was 6 m long and up to 2 m high and wide (STÁRKA 1982). Syngenetic
pseudokarst also includes certain “tree mould caves”, i.e. cave cavities formed when parts of plants –
generally the roots, branches and crowns of trees – become sealed in the surrounding rock. BELLA
and GAÁL (2007) describe these caves as being formed from volcanic rock, volcanoclastics and also
young travertines. Also considered syngenetic in the sense of the above work are caves which were
formed after tree-trunks burned in hot lava, soon after it poured out. Caves like these exist in Japan,
for example, but none have been recorded in the Czech Republic.
Epigenetic pseudokarst originate as the result of processes occurring after the solidification of the
parent rock. BELLA and GAÁL (2007) differentiate between tree mould caves which are the result
of the mechanical disintegration or biogenic decomposition of wood after lava has solidified, and
caves formed through a combination of both processes. Caves which fall into this category in this
country are probably Jeskyňky skřítků (the Dwarf Cavelets) in the Doupov Mountains, the somewhat
ambiguous origin of which was in the past explained as the selective ventilation of tufa breccia and
the subsequent opening up of gas cavities sealed in the rock (KRÁL 1973). Nowadays a more likely
theory is that they were formed when fallen tertiary trees were covered by mud-flow following the
eruption of a nearby volcano (MLEJNEK et al. 2002). The biogenic origin of the caves in the Doupov
Mountains is also confirmed by BELLA and GAÁL (2007), who characterise these cavities as being
demonstrably epigenetic. HRADECKÝ (1997) adds that the present-day cavities are the result of suffosis and subterranean erosion in the original cavities.
However, typical examples of epigenetic pseudokarst are objects generally represented by tectonic,
gravitational and erosion cavities in a wide variety of different types of rocks, caused by mechanical
processes long after the formation of the rock massif (fore more details see in next chapter).
There is an abundance of these formations in the Czech Republic. They are common in the strata
formations of blocks of sandstone in the Bohemian Cretaceous Basin; exceptionally extensive systems have been described made from the sandstone and marlstones of the flysch zone of the Western
Carpathians (HROMAS and BÍLKOVÁ 1998a, b). More recent tertiary volcanic activity in the České
středohoří Mountains and the subsequent modulation of the georelief due to slope movements and
frost weathering during the ice ages has led to the formation of several caves of varying lengths, as
well as similar sites which will be described in detail in the following sections.
9
Pseudokarst Caves
Epigenetic pseudokarst caves can be distinguished from the point of view of their formation and
divided into several groups. They are caves of crack, cleft, talus, erosion and combined types. The
specific type is a tree mould cave, which will not be discussed here. The terminology is based on studies of VÍTEK (1981, 1979) supplemented with the findings of POKORNÝ and POKORNÁ (2007) and
ESZTERHÁS (2007). Although the studies of Vítek and Eszterhás are devoted mainly to sandstone,
their classification is so general that it can be applied, in addition to sediments, to both igneous and
metamorphic rocks. This chapter describes mainly those genetic cave types representatives of which
can be found in the neovolcanites of the Ústí nad Labem region.
The first two types, crack and cleft caves, are genetically connected to the movement of rock blocks
and according to ESZTERHÁS (2007) therefore belong to one category.
Crack caves represent vertical narrow spaces where the height significantly prevails over the width and at the same time they pass
the required criterion of necessary length. Characteristic signs are
a trapezoidal cross–section and a ceiling formed by rock material
connected with upper layers of rock. Their formation is always predisposed by the existence of a system of more or less vertical concurrently converging cracks – zones of substantial tectonic disruption.
The cave is identified as the crack type on the condition that the
rock mass affected with the fan of tectonic failures (cracks) remains
stable as a whole. If this condition is not met, the cave is categorized
as the cleft type. In these cases block movements of rock blocks up
or down the slope occur and underground spaces are gradually widened and opened as a result. The rock can be cracked in the same
way as in the previous case, however generally the steep relief of the
landscape will start the process of loosening individual blocks along
tectonic predispositions and their sliding to lower levels. The space
formed is usually narrow and high in cross profile – according to the
method of formation – in the classical form either roof-shaped, in the
shape of letter “A”; or wedge-shaped, in the shape of letter “V”. In
the second case mentioned the ceiling is formed by accumulated and
Fig. 1. A crack cave. Illustratiembedded rock
on from the authors’ archive.
blocks.
The
classic example of a crack type is Bílanina cave. Examples
of cleft caves in the Ústí nad Labem region are
the Komora cave in the Lužické Mountains or
Loupežnická Jeskyně (the Robber‘s Cave) near
the town of Ústí nad Labem.
The caves whose formation is connected only
with cambering along the planes of division
Fig. 2. A cleft cave of the “A” type. Illustration from
the authors’ archive.
of rockslides are according to PANOŠ (2001)
called rock-slide caves.
The North Bohemian region is quite rich in geomorphological objects during the formation of
which frost weathering played a role in the Quaternary period (RAŠKA and CAJZ 2008). Various
rock and boulder streams and also rock cliffs, frost cliffs and non-karst caves can be found here. The
10
Pseudokarst Caves
origin of these “frost caves”, whose classification is difficult, is connected with the intrusion
of ice and the opening of cracks. Consequently,
the shift of rock bodies in these cases is not influenced by a steep slope. Considering the fact
that their genesis is connected with at least the
partial movement of rock bodies, these caves
are classified in one category of the cleft type
although for some of them the classification of
frost caves could be used as well, as PANOŠ
(2001) states in his survey. Such cryoplanation
cleft caves can be found in neovolcanites of the
Ústí region on the hill Buková hora (e.g. the Ledová Cave) and the Velká Jeskyně Skřítků Cave.
Fig. 3. A cleft cave of “V” type. Illustration from the
authors’ archive.
Other types are caves of the block-field type or talus type. Although these structures are not considered by some authors (e.g. OZORAY 1962) to be caves in a narrower sense of the word and the
terms like pseudo-caves have been suggested, it is beyond any doubt that they are caves when all the
criteria such as length, if it is passable, etc. are met. The formation of talus caves is, in the same way
as cryoplanation clefts, often connected with frost weathering. The longest Czech talus caves are
several hundred meters long and it is typical that they are not passable for an extensive section. The
category of subsurface talus in the region
is represented by, for example, the cavelet
near Dobrná and also by a smaller cave on
the hill Buková hora Hill and around the
Jeskyně Skřítků Cave.
Other types are caves of the erosion type,
which includes caves formed by water and
wind erosion according to ESZTERHÁS
(2007); caves during the formation of which
internal stress in rocks or temperature variations played a role and also caves formed
by chemical erosion (dissolution, hydration, hydrolysis, oxidation), including bioerosion. VÍTEK (1979, 1981) in his studies
Fig. 4. A talus cave. Illustration from the authors’ archive.
comments on bedding type caves, which
are a large subset of erosion caves including
structures formed by weathering and the removal of less resistant bedding planes or the crumbling
away of layer banks. The formed spaces are low, relatively wide and of various lengths. According to
VÍTEK (1981) bedding type caves occur only in sedimentary rocks, whereas similar objects in eruptive rocks and metamorphites according to him are crack caves based on horizontal cracks or planes
of division. However, POKORNÝ and POKORNÁ (2007) identified within the České středohoří
Mountains several caves and small caves corresponding to bedding type caves, e.g. in sandstone at
Český ráj Protected Landscape Area. For more details see POKORNÝ and POKORNÁ (2007).
11
Definition of Types of Natural Underground Spaces
The word used by the general public as a synonym for underground structures formed as the result
of natural processes is cave. This type of underground cavity, however, is just one of the many possibilities, while a brief list of the most common geomorphologic formations is given below. The criteria
used to differentiate between the different types used in this section are how they were formed, their
inner habitus and their relationship to the geological situation of the region in question.
Cave
It is undoubtedly the cave that attracts the most interest from the specialist and lay public, despite
the fact that as a technical term this category is not precisely defined in the professional geological
and speleological literature and there are a number of parallel, although not completely exhaustive,
definitions.
The term cave is defined in various ways. Generally, it refers mostly to a naturally formed underground cavity accessible for people. It is obvious that from the point of view of understanding what
happens underground it is not very important whether or how the man fits into the cavity or does not.
Using a higher amount of definitions, we tried to set up the definition which corresponds with the
orientation of our work to natural underground structures in north-west Bohemia.
“Caves are cavities in the geological environment situated under the surrounding terrain level, parallel with it, or exposed at any height of the corresponding rock body. The geological environment is
a part of the lithosphere formed by bodies of igneous, sedimentary and metamorphicrocks in various
stages of strengthening. The cavity represents the body formed by the forces of nature (endogenous,
exogenous or their combination) without human interference.
The cave must be sufficiently long in the major part of its length so that an average adult can walk
or crawl through it. This fact does not eliminate the presence of a component of the cave or of the cave
system that cannot be crawled through – components in the shape of narrow to dying-away endings,
side branches, slots, etc. of any length.
To fulfil the definition of a cave, the total passable length of the structure should be long enough to
be able to significantly distinguish the inner environment from the surrounding environment, especially on the basis of the presence of a different microclimate, humidity, the presence of an aphotic
zone, cave sediments and troglobiont or at least troglophile organisms. In practice, this limit was empirically set to the distance of 5m from the outer environment.
Caves are also natural underground spaces corresponding to the dimensional criterion, but without
a passable inlet orifice, or without any connection with the outer environment, if their presence is reliably confirmed using survey methods.”
Depending on the length of the cave, BÖGLI (1978) uses the following classifications: small cave
(up to 50 m in length), medium cave (50 – 500 m), long cave (500 – 5000 m) and giant cave (length
exceeding 5000 m).
According to HROMAS et al. (2009), these cave categories could also include rock perforations
– i.e. cavities which pass through the body of rock, with a separate entrance at both ends. If the perforation is large and reaches to the foot of the rock, it is referred to as a rock gate. Rock windows are
smaller and do not reach to the foot of the rock, while rock tunnels are longer than they are wide. In
the case of rock bridges, the ceiling merges with the surrounding relief.
12
Cavelet
These are horizontal or sub-horizontal underground cavities with an entrance that can be crawled
through, formed by the same processes as a cave. However, they differ in terms of their length, which
is no more than 2 – 5 m (sites exceeding 5 m in length are termed caves; objects less than two metres
deep are referred to as cave niches.)
Chasm
If the incline of the bottom of a sub-horizontal cavity towards the interior falls below an angle of 3060°, this is classed as a sloping cave (or cavelet, depending on the length of the cavity). If the incline
of the bottom is 60-80°, such structures are referred to as vertical caves.
If an underground space is significantly more vertical, the incline exceeds 80° and the depth is more
than 5 m, it is known as a chasm; shallower spaces 2 – 5 m deep are referred to as little chasms. The
morphology needs to be taken into account in this category, too. HROMAS et al. (2009) classifies
vertical cavities as follows: those with a uniform width are wells, whereas cavities whose diameter
varies are shafts. The light-hole chasms are open vertical cavities lit by daylight (e.g. the Macocha and
Hranice Chasms), vertical or sub-vertical narrow passages known as avens (e.g. the Arnoldka Chasm
in the Bohemian Karst).
Rock overhang
This is a natural rock outcrop formed by a variety of weathering processes in less resistant rock
positions. Different types of overhangs include contoured overhangs, which form at any height of
a rock wall through simple erosion; footers, formed by weathering at the foot of rocks, with the
height depending on capillary lift. The process is accelerated by frost and vegetation; breakers, which
are concave shapes caused by rock blocks breaking away and falling down; and erosion overhangs,
caused by the sideways erosion of watercourses, or the retrograde erosion and evorsion of waterfall
cascades. LOŽEK (1965) extends this list by adding overhangs, created in sedimentary formations
built up by fine carbonate sinter.
The standard dimensions of overhangs are 5 – 20 m in width, 0.5 – 2 (in rare cases up to 10) m
high, and 2 – 5 m long. Overhangs more than 5 m in length can be considered caves with a wide-open
entrance.
Fig. 5. Overhanging rock (left), rock overhang (right). Illustration from the authors´ archive.
13
Superficial subterranean habitats
Rock massifs and rock systems often contain systems of small spaces, too small for a person to enter, only a few millimetres or centimetres high yet running for several metres in all directions. These
can originate in many ways, depending on tectonic processes, slope movements, chemical erosion,
etc.
JUBERTHIE et al. (1980) define the basic term – milieu souterrain superficiel (MSS), translated
into Czech as prostředí podzemních povrchů (LAŠKA et al. 2008). The English literature translates the term as Superficial Underground Compartment (SUC) (CHAPMAN 1993). JUBERTHIE
and DECU (1994) propose
the English term Mesocavernous Shallow Substratum,
which would allow the same
abbreviation to be used.
The biospelological literature also sometimes uses the
synonym Mesovoid Shallow
Substratum (e.g. LÓPEZ and
OROMÍ 2010).
MSS is primarily a complex of narrow, impassable
interlinked cracks and fissures, generally situated in
the uppermost part of the
bedrock, where it meets the
soil cover. MSS can be found
in the deepest parts of the soil
profile in a position referred
to by pedologists as where
the soil-forming substrate
comes into contact with
the parent rock, and generally feature cavities copying
Fig. 6. Mesocavernous Shallow Substratum (MSS), an example of the soil and
the cracks in the rock as it
bedrock interface. Free interpretation of Juberthie and Decu (1994).
gradually weathers. Another
place where similar, shallowly set spaces below the surface of the ground can occur, according to RŮŽIČKA (1993), are the
spaces between grains of sand, gravel and stones in stony scree and stone seas. Spaces like these can
also be classed as MSS. If the cavities between rock blocks in boulder scree are large enough to admit
a person, they can then be described as talus-type caves. Although MSS are completely inaccessible
to man, these cavities are very important for the circulation of underground water and the migration
of underground organisms.
Besides these shallowly set systems, many rock complexes feature systems of fan-shaped cracks
and fissures which often extend very deeply into the rock massif. These cracks are generally caused
by the buckling and gradual movement of lithospheric bodies. In soluble rocks, cracks can be the initial stages of a cave, while JUBERTHIE et al. (1980) call them milieu souterrain profondeur (MSP).
CHAPMAN (1993) refers to similar systems as Deep Underground Compartments (DUC). In Czech,
the translation would be something like “prostředí hlubokých podzemních povrchů”.
14
Stony accumulations
A further type of natural subterranean habitat is bound to various stony accumulations. Certain
types of these subterranean habitats can be classified into different categories – mainly debris caves
and small caves. Terminology of the formations generated by mechanical and chemical weathering of
the rocks of rock massifs is relatively complex and, in particular, heterogeneous. Therefore variously
worked terminological hypotheses can be found in the literature (e.g. RUBÍN et al. 1986, DEMEK
1972). Despite this, numerous authors use different terms, often within identical geomorphologic
areas.
The criterion for classification is mainly the material of the accumulation and its shape. Further
criteria can be used in greater detail, for example, the dominant direction such accumulation moves
if these are stony accumulations on slopes, or the depth of accumulation. However, to obtain these
characteristics is usually very demanding in practice. So, as alleged by RAŠKA (2011, 2007), the
geologic and geomorphologic research based on studying the more advanced classification criteria is
rather exceptional. What dominates recently in the research of accumulations is the approach based
on the methods of classical inventory of fauna and flora. It is then the studies founded on the inventory of invertebrate animals in the hillside stony accumulations that are relatively frequented. RUBÍN
et al. (1986) chose an English term – talus – for this type of accumulation on hillsides. However, as
we mention above, the terminology is not definite and various terms are used for identical formations
even within the same geographic area (e.g. rock debris, scree slopes, etc.).
Aside from the geomorphologic approach to classifying the accumulations, they can be classified
according to further criteria as well as, for instance, chemical status of rocks or the climatic aspect.
Here we can also use the more closely mentioned climatic criterion, which has the closest relation to
the formation of underground biota referred to in the caves. The same chapter then contains a description of the phenomenon of ice pits and streams of hot air – ventaroles. Today, the stony accumulations
are therefore not only significant geomorphologic elements in the countryside but localities with significant fauna and flora as well.
Stony accumulations in the Czech Republic are developed in various regions that show pronounced
superelevation. These are mainly the mountain regions, canyon-shaped river valleys, etc. The specific
significant regions include, for example, the České středohoří Mountains, the Doupov Mountains, the
Lužické Mountains, or even the frontier ridges, where, however, most of the scree is covered with
vegetation (RUBÍN et al. 1986).
The origin of these accumulations in connected predominantly with the Pleistocene Period (1.8 My
– 11.5 ky) when the periglacial climate, influenced by not so distant Scandinavian and also Alpine
iceberg, reigned over the Czech Massif territory. What took place were the pronounced manifestations
of frost-shattering, destruction of rock bodies into the striated frost cliffs under which the released
weathered material – stony accumulations – had built up.
Since most of the known accumulations originated during the quaternary glacial periods, the erosion processes and the frost-shattering during the winter season take place now as well and formation
and development of these geomorphologic formations still continues, even if in a limited form.
15
Fig. 7. Division of the main types of stony accumulations. Free interpretation of RAŠKA (2011).
Most Important Cave Regions in the Czech Republic
The region of Central Europe, including the Czech Republic, is characterised by its long geological
development and the diversity of its rock types. The wide range of sedimentary, metamorphic and
igneous rocks has allowed a number of diverse geomorphologic processes, leading to the formation of
several thousand caves of various genetic type – HROMAS et al. (2009) put the number of recorded
caves at 3988 as of 2008.
As caves and other naturally occurring underground structures can be found in virtually any part
of the country, here we will restrict ourselves to those regions which are prominent in speleological
terms.
Bohemian Karst
Speaking of karsts, Bohemia is dominated by the Bohemian Karst region. In terms of area, this is
the most extensive karst region in the Czech Republic. Here the karst is linked to the non-metamorphic carbonate rocks of the Barrandien Lower Palaeozoic period, specifically to the strata complex of
Silurian and particularly Devonian limestone which forms the core of the Prague Basin.
