I TA L I A N H A B I TAT S

Caves and karstic phenomena

Italian habitats Italian Ministry of the Environment and Territory Protection / Ministero dellAmbiente e della Tutela del Territorio Friuli Museum of Natural History / Museo Friulano di Storia Naturale Comune di Udine Scientific coordinators Alessandro Minelli Sandro Ruffo Fabio Stoch Editorial commitee Aldo Cosentino Alessandro La Posta Carlo Morandini Giuseppe Muscio

I TA L I A N H A B I TAT S

Caves and karstic phenomena Life in the subterranean world edited by Fabio Stoch

Text by Mauro Chiesi Luca Lapini Leonardo Latella Giuseppe Muscio Margherita Solari Fabio Stoch In collaboration with Paolo Forti Maria Manuela Giovannelli English translation by Gabriel Walton Illustrations by Roberto Zanella except for 89 (Enrico Zallot) and 90 (Marco Bodon) Graphic design by Furio Colman Photographs by Archivio Circolo Speleologico e Idrologico Friulano 14/2, 40, 41, 49, 50, 56, 60/1, 78 Archivio Museo Friulano di Storia Naturale 21, 22, 23 Archivio Unione Speleologica Bolognese 11, 12, 17, 27, 28, 29, 30, 43, 45, 47, 54/2, 125/1, 132 Adalberto DAndrea 8, 14/3, 19, 35, 39, 44, 62, 74, 133, 134, 140, 148 Sergio Dolce 90 Fulvio Gasparo 52, 94, 99, 103/3, 107, 109/2, 116/1 Salvatore Inguscio, 103/1, 103/2 Enrico Lana 95, 100, 104, 106, 109/1, 109/3, 118 Luca Lapini 54/1, 60/3, 60/4, 122, 123 Leonardo Latella 63, 64, 69, 112/1, 112/3, 113/2, 126 Giuseppe Muscio 60/2, 85, 127 Tiziano Pascutto 88, 113,/1, 117, 121/2, 125/2 Giuseppe Lucio Pesce 87/1 Mauro Rampini 54/3, 112/2 Federico Savoia 14/4, 156 Pino Sfregola 152 Margherita Solari 14/1 Fabio Stoch 54/5, 92, 93, 96, 101, 124, 143 Franco Tiralongo 130 Elido Turco 129 Stefano Zoia 54/4, 87/2, 105, 108, 111, 114, 116/2, 116/3, 119, 121/1 2002 Museo Friulano di Storia Naturale, Udine, Italy Even partial reproduction of texts and photographs is forbidden. All rights reserved. ISBN 88 88192 03 4

Caves and karstic phenomena

Life in the subterranean world

M I N I S T E R O D E L L A M B I E N T E E D E L L A T U T E L A D E L T E R R I T O R I O
Cover photo: Cave of Santa Barbara, Sardinia (photo: Unione Speleologico Bolognese)

M U S E O F R I U L A N O D I S T O R I A N AT U R A L E C O M U N E D I U D I N E

Directive 92/43 of the European Community, issued on May 21 1992, and commonly known as the Habitat Directive, represents a considerable improvement in overall environmental policy. The innovative approach of the Directive lies in its recognition of the complex and crucial role played by interactions between animal species, living plants, and the ecosystem surrounding them. It was within the perspective of the Habitat Directive that long-term activities could be planned and carried out. In order to inform the general public about the natural history of Italy and to publicize the results of scientific research, the Italian Ministry of the Environment and Territory Protection has planned publications focusing particularly on the environment. Italian habitats is a series aimed at improving present knowledge of environments at risk of degradation or even of disappearance habitats in which special and noteworthy fauna and flora can be found. The Ministry appointed the Friuli Museum of Natural History to prepare this series of monographs, starting with the present volume on karst phenomena. The flexible but scientifically correct structure of these volumes describes Italian habitats of international interest. This important series, while enhancing common knowledge about the environment, will contribute towards laying the foundations for proper management of natural resources - that great heritage belonging to us all.

Altero Matteoli Italian Minister of the Environment and Territory Protection

Italian habitats

Contents

Intoduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Giuseppe Muscio 1 Caves and karstic phenomena 2 Springs and spring watercourses 3 Woodlands of the Po Plain 4 Sand dunes and beaches 5 Mountain streams

Karstic phenomena and speleology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Giuseppe Muscio

Biospeleology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 Leonardo Latella Fabio Stoch

Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 Fabio Stoch Leonardo Latella Luca Lapini 6 The Mediterranean maquis 7 Sea cliffs and rocky coastlines 8 Brackish coastal lakes 9 Mountain peat-bogs 10 Realms of snow and ice

Protection and conservation of the subterranean environment . . . . . . 131 Mauro Chiesi Luca Lapini Fabio Stoch

Suggestions for teaching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149 Margherita Solari

Select bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 11 Pools, ponds and marshland 12 Arid meadows 13 Rocky slopes and screes 14 High-altitude lakes 15 Beech forests of the Appennines

Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159

Introduction
GIUSEPPE MUSCIO

Talking about caves to people who have never visited one is difficult. Once, during a meeting on cave photography, a discussion arose regarding the best techniques for the best results, and one well-known cave photographer reminded those present that a true photograph of a cave is ... completely black! And it is true: in caves, light is an extraneous element, a real form of pollution which man brings into the darkest environment imaginable, into a silence broken only by the dripping of water or the sound of sometimes rapidly moving waters. Few places other than caves have inspired mans imagination: in the past, he used them as dwellings and for protection, but he rarely penetrated further than light was able to reach - that limit beyond which mystery hovered, accompanied by fear and superstition. Caves have also been places of worship or of fortification, but only from the 19th century onwards have they been systematically explored and scientifically studied. Since then, man has understood how very singular the cave environment is: limited in extent, protected by rock, and with a particularly stable climate. But for this very reason, he also understood how delicate the equilibrium of the underground world is, and how easily it can be upset. Thus, in recent decades, we have realized the importance of respect for and protection of karstic phenomena. Although it covers the subterranean world, this type of protection must necessarily start from the surface of the earth, where karstism is widespread. The term karstism derives from Carso (Karst), the name of the geographic area between Italy and Slovenia, which has become synonymous with a certain type of landscape or, more simply, of everything morphologically linked with caves. Carso in fact derives from karren, a proto-European term which simply means rock - grast in Slovene, in use since 1177; kras in Croat, used since 1230; carso in Italian, and Karst in German, in which the original root is clearly preserved. It is not surprising that the root means rock, because in karstic areas rocks outcrop frequently and are wonderfully modelled. The landscape is thus characteristic, partly because it is not only superficial but also, indeed mainly, underground. The opportunity to increase our knowledge of karstism in Italy in order to
Subterranean lake in Grotta di Punta Galera (Palinuro, Campania)

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protect it better is fully justified: there are many endemic forms in the subterranean fauna and interesting minerals which are typical of the subterranean environment. However, there is also a very simple reason for protecting it which, alone, is abundantly clear to all: at least 50% of Italian drinking water is of karstic origin (the percentage is even higher in some southern areas of the country) and this figure is certainly destined to increase in the future, due to the various forms of pollution involving the steadily deeper and deeper watertables of the plains. Equally clear is the fact that few environments are so vulnerable and so slow to change as the subterranean one. Today, the karstic environment is not only intensively explored but is also a source of scientific information, a reservoir of prime importance. In the past, caves were the places of the imagination, the dwellings not only of prehistoric man but also of mythical beings which populated - and still populate - folklore. Thus, we have learned to observe natural cavities in the earth as a highly variegated source of information. Speleology - the science which studies them comes from the Greek spelaion (cave) and logos (study), and now attracts research-workers from a whole range of fields: geology, mineralogy, biology, ethnography, archeology, etc., and the speleologist is, basically, a naturalist in the widest sense of the word. There are many only slightly known, or completely unknown, aspects of caves, like that of mineralogy: for example, already in prehistoric times, men used the gypsum from caves in the TuscanEmilian Appennines (e.g., Calindri), and nitrates were exploited even earlier. More recently, many caves have been completely emptied of their contents of bat droppings, since the guano, as it is called, is an excellent natural fertilizer. Speleology came into being as a science in its own right at Trieste in the 19th century. It was the natural response of a large city which, then in full expansion, was searching for new water supplies in its hinterland. And its hinterland was the classic Karst, in which the only river of a certain size, the Timavo, mainly flows underground. One indication of the amplitude of speleology in Italy is that there are at least 10,000 speleologists, and more than 33,000 caves have been explored and surveyed. Although some are several dozen kilometres long or hundreds of metres deep, very many are small. But this does not mean that they do not contain very fine natural beauties. Natural cavities in the earth are often exploited from the economic viewpoint. Among the various Italian caves open to the public, four are well-known: Grotta di Castellana in Puglia (Apulia), Grotta di Frasassi in the Marches,

Grotta Azzurra in Campania, and Grotta Gigante in Friuli-Venezia Giulia, with hundreds of thousands of visitors every year. On one hand, the caves bring in local tourist revenue; on the other, the tourists themselves cause many environmental problems regarding the proper conservation of underground environments: not just pollution due to the waste they leave or the damage they do - more simply the disturbance created by the sole presence of man influences the delicate equilibrium of the subterranean climate (temperature and humidity). Writing about karstism is therefore not a simple matter: it is almost impossible to put down on paper the feelings we experience when we enter a cave and penetrate its absolute darkness. For a speleologist, whether the cave is long or short is not important: what counts is the fascination of exploration. There are only a few very large caves, richly decorated with concretions or concealing important findings, but no cave lacks its own hidden mystery. The true passion which urges the speleologist to adventure beneath the earth is knowledge, the exploration of an underground world which no-one or very few have ever violated, the desire to study a phenomenon which, perhaps only on a small scale, is always unique, completely different from its surroundings.

11

Concretions in Grotta di Valdemino (Liguria)

Karstic phenomena and speleology

GIUSEPPE MUSCIO

13

The natural environment which surrounds us, which we observe without thinking about it, is generally called panorama. When our gaze embraces a set of characteristics which are all somehow linked, we call it landscape. Thus, the dry karstic landscape is often compared with one containing superficial bodies of water of various kinds, because one of the peculiarities of a karstic area is precisely the absence of any clearly observable surface water network. The karstic process is that set of mainly chemical factors which lead to the dissolution of carbonatic rocks (limestone, dolomia), forming surface cavities (dolinas) or underground ones (caves), creating a subterranean water network, and modelling outcropping rocks in a very typical way. However, not all natural cavities owe their origin to karstism: some, for example, may form in or under glaciers, or in lava, or be of mixed origin. There have also been lengthy debates on what effectively constitutes a cave and what kind of empty space in rock, carbonatic or not, enters this category. For the Italian Cave Registry, listing natural cavities which have been officially surveyed and described, a cave is a cavity sufficiently large to allow access to humans and more than five metres long. This restriction is not applicable if we consider caves as developing morphological elements within which the widening of a small crack in the rock may lead to a vast subterranean system which is later perhaps destroyed by collapse or occlusion or, in the long term, involved in the various processes of erosion to which emerged land is subject. The karstic process Limestone is a rock composed almost entirely of calcium carbonate (CaCO3), a compound with very low solubility in water. However, in the presence of carbon dioxide (CO2), the water becomes more aggressive and acid, and dissolves the calcium carbonate, forming calcium bicarbonate, according to the following reaction: CaCO3 + H2O + CO2 Ca++ + 2HCO3 -. The equilibrium shifts right or left according to pressure and temperature.