Here, karstogenic rocks come to the surface in several unrelated areas, forming a band 32 km long
and 8 km wide, delineated on the south-western edge by Zdice and on the south-eastern edge by
Prague. The total area covered by the limestone is 144 km2. The core of the Bohemian Karst has been
declared the Bohemian Karst Protected Landscape Area, containing a number of specially protected
areas.
The karst phenomena in the Bohemian Karst region are linked primarily to outcrops of limestone
in river valleys, particularly those of the Vltava and Berounka, and the Prokopský, Dalejský and Radotínský Streams.
The Bohemian Karst has relatively few typical subsidiary karst phenomena such as grikes, sinkholes, plunges of surface watercourses, etc.; however, despite this, as of 1 January 2008 there were
677 recorded caves with a total length of 21.9 km, including the longest karst system in Bohemia
and also the only caves in the region that are accessible to tourists – the Koněprusy Caves (2050 m)
16
(HROMAS et al. 2009).
Moravian Karst
As the name implies, the Moravian Karst is in the eastern part of the Czech Republic, specifically
in the north of the South Moravian region. The Moravian Karst is the largest and most well-developed
karst region with the widest range of karst phenomena in the Czech Republic. The karst relief has developed in a 3 – 6 km wide and 25 km long band of Devonian limestone, which stretches north from
Líšeň and Maloměřice near Brno towards Sloup and Holštejn. The total area covered by karstogenic
rock is 78 km2. The most important parts of the region have been declared the Moravian Karst Protected Landscape Area.
Favourable geological, climatic and hydrological conditions have led to the development of a karst
relief with a marked presence of the surface and underground karst phenomenon known as holokarst
(unlike the incompletely developed merokarst of the Bohemian Karst).
A unique feature of the Moravian Karst is the longest karst system in the Czech Republic – the
Amatérské Jeskyně Cave System – 34,900 m in length.
As of 1 January 2008, the Moravian Karst contained 1133 caves, five of which are open to the public
– these are Sloupsko-Šošůvské Caves (total length 4890 m), Balcarka C. (1150 m), Kateřinská C. (950
m), Punkevní C. (4750 m) and Výpustek C. (2000 m) (HROMAS et al. 2009).
Fig. 8. The typical speleothems in Kateřinská Cave (Moravian Karst). Photo from the authors’
archive.
17
Branná Belt Karst
This karst area has developed in disjunct bands of metamorphic carbonates. These form a narrow,
elongated belt stretching NNE-SSW, 60 km long and 3 km wide between the towns of Ruda in Moravia and Vápenná in the north of Moravia (the Šumperk district). As has already been said, the karst in
this region has developed in rocks that are not sedimentary but metamorphic – varying from granular
white to bright greyish-white banded crystalline limestone, originally probably dating back to the Devonian. The Branná Belt Karst is particularly characterised by the development of underground karst
phenomena and active karst hydrography. Surface karst phenomena are less perfect and sporadic, and
include, for example, shallow sink-holes, imperfectly developed grikes, etc. The most extensive cave
system in this region is the Na Pomezí Cave (1320 m), which is open to the public and is also the largest karst cave formed from crystalline carbonates in the Czech Republic. A total of 67 caves have been
recorded in the Branná Belt Karst (HROMAS et al. 2009).
Nearby there is a small karst area known as the Supíkovice Karst, formed by relicts of contactmetamorphic Palaeozoic limestone.
This limestone is white to grey with
a medium to coarse grain, containing, besides grikes, sink-holes and
one exsurgence, just one cave – the
Na Špičáku Cave. Due to its long
length (410 m), it has been opened to
the public.
Hranice Karst
This karst formation is situated to
the east of the Olomouc Town, on
both banks of the deep valley of the
Bečva River, roughly between towns
Hranice, Teplice nad Bečvou and
Černotín. Several bodies of limestone
from the Devonian to the Lower Carboniferous protrude from the surrounding younger rocks in a band 5.5
km long and 4 km wide, running in a
SW-NE direction.
The Hranice Karst was formed
through two different processes; classic karstification and hydrothermal
karstification, which here is the result of tectonic turbulence in the deep
zone where the geological blocks
come into contact with intensive outflows of tepid acidulous waters with
a high CO2 level, causing increased
Fig. 9. The Hranice Chasm, the deepest karst abbys in Czech Repubcorrosion.
lic. Photo I. Žambochová.
The most important surface manifestation of this hydrothermal karstification is the Hranice Chasm (reaching a known depth of 273.5 m below ground level); as regards un18
derground formations, it is worth mentioning the Zbrašov Aragonite Caves (1240 m), which are open
to the public and known for their unique CO2 gas lakes and blooms of clear to milky-white aragonite.
So far 29 caves have been recorded in the Hranice Karst (HROMAS et al. 2009).
Javoříčko Karst
The Javoříčko Karst is situated in the north of Moravia in the northern part of the Drahanská Vrchovina highlands, centred on the village of Javoříčko, some 12 km WSW from the Litovel Town.
Protruding from the not very rugged landscape relief are the featureless wooded peaks of the Rudka
(589 m.a.s.l.) and Špraněk (539 m.a.s.l.) hills among others. The karst phenomena are linked to De-
Fig. 10. Large amount of very thin stalactites in the Javoříčko Cave system. Photo from the
authors’ archive.
vonian limestone, which covers an area of approximately 6 km2. Typical features of this karst are
grikes, sink-holes, plunges and exsurgences; canyon-shaped valleys and collapse sink-holes. Plunges
and exsurgences are common throughout this area; 52 caves have been recorded, one of which is open
to the public – the Javoříčko Cave system (approximately 4000 m long) (HROMAS et al. 2009).
Mladeč Karst
The Mladeč Karst is situated in the Olomouc district, in the broader environs of Mladeč on the edge
of the Bouzov highlands. The karst phenomena are linked to the limestone bodies of the Skalka (335
m.a.s.l.) and Třesín (345 m.a.s.l.) hills among others. It is separated from the nearby the Javoříčko
Karst by a transversal tectonic line. The limestone with karst phenomena is from the Devonian and
is characterised by the presence of surface and underground karst features (grikes, sink-holes, sand
pipes). So far 6 caves have been recorded in this karst area (HROMAS et al. 2009). The most famous
19
karst feature of the Mladeč Karst is the Mladeč Caves (1250 m), which are open to the public; these
caves are an important archaeological and paleontological site (WILD et al. 2005).
Pálava Hills Karst
The Palava Hills in the south of Moravia form a long, rugged ridge running SW-NE, 10 km long
and 7 km wide, rising from the lowland of the Dyjsko-Svratecký Úval ravine between Mikulov, Dolní
Věstonice and Pavlov. They are formed from tectonic relicts of Jurassic to Lower Cretaceous limestone belonging to the Western Carpathians’ system.
The limestone massifs here have a pronounced karst relief with massive rock walls, steep precipices, solitary boulders, isolated stretches of dry karst valleys, and the occasional sink-hole. Scattered
around are small caves, a fine example of which is the Na Turoldu Cave system of cave passages and
halls, open to the public (1650 m
long). A total of 23 caves have been
recorded in the Pálava Hills Karst
(HROMAS et al. 2009).
Chýnov Karst
This is a small karst region situated in South Bohemia, not far from
the town of Tábor. In the broader
surroundings of the town of Chýnov,
in the SE part of the Tábor district,
there is a series of bodies of metamorphosed crystalline limestone
and dolomites in thin bands, pockets, and larger massifs. The entire
area constituted by carbonate rocks
forms a belt running roughly east
to west, 17 km long and 3 – 4 km
wide. The karst is more markedly
developed in the massif of crystalline limestone that form the Pacova hora Hill (589 m.a.s.l.), 2.5 km
northeast of Chýnov. This also contains the largest cave – the Chýnov
Cave, whose passages run for a total
length of 1400 m. This cave, which
is open to the public, is unique for
the absence of classic sinter and stalactite decoration, instead featuring
a wide range of shapes formed by
Fig. 11. Erosion shapes “Purkyně’s eye” in in the Chýnov Cave. Photo
erosion, evorsion and corrosion; it is
from the authors’ archive.
also famous for its diversity of colours, with yellow, reddish and greenish colours predominating. So far a total of 4 caves have been
found at the Chýnov Karst (HROMAS et al. 2009).
20
Karst of the Kamenice and Železný Brod catchment basin
This is not a particularly extensive karst region, linked to isolated relicts of metamorphosed carbonate rocks in the Bozkov highlands, roughly between the towns of Vysoké nad Jizerou and Železný
Brod in north-eastern Bohemia. It is composed of disjunct bands, pockets and irregular bodies of
metamorphosed carbonates originally dating from the Silurian in varying quality and composition,
from highly pure crystalline limestone (99% CaCO3) through to dolomitic limestone and dolomites.
Most karst phenomena are on the edges of deep valleys. A total of 10 caves have been recorded, one
of which is open to the public – the Bozkov Dolomite Caves (1060 m) (HROMAS et al. 2009).
Železné hory Karst
Emerging in the Železné hory (Iron Mountains), particularly to the south of the Heřmanův Městec,
are slightly metamorphosed rocks from the Chrudim Palaeozoic, specifically from the Ordovician and
Devonian periods. In the central area, around Prachovice and Vápenný Podol in the western part of
the Pardubice region, there are some fairly mighty limestone bodies with numerous karst phenomena
and caves. The substrate of the carbonate rocks comprises dark-grey moderately crystalline limestone, on which there are massive greyish-white to white crystalline limestone with isolated chert and
veins of diabase. The limestone body covers an area of approximately 3.5 by 0.5 km.
In the past the relief has been disturbed by a number of small local quarries; nowadays what is
causing the most disturbance is the large-scale the Prachovice Quarry – most of the western part
of the massif near Prachovice has been quarried out, while the eastern part above Vápenný Podol is
buried under the dumping ground. Although the original landscape featured surface karst phenomena
(grikes, sink-holes, plunges and emergences), only fragments remain today. The only caves to have
been described are from the quarry areas, and most of them have already been quarried out. A total of
41 caves have been recorded; the only larger ones to have been preserved are the Podolská Cave and
the Páterova Cave. None of the caves are open to the public (HROMAS et al. 2009).
Tišnov Karst
The Tišnov Basin to the northwest of the Brno Town is flanked by several karst zones. The most
significant of these is the Květnice hill on the left bank of the Svratka River on the north-western edge
of Tišnov; there are other smaller karst regions on the slopes of the Dřínová and Dranč hills, as well
as around the villages of Heroltice, Lažánky, etc. The karst here is linked to the Devonian limestone
of relatively high chemical purity. A total of 35 caves have been recorded in a band 6 km long; none of
them are open to the public. The longest is the Králová Cave system (approximately 800 m). Surface
karst phenomena in the region include, for example, submerged watercourses (HROMAS et al. 2009).
Main pseudokarst areas in the Czech Republic
Pseudokarst phenomena, relief forms where karst and similar formations have emerged in non-karst
rock, are relatively abundant in the Czech Republic. An extraordinary quantity and diversity of forms
are linked predominantly to massive strata complexes of sandstone blocks in the Bohemian Cretaceous
Basin, spread across the central, northern and eastern parts of the Czech Republic. Prevalent here are
fissure, crevasse, contour, combined, alcove and block (scree) caves and chasms. Many of these natural underground spaces have been modified by man in the past (as cellars, shelters, etc.). The most significant occurrence of sandstone pseudokarst is undoubtedly in the Děčín highland area, where there
are more than 150 caves ranging in length from a few metres to around 160 m, including a number of
chasm-like caves (e.g. the Loupežnická – Pytlácká Cave system), as well as the Ještěd-Kozákov ridge
in the Krkonoše foothills, the cave in the Dokes hills, which are part of the Kokořínsko Protected
21
Landscape Area, the series of narrow
chasm-like caves in the Jičín hills, the
pseudokarst relief in the Broumov highlands, and the Adršpach-Teplice Rocks
(the site of Teplická Cave, at 1065 m the
longest pseudokarst cave in the Czech
Republic).
Caves in the Bohemian Cretaceous
Basin are not limited merely to sandstone locations; there are a number of
caves in arenaceous marl and marlstone
areas, e.g. near the River Ohře (Díra u
Kystry Cave), and there are many caves
around Ústí nad Orlicí in Eastern Bohemia.
There are extraordinarily extensive
systems of crevasse, fissure and block
caves and chasms in the sandstone
and marl flysch belt of the Western
Carpathians (for example Kněhyňská
Cave, which is approx. 280 m long, the
Cyrilka fissure and chasm system, 370
m long, etc.).
The processes which accompanied
the young relief formed during the Cenozoic period also led to the formation
of predominantly crevasse and fissure
Fig. 12. The Loupežnická – Pytlácká cave system in the Děčín highcaves in the vulcanites of the České
land area is created by narrow fissures and chasms. Photo from the
středohoří Mountains, the longest of
authors’ archive.
which, Loupežnická Cave near Ústí
nad Labem, is roughly 130 m long.
The occurrence of pseudokarst caves is not limited to the presence of sedimentary rocks. Scattered
around the country there are isolated natural underground spaces (mostly overlapping crevasses, extended fissures and scree cavities in rocks and rocky scars). They are predominantly linked to solid
rocks with block disintegration, particularly magmatites and metamorphites. These include, for example, the caves in the Šumava foothills and the Novohradské Mountains formed by frost weathering
in the Quaternary, the 400-metre-long Ledové sluje (Ice Cave) near Vranov nad Dyjí, and the caves
in the Palaeozoic orthogenesis of the Ore Mountains among others (POKORNÝ and HOLEC 2009)
22
Subterranean habitats from the viewpoint of protection and administration
Subterranean habitats from the viewpoint of protection and
administration
Cave utilization
Man has used underground, above all caves, long-ago. It was not only a man as a cave inhabitant. Caves were profusely used much later, too, by e.g. stalactite gatherers and merchants or skeletal
remains of Vertebrates etc. The skeletal remains used to sometimes be processed to a large volume,
e.g. for sugar industry purposes or for production of fertilizers from phosphatic clay (HROMAS et al.
2009). The caves have been the main goal as well as a source of pleasure to many tourists, amateur
speleologists, rock climbers etc. Speleoteraphy deals with beneficial effects of the cave climate on human health (for more details see e.g. ŠTELCL and ZIMÁK 2003, BOHÁČ et al. 2001, ŠTELCL 2000).
Importance of caves is also undisputed when talking about science, because even the underground
ecosystems are not entirely separated from above-ground as well as human environment. They represent a source of biodiversity of organisms, often unknown elsewhere, they are also a part of energy
flow, nutrient cycles etc.
Cave management and care
The two authorities that look after caves that are open to the public and also protected under the
law in the form of a small-scale protected territory are state organisations under the Ministry of
the Environment. These are the Agency for Nature Conservation and Landscape Protection of the
Czech Republic (AOPK ČR) and the Cave Administration of the Czech Republic. AOPK ČR, based in
Prague, is an organisational body of the state, set up by the Ministry of the Environment. Its primary
task is the care of nature and the landscape in the Czech Republic, i.e. including caves and other areas
below ground.
There are a total of 14 localities in the category of caves that are protected and open to the public in
the Czech Republic – the Koněprusy Caves (NMM Zlatý kůň), NMM Bozkov Dolomite Cave, NMM
Chýnov Caves, NMM Zbrašov Aragonite Caves, Javoříčko Caves (NNR Špraněk), Mladeč Caves
(NMM Třesín), NMM Na Pomezí Caves, NMM Na Špičáku Cave, Punkva Caves (NNR Vývěry
Punkvy), Balcarka Cave (NR Balcarova skála-Vintoky), Kateřinská Cave (Moravian Karst Protected
Landscape Area), NR Sloup-Šošůvka Caves, Na Turoldu Caves (NR Turold), and the Výpustek Cave
(Moravian Karst Protected Landscape Area).
Caves not open to the public yet protected as part of a small-scale protected territory are administered by AOPK ČR, or the Administration of Protected Landscape Areas and the National Parks
Administration. As of 1/1/2009 there were more than 100 similar sites in the Czech Republic and, according to conservationists, this figure is rising gradually. The great majority of them are mentioned
in the summary monograph by HROMAS et al. (2009). Some of the more well-known of these include, for example, the longest pseudokarst scree cave in the sandstone of the NNR Adršpach-Teplice
Rocks, the Pod Luciferem Cave in the NNR Broumov Walls, the combined Pod Jezevčí dírou Cave
in the NR Ostaš, or the Naděje Ice Cave in the natural monument of the same name. The Ústí Region
also contains several similar cave sites – NM Loupežnická Cave, where the main subject of protection
is the pseudokarst phenomenon, as well as the NNR Bořeň, NM Radobýl, and NM Hradiště, where
the primary focus of protection is generally unique plant communities and within the boundaries are
smaller non-karst caves.
Many important caves have also been selected as being necessary for safeguarding nature protec23
tion at the European Union level. The current list of localities which contain caves not open to the
public as protected habitats under the NATURA 2000 system (i.e. Special Areas of Conservation),
lists 36 sites for the Czech Republic.
As part of NATURA 2000 biotopes (i.e. biotopes listed in Annex I of the Habitats Directive (92/43/
EEC of 21 May 1992), in addition to caves there are other biotopes bound to habitats under scree. The
term scree is then part of three other specific biotopes and these are protected in 25 territories in the
CR under NATURA 2000 (www.nature.cz).
The aforementioned directive covers not only the protection of biotopes, but also species referred
to as “nature” species. Of the organisms which are associated with caves, and particularly the deeper
parts of their cores, the EU regulations only specify eight species of bats that live in this country. Special Areas of Conservation have been set up for the six most common species, yet the Lesser MouseEared Bat (Myotis blythii) and Greater Horseshoe Bat (Rhinolophus ferrumequinum), which occur
sporadically in the Czech Republic, do not have their own Special Areas of Conservation.