Extensive karstifiable rocks in Sardinia

14

The strict meaning of the term karstic process refers only to the chemical reaction which affects calcium carbonate and, consequently, only to pure limestone. Today, the term has been expanded and karstic phenomena is accepted to refer to everything regarding the corrosive activity to which carbonates are subjected (thus, limestone and dolomia) and also to gypsum (calcium sulphate) and any other solubilizable rock (rock-salt, quartzite). Although the above reactions are the main causes of the development of karstism, they are certainly not the only ones and there are many variables

A model cross-section of a karstic area

The superficial part of a karstic area is intensely modelled and shows only a few traces of watercourses. The action of water creates structures like karren and dolinas. It then flows into the subsoil through the network of cracks, some of which widen to form true shafts, large enough to allow people to descend them. Through this dense network, water reaches first the percolation or vadose zone where, for example, it drips slowly, or flows as a thin sheet down whole walls of rock.

Giuseppe Muscio

15

Superficial karst forms Extremely variable in size; the best known are limestone pavements (also called karren or karrenfeld), dolinas and solution pans (or kamenitza) Swallow holes Water reaches the subsoil not only through minute cracks in rock, but also through true natural shafts. These are considerably modelled by the water which passes through them, first creating a circular cross-section, when water flows under pressure and occupies the entire shaft, and later scouring out part when flow diminishes and water acts only on one wall Collapse deposits At the bottom of large shafts and in areas of extreme fracturing, large quantities of collapsed materials may accumulate. This phenomenon sometimes leads to the creation of large halls in subterranean karstic systems Subterranean rivers Entire stretches of caves may be occupied by sometimes large watercourses. The walls often show traces of how the water regime developed, both in the shape of the cross-section and in the presence of special forms of excavation on the walls (scallops) Conduits and siphons The lowest parts of karstic systems are typically filled with completely flooded tunnels, called siphons. According to the morphology of the area, these siphons may meet in areas in which water generally flows freely.

Deeper down are those stretches of the karstic system where water flows more regularly (galleries with subterranean streams, waterfalls, small lakes, etc.) until the so-called phreatic zone is reached. Still deeper inside the rock is the water-saturated area with completely flooded galleries, called conduits under pressure. A succession of this type is the result of long, complex evolution of the territory, in which external factors intervene, e.g., tectonics, or variations in water-table levels.

16

which come into play: from the characteristics of the rocks to the condition of the vegetal cover, from the local climate to a series of mechanisms accompanying the main chemical reactions. Starting from lithotypes, not all limestones, as already mentioned, are pure: as well as calcite (calcium carbonate), small quantities of various minerals are always found, in particular dolomite (carbonate of calcium and magnesium), forming passages between limestone and dolomia. Karstic phenomena may also develop in these rocks; indeed, a small percentage of magnesium favours them. Therefore, although waters rich in CO2 can dissolve carbonatic rocks, to what extent they may do so is difficult to establish. First of all, we must bear in mind the climatic conditions and in particular the local level of rainfall, since water is an essential component of the reaction. The quantity of mass which is removed (rate of ablation) over a certain period of time has been calculated experimentally, taking into account the fact that karstic corrosion occurs not only on the surface but above all in the subsoil. Therefore, reported values do not refer to the removal of superficial carbonate and do not correspond to a simple lowering of the topographic surface, but to the entire rock mass which is dissolved, including that below ground. For a mean precipitation level of 1000-1500 mm/year, mean ablation rates of 50-100 mm/1000 years may be hypothesized for the Mediterranean area; values for Alpine areas are slightly higher. Recent measurements made in Friuli-Venezia Giulia have defined carbonate erosion rates of 20-40 mm/1000 years. i.e., slightly lower, but they refer only to outcropping rocks and may thus be compared with those quoted above. This means that karstism alone, in 10,000 years, can remove the equivalent of one metre of thickness from an Alpine carbonatic massif. This is a very significant value; in gypsum, the figure is ten times greater. If we examine the classic reaction, the presence of CO2 clearly plays an essential role, since it makes water aggressive towards carbonates. CO2 is present in the atmosphere, but its quantities may increase considerably in the soil, according to the vegetal cover, and its solubility is greater in cold rather than in warm waters. This means that, other conditions being equal, cold waters are more aggressive than warm ones; but in hot, humid climates the production of CO2 due to decomposition of vegetal residues is very high and it is for this reason that, although the waters are warmer, local conditions favour more marked karstic phenomena. The question of water temperature is important if, for example, we think of high-altitude karsts, where snow meltwater easily generates superficial karstic forms - always, of course, if they flow on carbonatic rocks.

17

Grotta Paradiso at Fluminimaggiore (Sardinia)

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It is also clear that, although the role of CO2 is essential to make waters aggressive, the possible presence of strong acids enhances their corrosive action, giving rise to what are called hyper-karstic phenomena. Strong acids may be present in the external environment, but they are more commonly found in the rocks themselves: for example, H2SO4 is linked to the reaction of oxidation of pyrite or H2S of deep origin. But how do meteoric waters enter carbonatic rocks? Rainwater, which is enriched in carbon dioxide in the atmosphere and soil, reaches the rocky surface and is already aggressive. It may corrode the surrounding rocks when long lines of discontinuity meet, since they are the preferential directions of flow of superficial waters, and it begins to excavate the rock, filtering inside it. At the same time, however, the water becomes saturated in calcium carbonate and cannot continue its corrosive activity beyond a certain point: other waters then widen cracks further. We know that theoretical saturation levels may be exceeded by karstic waters according to conditions of pressure and temperature, and also that mixing of waters with different chemical characteristics favours the development of karstic phenomena. With this knowledge, we can reconstruct the physical elements which are required for the karstic process to begin. These are: 1. carbonatic rock containing discontinuities (fractures, faults, etc.); 2. water containing dissolved carbon dioxide; 3. differences in altitude, permitting water movement. The second point has already been analysed. It is clear that water movement is essential for the development of karstic phenomena and to remove those fragments which, not soluble by CO2-enriched waters, could fill the cracks and prevent further speleogenesis (i.e., the formation of natural cavities). So let us examine further the first point: discontinuities are the natural consequences of deformational events undergone by the rocks during their geological history. The surroundings are thus full of cracks, mainly only a few microns thick and often not distinguishable on the surface because they have been altered by modelling, but they are sufficient to allow water to percolate down. In the past, a theoretical limit to the downflow of waters was believed to exist, due to the idea that, at a certain depth (500 or 1000 metres), cracks tended to close as a result of the enormous mass of overlying rock, thus preventing the passage of water. We now know that enormous quantities of water are found during the excavation of tunnels, even if they are overlain by thousands of metres of rock - an example is the Mont Blanc tunnel.

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Grotta Nuova at Villanova (Friuli)

Until now, in examining speleogenesis - that set of processes which create karstic cavities - we have referred to the fundamental role played by chemical corrosion. Although this is essential during the first speleogenetic phase - that is, when the first subterranean conduits form - as they gradually become larger, erosion also comes into play. This is a purely mechanical action on the part of the more resistant granules which chemical reactions free from the carbonatic matrix (commonly fragments of flint, quartz or ferrous minerals) and which swirling waters scour against the rock, abrading it. As karstic conduits progressively widen, collapses favour the formation of large halls at points in which rock fracturing is especially intense, often at the point where lines of tectonic origin meet. Research undertaken during recent decades has definitely changed the original view of the fathers of karstism, who were, in a certain sense, purists. We now know that there are so many

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variables which enter into play in the origin and development of a karstic cavity, that they certainly cannot be reduced to the simple formula: water + carbon dioxide + calcium carbonate: although everything starts from a simple corrosive reaction, all kinds of morphogenetic agents are at work. Superficial karstic forms Karstism almost always begins on the surface, i.e., in outcropping rocks, leading to what is defined as a karstic landscape, composed of highly differentiated forms according to the various features of the outcropping lithotype, and the local climate, with particular reference to rainfall: we thus speak of tropical karst, temperate karst, and so on. Forms are also generally grouped according to size. In Italy, karstic zones occupy more than 27% of the entire territory and there are widespread high-altitude and temperate karst areas, including, for example, the area between Venezia Giulia and Slovenia, which is part of the so-called classic karst. The superficial forms are grouped into categories partly according to size (the original term is often based on German or Slovenian words; see table, page 21). Ponds form where rainwater collects in small depressions; overland flow may give rise to karren, solution flutes or rills, and solution runnels, distributed according to the gradient of the outcrop. Dolinas may have various origins and shapes: they are large areas where the original terrain sank, due to dissolution or collapse. They are circular or elliptical in shape, and range from a few metres to several dozen metres across. Larger forms, sometimes several kilometres in diameter, are called

polje, and are often used for agriculture because of their very fertile soil, with a superficial, seasonal, water network. In Puglia (Apulia), the word Pulo is used for similar shapes: e.g., the Pulo of Altamura and that of Molfetta.
CATEGORIA CATEGORY ACCUMULATION OVERLAND FLOW INFILTRATION FORME PICCOLE SMALL FORMS SOLUTION PANS KARREN CAVERNOUS KARREN PERCOLATION SHAFTS CREVASSES KARST SPRINGS SUBSIDENCE DOLINAS DOLINAS, SNOW SHAFTS CORRIDORS POLJE KARSTIC VALLEYS, UVALA BLIND VALLEYS, POLJE KARSTIC LAKES CLOSED VALLEYS FORME MEDIE MEDIUM-SIZED FORMS FORME GRANDI LARGE FORMS

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OUTFLOW

Special shapes are due to considerable superficial dissolution which, freeing blocks of limestone which may become isolated from each other, are often modelled into strange shapes and aspects, the so-called cities of stone. Among the superficial forms are springs, either free-flowing (in which water

Two types of dolina evolution

Surface karst landforms due to impact and streaming of raindrops

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grooves pool

comes from a partially well aerated natural cavity) and those of Vaucluse type (in which the whole cross-section of the natural cavity is occupied by water). Then there are karstic lakes, similar in shape to classic lakes but lying on karstifiable rocks, with mainly subterranean supply. One of the best-known example is the lake of Doberd in the Karst near the river Isonzo. Subterranean karstic forms The ideal cross-section of an already evolved karstic territory may be subdivided into: an outcropping belt, containing all superficial morphologies and lacking permanent bodies of water; and the immediately underlying section, which has a series of shafts and galleries of karstic origin, where water is present only sporadically after rain. Further down (vadose zone) are karstic systems with water and, lower still, near what is called the piezometric surface, all the cavities in the rock are filled with water (saturated or phreatic zone). The modelling action of water in a karstic subsoil creates forms which are commonly subdivided into karstic conduits and wall forms.