The majority of caves in the Czech Republic are not open to the public and also protected by law in
accordance with Sect. 10 of Law No. 114/1992 Coll. on general protection. Such caves are automatically protected under the law and the state has the duty to look after them through a specific regional
authority, the appropriate Protected Landscape Area Authority or National Parks Authority in largescale protected territories. Military training areas, including cave sites, are under the authority of the
appropriate Military Training Area Office. Sect. 10 of Law No. 114/1992 Coll. thus provides general
protection to the majority of our approximately 5 000 caves. Basically, Sect. 10 states that caves must
not be destroyed. However, research of caves now requires a permit from the appropriate nature protection body. The same applies in the case of organisms, where it states that all species are protected
against destruction, damage, collecting and hunting which endangers or could endanger the existence
of these species or threaten their degeneration, disturb the reproductive abilities of the species, depopulate the species or destroy the ecosystem they form part of (i.e. the general protection of species
in accordance with Sect. 5, Paragraph 1 of the law).
In practice it is of course difficult to prove such damage to ecosystems and their species when
there are no restrictions on physical access underground and to subterranean species, this is why the
laws on the special protection of territories and species are more stringent. MŽP Directive 395/1992
Coll. distinguishes between endangered, seriously endangered and critically endangered species and
small-scale and large-scale protected areas. With the exception of Chiroptera, i.e. bats and horseshoe
bats, which are all subject to special protection (either as seriously endangered or critically endangered species), there are no other specially protected species of animals that are closely associated
with cave habitats. In small-scale categories of territorial protection there is a differentiation between
the categories Natural Monument, Nature Reserve, National Natural Monument and National Nature
Reserve. While the first two categories allow free access throughout the reservation, even though any
kind of damage, including catching animals and collecting plants, is forbidden, in the remaining two
categories visitors are restricted to marked pathways. In all cases access may be restricted in line with
the needs of nature protection, and it is often restricted for underground areas, e.g. in order to protect
bats and horseshoe bats.
However, the laws relating to nature protection, i.e. in this sense Law No. 114/1992 Coll., on the
Protection of Nature and the Landscape, including the implementary directives, are relatively complicated and therefore there has not been space to go into the details of all their aspects and options. The
protection of species and habitats is also covered by a series of other laws. The insensitive use of the
“above-ground” landscape is covered by other regulations, which also incorporate relations towards
24
Subterranean habitats from the viewpoint of protection and administration
underground areas. However, the impact on subterranean habitats may be much greater, as a major
specific characteristic of such habitats is the presence of only small quantities of nutrients. Moreover,
it is practically impossible to really check the situation in these subterranean habitats. Besides agriculture, gas emissions, logging in forests, etc., excess tourism can also have an adverse effect. Once
again, this is often mere speculation and no thorough research into this field has been carried out.
Recording visitor figures is also clearly a problem. In many caves visits can be entered into visitor
books, although visitor records in caves that are not open to the public is a purely private matter, or
one dealt with by various associations. One practical and effective form of protection, although still
not completely adequate, is to block cave entrances using either grilles or by partially bricking them
up. However, considering the number of caves that exist, this is not a feasible option.
Cave research and evidence
Research activity of organisations as well as the individuals is generally being presented on the occasion of a number of scientific meetings in form of lectures, posters and in scientific and other special
journals. In relation to the subterranean environment we can recall that the Czech Republic was (1973)
and again is going to be (2013) a venue of a speleological congress, which proceeds once every four
years in some member country of the International Union of Speleology (UIS). So called “Speleoforum” is the other very important event for all speleologists. This annual meeting of the Czech Speleological Society members and other speleologists from the Czech Republic as well as foreign countries
takes part once a year in the Moravian Karst. Its main activity is a presentation of the results achieved
in past year and long-term research.
Caves, indeed, are the main subject of speleologists´ interest as much as the subterranean is from
the practical point of view. That is why the other aspects of the research, connected with caves and
subterranean environment, are significant for this branch. This fact extends the possibility of data
presentation, too. The relevant information on caves is possible to be expected in a range of other
technical (ecological, zoological, climatological, geological, mineralogical, archaeological etc.) meetings as well as in other technical and scientific journals.
Much like we remembered certain significant speleologists´ meetings at the national and international level, we must mention some national and international scientific and technical journals. To
prestigious ones, where the results in term of important international research are published, belong
e.g. Cave and Carst Studies, Acta Carsologica, Subterrannean Biology etc. When talking about national ones, these are Speleo and Speleoforum, aimed at speleology.
We cannot neglect even the research of people and organisations that do not publish their observation results or that do not use recommended techniques of scientific methodology, although they
make their valuable knowledge available, whether in form of journals or various final reports (so
called grey literature). Both types of these text outputs have their own advantages and disadvantages
but we are not going to follow them. However information evidence and availability can signify a
quite big problem. Some of the scientific journals can be checked in free or paid databases. Published
technical works, but also the unpublished ones, except the plans of landscape management which are
the subject of special preservation, are collected by the Nature Conservation Agency of the Czech
Republic (NCA CR, AOPK ČR in Czech) in so called reservation books, available in electronic form
of DRUSOP system. It is also possible to get there demanded scientific document in electronic form.
Database of speleological objects (so called JESO) exists and is still built up in unpublished form. The
NCA CR, the Cave Administration of the Czech Republic, the Czech Speleological Society and others
participate in its build-up process and they also contribute to discovering and evidence of new caves.
These organisations have the essential meaning when speaking about subterranean investigation, too.
25
Environmental Conditions and Their Measuring
Subterranean habitats differ largely from open habitats. The dominating difference is the absence
of both luminous and thermal energies of incident solar beams. This results in a differing microclimate character (temperature, humidity, air currents) and dissimilar velocity and type of exogenous
processes (erosion, sedimentation). Differences in the composition of the atmosphere (accumulation
of CO2 or, alternatively, radon) etc. can be observed in enclosed subterranean spaces. These and further biotic factors then influence the species composition of organisms which are in harmony with the
subterranean conditions that offer these organisms a suitable habitat for living.
In order to be perfectly capable of understanding and interpreting the operation of processes and
ecosystems in subterranean habitats we have to know the character of the above-mentioned environmental factors, learn the accurate method for measuring them and, consequently, interpret the
acquired data correctly to yield functional connections. The next chapters therefore specifies the basic
characteristics of the selected most important factors, the methods of their measurement and evaluation in relation to the subterranean habitats.
Subterranean Microclimates
PŘIBYL et al. (1992) define three basic types of caves in the relation to their microclimate, where
the nature of the circulation of air masses in the system of inner/outer environment is considered to be
the main classification criteria. The first type is a static cave. There is generally one entrance to these
caves or if there are even more entrances, they must lie at the same altitude.
As long as the floor flow comes down in the direction of the entry below its level, in winter season
the lighter warm air begins to escape up out of the cave and at the same time the outer cooler air is
sucked in. In summer the circulation stagnates, since the heavy cool air is kept inside the cave and
warming in the direction of the depths from the entrance is only slow. Static caves with the floor falling from the entrance down are called cool caves.
In the case of the cave cavity being
situated above the entry level, a quite
opposite effect occurs. In summer season the cool air spontaneously flows out
of the cave and the warmed outer air is
sucked in. Later, in winter season, this
reservoir of warm air is cooled down by
the surrounding rock only slowly and in
winter, when the stagnation of circulation occurs, the temperature in these
static warm caves will to be markedly
higher than outside the cave cavity.
Static caves are thus generally characterised by air circulation only during some defined part of the year (cool
ones in winter, warm ones in summer).
However, similar caves are rather rare,
Fig. 13. Top left is the static cave with the upper entrance mainbecause most of the caves, karst or nontaining the cool climate during the whole year. Top right is the
static cave with the lower entrance where the cool air accumulates
karst, are established in a rock mass
during the whole year. Bottom left is the dynamic cave with air
interwoven by sets of cracks, bedding
circulation during the whole year. Free interpretation of MUSCIO
joints and slots. Small cavities generally
(2002).
26
allow passage to the surface and due to that continual circulation begins and continues for the whole
year. Relatively frequent are also cases when there are multiple inputs to the cave (both passable and
non-passable) at various altitudes. These caves are categorised as dynamic.
The air circulation in dynamic caves proceeds according to the scheme whereby in summer the air
circulates from the upper entrance in the direction of the entrance situated below, whilst in winter the
opposite occurs. In summer the driving force is the cool air inside the cave which, due to its higher
weight, flows out of the cave through the lower entrance and from above is substituted with the warmer air from outside. The intake air is cooled down through the heat transfer to the rock due to which
the relative air humidity is increased. When the air is 100% saturated, the subsequent precipitation of
humidity occurs. In contrast to static caves, the circulation cycle in winter is not interrupted, but due
to the outer temperature decrease it reverses. The inner warmer thus lighter air exits through the upper orifice and the cool air is continually sucked in through the lower entrance of the cave. Along with
the cooling of the rock from the cool air in winter, the air humidity in the cave when compared to the
summer heating of the rock is decreased and is supplemented by evaporation from the walls. So it can
be generally said that in dynamic caves the walls in summer are dewy and damp, and conversely, in
winter these caves become significantly drier.
According to PŘIBYL et al. (1992), statodynamic caves are sporadic and specific. Their basically
dynamic nature is interrupted for some part of the year due to the closing of one entrance, e.g. by
temporary glaciation or flooding. This place then acts as a stopper that prevents the circulation of air
masses. Similar air circulation occurs also in cracked massifs or rocky accumulations.
Places with warm winter flow streams were in the past incorrectly connected with volcanic activity and the improper term “fumarole” was used (see e.g. SOMMER 1833, POKORNÝ 1931), which
comes from the Latin word fumus, meaning smoke. Earlier authors speculated about the existence of
a crack system extending to such depths that the air could be heated from thermal groundwater bodies
extending from the foothills of the Ore Mountains (e.g. ŠIMR 1957).
Only later was it found (specifically by VÁNĚ 1992) that they are manifestations of air circulation
and therefore the term ventarols (ventus = wind), which better represents the principle of the phenomenon, was suggested.
Places where cool air and even snow and ice remain till late spring are called ice pits. Provided
that it is a cave, the term ice (where ice remains in the cave for the whole year) or pseudo-ice cave is
used, according to the time for which ice remains in the cave. According to HROMAS (1971) there
are no real ice caves in the Czech Republic. The various “ice caves” are therefore pseudo-ice caves.
Pseudo-ice caves are definitely more frequent. With the character and nature of the climate they are
very similar to ice caves, but due to a coincidence of factors (small size and length, partial insolation
of the entrance, periodic through-flow of water, etc.) the ice is present in these structures only until
spring or, at most, until summer. For some part of the year, these caves are completely without ice. For
pseudo-ice caves thick snowdrifts of firn snow in the cave portal are characteristic. The snowdrifts are
reservoirs of snow and at the same time they maintain the cool air inside the cave. Among the karst
pseudo-ice caves in the Czech Republic that can be named are Ledová Cave in the Moravian Karst or
Blátivá Chodba Cave in the Chýnov Cave system.
In the category of non-karst caves, Ledové Sluje (Ice Caves) near the town of Vranov nad Dyjí
should be mentioned. It is probably the longest cave in the Bohemia, well explored by professionals,
with the total length of up to 400m (HROMAS and BÍLKOVÁ 1998a, b). About 10m long, Ledová
Jeskyně cave in the body of Velký Bezděz is only briefly described by CHVÁTAL (1996) and HROMAS (1971) and is substantially less well-known.
In the Ústecký kraj region there are also structures with the character of pseudo-ice caves. In particular, there are cleft caves on the hill Buková hora, some of which have their cool nature already in
27
their name – Ledová jeskyně (Ice Cave) and Sněhová jeskyně (Snow Cave). The former has already
been mentioned in literature almost two hundred years ago, when (SOMMER 1833) spoke of the “30
feet deep crack where snow can be found even in late summer”. PLEISCHL (1838) published the
discovery of several deep clefts near the hill Buková hora and stated that during his visit at the end
of August there was already no snow. ZAHÁLKA (1890) describes in the vicinity of the peak of the
hill Buková hora the 12m long cavity with a NS orientation where an icy glaze forms even in the early
summer months. It can be found in travel literature that in the vicinity of the Ledová Cave cryophilic
species of plants grow due to the long presence of ice and snow, e.g. Rosa pendulina and Stachys alpinus (BREDSCHNEIDER 1928).
The presence of cool air to a certain extent simulates the character of the northern climate which is
confirmed by the presence of various cryophilic organisms – e.g. the mite Rhagidia gelida (ZACHARDA et al. 2005) and also the northern European spider Semljicola faustus (ČEŘOVSKÝ and HOLEC
1996). Conversely, the existence of ventarols and thus places where the temperature even in winter
doesn’t drop below freezing enables on the Borečský Hill the occurrence of the thermophile Mediterranean liverwort Targionia hypophylla, only at this one locality in the Czech Republic (KUBÁT
1971).
Probably the best known places where ventarols and ice pits occur are the National Nature Monument Boreč Hill (esp. ventarols), the Nature Monument Plešivec Hill in the district of Litoměřice (esp.
ice pits) and also the Nature Reserve of Kamenná Hůra east of Děčín (well developed ice pits and
ventarols). These structures are so significant that their conservation is protected by the declaration of
small-scale conservation areas where the main object of conservation is the occurrence of ventarols
and ice pits.
Light
The main source of light in subterranean habitats is the Sun, which continuously releases energy
quanta in the form of photons. These energy particles are radiated in all directions from the surface
of the Sun into free space. Photons are carriers of predominantly luminous and thermal energy. What
is defined in relation to the incident solar energy is a solar constant, which represents the amount of
energy that strikes during a day upon the Earth’s atmospheric surface of an area of 1 m2 perpendicular
to the direction of incident beams per unit of time. For the Earth, this constant attains a value of 1.38
kJ.m-2.s-1. However, the solar radiation is bounced off the Earth’s atmosphere to be absorbed with its
quality changed by dispersion. Only 51 % strike the earth’s surface, more than half of which is represented by visible light – that is, radiation of 400-750 nm wavelengths, where the remaining half falls
on ultraviolet and infrared radiation (SCHLESINGER 1997).
To measure the luminosity conditions in subterranean habitats, a convenient physical quantity must
be defined, that is, illumination defined as luminous flux Φ incident on specific area.
The illumination unit is lux (lx) and the illumination intensity in the open unshaded space during
a sunny summer day (in the conditions of the Czech Republic) is within 70000–100000 lx. The common value of diffused natural illumination inside buildings reaches 100–2000 lx. A bright moonlit
night during a full moon represents illuminance of max. 0.5 lx, and a moonless starry night approximately 3*10-4 lx. The limit of perception of human sight for the light stimuli is within the order of 10-9
lx (PEŠA and MAJER 2003).
The easiest method of measuring the illumination intensity underground is by using the apparatus
referred to as a luxmeter. Luxmeter designs may differ and they are associated with the luminous flux
measurement method and the principle of the actual light-sensitive receptor (photocell). The luminous
flux is usually measured using a suitable circuit with an operating amplifier, and the luxmeter must,
in addition to the receiver and photocell, also contain the actual measuring and evaluation system,
28
which will convert the established illumination intensity to a relevant absolute numeric value within
the calibration curve.
To be considered a first-rate luxmeter the apparatus must have a spectral sensitivity that corresponds to the spectral sensitivity of the human eye (or it is alternatively higher in orders) and whose
light-sensitive receptor is simultaneously equipped with a so-called cosine corrector to eliminate directional (angular) error at the oblique incidence of light (JENČÍK and VOLF 2003).
The luxmeter measuring process in subterranean habitats – most frequently in caves – takes place
in the following way. The process always starts outside the cave where a measurement is taken of the
illumination intensity in the outer space and it is used as a comparative value. The next step leads into
the cave where the helper holding the apparatus turns in such a way that the photocell is oriented at a
right angle to the cave mouth. The values are always read from the apparatus at a regular interval that
is chosen by helper in the methodology (or by one metre length, etc.). Reading the illumination value
requires utilization of the apparatus memory and working “in the dark”; supplementary lighting by,
for example, forehead torch, is inadmissible. Thanks to the acquired numeric data, it is possible to
delimitate, in the subterranean habitats, the following three basic sectors – luminous part (ca. >80%
of the outside illumination), transition part (80-20%) and aphotic part (<20%) (see Fig. 14).
It is a matter of fact that the character of solar radiation entering the subterranean spaces is significantly influenced by the morphology of the outer environment and the condition of vegetation. Another important factor is the plasticity of actual subterranean habitats – orientation of the cave mouth
to the points of the compass, the cave mouth’s size of a cave on a slope, the morphology of corridors,
Fig. 14. Delimitation of the luminous, transition, and aphotic part in the example of the Malá Jeskyně skřítků Cave.
Illustration from the authors’ archive.
29
the rock material, etc.
Since the illumination intensity is significantly dependent on the influence of direct light exposure,
it is recommended to perform measuring with an overcast sky when only diffused light penetrates
under the ground. However, as alleged by PEŠA and MAJER (2003), adherence to this condition
fails to completely solve the issue of distortion of the measured values in dependence on the direction of incoming light. The referred authors then recommend applying the modified luxmeter whose
photocell is placed in the centre of a frosted glass bulb. The light striking upon the bulb penetrates its
glass to form a homogeneous diffused light inside this bulb after multiple bounces off the bulb’s lightcoloured walls. Since then the photocell senses the value of the illumination incident in all directions.
Luxmeters equipped with continuous recording of the read values are available for continuous measurement of illumination. However, the disadvantage of these apparatuses with relatively high cost is
their relatively large dimensions (at least the size of a mobile phone) (see Fig. 15). It could then be a
problem in a series of caves to hide such a large apparatus from possible casual visitors.