23

runnels

karstic hole

pool

large runnels

interlayer cavity

Names of some common karst forms on rock

Superficial karst forms: small meander, solution pans, grooving

Karst springs: free-flowing water (left) and Vaucluse type (right)

Lake of Doberd (Isonzo Karst): one of the best classic examples of a karstic lake

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Dolinas
Dolinas are typical superficial karst forms found in temperate climates. A dolina is defined as a closed hollow, between a few metres and 1 km in diameter, and composing a simple hydrographic unit. The perimeter is generally circular or subelliptical, although there are also irregular or complex forms resulting from the fusion of a variable number of dolinas (uvala). According to the ratio between mean diameter and depth, dolinas are classified into three categories: flat, when the ratio exceeds 2; funnelshaped, when it is less than 2; and deep, when the depth is greater than the diameter and the slopes are subvertical. Dolinas are generated by normal dissolution phenomena by fast-flowing water moving towards a topographically depressed point, leaching and corroding the rocky slopes. The central point is often absorbent, due to microcracks in the rock or swallow holes, not always visible because they are covered by debris or soil which, steadily deepening, makes up the bottom of the dolina. Superficial dissolution also occurs under the soil cover and is due to normal solution, but it may also be related to periodic variations in the concentration of carbon dioxide produced by the biological activity of micro-organisms or vegetation. At depth, the effects of corrosion due to mixing of waters, cooling, or enrichment in magnesium prevail. The soil on the bottom of a dolina is often residual, the result of accumulated insoluble minerals which compensate the basic pH in soils deriving from carbonatic rocks. Dolinas which developed according to this model (normal solution) are often hemispheric or funnel-shaped, and may occur both in limestone and in rocks with various degrees of solubility (dolomitic limestone, etc.). They may also form in coherent non-soluble rocks like sandstone and, if they rest on soluble materials subject to karstism, may cause the collapse or subsidence of underlying formations. Very different is the evolution of collapse dolinas, which have subvertical slopes and originate from the fallen ceilings of caves: in this case, dissolution is a factor of secondary importance. Although the topographic position of a dolina may be random, it often reflects the presence of lithological or tectonic structures, such as the dip of rock layers or the directions of faults. Morphology too (e.g., perimeter lengthening) may be correlated with these factors. Large dolinas, with diameters exceeding 100 metres, may have their own microclimates, with thermal inversions accompanying inverse stratification of vegetation and the presence of more markedly cold-loving plant species. The micro-climate of a dolina is due to several factors. In the first place, cooling by radiation on a concave surface is greater than on a corresponding flat area, being directly proportional to the surface exposed. On the bottom, the soil is usually saturated in water and evaporation causes further cooling. In addition, in autumn and winter, from the first slope to enter shadow (the western one) cold air descends towards the bottom and lowers the temperature in the hollow even before sunset; conversely, in the morning, it is reached by the sun much later than the southfacing slope. The consequence of these

Margherita Solari

25

1150

1100

ice
1050

beech

Oriental spruce
1000

silver fir

dwarf pine

willow

rhododendron

Alpine flora

Seriation of vegetation in a steep dolina in the Italian Karst. Section illustrates phenomenon of inverse vegetation in relation to microclimates occurring inside cavity

factors, partially attenuated during summer, is a strong average thermal gradient - about 7 per 100 metres of depth, i.e., about 12 times greater than the external gradient in temperate climates. This means that, temperaturewise, descending 50 metres into a dolina is equivalent to climbing a hill 600 metres high. In these conditions of microthermy on the cooler southern slope (exposed to the north), the resulting vegetation may be a mixture of local species which prefer cool environments, or ones generally typical of higher altitudes. In this case, the four slopes of the dolina are covered with different plant associations, having intermediate conditions on the western and eastern slopes and extreme ones on the

northern and southern ones. These differences are also evident when the gradient of the slopes are symmetrical, and depend only on topography: this is called climatic continentalization. In very large dolinas, the inversely stratified vegetation is very clear and particularly interesting: in some cases, a complete series of plant associations may be found, typical of gradually higher altitude belts until we come to cenoses with dwarf pine typical of high-mountain and sub-Alpine belts. The peculiarities of dolinas thus represent an element of discontinuity which is not only morphological: they introduce diversity and enrichment into the environment by means of the selective organization of flora into different types of vegetation.

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Karstic conduits are created by the chemical action of water along discontinuities: at the beginning of speleogenesis, the conduits are completely filled with water under pressure and may later develop into vadose conduits in which water flows freely, with a consequent increase in its mechanical action. Vertical shafts are forms created by percolation. Water also models the walls of karstic conduits, e.g., in scalloped forms. Subterranean karstic systems are thus composed of galleries and shafts, sometimes filled with water (active) or abandoned (fossils). However, these two terms seem to be inappropriate, because the same system often has both active and inactive parts, or parts which are active during periods of intense rainfall and not active during the normal water regime. One particular aspect of speleogenesis in karstic areas is due to hydrothermal fluids, which are highly corrosive, favouring the formation of large halls, the development of which is often not influenced by cracks. In any case, speleogenesis requires lengthy periods of time - in the karstic environment, from hundreds of thousands to millions of years. Evolution is more rapid if hyper-karstic or thermal factors intervene; even more rapid is erosion of gypsum deposits; the filling-in of some caves in the EmiliaRomagna Appennines has been dated, and only lasted about 5,000 years. The whole karstic phenomenon of the Gessi del Bolognese is believed to have developed over the last 100,000 years, and most of the caves are less than 18,000 years old. Non-karstic cavities Not all natural cavities are due to true karstism: there are some caves of a certain importance which open inside conglomerates and which may be very large indeed. This happens because waters act by dissolving the carbonate which cements conglomerates, thus freeing pebbles and matrix, which are then removed by mechanical action. These caves often contain thick deposits of clay and, being subjected to frequent collapses, change their shape relatively rapidly. Among the most interesting are the caves in the Miocene conglomerates of Montello (Veneto), some of which are more than 1 km long (the Busa di Castel Sotterra extends for nearly 7 km). Some caves which develop at the contact between different lithotypes are considered of mixed origin: there are many which originate at the interface between carbonate and flysch. The first speleogenetic phase is due to corrosion of carbonatic rocks, but then the cave expands due to strong erosion of the flyschoid complex (composed of alternating marl, sandstone

27

Lava conduit on flanks of Mount Etna (Sicily)

and clay). Caves of this type have been studied in detail in Friuli: the complex of the Grotta Nuova di Villanova, more than 7 km long, develops precisely at the contact between a conglomeratic bank (in which both clasts and matrix and cement are all composed of calcium carbonate and thus the lithotype behaves, from a chemical viewpoint, like limestone) and Eocene flysch. Other types of mixed cave are sea grottoes. Their terminal portions, having originated from mixtures of salt and karstic waters, have often been widened by the destructive force of the sea. Instead, very far from any kind of karstic origin are caves resulting from lava flows: widespread mainly around Mount Etna in Sicily, they also occur in other Italian regions. These caves are called syngenetic, since they usually form over very short periods of time (from a few days to a few weeks), together with the rocks which contain them. They are also highly unstable and often do not last long, being rapidly destroyed if their ceilings collapse. However, it is relatively easy to date them, when the age of the various lava flows is precisely known. There are several morphologies, due to both type of lava and mode of

28

Grotta della Spipola, in gypsum near Bologna (Emilia-Romagna)

formation, but the Etna lava caves are grouped into two large categories: rheogenetic (superficial), and of fracture type. Rheogenetic caves are simply the empty spaces left by the flow of magma in lava tunnels. They are generated by liquid lava flowing on a solid surface: the upper part, in contact with the air, cools and solidifies quite rapidly, while the part underneath continues to flow, and then leaves an empty space once the supply of lava ceases. Fracture caves extend in depth along discontinuities sometimes created by seismic events. Intermediate or mixed forms may also exist, created by a combination of the two types. Although it may appear strange, as regards their genesis, lava caves are similar to glacier caves, which also form as a result of flow. In glacier caves, water from melted ice excavates galleries and shafts, often extremely short-lived. Climate in subterranean environments The problem of subterranean climates is often underestimated. However, if we consider to what extent it influences, for example, the fauna living in this particular environment or, from another viewpoint, the fact that caves were

once extensively used as places in which to preserve food, we realize that this is not simply a secondary aspect of speleology. The first point to examine is temperature: in general, apart from the first few yards of a cave from its entrance inwards, which is influenced by the external climate, the temperature deep inside is the same throughout the year and is very close to the mean value of the temperature in the area where the cave itself is located. This is because fluids (water and air) circulate in caves, and they determine the temperature. If we base our calculations on the thermal gradient of the earth, we should have an increase of about 3C for every 100 metres of depth (values measured during the excavation of mine shafts, for example) but, apart from very small variations, vertical caves such as shafts maintain their temperature even at depth. For example, in the depths of the high-altitude carsi (i.e., in carbonatic massifs at altitudes exceeding 15002000 metres), the temperature is generally close to zero, since the caves were created mainly by the snow meltwater which flows through them. Due to their isothermy, caves are thus relatively warm in winter and cool in summer. In low-altitude areas, temperatures range from about 10C in the preAlpine belt to 16-18C in caves in Sardinia or Puglia (Apulia). There are, of course, special cases like those of lava caves, which have their own special climatic conditions, with very high temperatures if they have been recently formed. But temperature differences do exist in karstic caves, due to the temperature of their subterranean rivers, the waters of which may come from several places. More constant than the temperature inside natural caves is relative humidity, ranging from 95 to 100%. Lower values are only found near the entrances and in particularly dry climates. Air currents inside karstic systems are of considerable interest. They are usually quite slow, but may be significant if the cave has several entrances or other kinds of links with the external world. Currents depend on season since, as mentioned Ice concretions. Grotta della Spipola above, the temperature of the air in (Emilia-Romagna). Swellings indicate moments of relatively higher temperature (shortly after caves is lower than the outside midday) when ice starts to melt and then temperature in summer and higher in freezes again

29

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winter. Draughts may be felt inside underground systems, particularly when they blow through narrow galleries from one part of the cave to another. Another great difference between the inside of a cave and the outside is the composition of the atmosphere: inside a cave it is highly variable, while outside it is much more constant. What changes significantly is the amount of CO2 which, in ordinary air, is normally between 1 and 3%, whereas inside a cave it may be up to 100 times greater. These values, which are due to biological activity in the soil and water transport, vary according to season and type of cave development, but they are in any case within our tolerance range. Only rates exceeding 10% of CO2 are dangerous: they are found in the lowest layers of some volcanic caves or ones in which hydrothermal fluids are present (the phenomenon was known even in the 18th century in the Grotta del Cane at Pozzuoli, near Naples). Breathing problems arise in some caves which are, or were, affected by sulphur exhalations: examples are the Grotta Azzurra and Grotta di Cala Fetente at Palinuro (Campania) and Grotta del Dragone at Maratea (Basilicata). For instance, sulphobacteria present in water greatly reduce the contents of the oxygen in favour of other

Cave deposits
Waters in caves always contain mineral salts. When they flow inside a cave, they may become oversaturated and give rise to chemical deposits. These deposits have always been divided into two subgroups: concretions and mineralizations, the difference essentially being based on the degree of crystallinity: high in mineralizations and low in concretions. In fact, in nature there are many macrocrystalline or even monocrystalline concretions, just as there are cryptocrystalline or definitely amorphous mineralizations: in practice, the possibilities are infinite. Thus, although subdivision based on the degree of crystallinity is not scientifically valid, it is useful for better understanding of the various aspects of the phenomenon. One particular type of cave deposit is the result of accumulations of bat droppings, called guano, used in the past thanks to its excellent properties as a fertilizer. It plays a significant role in the subterranean ecosystem and may sometimes even influence the genesis of certain minerals (e.g., brushite). Concretions This term was once given to deposits of calcium carbonate, but was then extended to gypsum. Then, as mineralogical knowledge of the subterranean world expanded, it was discovered that many other minerals could give rise to concretions which were absolutely identical in form and evolutionary mechanism to those of calcium carbonate: in Italy, for example, there are many well-known concretions of sulphur.

Paolo Forti Giuseppe Muscio Although several factors may concur to modify the form and external aspect of cave deposits, water movement undoubtedly most greatly influences their external aspect, so that the most logical classification of concretions is based on this parameter. Concretions resulting from water movement: slow dripping: stalactites, tubular forms, draperies and curtains dripping by impact: stalagmites, conulites, splash concretions, circles water flow: encrustations, columns, barriers, moonmilk submersion: pisolites, cave clouds, coralloids, moonmilk capillarity: eccentric and disc-shaped forms evaporation: floating crystals, coralloids, trays condensation: rims, boxwork, oriented coralloids, moonmilk salient: geysermites The first group contains the most common and numerous concretions, which is why they are further divided into two subgroups: the effect of the gentle detachment of a drop of water is very different from that of its impact on a surface below, and gives rise to very different forms.

31

Eccentric concretions of aragonite and calcite

Concretion of sulphur and gypsum flakes

Concretions of barite

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Cave deposits Mineralizations About a hundred years ago, known secondary cave minerals numbered less than 45, for two main reasons: except for calcite, aragonite and gypsum, which alone make up more than 99.5% of all cave deposits, other secondary minerals are very rare, scattered, and difficult to observe. However, they now number more than 260 and new ones are found every year. By cave mineral we mean a secondary mineral which formed in a cave and resulted from a chemico-physical reaction which involved one or more primary minerals existing in the rock or in physical and/or biological deposits inside the cave itself. These rigorous limitations are necessary to avoid secondary cave minerals being considered together with all minerals existing in nature. Caves are not generally environments particularly favourable to minerogenesis, and very many caves do not contain any secondary minerals at all. But they may contain a large variety of rocks: limestone, dolomia, gypsum, rocksalt, quartzite, basalt, etc.. Their chemicophysical degradation brings to the system a large number of ions which may be deposited as secondary mineralizations. In addition, speleogenetic evolution may cause primary mineralizations to outcrop on cave walls which, in contact with percolating waters or atmospheric oxygen, may give rise to new chemical compounds. Lastly, caves may contain deposits of clay, guano, bones, etc., which in turn bring in ions which produce further secondary minerals. On this great variety of substrates, waters of various origins - meteoric, marine, thermal - with very different contents of dissolved salts, in terms of both quality and quantity, then act on this great variety of substrates. In special cases, it may not even be water which acts directly, but other fluids, such as fumarole fluids in volcanic environments. The complexity of substrates and the variability of water chemistry, combined with the great differences in temperature which may exist in caves, give rise to a large number of phenomena and potentially active minerogenetic mechanisms. Some of these act in all types of caves and over a wide temperature range; others are only active in special types of caves or need particular temperatures in order to become active. So it is not surprising that many minerals were observed for the first time in cave environments. Just one example is that of francoanellite, a phosphate found for the first time in the cave of Castellana and named after the famous Italian speleologist Franco Anelli. Nine minerals are still today exclusive to the cave environment. The scientific importance of cave deposits Of all cave sediments, chemical ones and in particular concretions of calcium carbonate are highly flexible and powerful instruments for paleo-environmental and paleoclimatic reconstructions. This is because, as they grow, they incorporate various trace minerals, fragments transported in suspension during floods, dust carried in by air currents, and organic material, varying from molecules of humic acids to spores and pollens. The laminate and necessarily ordered structure of concretions immediately provides a relative chronological reconstruction of the events corresponding to each single growth band. Then there are a series of methods which, more or less easily,

Paolo Forti Giuseppe Muscio

33

B A

4
C

5 6 3 14
A E

7 10 9 8 15

2 12
B

13

11

18
D

17

19
F

16

furnish an absolute chronological scale. Thus, by analysing in detail the evolution of stalagmites, we can gain information on large-scale earthquakes over the last 500,000-600,000 years, and improve our assessments of seismic risk. Data on the deposition temperature of concretions and on the conditions and mechanisms of their formation can lead to detailed paleoclimatic and paleo-environmental reconstructions, providing much information on how climates evolved and, for example, what the landscape must have looked like in the second half of the Quaternary era. For instance, thanks to concretions in coastal grottoes (both emerging from the water and submerged) around Palinuro (province of Salerno, southern Italy), we can date the variations in sea level in that sector of the Tyrrhenian Sea over the last few thousands of years. Thanks to measurement stations located in caves, we can quantify movements connected with neotectonic phenomena.

A B C D E F

flowing water dripping water capillary water water in solution pans water forming by condensation warm rising water flowstone draperies, curtains column drop stalactite hollow stalagmite curved stalactites moonmilk disc-shaped forms macrocrystalline eccentric forms inflorescences microgours pisolites boxwork rims geysermites rounded concretions encrustations mud stalagmites large crystals

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19

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A B

35

6C
20 C 18 C 16 C

4C 2C

Air circulation in natural caves: wind tunnel in summer (A) and winter (B); cold (C) or warm (D) air trapped; in caves with only one opening, air circulation depends on atmospheric pressure (E)

compounds (mainly H2S): while there are no problems in areas well connected with the outside world, with good air supply, special attention must be paid further inside. Neglecting this phenomenon has cost the lives of more than one speleo-diver. Karstic areas and the major subterranean systems in Italy Italy occupies a prominent position both in the field of cave study and in more purely exploratory activity. This is for historical reasons, and because karstic phenomena are so widespread in Italy: 27% of the territory is composed of karstifiable rocks, and well structured and highly developed research on caves has been going on for more than a hundred years. The various regional registries list more than 33,000 caves, which is a somewhat low percentage of those actually present in Italy: for example, the cave registry of Friuli-Venezia Giulia contains more than 6,300 caves over an area of about 7,800 sq.km., and every year this number increases by about 200 new entries. There are as many as 180 subterranean systems in Italy (1999 data) which exceed a depth of 300 metres, and of these six even exceed the threshold of

Grotta del Dragone at Maratea (Basilicata)

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TN AO VE MI TO BO GE FI AN PG AQ ROMA TS

1000 metres. The record depth is that of the Abisso Paolo Roversi in the Apuan Alps, which goes down as far as -1250 metres. There are 92 caves (again, 1999 data) which exceed 3 km in length, of which 20 exceed 10 km. The longest Italian cave is the Corchia Complex (Apuan Alps), more than 52 km long. This is quite a feat, but it pales when compared with the length of the largest underground system in the world, the Mammoth Flint Cave in the United States, with more than 500 km of galleries! Comparing tables of the longest and deepest caves with cave registry entries, we see that Friuli-Venezia Giulia and Veneto, which have 9% of karstifiable territory, have more than one-third of known caves, and the same regions, together with Tuscany, Piedmont and Lombardy, contain three-quarters of all large Italian caves. These are the most important karstic areas (but it is also true to say that they have been systematically studied for longer, and have thus given rise to a greater quantity of data). Piedmont and Val dAosta. The Val dAosta is almost totally composed of metamorphic and magmatic rocks, so that it does not contain any karstic phenomena of significance. However, Piedmont has some of the largest karstic systems in Italy, mainly concentrated in two large areas of the Maritime Alps, Marguareis and Mongioie. The Piaggia Bella complex opens in the Marguareis valley, and contains about a dozen different caves which, taken together, form a subterranean network of more than 35 km, reaching -950 m. Slightly further south is Labassa, a downvalley continuation of Piaggia Bella, more than 14 km long and with a maximum depth of -609 m. In the same massif, the Conca delle Carsene complex extends for 13.5 km and reaches -759 m. The Abisso dei Perdus, Abisso Scarasson, Abisso Cuore di Pietra, Abisso Libero, Abisso Ferragosto and the Colle dei Signori Complex (partly in Italy and partly in France) all exceed 500 m in depth. In the Mongioie massif are the C1-Regioso complex (more than 6 km) and the Grotta delle Vene (4.7 km). The Abisso Ngoro-Ngoro and Abisso M16 in Val Corsaglia both exceed -470 m. The Risorgente della Mottera has been explored to a depth exceeding 600 m (14 km), almost as far as the karst surface basin. The largest cave open to tourists, near Cuneo in Piedmont, is the Grotta di Bossea, also equipped as a scientific laboratory and well-known for its remains of the cave bear Ursus spelaeus. The Grotta dei Dossi, near Mondov, was one of the first Italian caves opened to tourists, being equipped to allow them to visit in the early 19th century.