Fig. 15. Luxmeter PHYSICS Line C. A 811 with the measurable range 0.01 – 99990 lux. Photo from the authors’
archive.
Heat (temperature)
About 45 % of the solar beams incident upon the Earth’s surface has a shape of radiation with 750
– 5000 nm wavelengths (the radiation with higher wavelength is filtered by the atmosphere). Such a
radiation of wavelength exceeding the wavelength attained by the visible light is referred to as infrared radiation or, as case may be, thermal radiation (SCHLESINGER 1997).
At the incidence of this radiation on the Earth an energy is conveyed; this incites acceleration of
molecular oscillations of the matter exposed to the thermal beams = the Earth is heated. The physics
strictly differentiate between two quantities – heat and temperature. While the heat is an aggregate
energy of all the moving molecules of specific matter, the temperature is a measure to express how
quickly the matter molecules move. In the context below, the heat will be considered the energy condition and the temperature the measured characteristics (JENČÍK and VOLF 2003).
30
It is possible to define several basic sources of heat in the subterranean habitats. In the places that
are exposed to direct solar radiation for a part of the day or year, the main source of heat can be seen
just in the Sun and heat is conveyed by radiation here. These places are, for example, the front parts
of caves with a wide entrance or oriented to the direction S or SW.
The Sun can also warm the subterranean habitats vicariously, that is, via a medium. A typical example is stony accumulations (stony debris, stone fields). In summer, their surface is often heated up
to a temperature of 50°C (HOLEC, not published) and the thermal energy then passes through the
rock under the ground by conduction (conveyance).
Specific air circulation takes place in the underground systems of a shape of a fissure or slot system,
e.g. faulted rock massifs, extensive stony accumulations, etc. during a year (see Chapter Subterranean
Microclimates). The convection (circulation) principle is applied here then wherein the subterranean
habitats get warmer from the warmer air penetrating here from the outside.
Intensive microbial decomposition of the organic matter at sufficient supply of oxygen, so called
mouldering, takes place in the inclined caves or abysses wherein large quantities of the organic matter
Fig. 16. The Lesní díra Cave in the Kaňon Labe National Natural Area (Děčín District) is a typical place where
organic matter gathers underground to decompose and this leads to the generation of large quantities of heat.
Photo from the authors’ archive.
(litter-fall, leaves, etc.) accumulate and, at the same time, the influence is combined here of a large
volume of penetrating water and dynamic circulation, which manifest themselves by heating the air
passing through the rock cracks. These are processes similar to those that take place in a common leaf
mould which result in an exothermic reaction that is capable of warming through the decomposing
matter up to 20-30°C. The released heat then warms the air in the cave and this air rises towards the
31
surface. The streams of hot air are apparent in the winter season when snow usually melts near the
cave mouth or small clefts leading to the surface (in the speleological terminology, so called “greasy
spots”, (see Fig. 16). “Nevertheless the term “greasy spots” is not clearly defined by hot air circulation
due to microbial microbial organic matter decomposition and can be synonymized or at least particularly synonymized with the more general term “ventarols” (see also on the chapter “Subterranean
Microclimates”).
The warmth from the earth’s surface already fails to penetrate into the very deep underground and
the only source which brings warmth to the rocks and their cavities is the geothermal energy. It is supposed that this energy has its origin in the radioactive disintegration of unstable isotopes in the Earthmantle rocks and it leads namely to the generation of a layer of melt in the upper Earth mantle – a so
called astenosphere and then, as a result, to the heating of underground waters, volcanic manifestations, etc. Geothermal energy warms through the Earth’s crust and mantle according to the so-called
geothermal gradient that says by how many centigrade the Earth’s internal temperature rises per 100
metres of depth (SMITH and SHAW 1975).
Just as the subterranean habitats can be segregated into individual parts in relation to the illumination intensity, the zones with a specific temperature pattern can also be delimitated. The thermal
mode of the earth’s surface show both circadial (day/night) and circannual (summer/winter) changes.
The temperature changes between day and night several cm to dm under the surface start to coincide, the same takes place in the depths in the order of tens of metres with temperature differences
between summer and winter, and the subterranean habitats acquire a seemingly stable climate. This
near-surface zone with measurable, periodically repeating changes within one year, which correspond
with a mild delay to the on-surface climate behaviour, is referred to as a so-called heterothermic zone.
Below this zone, in the depths reaching to the order of hundreds of metres, lies a so-called homothermic unsaturated zone. This is typical of a stable temperature course within a year. What already
manifests here is the geothermal gradient that varies within a range of 0.3 and 0.6 °C/100 m for reasons of the seepage of surface moisture.
Fig. 17. Circulation of air and water in subterranean habitats influence temperature distribution and influence
of the geothermal energy. Free interpretation of JEANNIN et al. (1998).
32
However, as discovered by STOEVA and STOEV (2005) on the example of long-term temperature
measurements in Bulgarian caves, even such depths allow observation of the temperature variations
that show a long-term but still regular cyclicity. It is periodical variation of the average annual temperature within the range in the order of 1°C while these variations have a frequency of about 10-11
years. Authors of the contribution correlate these repeated variations demonstrably with the 11-year
occurrence cycle of sun spots as well as with the changes in the geomagnetic activity of the Earth.
However, the authors do not discuss a possible influencing mechanism of these factors in detail.
The low limit of the heterothermic zone depends largely on average annual temperature of a specific spot
on the Earth while the low limit of homothermic non-saturated zone is given by the level of underground
water, quantity and character of tectonic faults (fault, clefts, etc.), thermal conductivity of the rocks, etc.
What is already located in the area below the underground water level (in the so-called aquifer) is the socalled homothermic saturated zone where the geothermal gradient attains a value of 2°C and then, gradually, the constant values of 3°C/100 m (BADINO 2005, JEANNIN et al. 1998) (see Fig. 17).
In measuring the temperature in subterranean habitats it is important to pay attention to two characteristics – air temperature and rock surface temperature. The simplest way to measure the air temperature is by
using a thermometer. There is a series of designs (liquid, bimetallic, gas, etc. types) nevertheless, what is
more important than the used medium with known thermal expansion is the condition that such thermometer features the reading recorder function.
What is sufficient for reference measurement is the so-called maximum-minimum (or extreme) thermometer, which is fitted with a U-letter shaped glass tube whose both branches contain a moving rider displaced by the liquid extending in the tube (see Fig. 18.1). If the temperature rises, the right mercury column
displaces the relevant rider upwards. The right mercury column goes down with decreasing temperature;
however, the rider remains in its original position to indicate, with its lower edge, the attained temperature.
The right mercury column goes down constantly while the left column goes up along with the other rider,
which indicates, with its lower edge, the attained temperature. The thermometer thus always indicates three
temperatures at one moment – the maximum and minimum temperature for the entire measuring period
and, at the same time, the actual temperature. After the maximum and minimum temperatures have been
read, the thermometer requires resetting each time, i.e. pushing the rider to the actual temperature. This
resetting is performed with the magnet or the button located in the apparatus’ centre and the thermometer
is ready to take measurements in the next time period (See Fig. 19).
So-called dataloggers are very convenient (see Fig. 18.2, and Fig. 18.3). They are digital instruments
which enable continuous measuring of temperatures at the chosen time interval, instruments that are
equipped with a memory for storage of readings. The data from dataloggers can be easily uploaded to a
computer using software (see Fig. 20). These dataloggers were used during 2007 – 2011 to measure temperatures in the karstic caves of the Czech Republic within the VaV Project of the Ministry of the Environment titled “Determination of Cave Microclimate’s Dependence on External Climatic Conditions in the
Accessed Caves of the CR” (HROMAS et al. 2009).
The rock surface temperature is usually measured using so-called radiation thermometers. They operate
on the principle of measuring the infrared radiation emitted by the examined material (a rock, in this case).
Radiation thermometers are usually equipped with a laser locator that facilitates aiming at the measured
point on the surface (see Fig. 18.4). Use can be made of the surface-temperature contact path indicators
with two electrodes fitted with a thermocouple.
In the case that a layer of the organic or at least loamy erosional sediment is present in subterranean
habitats, temperature can be measured even inside of this material. A suitable type of instrument is the socalled puncture thermometer that features a measuring element which can be inserted under the sediment’s
surface. A frequently used type of puncture thermometer is a gauge with resistance sensor operating upon
the dependence of material’s electrical resistance on temperature. A further possibility is employment of,
e.g., a bimetallic thermometer, etc. (JENČÍK and VOLF 2003)
33
Fig. 18. One of the possible designs of a maximum-minimum thermometer, suitable for reference measurements
of temperature in the subterranean habitats. 2, 3 - Dataloggers providing for the continuous and long-term measurement of humidity and temperature. 4 – Digital thermo-couple thermometer. Photo from the authors’ archive.
Fig. 19. Temperature behaviour inside the Velká Jeskyně skřítků Cave (Děčín District), measured with a creosot-filled maximum-minimum thermometer (POKORNÝ and HOLEC, not published).
34
Fig. 20. Temperature changes inside and aboveground the Velká Jeskyně skřítků Cave (Děčín District), measured with digital datalogger OMEGA® OM-43 with the electric resistence sensor (POKORNÝ and HOLEC, not
published).
Humidity
Humidity is a physical quantity representing one of the basic properties of the air. Air humidity
specifies what amount of water in gaseous form (water vapours) is contained in specific air volume.
The amount of water vapour is very variable in terms of time and space.
Two basic, mutually associated quantities are distinguished – absolute humidity and relative humidity. The absolute air humidity usually expressed as Φ specifies the weight of water vapour contained
in the air volume unit that is ordinarily given in grams of water vapour per cubic metre of air. The
volume of water vapour in air is limited. The air becomes saturated with water vapour, no longer receiving further humidity – water starts to condensate. Specific value of the absolute humidity depends
mainly on temperature and it grows with rising temperature. However, it is important to say that absolute humidity fails to describe to what extent the air is dry or wet. This information is contained in
the relative air humidity.
The basic principle to measure the absolute humidity of air is to capture water vapour physically
from the predetermined volume of air. One possibility is the employment of a so-called gravimetric
hygrometer. This is filled with a substance capable of integrating water molecules into its structure
(e.g. calcium chloride). After the passage of air through the hygroscopic medium, water is absorbed
and thereby the medium weight increases. The absolute air humidity is then calculated as a weight ratio of the trapped water and the volume of aspirated air. However, this measurement is time-demanding and it is predominantly used to verify and calibrate other instruments (JENČÍK and VOLF 2003).
What is most frequently measured in practice is relative humidity – φ, which indicates the ratio
between the actual and maximum saturated content of water in the air. It is specified in percentage.
Thus, relative humidity expresses percentage saturation of dry air with water vapour at a given temperature. It is apparent that the dry air has a relative humidity of 0 % and the air saturated with water
vapour 100% humidity then.
The relative humidity measuring may employ the principle when a convenient solid substance alters
its properties due to the influence of adsorbing water (e.g. electrical resistance, length, etc.). The hygrometer most commonly used in practice that operates on this principle is a so-called expansion hygrometer. The fact is employed here that degreased human hair changes its length with changing relative humidity from 0 % to 100 % by 2.5-3 %. Linear expansion of the hair can be converted by simple
leverage into the indicator placed on a calibrated graduation. The advantage of these hair expansion
hygrometers is their simple structure, mechanical principle that requires no electronic components,
35
relatively high accuracy (error is about 3 %) and also the fact that these hygrometers can operate even
at temperatures below freezing point (see Fig. 22.1).
Resistance hygrometers already represent electronic instruments while, just like in the case of temperature measuring, the humidity measuring dataloggers provide for performing long-term continuous measurements and, at the same time, saving the readings to the internal memory.
In the electronic resistance hygrometer, the relative humidity is calculated from the amount of adsorbed water on a convenient sensor – usually a ceramic plate coated with a layer of ion salt (e.g. lithium chloride) and with connected electrodes, with ceramic used in this phase to measure the amount
of adsorbed water on the basis of changing electrical resistance as a result of the water contained in
the sensor (see Fig. 21). The advantage of these hygrometers is very high accuracy (tenths of %).
A structurally similar device is the capacity hygrometer where the moisture sensor, composed of a
dielectric capacitor that is formed by a slight layer of material that adsorbs moisture from the ambient
environment, measures not only the change in electrical resistance but in capacity (= impedance) as
well (TESAŘ 2009).
The underground generally represents a
very specific environment whose microclimate is typical of a series of dissimilarities as
against closed spaces of anthropogenic origin
as well as the open. The previous chapter already mentioned the stable or at least metastable character of temperatures, and a further characteristic is high air humidity being
frequently in the vicinity of 100 %, and even
100% saturation of air with water vapours is
not an exception, which leads to the condensation of liquid water on the rock surface inside
the subterranean spaces. This fact imposes
high demands on the quality of equipment
which can be used to measure the humidity in
subterranean spaces.
Absolutely unsuitable examples include resistance hygrometers the sensor become wet
with seam up, which renders the sensor temporarily inoperative so the instrument must be
dried before further measurements. Negative
experience with the application of resistance
hygrometers is substantiated by, for example,
Fig. 21. Resistance sensor of the electric hygrometer – daPOKORNÝ and HOLEC (2009) on the extalogger OMEGA® OM-43. Photo from the authors’ archiample when measurements were taken in the
ve.
environment of neovolcanic caves of the Ústí
nad Labem region.
Substantially more convenient instruments for performing measuring in the environment with high
humidity include the capacity hygrometers the sensor of which is substantially more resistant to water
condensation (TESAŘ 2009). However, as alleged by HROMAS et al. (2009), the capacity sensors of
standard design are not totally suitable either.
Just like in the case of temperature, also humidity necessitates that, in the subterranean habitats,
a line is drawn between the air humidity and the sediment humidity (if sediment is present in the
specific locality). If we intend to measure the sediment humidity, we can apply the procedure used to
36
measure soil humidity. This consists in taking a sediment specimen of a known weight. The specimen
is then dried in a drying plant at 105°C until a constant weight is obtained (drying period of approximately 6 hours) to be consequently re-weighed. The difference between the dry and humid weights
divided by the dry weight is then used for the calculation of so-called weight humidity of the sediment
expressed in %wgt. If we need to know the volume humidity of sediment, a so-called undisturbed specimen must be taken, which is a specimen with the original porosity intact. This can be attained by taking the sediment into a hollow metallic cylinder of a known volume (so called Kopecky cylinder). The
calculation of volume humidity (%vol) then consists in the difference between dry and humid weight
divided by the specimen volume. Naturally, we can measure the sediment (soil) humidity e.g. using
the resistant hygrometer directly in the subterranean (see Fig. 22.2).
What is also important for the characteristic of subterranean habitats in relation to the organisms,
apart from the actual sediment’s humidity, is the observation of in what form the water is present in
the sediment. It is predominantly so-called hygroscopic water (water percent contained in the soil
dried in the air, maintained by adsorbing forces), capillary water (water retained in the sediment
by capillary forces), and gravitational water (water running through the pores in the sediment by
gravity). Several of the above-stated forms are usually detected together by means of, for example,
maximum water capacity – the amount of water that the water-saturated soil is capable of retaining,
total water potential – static availability of water in sediment for the plants (so called pF curves), etc.
Fig. 22. 1 - Hair hygrometer, also combined in this case with the bimetallic thermometer. 2 - Puncture resistance hygrometer allowing simultaneous temperature measuring. Photo from the authors’ archive.
37
(HORÁČEK et al. 1994).
Organic matter in subterranean habitats
Organic matter is a source of energy and the building block for most organisms, but it can also
provide a habitat for these organisms. The nature of organic matter and the stability of the processes
associated with the transformation of organic matter depend on the properties of the surrounding
environment. The basic crucial factors are particularly temperature, pH, quantity of water, soil granularity and texture, etc. Organic matter itself also plays a part in changes in the environment of organisms (e.g. through the absorption of nutrients, water and air conditions in the soil, the movement of
substances in the soil, pH). These organisms then again play a role in changing the environment in
which they live.
Organisms partly break down the organic portion of the soil, and also partly help to create new
matter; this particularly includes the actual bodies of the organisms and the waste products of their
metabolism. Excrements of soil invertebrates are very important in the formation of the soil microstructure. Organic matter is therefore continually interacting with organisms and so can be considered
to be one of the basic factors in the environment of these organisms. The study of organic matter is
relatively difficult due to the volume of feedback. Despite this, a number of chemical, physicochemical and biological methods have been developed to quantify or determine the quality of organic matter. These are described in many textbooks on soil biology and other treatises. The aim is therefore
particularly to highlight the importance of organic matter as a factor in the environment of organisms
and to describe certain means of measuring organic matter, either direct approaches, where the actual
quantity and quality are measured, or indirect methods, where we ascertain the organisms, for example (quantity, species spectrum, etc.), and their activity, and this can indicate the nature and volume
of organic matter.
Due to the absence of primary producers, except for a few chemosynthetic autotrophic bacteria (chemosynthetic bacteria used sulphur and iron as a donor electron – i.e. CAUMARTIN 1963, CULVER
and PIPAN 2009), underground areas and the organisms found there are mostly dependent on organic
matter supplied from the outside. DUDICH (1932) designates caves based solely on the movement
of nutrients (and thus also organic matter) from outside as monotrophic, as opposed to amphitrophic
caves, where the source comes from the external environment and is its own, originating from the
cave, i.e. autochthonous. However, Dudich’s terminology is not widely used in the literature. Organic
matter generally finds its way underground more or less in the form of fluvial deposits, underflows, or
together with organisms in the form of their bodies (bats, butterflies, crickets, and the dead remains of
such animals, etc.). In places, for example in the form of bat guano or in depressions formed by rains
carrying organic matter, it can accumulate in greater quantities. The nature and quantity of organic
matter in subterranean habitats thus depends on the nature of the entrance hole, or holes – particularly
on the area and frequency of the holes, as well as on the geometry of the passageways through which
the organic matter gets underground, on the nature of the surrounding land as regards its susceptibility to erosion, the quantity and nature of organic matter around the entrance leading below ground,
the attractiveness of the subterranean habitat for the organisms, etc.
Despite the fact that in these texts we describe the terrestrial environment, it is clear that the division into terrestrial and aquatic environments is actually very formal and imprecise. Many organisms
in caves are bound to a semi-aquatic environment, or one with high levels of relative humidity, or
are organisms wholly bound to an aquatic environment. Real aquatic fauna is a normal part of the
soil micro-environment. Therefore at this point we should also mention the following way in which
organic matter can get into subterranean habitats. This involves the transmission of organic matter
dissolved in water, which infiltrates the soil and trickles or leaks underground and in this form is con38
ditional for a specific form of life. In some caves or parts of caves dissolved organic matter may be the
only source. SKET (2004) defines caves with permanently soaking-wet walls as hygropetric caves.
This term is not conditional upon the presence of organic matter in water, although it may be present.
For practical purposes BOT and BENITES (2005) divide organic matter in the soil into aboveground matter and below-ground matter. According to this system, the above-ground organic fraction
contains the residues of plants and animals. Below-ground organic matter consists of living animals
and microflora, the partially decomposed bodies of plants and animals, and humic substances. The organic portion of soil is generally divided up into the live element and the dead element. The live element consists of soil organisms (edaphon), while the dead element comprises organic matter formed
in the soil when these organisms die off.
There are many possible approaches and methods for determining dead organic matter in the soil
and soil organisms. Animals which are associated with subterranean habitats are covered in separate
sections – partly in the section where certain methods used to detect them are described, and partly in
the overview of organisms, which is also obviously just a selection. In these sections we look at easily
available techniques and also at groups of organisms that are easily observable, with the exception of
roundworms (nematodes). However, no reference is made to techniques which are essentially chemical processes used to quantify and determine microorganisms and which require laboratory facilities,
therefore some of these techniques are mentioned briefly here.
The nature of dead organic matter (organisms, the remains of dead organisms, the intermediate
products of their decomposition and humus) can be described using a number of chemical, physical,
physicochemical, and biological methods. An important property of the organic dead portion, or its
humus portion, is particularly the degree of polymeration and humification, its absorption properties,
chemical composition, etc. In the case of organisms we are especially interested in the number of
organisms, biomass, or activity.
Just one example of an indicator of the quality of organic matter is the monitoring of the ratio of
organic carbon to nitrogen (=C:N). The high ratio of organic C:N, e.g. in wood, hinders the microbial
decomposition of organic matter. A lower C:N ratio, e.g. in leaves, is more favourable for decomposition. But not as much as the generally low ratio in animal tissues, where the proportion of tissue carbon to nitrogen is lower than in plants, thus facilitating decomposition. Another way is to determine
the ratio of humic acids and fulvic acids. The quality of the humus and the soil decreases with the
ratio of humic acids.
One of the fundamental methods used to determine the quantity (weight of biomass, number of
colonies) of microorganisms is the technique based on soil incubation and the use of fumigants. A fumigant is a volatile chemical which during incubation breaks down the bodies of microorganisms and
releases the contents of their protoplasm. One substance that is used is chloroform, and the method
is known as “the chloroform fumigation and extraction technique” (e.g. COLEMAN et. al. 2004). If
we determine the quantity of carbon in the soil before and after fumigation, we can then work out the
quantity of microbial carbon, or nitrogen and phosphorus.
Other techniques can also be used to quantify microorganisms or to classify them into groups (e.g.
cultivation methods, determination of phospholipides, molecular biological techniques, using dye and
a microscope to assess microorganisms, etc.), although these often require well-equipped laboratory
facilities and we will not go into those here on principle.
An indirect and relatively easy way of estimating the quantity of microorganisms is the method
based on measurement of respiratory activity, i.e. the release of carbon dioxide. Measurements can
be taken in the field or in the laboratory. A simple field test for determining microbial activity in
soil, which also indicates the quantity and quality of the organic matter in it, is the decomposition of
organic matter in perforated sachets (=litter bags). Either the organic matter available at the locality
39
(autochthonous material) is used, or another material, e.g. cellulose (see Fig. 23).
A good example of a study into relations between animals and organic matter from caves may be
the work of the authors DUCARME and LEBRUN (2004). In their monitoring work the authors expected to find a strong correlation between the quantity of organic matter in cave sediment and the
abundance of acari. They based this logical assumption, which is corroborated by the published data,
on the fact that the organisms’ relationship to organic matter which is food has a positive effect on the
porosity of the sediment, pH, etc., as stated above. However, their observations of the distribution of
acari and organic matter did not confirm this dependence and the authors found only a weak correlation. The authors assume that in the soil acari have less freedom of movement and are therefore more
dependent on the source of organic matter than in the cave, where there is more space available, allowing them greater mobility in searching for sources of organic matter. In another study DUCARME
et al. (2004a) consider the quantity of organic matter as being limiting for mites from deeper within
the soil, although not for mites from a cave where, according to the authors, floods likely had a negative effect. Verification of this hypothesis was complicated by the difficulties
in accessing flooded places, the fact that
floods are unpredictable, and by newlyformed sediments.
Another example comprises works
from the hygropetric caves mentioned
above. SKET (2004) hypothesises that
bound to the damp environment of the
stony surfaces of these caves are specialised species of leptorine beetle of
the genus Cansiliella (Coleoptera: Cholevidae: Leptodirinae) and some others
(occurrence: caves along the Dinaric
and Italo-Dinaric karst between W
Montenegro and NE Italy), which could
live by filtering the fine particles of the
allochthonnous organic material present in the thin film of water covering
the rock. They also did more detailed
research into the food relationships in
these ecosystems and in the environment known as moonmilk. According
to HILL and FORTI (1997), this is a hydrated, spongy to powdery assemblage
of microcrystalline carbonate minerals,
resembling e.g. toothpaste. Only very
small or no quantities of organic matFig. 23. Litter bags with cellulose (Velká Jeskyně Skřítků Cave).
This method is commonly used to determine the speed at which
ter from the surface have been found
organic matter decomposes in laboratory and field conditions. The
in moonmilk. In their study the authors
general principle is that the size of the mesh only permits microimply that, for example, the species
bial decomposition, or colonisation and decomposition through the
Cansiliella servadetii gets the nutrients
action of mesofauna – particularly springtails (Collembola) and
it needs from moonmilk microbial commites (Acari). Photo: Vojtěch Kadavý.
munities, i.e. in a completely different
40
way to other troglobiotic cave beetles and therefore, for entomologists, beetles of the genus Cansiliella are not species which are easily caught using various types of food bait.
The study of the links between subterranean organisms and organic matter is only just beginning.
Study methods, target groups of organisms and the nature of subterranean habitats can all vary. Even
underground areas which are generally accessible to man, i.e. particularly many caves, are often hard
to get into and require climbing skills. In technical terms research by proficient speleologists may be
difficult to carry out, especially if such research involves collecting and transporting large quantities
of sediment from convoluted underground labyrinths.
Atmosphere in subterranean habitats
There are many differences between the atmosphere of subterranean habitats and that of the outdoor
environment. Above all there is an increased concentration of certain gases (CO2, CO, H2S, Rn, etc.),
which is partly caused by the very limited air circulation underground, and partly by chemical processes which result in the production of a particular gas.
The movement of air underground is caused by differences in air densities inside and outside subterranean spaces, where the air density is generally determined by the temperature, humidity and atmospheric pressure. Other important factors are the geometry of the underground area and the incline,
number and size of entrance holes (KOWALCZYK and FROELICH 2010; CIGNA and FORTI 1986).
In general it can be said that air flows very slowly underground, usually at a speed of < 0.1 m.s-1
(CIGNA 2004). This means that it takes a long time for air to be exchanged and some gases with a
density higher than the normal outside atmosphere have the tendency to remain and collect underground.
Carbon dioxide (CO2)
On Earth this gas is created primarily by the chemical decomposition of carbonates in the upper
parts of the lithosphere and also by magmatic, volcanic, post-volcanic and metamorphic processes
(LOWENSTERN 2001). One crucial source is the breakdown of organic substances in the biosphere,
caused particularly by the microbiological degradation of organic matter and the respiration of plant
root systems. Only a small percentage of CO2 enters the atmosphere through the burning of fossil
fuels and the combustion process in general (LAVELLE and SPAIN 2001).
In the Earth’s atmosphere CO2 reaches an average volumetric concentration of 0.03-0.04%vol.. This
gas collects underground and reaches higher concentrations – in the soil, approx. 1%vol. (Lavelle, Spain
2001), in caves generally 0.1-10%vol. (e.g. BATIOT-GUILHE et al. 2007; BALDINI et al. 2006; EK and
GEWELT 1985, etc.). In the caves of the Czech Republic concentrations of CO2 range from 0.12 to
0.49%vol. with the occurrence of local anomalies, such as in the Zbrašov Aragonite Caves, where levels
of 93%vol. have been measured. The figures from caves in the Czech Republic come only from karst
environments; no detailed measurements of CO2 in non-karst caves and other types of subterranean
habitats have yet been taken (HROMAS et al. 2009).
Subterranean, particularly in carbonate rocks containing karst features, CO2 is formed particularly
as it is released from underground waters, which may contain tens of mg.l-1 of dissolved CO2. This is
based on the principle that rainwater is enriched with dissolved CO2 as it passes through the atmosphere and especially by the soil profile to form carbonic acid (see Equation I on p. XY – upravit až
po vysázení textu). When this water comes into contact with carbonate rock, it dissolves the rock, as
although carbonates show minimal solubility in pure water, in acids they are highly soluble.
After entering the cave, where the concentration of CO2 in the air is generally lower and thus
the partial pressure of this gas is lower than in the soil, the CO2 is retroactively released into the
41
atmosphere of the cave. The result is a supersaturated aqueous solution of dissolved carbonate and
re-precipitation of the calcite in the form of speleothems (see Equation II on page 5). In subterranean
habitats the production of CO2 through the condensation of calcite leads to concentrations that on
average range around 1-5%vol. (PŘIBYL et al. 1992).
The concentration of CO2 in subterranean spaces in the mild zones of the northern and southern
Earth hemisphere changes depending on the season. High concentrations are typical for the summer,
when the outside temperature is higher than underground, and this suppresses air circulation. In the
winter the concentration of CO2 tends to be low. Another reason for this seasonal periodicity is the
decline in microbial activity in the soil, meaning that less CO2 is produced during the winter (KOWALCZYK and FROELICH 2010).
It should be pointed out here that while the release of CO2 into the atmosphere of the cave leads to
an increase in speleothems, in certain conditions it may also cause the opposite to happen, i.e. when
water condenses in the cave from the highly humid cave air (100% humidity). This water, which
condenses on the surface of the cave walls, reabsorbs the CO2 and may actually dissolve carbonate
speleothems – this is known as condensation corrosion (BALDINI et al. 2006).
According to BALDINI et al. (2008), another important source of underground CO2 is the activity of microorganisms in karst soils and
in places where organic matter collects
underground, e.g. bat guano, washedup plant detritus, etc. These respiratory
biogenic processes are the cause of CO2
concentrations of 5-10%vol. e.g. in the
Nerja Cave in Spain or Les GrandesCombes in France (VADILLO et al.
2010, BATIOT-GUILHE et al. 2007).
We know of much higher concentrations, for example those from Zbrašov
Aragonite Caves in the Czech Republic
are caused by emanations of juvenile
gas leaking down from the upper layers
of the lithosphere along faults on the dividing line between the Carpathian Flysch and the Bohemian Massif (CÍLEK
and ŠMEJKAL 1986).
In caves that are open to the public,
the presence of humans is a significant
source of CO2. HROMAS et al. (2009).
In the Císařská Cave in the Bohemian
Karst, for example, it has been shown
that the presence of 120 people in the
cave doubles the concentration of CO2.
Fig. 24. Hand-held CO2 analyser Crowcon working on the prinAn interesting study is that by JAMES
ciple of infrared spectrometry. Photo from authors’ archive. Handheld CO2 analyser Crowcon working on the principle of infrared
et al. (1998), which deals with the comspectrometry. Photo from authors’ archive.
position of the cave atmosphere in the
Jenolan Caves in Australia. Increased
concentrations of CO2 have been recording in this locality, caused by high visitor figures and the associated exhaust fumes from the nearby car park and busy road.
42
CO2 is a clear, unbreathable odourless gas, which in higher concentrations may leave a weak acidic
taste in the mouth. People experience the first symptoms of poisoning at concentrations of around
1%vol. – tiredness, sweating, and hot flushes. Concentrations exceeding 5%vol. cause headaches, nausea,
and visual disorders. Concentrations higher than 10%vol. result in the impairment of the carbonic balance in the blood and subsequently acidosis, loss of consciousness, and death (SMITH 1997).
As there are a number of industrial processes producing CO2 which could be harmful or fatal
(breweries, wine cellars, silage pits), in the Czech Republic these levels are covered by Governmental
Directive No. 361/2007 Coll., which defines the conditions for health and safety at work. This directive specifies, amongst other things, the maximum permissible concentration of CO2 in the workplace
– 9,000 mg.m-3 (approx. 0.5 %vol.) as the average concentration for the entire duration of the working
time and 45,000 mg.m-3 (approx. 2.5%vol.) as the maximum one-off concentration. These levels are
also recommended for people working underground (old mine works, caves, etc.).
There are a number of methods used to determine the presence of CO2 in subterranean habitats.
The simplest of these is to use the principle that a flame will be extinguished in the absence of oxygen
(and thus logically in the presence of any other gas that replaces it). The mining industry used a gas
indicator lamp, which is extinguished when CO2 levels reach 11%vol.. However, this method is only
rough and very imprecise. More accurate devices are detector tubes, which work on the principle of
a chemical reaction between indication chemicals and the gas in question, in this case CO2. The disadvantage of this method is that these tubes can only be used once, which makes it a costly method
when frequent measurements are needed.
Interferometers are optical devices which determine the components of the air (including CO2) using different refraction indices of light passing through a gaseous environment. The light rays pass
through a system of glass prisms, split partly through a comparison chamber containing pure gas, and
partly through a chamber into which the air to be measured is sucked. The difference in the refraction
of the rays is manifest as a shift in the interference image as seen through the eyepiece (GERŠL and
VITOVJÁK 2003).
The devices that are currently most commonly used are electronic CO2 analysers, which generally
work on the principle of the absorption of a certain light wavelength by the gas being measured (infrared spectrometers), the different head conductivity levels of various types of gas (katharometers),
the ability of gas to oxidise on a metal catalyser (galvanometric sensors), etc. (KABEŠ 2005).
Other potentially hazardous gases
Higher concentrations of other unbreathable or poisonous are very rare in subterranean habitats, but
they do exist. The most hazardous is probably carbon monoxide (CO). This is a non-irritating colourless gas with no taste or odour, which is lighter than air (ρCO = 1.25 kg.m-3, ρair = 1.29 kg.m-3 at 101,325
kPa and 20°C).
Upon inhalation, carbon monoxide enters the circulatory system (through the pulmonary alveoli),
where it binds to the blood pigment haemoglobin much more strongly than oxygen, blocking the distribution of oxygen to the tissues and organs of the body. Symptoms of poisoning are the skin turning brownish-red, followed by coma, spasms, and death. According to Governmental Directive No.
361/2007 Coll., in the Czech Republic the highest permissible concentration is 30 mg.m-3 (0.0025%vol.)
and 150 mg.m-3 (0.013%vol.) as the maximum one-off concentration.
In nature carbon monoxide is present in negligible quantities in the atmosphere, where it is created particularly through the photolysis of carbon dioxide as a result of ultraviolet radiation. Another
source is the imperfect combustion of fossil fuels and biomass. A further major producer of carbon
monoxide is volcanic activity, particularly discharges of volcanic gases). Assuming there is sufficient
humidity and quantities of organic matter, CO is created through the chemical oxidation of organic
43
oxygen in the soil and also in MSS (Mesocavernous Shallow Substratum) (GÖDDE et al. 2000).
The concentration of carbon monoxide in pure natural air is 0.1-0.2 mg.m-3, while this level is rising slightly over the long term. It is degraded from the air as it oxidises into CO2 and also through the
metabolism of certain species of soil bacteria and plants.
Underground, carbon monoxide is produced particularly through the spontaneous thermal decomposition of coal, which cases major problems in deep coal mines; higher concentrations can also be
seen in old mine works where coal was mined in the past (equations III – VI) (SHADLE et al. 2002).
III.
IV.
V.
VI.
C + O2 → CO2 (+ heat)
C + CO2 (+ heat) → 2CO
C + H2O (+ heat) → H2 + CO
C + 2H2→ CH4 (+ heat)
In caves and other types of natural subterranean habitats carbon monoxide is very rare. In exceptional cases it may originate autochthonously – CO emanations have been recorded in the Cueva de
Villa Luz in Mexico, associated with the circulation of thermal waters. In the lower reaches of this
cave CO concentration levels of 55-97 mg.m-3 have been measured (HOSE and PISAROWICZ 1999).
However, a much more common cause of CO underground, or around the entrances to subterranean areas, is human activity. BREGANI et al. (1999) highlight the risk of increased concentrations
of carbon monoxide which may accumulate in caves when speleologists light their way with carbide
lamps. This type of lamp works when gaseous acetylene is released from calcium carbide and then
combusted. If the burner of the lamp is not adjusted properly, it leads to the creation not only of carbon dioxide
and water, but also products from imperfect combustion
– particularly carbon monoxide.
Carbon monoxide produced in a similar manner may
also penetrate underground from outside, such as from
traffic on a road near the entrance leading underground.
Another major hazard is when people camp or build fires
in cave entrances, or use burning torches. Carbon monoxide is also contained in cigarette smoke, which in unventilated subterranean areas may increase the concentration
of this gas.