37

CB BA NA PZ

CA

PA

RC

LIMESTONE AND DOLOMIA

GYPSUM

L AVA

Main karstic areas in Italy

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Lombardy. Under the plateau of Cariadeghe (Brescia) is one of the most important karstic cave phenomena, the largest of which is the Omber en banda al Bus del Zel, more than 15 km long and about 420 m deep. The Bergamo mountains contain many karstic areas of great interest: the Grotta Maddalena in Val Taleggio is about 10 km long. From the exploratory viewpoint, of considerable potential are the caves around Lake Como: the Tacchi-Zelbio complex at Piani del Tivano extends for almost 10 km and the cave at Capanna Stoppani more than 8 km; the BlGuglielmo complex reaches -557 m. The Grigne mountain group contains some of the deepest caves in Italy: speleologists have descended the Abisso Viva le Donne to -1170 m and the Abisso Capitan Paff to -795 m. Almost at the border with Piedmont, near Varese, the Parco di Campo dei Fiori is another karstic area of great potential: the Nuovi Orizzonti and Marelli caves extend for almost 6 km, and the Abisso Schiaparelli goes down to -600 m. Some karstic phenomena in the region were visited by illustrious personages in ancient times: the intermittent spring of Torno (Como) was described by Pliny the Elder in 50 A.D., and the cave of Fiumelatte and the Ghiacciaia di Moncodeno (Como) were described by Leonardo da Vinci. The Ghiacciaia (ice-box) was again described in minute detail by the Danish polymath Stensen in 1627, in a letter addressed to Nicol Cosimo III, Grand Duke of Tuscany. Trentino-Alto Adige (South Tyrol). For geological reasons, the largest caves are to be found in the southernmost area of the region. Although this territory was the cradle of mountaineering, after much research around the 1930s, karstic phenomena were only properly studied after the 1960s. One of the largest Italian caves, Bigonda, is in the Trentino region, and runs for about 26 km in Valsugana, collecting the waters of the Asiago plateau, where the Grotta del Calgeron (5.3 km) also opens. Again in Valsugana are the still active complexes of Fosca and Castello Tesino. There are about 700 caves in the Brenta Dolomites and the adjacent Gazza-Paganella group, including the Grotta di Collalto (5.2 km), Torrione di Vallesinella, Abisso Popov, and Abisso di Lamar. In Val Daone, south of the Adamello group, is the Aladino cave (7 km), and the Abisso di Val Parol (1.6 km, -430 m) opens in the northern slopes of Monte Baldo. Superficial karstic phenomena are marked and widespread in the low Sarca valley and on Mt. Baldo. Lastly, explored only during the last 20 years are the karstic phenomena of the

Alto Adige (South Tyrol). Some very long caves have been explored in the Marebbe plateau in the north-west of the region. Of great importance are the remains of bears in the Grotta delle Conturines in the high Val Badia. Veneto. With more than 6,500 registered caves, the Veneto is one of the Italian regions in which karstism is not only extensive but also best studied. Apart from small sectors in which magmatic or metamorphic rocks outcrop, the entire mountain belt, made up of mainly carbonatic rocks, appears as a single uninterrupted karstic area. The Lessini and Beric Hills, the plateaus of Asiago and Tonezza (Vicenza), and Grappa, Montello and Cansiglio are all karstic areas par excellence, and important pages in the history of Italian speleology have been written about them. Although the Belluno area has practically been explored only in recent years - with completely unexpected results - many Dolomitic mountains are still totally unknown as regards karstism, and the Dolomites may be considered a new frontier in Veneto speleology. The Buso della Rana (in the Vicenza Lessini Hills) is the largest Veneto cave, more than 25 km long, and it holds the record (which may soon be broken) of being the longest Italian cave with only one entrance. The Busa di Castel Sotterra near Montello is one of the longest caves in the world in conglomerates (more than 4 km). Two large Veneto caves nearly reach 1000 m in depth: the Abisso di Malga Fossetta in the Asiago plateau (-974 m) and the Piani Eterni complex in the Vette Feltrine (-966 m). The Spluga della Preta in the Verona Lessini Hills is third on the list of large Veneto caves. It remains symbolic for its exploratory history, a true archetype and source of historical knowledge, not only for the Veneto but for the whole of Italian speleology. Friuli-Venezia Giulia. The Karst, particularly near Trieste, was the cradle of Italian speleology and, although systematically researched for 150

39

Grotta di Castel Sotterra (Montello, Veneto)

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years, continues to supply new revelations. The density of known caves is extremely high: 3,000 of them over a surface area of only 200 sq.km. And the mystery of the subterranean course of the river Timavo has not yet been completely solved. Among the best-known of the Carso caves is certainly the Abisso di Trebiciano which, by means of a series of shafts, reaches the course of the Timavo at a depth of more than 350 m. The Grotta Skilan is the largest in the province, being more than 6 km long and almost 400 m deep. The best-known cave in this area is certainly the Grotta Gigante, one of the most popular with tourists. The Grotta Gualtiero Savi extends for almost 4 km. In Friuli, there are some interesting caves in the Julian Prealps, mainly extending inside alternating carbonatic and flyschoid layers. For this reason, most of them are horizontal: the Grotta Nuova di Villanova, partly equipped for tourist visits, is more than 7 km long. Four km is the length of the San Giovanni dAntro cave, which has at its entrance a small church and Medieval fortifications resting on other structures going back to Roman times. The Canin massif in the Julian Alps, composed of Upper Triassic deposits, was the first high-altitude karst to be systematically explored, and the Col delle Erbe system (Abisso M. Gortani and others), 880 m deep and 22 km long, was for many years the deepest in Italy. Now the area counts dozens of caves exceeding -500 m (Foran del Muss complex, -1100 m, 15 km; Abisso Led Zeppelin, -960 m, Abisso ModonuttiSavoia, -805 m, etc.). In the Devonian Calcari di Scogliera (limestone cliffs) One of the very many high-altitude shafts in of the Carnian Alps is the Monte Mount Canin (Friuli) Cavallo di Pontebbe complex, -690m, which extends into Austrian territory. In the Cretaceous limestone of the Carnian Prealps, the largest caves are the Risorgiva di Eolo, more than 5 km long, and the various subterranean systems of the Pradis area (La Val-Noglar-Mainarda complex, nearly 7 km). New zones of interest have been identified inside the Parco delle

Grotta Nuova at Villanova (Friuli)

Dolomiti Friulane (e.g., Landri Scur di Claut, 4.8 km). Other interesting phenomena are those of the Cansiglio plateau, bordering the Veneto, where the Bus de la Lum was explored between the late 19th and early 20th centuries. However, the most interesting feature is the Gorgazzo spring, of Vauclusian type (i.e., its waters entirely fill the conduit), which collects the copious waters from the plateau and which, although starting at 47 m above sea level, has been explored to a depth of more than 100 m without the bottom being reached. It is a very dangerous cave, and several speleo-divers have lost their lives in it. It is now explored by means of small submarine robots.

41

Liguria. For its climate and geographic position, Liguria has many caves with archeological and paleontological deposits, some of great significance. At the frontier with France, the famous Balzi Rossi caves are among the most important prehistoric sites in Italy. Equally interesting are the Toirano caves, which may be visited by tourists. One curiosity is certainly the Grotta del Treno di Bergeggi, which opens off a railway tunnel! In the limestone of Mt. Tampa, near Giustenice (Savona) is the Grotta degli Scogli Neri (Black Rock Cave), the longest in Liguria (almost 5 km). Near Finale (Savona) is the Arma Pollera-Arma do Buio complex, two caves linked by a siphon. The former cave is also an important prehistoric site. At the frontier with France is the Grotta della Molesa (Imperia) which, at -253 m, is the deepest in this area. Large-scale explorations are currently in progress in the Balbiseolo cave (Bardineto), a downstream collector of the Buranco Rampion: speleogists have already discovered almost 5 km of galleries and are enthusiastic about future prospects. Emilia Romagna. In the Appennines, a long but discontinuous belt of Messinian and Triassic evaporitic deposits outcrops. Gypsum, hosting many very interesting karstic phenomena due to its variety of environments,

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mineralogical aspect and insect fauna, does not contain many concretions and thus is often not particularly attractive from the classical speleological point of view. The most interesting system is that of Spipola-Acqua Fredda, not far from Bologna, more than 10 km long inside an area of such interest that it has been established as a natural park. It is the longest cave in gypsum in Western Europe. The high valley of the Secchia (Reggio Emilia) contains extensive outcrops of Triassic gypsum, in which karstism develops in underground meanders, i.e., watercourses penetrate evaporatic layers and then return, after an underground course of greater or lesser distance, to their natural aboveground beds. The Fonti di Poiano are some of the most copious karstic springs in the Appennines. The deepest gypsum cave in the world, the karstic system of Mt. Caldina, with a difference in altitude of 265 m and a length of 1 km, has recently been explored. In the gypsum outcrops of Brisighella are the Abisso Fantini, Grotta della Tanaccia, and Rio Stella-Rio Basino complex, one of the few fully protected and conserved Italian caves. Tuscany. Speleologically, Tuscany means the Apuan Alps: the Mecca of cave exploration in recent years, but also one of the places where protection of the subterranean world causes some of the most serious problems. Many caves are threatened with destruction due to local quarrying for Carrara marble, some of the highest-quality stone to be found anywhere in the world. The Abisso Paolo Roversi, more than 1250 m deep, conceals the Black Hole, a single shaft more than 300 m deep and so wide that its walls cannot be seen while speleologists descend it. The Abisso Olivifer reaches -1215 m, and the Monte Corchia complex which, at more than 50 km, is the longest in Italy - goes down to -1190 m. The Saragato-Aria Ghiaccia complex is another of the small number of caves exceeding -1000 m. Of historical interest is the Tana che Urla (Screaming Lair), near Fornovolasco. Although this cave is not particularly long, through it flows a watercourse which was studied by Vallisneri during his work on subterranean hydrology in the 18th century. In the same area is the Grotta del Vento, open to tourists. The hydrothermal Giusti cave, near Monsummano Terme, contains waters at a temperature of 35C.

Umbria and Marches. The Appennines in this area have extensive outcrops of Mesozoic limestone. In the Monte Cucco massif is the cave of the same name, 920 m deep and more than 31 km long. But the most famous caves here are those of the Gola di Frasassi, not far from Jesi. The Fiume Vento complex (more than 23 km) is one of the most beautiful subterranean systems in the world: in part accessible to tourists, more than 400,000 visitors come to admire it every year. This is a cave in which recent works aimed at respecting as much as possible the requirements of the underground environment and above all at safeguarding the peculiarities of its microclimate. In the same area is the Grotta di Frasassi, at the entrance to which is a fascinating little temple built in the early 19th century to a project by the architect Valadier. Abruzzo and Molise. Until now, the carbonatic massifs of the Gran Sasso and Maiella have only partially revealed their enormous potential. One example is the Cavallone cave (open to tourists), in which a keen nose can smell the hydrocarbons present in the rock. Near the city of LAquila is the Risorgiva di Stiffe, more than 1.5 km long, open to Fiume Vento complex (Marche) tourists. Larger caves are in the Matese, a carbonatic complex south-west of Campobasso, where the Pozzo della Neve reaches the respectable depth of 1045 m, and the Abisso Cul di Bove extends for almost 4 km. Latium. The massif of the Monti Lepini hosts the Grotta del Formale (more than 4 km) and the Abisso Consolini (about -600 m). Other karstic areas of great potential are those in the Monti Aurunci, where the Abisso Vallaroce reaches -565 m; Monti Ausoni, whose waters are collected by the Pstena caves, more than 3 km (open to tourists); Monti Simbruini, with the Grotta di Bellegra; and Monti Ernici, with the Abisso degli Urli (-610 m, more than 3 km). Some remains of Neanderthal Man were discovered in 1939 in the Grotta

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Guatteri on the Circeo promontory. Other caves of paleo-ethnological interest also occur, mainly near Latina. Campania. The Alburni massif was the first karstic complex to be systematically explored in Southern Italy. Extensive karstic areas contain the Grava dei Gentili (-484 m), Grava del Fumo, Grotta del Casone Vecchio, and others. At the foot of Monte Alburno is the Grotta di Castelcivita (open to tourists) recently found to be linked with the active system (Ausino) of water collection of the overlying massif, forming a complex more than 6 km long. Grottoes, or sea caves, are of great importance: the best-known in Italy are the famous Grotta Azzurra on Capri and that of Cape Palinuro. Again at Palinuro is the Grotta di Punta Galera, more than 1 km long, crossing the narrow promontory almost completely. Of particular interest is the fact that many of the grottoes of Cape Palinuro have been, and partly still are, subject to the circulation of sulphur-bearing fluids which was the main factor influencing the formation of their cave deposits. Interesting prehistoric sites are found in many caves between Palinuro and Camerota. In the Cilento area is the underground course of the river Bussento which, near Casella in Pittari and Morigerati, enters the carbonatic massif and leaves it again more than 4 km downstream. Until a few years ago, this stretch of subterranean flow was used to transport sewage!