There are no data about concentrations of CO in Czech
caves, although levels are likely to be very low.
Hydrogen sulphide (H2S) is another gas which in rare
cases may accumulate in higher concentrations underground. This is a colourless gas which is heavier than
air, with an odour of rotten eggs even at very low concentrations. It is highly toxic, and fatal at concentrations
exceeding 0.01%vol.. In the Czech Republic Governmental
Directive No. 361/2007 Coll. sets the highest permissible
concentration of hydrogen sulphide in the workplace at
10 mg.m-3 (0.0007%vol.) and 20 mg.m-3 (0.0014%vol.) as the
maximum one-off concentration.
Underground, hydrogen sulphide tends to originate
autochthonously. Most often it rises up from the deeper
44
Fig. 25. Digital CO analyzer, allows measuring in mg.m-3 or in ppm. Photo from authors’
archive.
layers of the Earth’s crust, particularly in regions which have seen recent volcanic activity. A typical example is the Harghita mountain range in Central Romania. The landscape of this region was
formed by volcanic processes during the Miocene – Pleistocene periods (10 – 0.03 My) and a common
phenomenon here are mofettes – volcanic gas exhalations, often with a higher H2S content. Many
mofettes are discharged into the caves there, such as Peştera Sulfatara, etc., from where BARTI and
VARGA (2004) describe concentrations of hydrogen sulphide reaching as high as 515 mg.m-3. Hydrogen sulphide has not been recorded in any caves in the Czech Republic.
There is an increased concentration of hydrogen sulphide emanating in Cueva de Villa Luz in Mexico, where measurements have shown levels of up to 210 mg.m-3 (HOSE and PISAROWICZ 1999). No
clear explanation has yet been found as to the origin of this gas in the caves there, although reasons
discussed by the authors include the oil deposits a few dozen kilometres away in Villahermosa, or the
volcanically active region of El Chicón, roughly the same distance away. Based on analysis of the 14S
isotope, SPILDE et al. (2005) show that sulphur originate in the reduction of sulphate rocks and the
subsequent scouring by thermal water.
In the specific conditions of subterranean habitats hydrogen sulphide may also result from biological processes. This is conditional upon the influx of organic nutrients and sulphur in the absence of
oxygen. The result is anaerobic sulphuric respiration, typical, for example, for bacteria of the genera
Desulfovibrio and Desulfotomaculum. Something that is much more common than in natural subterranean areas is the biological production of H2S that is typical for drainage conduits, for example,
amongst others (LUPTAKOVA et al.
2009).
Carbon monoxide and sulphur are indentified and measured using the same
methods used for CO2 – i.e. detector
tubes, interferometers or digital gas analysers.
One rather specific gas is radon (Rn).
This is an inert colourless gas with no
taste or odour, the most widespread isotope of which is 222Rn. Radon is a product of the radioactive decay of the uranium series and thanks to sources in the
lithosphere it is gradually released, or
emanated, into the atmosphere. The distribution of radon depends on the lithology of the rock; increased radon content
is particularly associated with igneous
rocks (granites, granodiorites, etc.). As
Fig. 26. Diagram showing the breakdown of the radioisoto222
radon is highly soluble in water, the ocpe Rn into subsidiary products. Free interpretation of KAHN
currence of this gas is often linked with
(2006).
the emergence subterranean waters. Also
associated with this is the increased presence of radon in regions with highly water-permeable sandy
basement (CIGNA 2005).
The amount of radon in the atmosphere is not specified in units of concentration, but considering
the emission of radioactive radiation, specific activity is used – the number of atoms of the radioactive
substance which is transformed in one second relative to the unit of volume or the unit of mass of this
substance. This unit is then Bq.m-3.
45
Radon is not toxic in the chemical sense of the word; the main hazard is posed by its radioactive
properties. When inhaled it can increase the risk of cancer, particularly lung cancer. The great majority of radon isotopes are α-emitters with a short half-life (days-seconds). After the termination of
short-lived isotopes of radon, the risk does not disappear, in fact just the opposite – decay releases
other radioactive and, due to their higher energy, even more hazardous fissionable products (e.g. 218Po,
214
Bi, 214Pb and 214Po) (NAVRÁTIL 2000).
One major factor with a direct proportional influence on the specific activity of radon is the dustiness of the atmosphere, or the concentration of aerosols. The atoms of subsidiary α-emitters in solid
form cling to dust particles and when inhaled may concentrate in the respiratory passages and, due
to their longer life, may seriously impair the DNA structure inside the cells and trigger a cancerous
growth. OSINA et al. (2010) corroborates this fact by referring to the significantly higher number of
chromosomal aberrations in the group made up of employees of the Slovak Caves Administration,
who spend time on a daily basis in caves with increased radon activity, compared with a randomly
selected control group of people working in normal conditions. CRAVEN and SMIT (2006) also came
to a similar conclusion, referring to the increased risk of cancer for speleologists exposed to higher
doses of radiation. CRAVEN and SMIT (2006) also add that this risk is increased approximately tenfold if the speleologist is a smoker.
Dust levels are very low in caves, partly due to the fact that there is only a gradual or even zero
exchange of the inside and outside atmosphere, and partly because dust particles cling to the water
on the damp walls. Another typical feature of caves is minimal air circulation, which is a factor
greatly affecting the specific activity of radon in indirect proportion. The reason for this is the fact
that radon is heavier than air – it is the heaviest gas found in nature – and thus has the tendency to
collect in natural underground cavities, particularly caves. The result of this is that the
more the air circulation drops, the more the
radon activity increases as there is no way for
it to be discharged outside (ROVENSKÁ et
al. 2007).
Nevertheless, as regards radon activity
not even the cave environment is completely
stable, as shown by HROMAS et al. (2009)
based on the monitoring of Czech karst caves.
There, circannual fluctuations of Rn activity
were seen depending on the mutual interchange of the inner and outer atmosphere of
the cave as temperature differences evened
out.
This implies that the unventilated environment of a cave is a locality where there is an
increased likelihood of the risk of radon levels which exceed the limits. This fact is also
reflected in the Czech legislation. Section 87
(b) of Directive No. 307/2002 Coll., on Radiation Protection, designates caves as places
where individuals can be exposed to significantly higher levels of radiation from natuFig. 27. Principle of a simple radon trace detector. Free inral radioactive sources. This obliges caves to
terpretation of KAHN (2006).
monitor radon levels, assuming the cave is
46
classed as a workplace – monitoring is therefore obligatory for caves that are open to the public and
where tourist guides and other employees work. Levels detected range from 135 to 22540 Bq.m-3,
while the highest radiation levels were measured in the Bozkov and Zbrašov dolomite caves (an average of 9,700 Bq.m-3); the lowest levels were detected in the Chýnov Caves and in the Na Špičáku Cave
(an average of 960 Bq.m-3) (e.g. HROMAS et al. 2009, ROVENSKÁ et al. 2007, ZIMÁK and ŠTELCL
2004, BURIAN and ŠTELCL 1990).
Somewhat lower radon activity levels were found in selected karst caves in Slovakia, which is due
to the different structure of the rock and the geological composition of the region. Levels measured
there range from 59 – 11,280 Bq.m-3, with the highest radon activity in the Bystrianska Cave in the
Low Tatra Mountains (an average of 5,785 Bq.m-3) and the Gombasek Cave in the Slovakian Karst
(4,615 Bq.m-3), and the lowest in the Jasovská Cave in the Slovakian Karst (795 Bq.m-3) (VIČANOVÁ
et al. 1997).
Probably the highest radon activity in a natural environment has been measured in Great Britain.
In the Midlands of this island there is the Peak District National Park, which is a karst region formed
in Lower Carboniferous limestone. In the precipitous unventilated Giant’s Hole Cave GUNN et al.
(1991) recorded radon activity of 155,000 Bq.m-3. Very high radon activity levels – 88,060 Bq.m-3
– have also been described by PAPASTEFANOU et al. (1986) from the Petralona Cave in Western
Greece. According to HYLAND and GUNN (1994), radon activity levels in caves around the world
are generally within the range of 450-8,850 Bq.m-3.
Radon activity can be measured using trace detectors, for example, which contain a sensitive film
in a plastic case placed in the site in question. The irradiated places on the film caused by α-particles
are then evaluated. Another possibility is electret dosimeters, which work on the principle of the
discharging of a special, positively-charged material placed in the measuring chamber. Continual detectors filter air containing subsidiary products of radon bound to dust and aerosol particles through
a microfiber filter. The decay products trapped in the filter release γ-radiation, which is detected by
silicon detectors located next to the filter (NAVRÁTIL 2000).
47
Methods of animals’ collection
As it was mentioned above, ecology addresses the relationships of organisms and habitats in a wide
range. We will deal here only certain measuring methods and procedures that can be applied in the
subterranean habitats. Nevertheless, emphasis will be put namely on such methods that can be supplemented with practical examples.
The basic measurement procedures used by ecologists include the observation of the occurrence
influencing factors (this bears relation to the factors which influence birth-rate, mortality, migration,
relationships between organisms, and so on) and the abundance of organisms and, further, monitoring
of the abundance of organisms. As a matter of course, ecologists also deal with further issues such
as, for example, how the abundance of organisms changes with time (e.g. study of succession, study
of the changes in abundances during a day or a year or even longer periods, etc.) and what influences
such changes, the way they change in space, etc. To answer these questions, however, there is the basic
question of how to characterize the habitat and how to count organisms. The measurement examples
of the organism collection methods are specified in the following text.
Direct searching for animals
The main aim of direct or hand
searching is to collect as many species as possible but usually not in
large numbers (HUNT and MILLAR
2001). Unfortunately, this method is
very time consuming and in case of
most taxonomical groups of animals
it can take several hours to record at
least some specimen. Nevertheless
the same or close result can be used
by others in traps based on monitoring methods. Searched small invertebrate animals are usually collected
directly by hand, to tweezers, sucked
by exhaustor, stick on wet brush-pencil etc. (Fig. 28). The collected animal
should be put into plastic vial with the
solution of ca. 80% ethanol. It is important for each vial to be labeled at
the time of collecting.
However, the direct searching is
good for species easy to record and
for endangered species, especially
bats in caves. Details for measuring
with detectors and other information
about bat monitoring in the Czech
Republic can provide Czech Bat Conservation Society (www.ceson.org).
48
Fig. 28. An exhaustor (=aspirator) is a device which is used for collecting of different small terrestrial animals, especially spiders, terrestrial soil crustaceans, insect etc. Different types of exhaustors are
used but principally they are similar. Animals are sucked in by a collector through an opening into a container, which is filled by conservation liquid (usually ethanol) or free and where they are not killed.
The container is connected to a tube on the other end, to which investigated animals are sucked. Illustration from the authors’ archive.
Methods of animals’ collection
Pitfall trapping
The pitfall traps are widely
used traps in terrestrial ecology. There are many variations
of pitfall traps but they are principally similar. All of them are
exhibited by container, which
is sometimes free, sometimes
filled with a preservative (e.g.
formaldehyde, ethanol, propylene glycol etc). Entomologists
often use some attractants, e.g.
beer, cheese etc. to attract invertebrate animals. Their utilization in ecosystem’s study is
disputable because abundances
and species richness can be influenced by animals attracted
from different habitats or with
Fig. 29. The pitfall trap (e.g. plastic cup) with the preservative solution is
unknown distance from subtercommonly used above the ground but it can be also used in subterranean
ranean habitats. On the other
habitats. Rare animals can become extinct while using this technique, espehand, especially in subterracially when it is left out for a long period. Photo from the authors’ archive.
nean habitats where animals
are rare, their usage can be convenient or fundamental (HUNT and MILLAR 2001). Nevertheless,
different specialists have often self-experience with improvement of different baits or baited pitfall
traps. In the same case only baits without traps can be used. For winter trapping antifreeze alcohol
mixed liquid must be used outside caves only. For increasing of wettability several drops of detergent
are usually added (Fig. 29).
Pitfall traps cannot be modified only by their content but also by their general features. For example
RŮŽIČKA (1982) has equipped traps with a board. Mentioned improvement can be valuable, for example, for surface or subterranean trapping among stones on screes, where the gripping is difficult.
SCHLICK-STEINER and STEINER (2000) have modified pitfall trapping for subterranean investigation of animals in different depths of soil or stony accumulations. Traps – cups with preservatives are
situated inside a perforated tube that is vertically located in soil or accumulation.
Core sampling of soil, sediments or litter and fauna extraction
Sampling of inhabitants in soil and in sediments or naturally sampling for measurement of abiotic
characteristics of soil sometimes need to use coring tools. For example, the O’Connor split corer can
be used. Soil fauna can be sorted from soil monolith manually, or it can be prepared with higher efficiency using heat and dryness extraction in e.g. Tullgren funnels. This method use special arrangement for animal isolation from soil. Material (sediments, litter etc.) is placed on a piece of gauze and
it is given to a funnel. The funnel with material is placed under light, usually light bulb. Soil animals
try to move downwards away from heat and light and crawl deep into material. Under the funnel there
is situated a cup containing conservation, e.g. alcohol. The soil dries usually several weeks (for more
details see SUTHERLAND 2006, COLEMAN et al. 2004, HENDERSON 2003) (Fig. 30).
49
Fig. 30. An schematized example of the Tullgren funnels. Illustration from the authors’ archive.
50
Ecological and Evolutionary Classifications
The goal of various classifications is to organise knowledge into a well-arranged and practically
usable system. It is often a simplification at the expense of precision. In practice, then, we can either
use more simple, but sometimes less precise classification, or more detailed classification, which often
demands more information about the classified object, on the other hand. The situation is not different
for the classification of cave organisms.
For the classification of cave, or cavernicolous (Latin: caverna – “cavern“, colere – “to inhabit“)
species, the simplest approach seems to be to divide them into cave and non-cave organisms, or into
those that cannot be unambiguously classified as one of these two categories. Traditionally troglobionts, troglophiles and trogloxenes are distinguished. With regards to the often less well-known
bionomics of various species and a variety of classifications where the meanings of the same terms
overlap or even new terms are used, the use of classifications is rather problematic. In contemporary
literature, the exact meaning of the term according to the sense in which the author meant it or in
which it is used should always be mentioned.
Troglobionts (Greek: trogle – “cave“, bioteuo – “dwelling“)
Troglobionts are real cave animals, or more simply – just cave animals. The presence of troglomorphisms (morfological adaptation for life in subterranean habitats) plays an important role in the definition of troglobionts in speleobiological literature. Other specific characteristics (e.g. metabolic rate,
presence of humidity detecting sensory structures, size of appendages and size of sensoric organs
etc.) are unfortunately often unknown for particular subterranean species. In the case of soil species,
the term euedamorphism can be used , althought is the less frequently used term nowadays (see e.g.
DUCARME et al. 2004b). Problems concerned with adaptation concepts are mentioned more detailed
e.g. CULVER and PIPAN (2009).
CHRISTIANSEN (1962) defined:
a. troglomorphic taxa, i.e. highly modified forms when compared to epigeic taxa,
b. ambimorphic taxa, partly modified and similar to epigeic taxa,
c. epimorphic taxa, i.e. dwelling and reproducing in cave, but without morphological modifications,
d. trogloxenous taxa, i.e. taxa in random caves or exploiting caves, but without specific adaptations.
Morphological adaptations as a criterion for
the delimitation of spider species dwelling in
underground environment of stony accumulations were used by RŮŽIČKA (2001).
Some species, however, do not manifest
characteristic morphological properties, or
these properties are manifested also by species living outside the caves (see e.g. CULVER 1982, MATILE 1970). It is not always
important if the biotopes are “similar” (e.g.
in the sense of similar humidity, temperature,
dynamics of environment changes, etc.) or
not. For instance SKET (2008) reports the example of the freshwater crustaceans (shrimps)
Fig. 31. Example of the reduction of the eyes at the spiders
Porrhomma egeria. The independent reduction of individual
eyes and variability of the reduction are interesting. Free
interpretation of SANOCKA (1982).
51
Niphargus where the reduction of eyes is not dependent on life under the ground. Moreover, differences can be also found in animal behaviour, their physiological processes, etc. On this account, a
purely morphological approach is rejected by some authors for the definition of real cave animals. In
addition, for etymological reasons, the term troglomorphy should be substituted by the term troglobiomorphy according to JUBERTHIE and DECU (1994). Many authors respect this, although other
authors use more traditionally established term. CHRISTIANSEN (1992, 1962) distinguished sensu
stricto troglomorphisms and analogically sensu lato troglomorphisms based on all responses of organism to the environment.
Whatever the criterion selected, i.e. whether it’s morphological, physiological or other, we always
look for the properties that we expect in the organism and only on their basis we determine whether
it is a cave animal or not. This means that we tend to determine its relationship to caves indirectly.
It can be true that a given species or population is permanently connected with caves, i.e. it is a real
cave organism, but the time when selection operates is short and no obvious and expected adaptations
occur, or that these properties are not influenced by any selection in this case. Many cave species with
unknown surface populations are categorised as troglophiles due to the fact that they show only indistinct signs of regressive evolution (pigmentation loss, eye reduction, etc.) (CULVER 1982).
The intuitive explanation of adaptations “apparent at first sight” may, with some problematic exceptions, meet another problem. As was mentioned above, some cave species have long limbs “because”
there is little light in the cave, so the food cannot be seen and must be felt for, which is probably easier
with long limbs than with short ones and this fact, from the point of view of evolution, is so significant that this selection really occurs. But it is based on the general presumption of the lack of food in
caves. Locally, however, there can be enough food, e.g. in the environment of bat faeces. The generally used term of an abundance of food is then also problematic. Theoretically, this adaptation should
be advantageous only in dark and food-poor caves. This means that even short leg forms should be
considered troglobionts. It could also be the problem of various morphometric measurements that
underground species often have larger bodies. For example DUCARME et al. (2004b) confirmed that
the cave mites he studied generally have larger bodies and at the same time greater length of some
limbs, trichobothria, etc. than soil species. After taking into account the lengths of these mentioned
dimensions relative to body length, the differences between cave and soil species disappeared. The
result from this approach was that only one sign remained from several possible signs. Other evidence
for relative character of some adaptations and examples of inappropriate comparisons in some morphometric studies mentioned i.g. CULVER (1982).
Troglophiles (Greek: filéo “to love“)
In the sense of the classification proposed by Schinner-Racovitza this is a transitional group between entirely cave-dwelling species (troglobionts sensu SCHINNER 1854 and RACOVITZA 1907)
and species dwelling in the outer environment that only occasionally visit caves (traditional trogloxenes sensu SCHINNER 1854 and RACOVITZA 1907). According to these authors, troglophiles are
regularly found in caves with daylight rahter than outside them. Other classifications have been offered by many authors, among which it is important to mention RUFFO (1957), PAVAN (1944), DUDICH (1932), etc. For example, according to the Italian author PAVAN (1944), troglophiles are those
species that prefer the underground environment, but are not adapted to it, i.e. do not show special
adaptations. Another Italian author, RUFFO (1957), defines so-called eutroglophiles, species able to
live externally, but preferring to live underground and reproduce here as well, i.e. they are surface
species able to form more or less stable populations underground. However, this is what BARR (1968)
considers “troglophiles”. In the traditional system of Italian speleobiologists, in addition to eutroglophiles, the group of subtroglophiles is used. These are the species that exploit the cave environment,
52
Fig. 32. A possible schematic expression of the classification of cave animals (organisms). Illustration from the authors’ archive.
but the non-cave environment is used by them for at least one life function (food, reproduction etc.).
Trogloxenes (Greek: xénos – “foreign“)
In the sense of the traditional classification proposed by Schinner-Racovitza (Schinner-Racovitza’s
system), these are “foreign to caves” animals, i.e. species that get into a cave more or less randomly.
In spite of these facts this is a group over which there are the smallest conflicts, even here we face
here nomenclature troubles. Whereas the above mentioned Italian authors use the term trogloxene in
the sense of Schinner-Racovitza’s system, BARR (1968) did not use this term in the sense of subtroglophiles and instead of the term trogloxene used the term “accidental”. He thus introduces a category
which is underground even less than trogloxenes.
In principal the same classification, based on a classification according to the environment, is also the
recently published one proposed by SKET (2008) with the following categories: troglobionts, eutroglophiles, subtroglophiles and trogloxenes. With regard to variety of terminology and classification problems, the use of the chosen unified scale (e.g. the one proposed by Sket) would facilitate the orientation
in the problem.
Troglobionts (troglobites, eutroglobiontes, obbligato troglophiles, in water environment so-called stygobionts, etc.) should, according to the author, be the most strongly related species to the underground
environment. A population or subspecies that forms a part of an eutroglophilous species can even be
troglobiotic.
Eutroglophiles (partial troglophiles, facultative troglophiles and hemitroglobionts) are species which
are essentially surface ones, but they are able to maintain constant underground populations that can be
troglobiont.
Subtroglophiles (partial troglophiles, pseudotroglobionts and trogloxenes) are inclined to live (permanently or temporarily) underground, but they use the outer environment for some biological functions
(daily – food; seasonally or throughout their life – reproduction).
Trogloxenes (“accidental”, eutrogloxene, tychotroglobiont) are the group of species that sporadically
occurs in the underground environment and is not able to form underground populations (SKET 2008).
53
Tab. II. Clasifications of ecological categories (according to CULVER and PIPAN 2009)
54
Category
Definition
Synonyms
Troglobiont
Obligate, permanent resident of subterranean habitats
Troglobite
Eutroglophile
Facultative, permanent resident of
subterranean habitats
Troglophile
Subtroglophile
Obligate or facultative resident of
subterranean habitats but associated
with surface habitats for some part of
its life cycle
Troglophile
Trogloxene
Trogloxene
Sporadically in subterranean habitats
appearing species
Accidental
Trogloxene
The origin and emergence of subterranean terrestrial fauna
The origin and emergence of subterranean terrestrial fauna
Soil is generally considered the basic environment from which terrestrial animals get underground
(VANDEL 1965). However, the parts of the soil that have been investigated the most are only those
top layers right below the surface. Just a few dozen centimetres lower lurks the biota of the soil, in
general referred to as the edaphone, the focus of soil biologists’ attention. Yet life here is known to be
relatively rich (e.g. DUCARME et al. 2004a).
In the Czech Republic the vertical composition of fauna of stony screes, which is also one of the habitats considered for subterranean settlement (see e.g. GULIČKA 1978) and the structure of subterranean stony biotopes has been covered in general particularly by V. Růžička in his studies focusing on
spiders (see e.g. RŮŽIČKA and ZACHARDA 2009, RŮŽIČKA 1999, RŮŽIČKA 1993). LAŠKA et
al. (2008) has explored the organisms of several selected soil profiles, Superficial Underground Compartments (SUC) and caves by various groups of invertebrates, to a depth of 95 cm, using specially
modified land traps (SCHLICK-STEINER and STEINER 2000).
Another source of subterranean species may be the burrows of mammals, mountainous habitats,
especially on the edges of glaciers, moist mossy habitats, etc. This is why species that we also find
living underground, besides cave species, are those bound to a cold or wet climate, i.e. in mountainous
or northern regions.
Besides questions concerning the origin of the habitat, speleobiologists are also interested in the
development of cave fauna in the broader geographic scale and sense. Here, the development of Czech
fauna tends to be associated with the influence of the ice ages in Quaternary. The generally accepted
theory explaining the relative scarcity of cave life forms in this country is the extreme climate here
during the ice ages. In the aftermath of the iceberg, although there was again the possibility that our
subterranean areas would be colonised from regions with a more favourable climate, it is assumed
that the time needed to colonise the caves from the end of the last ice age to the present day was too
short. According to this theory, the species diversity of subterranean life forms would not have had
sufficient time to develop into the form we assume from the time prior to the coming of the ice ages.
The greater distance between glaciers in the Quaternary is also seen as the reason why, for example,
the Southern European karst is seen as one of the most diverse locations in the world in terms of troglobiont subterranean fauna.
However, not all cave systems in warm regions are species-rich. On the contrary, it seems that
tropical regions, i.e. regions which are generally the most varied on Earth as regards the diversity of
species living there, tend to be less diverse in terms of specialised troglobiont subterranean fauna in
comparison, for example, with the karst regions of Southern Europe; in fact it can even be said that
species-rich caves are rare (e.g. DEHARVENG 2005). According to CULVER and SKET (2000), the
rarity of species-rich caves in the tropics is a mystery that has not yet been resolved.
It must be said, however, that there is considerable dispute over the detailed nature of the effect that
the ice ages had on our fauna. Earlier mention was made of the assumption that at that time something
occurred that had a powerful influence on fauna as a whole, and in the case of cave fauna led to its
complete destruction. Some works, however (see e.g. LOŽEK and HORÁČEK 2004, HORÁČEK and
LOŽEK 2004) imply that life during the ice ages and in caves could have been more diverse than is
generally expected, yet this does not explain the low species diversity seen today.
However the cave fauna in this country developed during the Tertiary before the coming of the ice
ages, i.e. whether it was well developed and then decimated or only partially decimated, or something
completely different, in certain respects at least, due to the periglacial climate, a climate influenced
by the vicinity of a glacier (LOŽEK 1973), there was a demonstrable increase in the diversity of our
subterranean fauna. Species from cold periods, known as glacial relicts, have been preserved to the
present day in certain subterranean habitats. This was caused by the cold temperature shock and in55
tense process of frost weathering, which in the long term led to the creation of many different types
of natural subterranean non-karst spaces, from small fissure systems to cavities which match the definition of a cave. In many cases rock blocks disturbed by the frosts caused landslides, creating both
shallow and deep subterranean systems which, due to the absence of frost penetration and greater
temperature stability, were probably soon colonised.
Fig. 33. Distribution of the most species-rich underground localities, known as underground biodiversity hot spots.
Free interpretation of CULVER and SKET (2000).
56
Survey of selected groups of organisms
Although caves and other subterranean environment represent for many groups of organisms extreme habitat, all typical above-ground organisms live here, too. Lack of light limits development of
photosynthetic organisms; nevertheless some of them are able to colonize such environment, where
small amount of light occurs at least.
Algae surrounded by artificial lighting are also characteristic in tourist accessible caves (in the
Czech Republic often German term “lampenflora” is remarked for green organisms dependent on artificial cave lighting). Some species of bryoflora are also characterized for the entrance parts of caves
and similar entrance to subterranean habitats (e.g. Conocephalum conicum, Leptobryum pyriforme,
Tetrodontium brownianum or T. repandum) (SÁDLO 2001). From the Labské pískovce Protected
Landscape Area luminous moss Schistostega pennata is known (WINKELHÖFER 1998). Roots of
trees are also very typical for some caves. In some relatively rare localities roots look like true calcite
stalactites were recorded (Fig. 34). They can be called root stalactites; concerned important literature
was summarized in POKORNÝ and HOLEC (2010).
Studies on microorganisms in subterranean habitats are relative rare (e.g. BASTIAN et al. 2010) and
we will not be interested in these different groups of organisms, although they are very important,
too. Their importance lies in interactions with other organisms, which means they exhibit important
part of food web, decomposed organic matter. Others microorganisms can be also human or animals´
pathogens. For example of some microscopic fungi (see Fig. 35 and Fig. 36). Investigation of animals,
especially study of some generally attractive species, eg. beetles or bats, have a long tradition in spe-
Fig. 34. Root stalactites in Velká Jeskyně skřítků Cave near Dobrná (Děčín district) (POKORNÝ and HOLEC 2010).
57
Fig. 35. Insectivorous fungi colonized body of different
insects or other invertebrates. Photo from the authors’
archive.
Fig. 36. Mucor fungus colonized fecal pellets of bats.
leology. List of the characteristic groups with
the short characteristics is described in the text
below.
Nematodes (Nematoda)
Nematodes (Greek: nematos – “thread”) are
very abundant animals occupying different
habitats. Some members are known as important parasites but there also exist many, usually
microscopic (0.2-3 mm), free-living species. It
is known about 14 000 species in the word and
about 1/3 from them live in soil (SEGERS and
MARTENS 2005). Free living roundworms inhabit also water and soil sediments. NICHOLAS (2001) has estimated cca. one million
species in the word. In the Czech Republic,
505 free living species are recorded (HÁNĚL
2005), but only one specialist has been systematically interested in free-living soil nematodes
and species richness has been estimated as
Fig. 37. External features of generalized nematodes. Ilhigher.
lustration from the authors’ archive.
Their body is slender and elongate, tapered
at both ends. It is covered by smooth or structured cuticle which maintains hydrostatic pressure and provides mechanical protection. The cuticle is
molted and it is an important common feature of arthropods. Nematodes with arthropods are grouped
by fylogenetics to a group called Ecdysozoa (animals that molted cuticle).
Free living species feeding on organic matter are known as decomposers or as predators. DUMNICKA (2005) distinguishes only 20 described troglobiotic species. There are no published data
58
about cave specialists in the Czech Republic.
Annelids (Annelida)
There is about 10 200 species of annelids in the world (SEGERS, MARTENS 2005) and the best
known members are earthworms. Their external structure is characterized by long segmented body
with typical clitellum that is used for mucous secretion during copulation and cocoon formation.
Earthworms are important burrowers and they are important for soil horizon mixing and soil organic
matter decomposition. About 52 earthworm species are recorded in the Czech Republic, some of
them live also in caves but those species are not cave specialists (PIŽL 2008). Frequently observed
earthworm species are e.g. Dendrodrilus rubidus, Aporrectodea caliginosa, A. rosea, Dendrobaena
octaedra, Octolasion lacteum (PIŽL 2008). Some species can form troglobiotic populations. Popula-
Fig. 38. External features of generalized Annelid. Illustration from the authors’ archive.
tion of Aporrectodea rosea and Dendrodrilus rubidus are known from some Moravian karst caves
(MLEJNEK and TAJOVSKÝ 2008), where their fecal pellets deposits are very interesting.
Molluscs (Mollusca)
Molluscs are not externally segmented and
their body is usually covered by a shell, although
shell of some species is reduced. In the Czech
Republic we can recognize two molluscs’ groups.
Bivalves (Bivalvia) are not the subject of our interest in this text aimed on terrestrial habitats
because their members live in water only. Gastropods (Gastropoda) have some species closely
related with caves, although those live outside
the ecosystems, too, and are not specialized.
There is about 80 – 135 000 especially marine
species in the word (SEGERS and MARTENS
2005). In the Czech Republic live 240 prevailFig. 39. Oxychilus glaber from the Valkeřická jeskyňka
ingly terrestrial species (BERAN et al. 2005).
Cavelet (Děčín district). Photo from the authors’ archiFor example some Oxychilus species, e.g. O.
ve.
cellarius, O. glaber or Limax cireneoniger are
also known from deeper caves. Those species are
not cave or subterranean specialists but their populations can specialize to subterranean habitat. For
example above-ground populations of Oxychilus cellarius are known as litter feeders, but according
to JEUNIAUX and TERCAFS (1961) cave populations are predators feeding on different hibernating
or aestivating invertebrates’ fauna.
59
Fig. 40. Metellina merianae is very common spider of
many subterranean habitats including artificial ones,
e.g. old mines (Photo in artificial gallery on Deblik hill
in České středohoří Mts.). Photo from the authors’ archive.
Fig. 41. Nesticus celullanus from the cave on the Špičák
hill (Krušné hory Mts.). Photo from the authors’ archive.
Spiders and other related groups
Spiders, mites, harvestmens and false scorpions are four members of Arachnida group that live
in the Czech Republic and they are grouped with other related animals with chelicerae to Chelicerata group. Chelicerates (Greek: chele – “claw”, + ata – plural suffix) have two tagmata – prosoma
(cephalothorax) and opisthosoma. Prosoma is a sensory, locomotors and feeding part of body. To the
prosoma paired appendages are attached – one pair of chelicerae, one pair of pedipalps and usually
four pairs of walking appendages – legs (all living in the Czech Republic have 4 pairs of legs, except
young mites, where only 3 pairs of legs). Chelicerae are usually used for feeding. External body
features differ and we will describe only four mentioned groups. In our country also Scorpiones are
well-known, but their records were unique. Nevertheless, some authors have postulated relict population of scorpions living in the Czech Republic (FARKAČ and KRÁL 2005). Also records of one
species of Schizomida group in the Czech Republic were unique and moreover record of this species
was connected only with glasshouse (Sentenská and Líznarová 2010). The Scorpiones and
Schizomida have no importance for subterranean habitats in the Czech Republic.
Spiders (Araneae)
The prosoma bears chelicerae with poison glands. Pedipalps are, in case of males, modified for
sperm transfer. Spiders are also conspicuous for six or eight eyes, although typical cave spiders have
their eyes reduced. Opisthosoma have several conical projections associated with silk glands. Those
are called spinnerets and the imitated silk is used for different activities (ballooning, prey capturing,
communication, etc.). There live more than 850 spiders´ species in the Czech Republic (BUCHAR
and RŮŽIČKA 2002).
The occurrence of 1 mm small pale yellow or ochre colored spiders is connected with deep parts
of caves, debris stones and other, including artificial subterrranean habitats. Most of them belong to
genus Porrhomma and their records are usually rare. Nevertheless, some Porrhomma species are associated with above-ground habitats, too. Porrhomma egeria is the most common subterranean species, although sometimes it is observed above-ground, usually in stony habitats connected through
60
galleries with underground. Pigmented specimen P. egeria is known from above-ground mountain
forest litter, too (RŮŽIČKA 2007).
Several obvious and typical spiders are connected with enters of caves, celeries and mines. The biggest and, for cavers or owners of wet celeries, generally known spider is Meta menardi. Smaller but
still distinct and common are also e.g. Metellina merianae or Nesticus cellulanus.
Harvestmen (Opilionida)
The prosoma is broadly connected with opisthosoma, apart from spider body, where prosoma is
attached to opisthosoma by slender pedicel. Harvestmen have no spinnerets, no silk and no poison
glands. Long legs are typical for most specimens. Harvestmen can be predators but many of them are
omnivorous. 36 species is known in the Czech Republic (BEZDĚČKA 2010).
Specialized subterranean harvestmens are connected with extensive cave regions only and they are
not known in the Czech Republic. Although for extensive cave regions in south Europe, some species
of genus Holoscotolemon are typical, H. jaqueti has been relatively freshly recorded in Slovakia, too.
Some species are conspicuous for their big chelicerae. Ischyropsalis species lives also in stony habitats and are sometimes recorded from caves, too.
False scorpions (Pseudoscorpionida)
A false scorpions (pseudoscorpions) are small, usually in forest litter living organisms. Chelicerae
are small and they are not connected with poison gland. They have long palpal pincers which resemble
scorpions. A poison gland is located in pincer’s fingers. Chelicerae are connected with silk gland.
There is recorded 34 species in the Czech Republic (ŠŤÁHLAVSKÝ 2006). Pseudoscorpions are
predators. Cave or underground specialists are not known in the Czech Republic. Specialized Neobisium slovacum from the Neobisium genus, with many species characterized for subterranean habitats,
has been relatively freshly recorded in Slovakia (DUCHÁČ and MLEJNEK 2000).
Fig. 42. Ischyropsalis hellwigi feeding on some snail (in caves
Oxychilus species are typical). Free interpretation of ARENAZA
(2008).
Fig. 43. Some cave specialists with very
long pedipalps belong for example to genus Neobisium. Illustration from the authors’ archive.