Puglia (Apulia). This is one of the regions where carbonatic rocks outcrop most extensively. Three separate karstic macro-areas have been identified: Gargano, Murgia, and Salento. Superficial forms are very widespread on the Gargano promontory, which also contains one of the largest dolinas in Europe, the Dolina Pozzatina, more than 600 m across and 100 m deep. The Pulo di Altamura and Pulo di Molfetta lie on the Murgia plateau. Also of importance are the Gravine, long karstic valleys which mainly cut the Murgia Tarantina plateaus. The best-known cave, the Grotta di Castellana, discovered by Prof. Anelli in 1938, is today one of the most popular Italian caves open to tourists. The deepest cave in the region is the Grava di Campolato (San Giovanni Rotondo),

45

Grotta di Castellana (Apulia)

Grotta di Porto Badisco (Apulia): wall paintings

One of several grottoes along Cilento coast (Campania). Note concreted paleosoils at entrance: many caves were used by prehistoric man

more than 300 m deep. Many Apulian caves are interesting not only from the purely speleological viewpoint, but are also of special importance for the paleo-ethnological evidence they reveal. Examples are the Grotta Paglicci (Gargano), frequented by man since Paleolithic times, Grotta dei Cervi at Porto Badisco in Salento, and the Scaloria-Occhiopinto complex at Manfredonia, used in Neolithic times for water worship. Of great significance is the recent discovery of human remains in the Grotta di Lamalunga near Altamura. Lastly, mention must be made of caves used by man for purposes of worship: from paleo-Christian until Late Medieval times, caves were lived in by hermits, used as crypts, and even devoted to the cult of the Archangel Michael. Unfortunately, much of the deep karstism in Apulia has been profoundly and severely altered by man in modern times. As well as total destruction of caves due to quarrying, there are abundant cases in which city sewage has been sent into natural cavities. Very many caves have been used as dumps for all

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kinds of materials, sometimes even toxic waste. A significant example is the Grava di San Leonardo (San Giovanni Rotondo), full of tons of expired medicines and other hospital material, which could very easily pollute the underlying water-table. Basilicata. Near Maratea, in the narrow Tyrrhenian belt of this region, is the Grotta del Dragone, with about 3 km of galleries, a fossil exit of the karstic complex of Monte Coccovello. Although there have been reports of exhalations which might have made exploration dangerous, due to the presence of a waste dump in a dolina, the problem now appears to have been solved. Particularly well developed here is exploration of grottoes, and the Centro Europeo di Speleologia Marina is located here. Along the coast is the Grotta delle Meraviglie, partly open to tourists. Nearby, at Trecchina, is the Grotta di SantAngelo, about 500 m long, full of decorative concretions and fascinating details. Also important is the Grotta di Castel Lepre near Marsico Nuovo (almost 2 km), and even larger is the Grotta I Vucculi at Muro Lucano. Particular mention must be made of the Matera area, where natural caves were extensively used as dwellings for humans - in fact, some are still occupied! Calabria. The Abisso del Bifurto in the Pollino massif reaches a depth of -683 m (once the Italian record). Another cave of interest is the Grotta di Serra del Gufo. On the Ionian side of Calabria are the Grotte di SantAngelo near Cassano: described as long ago as 1571, they extend for more than 2 km and are linked to a system of sulphur springs. Hot sulphur-bearing waters are exploited as thermal waters in the Antro delle Ninfe near Cerchiara. The Tyrrhenian side contains many caves of archeological interest. Near Crotone is the Grotta di Samouri Tour, more than 2 km long, through which an underground river flows. Sicily. The carbonatic massifs of the island (Nebrodi, Iblei, Madonie) contain the Grotta di Pantalica and many other karstic systems, but most characteristic of all are certainly the lava caves, many of which open on the flanks of Mount Etna. The best-known is the Grotta dei Tre Livelli (more than 1300 m long, more than 300 m deep), but the presence - at the foot of a volcano - of a subterranean

glacier in the Grotta del Gelo (frozen cave) is truly out-of-the-ordinary. The Abisso Profondo Nero has more than 1 km of galleries. The Sicilian speleological potential is completed by caves in gypsum, like the well-known Grotta di Santa Ninfa, with more than 1 km of galleries richly covered with concretions, and SantAngelo Muxaro. Mention must also be made of the Grotta di Monte Kronio at Sciacca. This is an important prehistoric site, used in the past as a place of worship and now practically closed to exploration, due to hot vapours which bring the internal temperature to more than 38C. This cave can now only be visited with special equipment. Sardinia. Extensive outcrops of carbonatic rocks mean that Sardinia is one of the most interesting areas for speleologists, with often very long horizontal caves decorated with concretions. In Supramonte di Oliena and around Dorgali, there are large subterranean systems like Su Bentu-Sa Oche (15 km) and San Giovanni Su Anzu (11 km). But the largest is Codula de Luna, formed of a join between Su Palu and Su Spiria, more than 38 km long. There are also many sea grottoes, very popular with tourists, such as the Grotta di Santa Barbara (Sardinia) Grotta di Nettuno at Alghero and the Grotta del Bue Marino at Dorgali. Short history of speleology and speleological structure in Italy A brief history of Italian speleology is extremely difficult - above all when we realize that, since the Second World War, Italy has been the country in which this science developed most. For thousands of years, caves were lived in and used for protection by man and animals. Later, they were privileged places for religious or pagan rites, and have always been treated with respect - perhaps with more fear by Medieval people than by prehistoric man, who transformed sometimes highly inaccessible caves into true temples of art (e.g., Grotta dei Cervi, at Porto Badisco in Apulia).

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Dante himself writes that he entered Hell through a cavern, and later Leonardo da Vinci explored a cave with fear and desire ... to see if there were something to admire inside it. Definite evidence of explorations between the 13th and 15th centuries is given by dates carved in the famous Grotte di Postumia (Adelsberg, now in Slovenian territory). But it was in the 16th century that more attention began to be devoted to describing caves: Piero Coppo and Alessandro Alberti studied the course of the Timavo, the underground river flowing between Slovenia and Italy. The following century saw research concentrated mainly in the Karst area. The 17th century ended with the masterly work by Athanasius Kircher, a German Jesuit, who published his Mundus subterraneus in Rome. This was a sort of manual of speleology, describing not only exploratory methods but also presenting speleogenetic theories, classifications, and much else besides. Kirchers ideas were sometimes bizarre or highly imaginative, but they did represent an enormous step forward in their attempt to interpret phenomena and classify forms. It was the following century, with the work of Vallisneri on the origin of springs, that saw the first scientific approach to karstic hydrology. But karstism as such still did not exist. The great schools of karstism came into being in the 18th century: French, Austrian, Slovenian and Italian. The last three found a natural field for comparisons in the area near Trieste, then a port of the Austro-Hungarian Empire. The authorities were faced with the problem of guaranteeing proper water supplies to a rapidly expanding city. The most logical solution appeared to be that of exploiting the waters of the Timavo, but this could not be done without knowledge of its true underground course. The river disappeared into the Abisso di San Canziano and reappeared on the surface near Duino - but how, and where, did it flow for the dozens of kilometres which separated these two places? The main explorer was Antonio Federico Lindner. He first touched the bottom of the Abisso di Padriciano at -227 m and later, in 1841, established the then world record by reaching -329 m in the Abisso di Trebiciano, where the subterranean stretch of the Timavo passes. However, the true father of Triestine speleology was the great Eugenio Boegan who, in 1883, founded the Caves Commission of the Societ Alpina delle Giulie, the first speleological association in the world, which attracted first-class researchers like Bertarelli and Timeus. But Trieste was then in Austria, and the flag of speleology in the Kingdom of Italy was that of Friuli. In 1897, at Udine, the Circolo Speleologico e Idrologico Friulano was founded by Marinelli and supported by famous figures as Gortani, Feruglio, Desio and De Gasperi. The Circolo published Mondo Sotterraneo, the first issue of which appeared in 1904 and which is still today

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Giovan Battista de Gasperi during 1911 exploration of Grotta Doviza (Friuli)

regularly published. The association focused on the Julian Prealps area and supported the exploration, among others, of the Grotta Doviza which, until 1914, was considered the longest in Italy. Today, reading Grotte e Voragini del Friuli (Caves and chasms of Friuli) by De Gasperi (who, when he wrote it in 1915, was only 24 years old) or Duemila Grotte (Two thousand caves) by Boegan and Bertarelli, published in 1926, we are struck by the quantity of high-quality information which these volumes contain. In the meantime, speleological associations were founded in Milan and Rome and, in 1903, Michele Gortani founded the Societ Speleologica Italiana in

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Bologna, mainly as a tool for scientific research, which published a few numbers of the Rivista Italiana di Speleologia. But the moment was not yet right for national coordination of speleology, which passed through a period of dualism between exploration and research. Although the First World War greatly affected many associations (for instance, the Circolo di Udine emigrated to Florence after the defeat of Caporetto), it did lead at last to the conquest by Italy of all the classic Karst territory, enabling Friuli-Venezia Giulia to maintain its position of speleological preeminence on a world scale. But gradually successful explorations throughout North Italy followed: from the Abisso Bertarelli (Karst; 450 m), to the Spluga della Preta (Veneto), the Abisso Guglielmo in Lombardy, to the -541 m of the Antro del Corchia in Tuscany, reached in 1934. At that time, limitations were certainly not dictated by the capacities of the explorers, but only by logistic problems and the weight of all the necessary materials which had to be transported into the bowels of the earth. Explorations which nowadays can be undertaken in a few hours by a team of three speleologists, at that time required months of preparation, dozens of men, and several days hard work by the actual exploring team. These