61
Mites (Acari)
Mites, perhaps evolved from various arachnid groups, are very diverse group of animals. Their brief
and clear definition is very difficult. Primary segmentation is reduced and abdominal segmentation
often disappears. Their prosoma and opisthosoma are fused, body is covered with single carapace. It
is usually divided to two regions; head region with the mouth parts which is called the gnathosoma,
and the rest of the body called idiosoma. Chelicerae and pedipalps are variously modified for biting,
anchoring, sucking, piercing. Adults have usually four pairs of legs but some species can have fewer.
Larval stages have a maximum of three pairs of legs. Many of mites are free living (herbivores, scavengers, predators) but there are also many parasites. Diverse are also mites adapted to an aquatic
existence.
Within the subterranean habitats parasites are usually connected with bats, predators and decomposers. They are connected with sediment, especially rich in bat guano. Some parasites can be also
dangerous for human, too. In the Czech Republic mite parasite Argas vespertilionis can transfer encephalitis from bats to human (DUSBÁBEK 1979). Interesting in the Czech Republic is the presence
of predatory mites of Rhagidia gelida, whose existence is connected with cold climate and it is known
from ice holes (ZACHARDA et al. 2005, ZACHARDA 1993). Except the examples of parasites or
predatory mites, free living oribates (order Oribatida) are studied in the Czech Republic, too. STARÝ
(2008) has recorded 106 species of oribatid mites in karst caves. To very important findings also belong troglobiotics Kunstidamaeus lengersdorfi or Pantelozetes cavaticus. Troglophilous rhagiid mites
(Rhagidiidae) are relatively common and obligatory components of cave fauna worldwide. Also in
the Amateurs´ Cave, Moravian Karst, Poecilophysis spelaea, P. wolmsdorfensis, P. wankeli (Rhagidiidae), Bonzia halacaroides (Cunaxidae), Alicorhagia clavipilus, A. usitata (Alicorhagiidae), Speleorchestes pratensis (Nanorchestidae) and Riccardoella sp. (Ereynetidae) were recorded (ZACHARDA
1978).
Fig. 44. The genaralized oribatid mites in the left picture, Argas verpertilionis in the right picture. Illustration from
the authors’ archive.
Crustacea
Crustacea, mainly water groups, are very diverse animals. Although their diversity is concentrated
in marine ecosystems, many species can be found in freshwater ecosystems, too.
Terrestrial isopods (Isopoda, Oniscidea) are animals occurring under logs, stones, wet litter etc.,
62
because they are susceptible to desiccation. Some of them are able to roll their body up into ball
(Glomerida) and remind some diplopods species from the Glomerida order. Animals are active usually during the night. Isopods are saprophagous, but they can also feed on roots or seedlings foliage
and due to this fact they are very important for litter fragmentation and nutrient cycling. 42 species
are known in the Czech Republic.
There are known three more or less troglophilous species of terrestrial isopods in the Czech Republic – Androniscus dentiger, Trichoniscus pygmaeus, Cylisticus convexus (MLEJNEK and TAJOVSKÝ 2008). Also other species can be recorded in underground habitats. For example DVOŘÁK
(2002) has recorded several regular inhabitants in investigated celeries – Oniscus asellus, Porcellio
scaber, Porcellio spinicornis and above all mentioned troglophilous Cylisticus convexus.
Millipedes and centipedes
Centipedes and millipedes and several other related, but in presented text not mentioned, small
groups of soil animals are grouped into convenient taxon called myriapods (Greek: myriad – “ten
thousand”, podus – “foot”). All have long annulated body with high number of legs. Only young millipedes have three pairs of legs.
Millipedes (Diplopoda)
Fig. 45. Generalized millipede. Free interpretation of different authors (e.g. BLOWER 1985).
Millipedes have fused most of trunk segments in pairs to form diplosegments (Greek: diploos –
“twofold”, podus – “foot”). Each diplosegment bears two pairs of walking legs, relatively short head
bears short antennae. Some millipedes are tubular, round – backed, others are dorsally flattened and
some resemble to terrestrial isopods and can roll their body into a ball. Some millipedes can secrete
dangerous repellants that protect them against predator.
Millipedes are susceptible to desiccation in low humid habitats and their skeleton is calcareous.
That is why they inhabit especially, but not only, wet and calcium rich habitats. We can usually find
them under logs, in leave litter, under stones etc.
Millipedes sometimes concentrate in thousands, especially during outbreaks and migration. In ecosystem millipedes are very important consumers of dead organic matter.
There are 20 000 millipede species in the world. In the Czech Republic 73 species have been recorded (TAJOVSKÝ 2005, TAJOVSKÝ 2001b, KOCOUREK 2001).
True troglobionts are not known in the Czech Republic, although two millipedes Brachychaeteuma
bradeae (5-8 mm) and Macrosternodesmus palicola have been recorded in hypogean environment
only, but their records are rare and both are not typical for cave habitat in other countries. More or less
63
troglophilous millipedes are Brachydesmus superus, Trachysphaera costata, T. gibulla and Blaniulus
guttulatus. Those penetrate to caves more frequently than others.
Centipedes (Chilopoda)
Centipedes are dorsally flattened with relatively long
legs and antennae. Almost each segment carries one pair
of walking legs. The characteristic features of centipedes
are often quite huge maxilipeds (modified first trunk of appendages) with poison claws. Centipedes are usually running surface dwellers, some are burrowing. Most are fast
predators, several species are herbivorous. TAJOVSKÝ
(2001a) and LAŠKA (2004) registered 73 species in the
Czech Republic.
No centipedes are closely associated with subterranean
environment but some species, e.g. common Lithobius forficatus, have been recorded in caves or celeries more frequently than the other ones.
Hexapods (Hexapoda)
In spite of enormous diversity there are common insect’s
Fig. 46. Examples of various types of cenfeatures. Their body is usually divided into three parts:
tipedes. Free interpretation of different auhead, thorax
thors and KRATOCHVÍL (1959).
and abdomen.
The
head
bears a pair of antennae, thorax carries three pairs of
legs. Many insects have wings, but not all of them.
Springtails (Collembola)
Fig. 47. Several species of springtails more
or less associated with subterranean habitats
(1 - Heteromurus nitidus, 2 - Mesogastrura ojcoviensis, 3 - Pogonognathellus flavescens, 4
- Pseudosinella sp. Photo: V. Papáč.
64
They are small, very abundant soil animals. The most
obvious feature of springtails is a jumping organ called
furca, which is reduced in case of soil confined species.
Those animals are abundant in many caves, although not
as much as in organically rich horizons. Springtails are
very important for soil structure development by faecal deposites. Their effect on growth mycorrhyzae and
control fungal diseases of some plants is also known
(DROMPH and BORGEN 2001). Some species feed on
fungal hyphae, some are carnivorous and feed on nematodes, rotifers etc. Fore more details on springtails biology see HOPKINS (1997). In the world 7 500 species
are known (BELLINGER et al. 1996-2009) and in the
Czech Republic 334 species (RUSEK 2005). Some species, e.g. Arrhopalites ruseki or Onychiurus rauseri
(NOSEK 1975a, b) have been described in the Moravian
karst caves. Nevertheless little data were still published
from subterranean habitats in the Czech Republic. Several examples include photos of above-ground
and subterranean species, which were based on personal information of Vlado Papáč from Slovakia.
Pogonognathellus flavescens is a forest species that is often recorded in cave entrances. It is an example of species with well developed eyes (6+6 eyes) and body pigmentation. Heteromurus nitidus is
frequently recorded in caves and also in deeper soil layers or under stones. Intermediate forms with
reduced eyes and pigmentations are known from caves. Mesogastrura ojcoviensis is known from
caves and from nests of small mammals. Its pigmentation is reduced, but eyes are well developed.
Pseudosinella species are the example of animals with well developed adaptations to cave environment (prolonged claws on the legs, bigger and depigmented body, eye reduction), where they live.
Orthopterans (Orthoptera)
In some subterranean nature and artificial habitats
(especially sandstone caves and old mines) near Děčín
Town in NW Bohemia and in old mines near Olomouc
Town in Moravia, grasshoppers Troglophilus neglectus and T. cavicola are very characteristic. According to the first literature data (CHLÁDEK et al. 2000,
HOLUŠA et al. 1999) it seems two mentioned species are known from the Czech Republic, but the last
literature data prove there is probably only one species in the Czech Republic – Troglophilus neglectus
(KOČÁREK et al. 2005).
Fig. 48. Troglophilus neglectus from the sandstone caves in Labe Valley. Photo: J. Kukla.
Beetles (Coleoptera)
Beetles are generally known and very diverse insects. Ca. 500 000 species are recorded in the world and ca. 6 000 species
in the Czech Republic (SEGERS and MARTENS 2005, LAŠTŮVKA et al. 2001).
Some are associated with caves in the
Czech Republic but true troglobionts probably absent in here. Some species, for example Trechoblemus micros (Carabidae),
Laemostenus terricola (Carabidae), Quedius
mesomelinus and Omalium validum (Staphylinidae) or some Leiodidae ones, are closely associated with different subterannean
habitats (undeground nests of mammals, old
mines, caves, stony accumulations).
Fig. 49. Trechoblemus micros (Loupežnická Cave near velké Březno near Ústí nad Labem) is one of the ground beetle
(Carabidae) species more or less connected with different
subterranean habitats. Illustration from the authors’ archive.
Dipterans (Diptera)
Very diverse insects are also dipteran flies. In the world there are known ca. 100 000 species and
in the Czech Republic live nearly 8 000 species (see JEDLIČKA et al. 2006). The most characteristic
feature of adults is one pair of well developed wings and second pair of reduced wings. Their live
is associated with terrestrial habitat where they feed on different organic mater, e.g. decaying plants
65
or animals´ tissues, pollen, etc. Adults of some species do not feed. Some groups are also predators.
Larvae live in different terrestrial or water habitats or are associated especially with the tissues of
dead animals and plants.
In caves and similar habitats, species of different groups, for example Limoniidae, Keroplatidae,
Sciaridae, Culicidae, Heleomyzidae, Phoridae etc. can be recorded. Some species represent only summer visitors, others spend all their life here. Quite interesting is for example species Speolepta leptogaster (Mycetophilidae). Its larvae look like small nematods. They are also the example of silk
produced species that can resemble spiders net. According to different authors threads can serve to
larvae as protection against often disturbance in subterranean habitats; or larvae can be prevented
from touching of the wet walls and consequently attached by fungi.
Fig. 50. Limonia species can be very numerous in some underground habitats (Mining gallery on Buková hora near
Děčín. Photo: B. Franěk.
Caddisflies (Trichoptera)
Caddisflies are moth-like insects with two pairs of hairy membranous wings. Larvae are typical
for water, especially, stream environment. Larvae of several species are associated also with wet soil.
252 species is known in the Czech Republic (CHVOJKA and KOMZÁK 2008). In caves or similar
habitats in the Czech Republic adults of two genera – Stenophylax and Micropterna can be observed.
Caddisflies are especially summer visitors of caves.
Fig. 51. Trichoptera Micropterna nycterobia. Illustration from the authors’ archive.
66
Butterflies (Lepidoptera)
Butterflies, similarly to beetles and dipterans, are among the most diverse group of
insects. Many species are generally known
and also their larvae, called caterpillars, are
well known. Despite this fact, determination
of many species is very difficult. There is
known ca. 3 500 species in the Czech Republic
(LAŠTŮVKA and LIŠKA 2008).
For many caves and mines at least one of two
common species Scoliopteryx libatrix and Triphosa dubitata are typical. Nevertheless also
some others, for example Inachis io, Aglais urticae, can be observed. Mentioned butterflies
are only winter or summer visitors in subterranean habitats.
Fig. 52. Triphosa dubitata. Photo from the authors’ archive.
Bats (Chiroptera)
They are the only mammals with well developed wings that enable them active flying. Wings are
formed by their skin which is stretched between their long fingers and the body. 26 species of bats is
known in the Czech Republic (ANDĚRA 2009, LUČAN 2009, REITER et al. 2007, ANDĚRA and
ČERVENÝ 2003, ZIMA and ANDĚRA 1996).
Bats eat different small animals, especially insects. They
are generally known as cave inhabitants but not all of them
are strictly associated with caves. Most of them prefer similar
habitats e.g. old mines. We can observe them here only for
general part of year, especially for wintering. Bats nesting in
trees hollow, many bats shelter in roof spaces in buildings.
Very characteristic is bats’ echolocation. Although bats
can see well, they use calls to detect their prey or to map terrain. Emitted signal with high frequency is returned and listened and sonic map is developed. Human can usually hear
frequencies of tones higher than 20 Hz. Some bats can hear
sounds frequency up to 110 kHz. Scientists and other people
interested in bats use detector to determine or more carefully
study bats (details for measuring with detectors and other
information about bat monitoring in the Czech Republic can
provide Czech Bat Conservation Society (www.ceson.org)).
Although bats are clean animals and spend a lot of time by
grooming themselves, they can be dangerous because they
can transmit EBL virus infection (European Bat Lyssavirus
1 a 2 EBLV1 and EBLV2). Only little is known about the susceptibility of the infection hosts and man contamination by
bats or by their guano is very rare. There have been infected
Fig. 53. Wintering Rhinolophus hipposionly several people in Europe and no one in the Czech Rederos in mining gallery near Jakuby near
public, although infected bats are known from our country,
Děčín. Photo from the authors’ archive.
too (HELEŠIC et al. 2007).
67
Althought all bats are protected by the law, the access to caves or to other subterranean resting
habitats (other habitats are not concerned here) is often free. Because wintering bats need to save energy, they must keep their body in the rest. Therefore it is important to protect the wintering localities
against excessive entry of visitors. Moreover, during last time specialists recorded presence of White
nose syndrom among our bats. White nose is caused by fungi. Fungus causes itching, blisters, etc.,
especially on their ears. This leads to the awakening of bats and they are often forced to fly. That is the
reason of their weakness which often leads to perishing. According to text version of prof. Horáček
discussion on Leonardo on the Czech radio station Leonardo from 24.1.2011 (www.rozhlas.cz\leonardo), the fungus will not exhibit so big problem for our bats in the future as it is predicted for the USA.
When the caves are officially investigated, investigators must have a special permission. Permission is
also needed in protected localities (e.g. reserves). Practical control of the visitors in terrain is difficult.
Nevertheless bats are not only resting during winter, it is also very important time for their reproduction. Young bats are born in summer but mating is running during winter months. Time to time bats
naturally wake up, too, and search for food and water.
Other vertebrates
Caves are used not only by bats but many other animals visit it, although their association with this
environment types is not so close. Among such animals belong e.g. salamanders (Salamandra salamandra), Pygmy Shrew Sorex minutus or some bigger ones such as fox (Vulpes vulpes) or European
Badger (Meles meles). Also the other big vertebrates as Ursus spelaeus, Crocuta spelaea used to
live in these caves in the past. Rich records are known e.g. from the Sloupsko-Šošůvské Caves (e.g.
DIEDRICH 2009, DIEDRICH and ŽÁK 2006) in the Czech Republic as well as in other countries.
These paleontological findings, including “humans cave cultures”, are not subject of our interest and
therefore we are not going to describe them here.
68
Future of underground research
Future of underground research
Whereas cave investigation is only a few associated with the conception of commercial exploitation
in the Czech Republic nowadays, the research is becoming a subject of so called basic research of
universities and research organisations supported by the state. That is why character and extent of a
research is determined not only by the scientists themselves but also by state, which sets the laws of a
research evaluation, that branch and form will or will not be supported.
Currently, there are some significant changes in the organization and support of research, education and even other areas necessary for state function in our republic. In addition, these changes are
not concern of our republic only; they are more of global character. We assume that, as well as it was
in the past, in the future there will also be possible to lay hopes on voluntary or state-supported associations, which participate in underground mapping much like basic morphological characteristics
obtaining. It is believed that the regional core, which will lead to fill many gaps in our knowledge of
nature, will be pursued in the future, too. It is known that some kinds of research have applied meaning rather than theoretic. “State rules” try to delimitate the difference between these two types, which
is still more likely formal. Thus it will take a long time before many regional practical problems of nature conservation will be sorted out – before we get to know what in particular cave occurs, where the
cave leads or even where the cave situated is. These regional practical problems can be also financed
by state administration bodies (Landscape Parks, regional authorities, etc.) however their budget is
much lower than of the Czech Grant Agencies, which support prestigious and commercial research.
It can be assumed that regional applied research, available for nature conservation authorities, will
be still more carried out rather than prestigious research oriented to international journals, and that
financing will be only a little dependent on state budget. However we believe that a basic research will
take place in a highest quality in the Czech Republic as well, although the amount of entering people
is going to be limited because of the number of job opportunities. The underground investigation is
maintained by methods of molecular biology, models of certain caves can be solved by 3D scanners,
underground sinuses are detected by geophysical methods, which can also help with the questions of
permanent ice existence in our conditions etc. Not only available underground but also the unavailable
places can be discovered, e.g. almost unknown epikarst environment (see for example PIPAN 2005),
etc.
Summarized, research of underground and its protection has not only a certain tradition in our
country but also a future, even its protection and investigation has not been a priority social issue. In
the Czech Republic new caves are discovered and many excited speleologists discover wide underground places in abroad as well. That is why we can make conclusion that underground research is a
quite interesting event and it will always be attractive and alive in our country.
69
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78
Název:
Autoři:
Mgr. Michal Holec, Ph.D.
Ing. Richard Pokorný, DiS.
Vědecký redaktor: Prof. Ing. Jaroslav Boháč, CSc.
Recenzenti:RNDr. Miloslav Zacharda, CSc,
RNDr. Karel Tajovský, CSc.
Vydavatel: Univerzita J. E. Purkyně v Ústí nad Labem
Místo a rok vydání: Ústí nad Labem, 2012
Vydání: první
Náklad: 80 výtisků
Rozsah:
80 stran
Tisk: PrintActive, s.r.o.
ISBN: 978-80-7414-416-5 (brož.), 978-80-7414-897-2 (online: pdf)

Subterranean Habitats - EnviMod

Transcription

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