Descent by rope ladder into Viganti shaft (1949)

problems, essentially the same, remained, although with improved materials and equipment, until the 1970s, when new climbing techniques with advanced kinds of ropes meant that transport of heavy materials could be greatly reduced while safety measures were respected. And high-altitude caves could be explored, in areas already well-known potentially by the fathers of modern speleology. Returning now to the period between the two world wars, an essential role was played by the Istituto Italiano di Speleologia, with headquarters in the Grotte di Postumia, which created a Cave Registry, published the important review Le Grotte dItalia, and coordinated exploration and research by various expert groups. This favoured the development of speleology in Italy and its expansion to the southern regions, but after the Second World War the assets of the Institute were dispersed, the great speleologists of the past died, and Italy also lost not only most of the classic Karst, but also its supremacy in the speleological field. The dense network of speleological associations started functioning again around 1950, when the Societ Speleologica Italiana was founded in Verona, viewed as an association for actively interested speleologists. A few years later, Gortani re-established the Italian Institute of Speleology in Bologna, this time focusing principally on scientific research: only after years of strained relations did the two associations manage to agree. In the 1960s, partly thanks to improved techniques, new explorations were undertaken, with the record for depth (-878 m) going to the Spluga della Preta (Veneto). These were the years in which the Apuan Alps, Canin and Marguareis were explored, still today the subject of constant surveys. However, in the same years, several deaths occurred which led, among other things, to the setting-up of the Speleological Rescue Service within the Alpine Rescue Service. In those years, speleology was only well-developed in some regions (FriuliVenezia Giulia, Piedmont, Emilia-Romagna, Tuscany) and was somewhat restricted: there was a sort of distance, a coldness, between the two branches of speleology, exploratory and academic. Fortunately, this state of affairs gradually subsided. Over the next few years, speleological structures and organizations began to develop and, although each group maintained its autonomy, regional federations were set up and all administrative bodies followed the path opened in 1966 by the Friuli-Venezia Giulia Autonomous Region which was the first to issue legislation for the protection of karstic phenomena, support speleological activity, and officially constitute a Catasto Grotte (Cave Registry). Now, almost all Italian regions have their own federations and functioning cave registries, and more than 300 groups are active.

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Biospeleology
LEONARDO LATELLA FABIO STOCH

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Biospeleology in Italy Origins. Initial interest in underground fauna in Italy goes back a long way: already in the second half of the 16th century the Vicenza-born Gian Giacomo Trissino wrote to Fra Leandro Alberti, describing some tiny shrimps found in the Beric Hills (he was speaking of amphipod crustaceans, now known as Niphargus costozzae). But he was simply curious, he was not carrying out scientific research. The first true mention of cave-dwelling animals was made in 1689, when Baron Johan Weichard Valvasor spoke of a ptit dragon found in a spring in Carniola. In 1768, Joseph Nicolaus Laurenti described it under the name of Proteus anguinus, and set it in the group now called caudate amphibians, although he did not recognize it as a true troglobite. The first scientific description of an animal really living inside a cave was made by Ferdinand Schmidt who, in 1831, described Leptodirus hohenwarti, a troglobic insect found in the Postumia caves. It is perhaps from that date that science made its official entry into the underground world. A long series of zoological studies began in the 1830s, aiming at knowledge of cave-dwelling animals. And it was in the Postumia caves, open to visitors for the first time, that the fulcrum of modern biospeleology may be said to rest. The first scientific works on subterranean fauna were published in Italy in the first half of the 20th century. Among these pioneering works were the studies of Ruffo on cave-dwelling fauna in the Verona region in 1938, and those of Denis on spring-tails in Italian caves in 1931. But a wealth of literature was soon created, mainly from taxonomists who were rapidly discovering the incredible variety of subterranean fauna. After approximately ten years of inactivity, principally due to the Second World War, work began again with renewed enthusiasm. The 1950s saw the works of Patrizi and Cerruti on cavedwelling fauna of Latium and surrounding regions (1950) and Latium and Sardinia (1953), of Conci on Venezia Tridentina (1951), Ruffo on Apulia (1955) and Franciscolo on the Savona region (1955), to quote only a few of the most important works of general interest. In the last 40 years, the number of Italian zoologists who have dealt with biospeleological aspects and subterranean

Sphaeromides virei

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Distribution of trogloxenes, troglophiles and troglobes in caves is influenced by variations in light, ease of access, and type of micro-environment

relics obliged to colonize the subterranean environment as a result of adverse living conditions, and at viewing the underground world as poor in species, simply did not correspond to reality. On the contrary: caves must be considered as habitats governed by the same rules as surface environments where only a few limiting factors prevail (in this case, the absence of light and reduced availability of resources). However, this does not diminish interest in caves which, due to the simplicity of their communities, are considered as excellent models on which to apply and test theories of general interest. Classification of subterranean environments and cave-dwelling fauna Terrestrial subterranean environments. The term cave ecosystem was long applied to that part of the subsoil accessible to man. In recent decades, however, it has been realized that caves understood only in a speleological sense are not the only environments in which animals live. Terrestrial cave habitats may be summarily divided into three large groups: true natural caves, i.e., accessible to man; the superficial subterranean environment (MSS for French authors), and artificial caves created by man. True natural caves are areas with spaces, sometimes very large, which may host quite large animals, like bats and amphibians. In them, various ecological factors influence the environment and consequently the distribution of the animals which live in it. The presence of several clearly defined faunal associations may be identified in many cases. Near the entrance, according to season and altitude (this sector is greatly affected by seasonal variations in climate), several animals find refuge in this damp, protected part: some are common birds, like pigeon, wren and chough (the latter mainly in vertical caves at high altitude), which often nest near cave entrances, or mammals like dormouse, caudate amphibians and anurans, molluscs, insects and other arthropods like isopods and diplopods. From the penumbral zone as far as the beginning of the dark zone, the walls of caves host animals belonging to various taxonomic groups making up the so-called wall association. This quite heterogeneous community normally occupies cave walls and ceilings and is made up of habitual cave-dwellers, even in the more internal zones (troglophiles) and others which only visit caves or use them as summer or winter quarters (trogloxenes). They include common dipterans, lepidopterans, rhaphidophorids and gryllids, trichopterans, spiders and phalangids (harvestmen). The composition of this association, to a lesser extent, is influenced by several factors such as the structure of the cave, the altitude at which it opens, and seasonal variations in climate. The internal zones of caves

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waters has grown; but this is recent history, and biospeleology has now reached the rank of Science. Many Italian researchers have given new impetus to biospeleology over the last decade. Their interest has essentially focused on three wide-ranging topics: taxonomy (discovery and description of new cave-dwelling species), evolution and biogeography (partly with the help of molecular biology techniques) and, to a lesser extent, ecology. After pioneering beginnings and the establishment of a separate discipline, the various branches of biospeleology began to be integrated within the wider themes of general ecology and the theory of evolution, and the subterranean environment is losing that definition of special habitat which was applied to it until relatively recently. It was realized that many models describing specialized life in caves as an evolutionary cul-de-sac, at treating cave-dwelling animals as scattered

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may be used by colonies of bats which, with their nutrient-rich droppings, leave guano, which is an important source of food for very many species (guanobes). Guanobes may be found around or above the heaps of guano which are built up under the colonies, the most common being oligochaetes, mites, spring-tails, beetles, dipterans, snails and microlepidopterans. Like the members of the wall fauna, the species making up the guanobe association may belong to several ecological categories, which live around an extremely important source of food in an environment which Concretions in large hall in Grotta Nuova at would otherwise be poor in resources. Villanova (Friuli) The guano populations change according to the movements of the bats which produce it and with which they are closely associated: the animals found in guano vary according to its freshness and quantity. In the deeper zones of caves, where temperature is constant, humidity high, and darkness total, we find the true terrestrial cave-dwellers. They may be found on the soil and under stones on the ground, on stalagmites or near organic debris. They are usually small, and often move into the superficial subterranean environment: the second of the subterranean areas mentioned above, formed of micro-cracks in fractured rock under the last layer of soil, often linking caves with the rest of the karstic network which we cannot enter. Recent studies have shown that some species of troglobitic beetles spend long periods of the year in this environment, only living in caves at certain seasons. However, the superficial subterranean environment does not always mean caves as such: there are many areas where numerous animal species manage to survive. The third subterranean environment is that of artificial caves. Not so many studies on their populations have been carried out as on natural caves, although much faunal work was carried out starting from the 19th century. Several new species living in and on materials collected from artificial caves have been described; one example is the rhaphidophorid orthopteran Dolichopoda geniculata, found for the first time in a basement in Naples in

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karst cavity

top soil layers

MSS

fissured rock

Diagram showing terrestrial subterranean environmnts

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1833 and described by Costa in 1860. Studying specimens collected from the cellars of the castle of Brescia allowed description of the leptodirine beetle Boldoria ghidinii. The foundations of the Certosa di Pesio, excavated in carbonatic rock, were chosen by Sella as the typical location of the trechine beetle Duvalius caranti. In 1977, Karaman and Ruffo described a new amphipod living in catacombs in the province of Siracusa, and called it Echinogammarus catacumbae (now Tyrrhenogammarus catacumbae). There are many other examples revealing the biospeleological interest of these environments. Among these, mention must be made of the particular importance of artificial caves in only slightly karstifiable carbonatic rocks (quarries, cellars, tunnels excavated for military purposes, etc.), which open a window on habitats which could not otherwise be visited by man. Aquatic subterranean environments. From the viewpoint of ecology, aquatic subterranean environments are divided into two large categories: waters circulating in porous rocks (alluvial terrains), and others circulating in fractured or karstified rocks. They are inhabited by separate fauna (only along transition zones, or ecotones, do we find both types) and the organisms which live in them have made many different kinds of adaptations in order to survive. Here, we will not describe in detail waters circulating in alluvial terrains, but will simply recall that they too contain fascinating organisms (called phreatobes when they are exclusive to that environment), still imperfectly known. Their bodies are considerably modified to allow them to live in spaces between grains of gravel or sand (interstitial environment). We consider here waters circulating in rocks - circulation which may occur as a result of permeability due to cracks (in non-carbonatic rocks like sandstone, or igneous or metamorphic rocks) or karstism (i.e., waters circulating in micro-cracks or karstic conduits). Within the framework of karstic water circulation, the three zones distinguished by hydrogeologists are of great ecological interest. In the vadose zone (with prevailing vertical flow of percolating waters), the fauna is adapted to life in micro-cracks, and may occasionally be collected in pools (the gours of French authors). These environments generally contain minute organisms sometimes only a few millimetres in size, with long or flat bodies, fitting their life-style (nematodes, oligochaetes, and above all copepod, syncarid and amphipod crustaceans), although larger species are also found, like some Niphargus which may exceed 2 cm in length. The fossil karstic areas in which we find only the vadose zone have undergone a high degree of geographic isolation,

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VADOSE ZONE

PHREATIC ZONE

INTERSTITIAL ENVIRONMENT

Classification of aquatic subterranean environments

which favoured speciation: the fauna is rich in endemic species. Instead, the epiphreatic and phreatic zones, where water flow is prevalently horizontal, generally host a higher number of species, some of which are adapted to life in large areas of free water: this is the reign of the large crustaceans (isopods, amphipods, decapods) and of the proteus. But there are also small benthic (i.e., bottom-living) organisms and, more rarely, plankton (living suspended in waters). Classification of cave-dwelling organisms. Not all organisms living underground have adapted to life in this particular habitat in the same way. For this reason, ecological categories which take into account the adaptive characteristics of each species have been created. For cave-dwelling organisms, we use the classification proposed by Schiner in 1854, adopted by Racovitza, Pavan and others, and revised by Ruffo in 1955. Cave-dwellers are subdivided into four categories. Animals which are found in caves only accidentally, like those brought in by waters or which fell inside shafts in vertical caves, are called trogloxenes; those found more regularly in caves are called troglophiles, a composite category which actually contains

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although they do not refer exclusively to cave fauna. So phreatobes, which live in the hyporheic interstitial environment and in phreatic layers of alluvial soils, are also stygobites. It is always appropriate, therefore, when using the term stygobite, which implies a series of specialized adaptations to the subterranean environment, similar to those of troglobites, to specify whether they are cave-dwelling organisms or not. It is not always easy to attribute all cave-dwellers to these categories, since there are many intermediate cases. But the classification has great practical value, and allows us to apply and interpret models widely used in biospeleology. Methods used to study organisms living in the subterranean environment Faunistic studies. Terrestrial fauna. Cave-dwelling organisms may be collected to determine which species populate a certain cave (faunal studies), or to select specimens of one or more species for laboratory work. Direct sampling uses entomological tweezers (with very soft tips, which do not damage specimens), an aspirator for the smallest and fastest organisms, and a brush for even smaller ones. For species which are difficult to obtain directly, baits may be used. Placed at several points of the cave and protected by stones, they attract not only cave animals but also their predators. Excellent baits are meat, fish, strongsmelling cheese and rotting fruit, left in place for not less than one week. Except in the case of special and fully justified scientific research, traps are definitely not recommended: if they are left in place for long periods of time, they can cause mass mortality and consequent considerable impoverishment of the fauna of a cave, resulting in disequilibria of the cave-dwelling community. A Berlese funnel may be used in the laboratory to collect small animals hiding in soil and guano. This device is composed of a sieve with a grid of 0.2-0.3 cm, on which the soil or guano is laid, above a funnel supported by a tripod. At the base of the funnel is a can containing a preserving fluid. As the superficial layers of earth or guano dry out, the animals, which are all hydrophilous (waterloving), descend to still damp layers and fall through the sieve into the funnel, and thence into the preserving fluid. Animals may be preserved and transported in vials containing 70 alcohol (which must not be of denatured type, or specimens become hard), or a can containing cork or wood shavings (sprayed with acetate ethyl) for beetles.

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Classification of organisms living in natural caves: trogloxene (frog), troglophile (bat), troglobite (olm)

two separate groups of animals. Those found in caves only during certain periods of their lives and which do not show particular adaptations are called subtroglophiles: examples are some dipterans (culicids, limonids) and bats, which only inhabit caves in some periods of the year or during the day. Instead, eutroglophiles are animals which, although definitely preferring the subterranean habitat and showing special morphological and physiological adaptations to it, sometimes live and in some cases even reproduce outside it: examples are cave-dwelling grasshoppers. Lastly, the troglobites show the most extreme adaptations to cave life: they are born, reproduce and die only inside caves or similar habitats, like the superficial subterranean environment: they are the true cave-dwellers. Organisms living exclusively in subterranean waters take the name of stygobites (by analogy with troglobites; also used are the terms stygophiles, eustygophiles and stygoxenes). These apparently difficult words are in fact evocative of the Styx, the river which, according to Greek mythology, mortals had to cross in order to enter the Underworld. They are now commonly used,

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Star Cedat spring (Natisone valley, Friuli)

It is essential that one vial should be used for each cave and that it should contain a card, written in pencil or even better in china ink (which does not fade), indicating the region where the cave opens, the nearest village or town, the name of the cave and, if known, its registry number, the date of collection and the name of the collector. Cave samplings are also made to study the ecology of cave-dwelling Measuring a cave-dwelling grasshopper before communities. Examples of this type of releasing it research are studies on the dynamics of certain populations over the year, and analysis of how the cave is used by one or more species over time and their interactions. For these quantitative samplings, some of the instruments mentioned above may be used, like Berlese funnels and traps. In this case, samplings must be standardized, so that they can be repeated in time and compared with each other. However, it is not always necessary to collect organisms for these studies, since it is often possible to carry out counts and monitoring directly in the cave, without disturbing the animals excessively. Aquatic fauna. Fauna in subterranean karstic caves may be collected with special methods which vary according to whether the study concerns vadose, epiphreatic or phreatic waters. Each sampling should always include data on the main chemico-physical parameters (at least temperature and, if possible, conductivity) and notes on the type of environment sampled. Emphasis is placed on the fact that any sampling lacking exact indications of date and place (again accompanied, if known, by the cave registry number) cannot be used in any way for research, and is therefore completely useless. Hand-fishing. This is the only effective method of capturing specimens of a certain size, which can be picked up with tweezers or small aquarium nets. However, the method only collects the largest species and neglects the most important fraction of the fauna, composed of organisms which may be invisible to the naked eye. Filtering. Indispensable for collecting very small animals is a plankton net, modified for cave use. This is a very fine (0.07-0.1 mm) nylon net, with a cup at the base for collecting detritus and organisms once the water has been filtered. Nets used in caves often have handles and are shorter than traditional

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ones (to stop them becoming caught up on rocky projections, etc.), with semicircular mouths about 20 cm across, and reinforced frames and corners covered with strong cloth or metal - for extra-long life in cave environments. In the vadose zone, water is filtered from pools or percolating flows with nets, and if necessary collected it with a small hand pump or a rubber syringe. In streams where water quantity permits it, nets may be used; detritus from upstream is removed, and only sediment brought down by the current is used. In siphons and large pools, the method is the same as those for surface waters, water being filtered through the net but care being taken to remove the bottom detritus and to scrape the cave walls. In view of the low density of specimens in underground water and the rarity of some species, samplings must be carried out over a long period of time and repeated in various hydrological conditions. Sometimes, in the thick gravel deposits of the streams of some caves, it is better to exploit techniques used for interstitial hyporrheic waters of rivers (Bou-Rouch pump, or Karaman-Chappuis technique). To collect molluscs in these conditions, enough sediment must be sampled. Trapping. The commonest traps are simply open cans containing the bait, generally a small piece of meat (liver is excellent, but salami, ham or cheese are also good). The method mainly attracts large crustaceans (isopods, amphipods, decapods) but, as they sometimes arrive in large numbers, the tins must be left open. Devices like lobster pots should not be used: if they are forgotten, like traps for terrestrial fauna, they could cause overkill of rare or local species. Ethics of biospeleological research. Occasional sampling of invertebrate fauna in a cave certainly does not damage subterranean communities, since a cave is only a window on an enormous system of crevices, and organisms which can be collected from it only represent a tiny fraction of the existing ones. For example, during floods, thousands of organisms per cubic metre of outflowing water are transported to the open air and die.

Manual sampling rarely yields more than a few dozen specimens, and its impact on the cave ecosystem is therefore insignificant. Repeated trapping or sampling from the same site is a very different matter. Traps are well-known to attract organisms from sometimes enormous distances, and may cause overkill of rare species and negatively influence the numbers of small populations. They must therefore be used with extreme caution and always under the supervision of experts. A different discourse must obviously be made for vertebrates, which are covered by legislation (e.g., the Habitat Directive, regulated according to Italian DPR no. 357 of September 8 1997, the Bern Convention, and various regional norms) which forbid them to be sampled, killed, kept in captivity, or marketed for profit, and which also forbid damage to their habitual sites of living and reproduction. These regulations impose total respect of all Italian cave-dwelling vertebrates, with particular regard to proteuses (olms), cavedwelling newts and bats, and their habitats. For these animals, useful study techniques - adopted exclusively by experts who have definite scientific aims in view - cover only censuses, photography, and observation of behaviour and feeding strategies. In any case, research is best conducted during the least delicate stages of the life-cycle - for instance, bat nurseries must not be disturbed. Research must be carried out with caution, avoiding not only excessive or inappropriate sampling, but also techniques which may alter the physical environment (excavations or excessive tampering with sites). This is not just for protection: it is also part of that research ethic which all biospeleologists or even just enthusiasts must follow, in order to respect the environment and the organisms they wish to study. Cave vegetation Vegetational seriation at cave entrances. Caves and dolinas have special vegetation: the rapid changes in environmental conditions give rise to the wellknown phenomenon of inverse stratification of flora. A dolina may be compared with a mountain turned upside-down, its summit representing the beginning of a funnel. A classic example of this is the large dolina of the Slovenian Karst, 90 metres deep, called Grande Paradana. The flora found on the walls of this giant funnel is similar to that seen on high-altitude slopes. The first 50 metres shows us a wood of Norway spruce; from 50 to 70 metres the trees are smaller and contorted, and we find abundant rhododendrons, typical Alpine plants, and ferns; to 80 metres, dwarf willows and mosses prosper; from

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Collecting samples of aquatic fauna in a plankton net

66

80 to 90 metres we find only mosses, like those inside caves; and then only ice. The mouth of a cave is a transition zone (an ecotone) between the superficial environment and the underground one. Botanists subdivide caves into three zones according to the intensity of light in them: the entrance zone (subdivided again into external and internal) where we still find phanerogams, with light reduced to one-500th of its intensity outside; the transition zone, again subdivided into an external sector, with cryptogams (mosses and ferns) with a light intensity reduced to onethousandth, and an internal sector, containing some algae and fungi, which extends from the zone with very little light to the beginning of the deep zone, characterized by complete darkness. This vegetational seriation, particularly clearcut in vertical shafts, was related by the Triestine botanist Livio Poldini with the reproductive mechanisms of the vegetal species making up the various belts, highlighting a special aspect of great educational interest. The reproduction of algae, both sexual (i.e., fusion of male and female sexual cells, called gametes) and asexual (spores, or simple cell division) undergoes complex cycles, all essentially linked to the presence of water. In mosses, the sexual phase prevails over the asexual one: mosses with small leaves (gametophytes) produces gametes, requiring water for fertilization, while spores are carried in an urn with a small filament (sporophyte) extending from the gametophyte. The spores, which give rise to new gametophytic plants, are windborne and therefore do not depend on water, with the consequence that mosses depend on water only for sexual reproduction. In ferns, the fronds we see are the sporophytes, carrying spores on their underside: the sexual phase, linked to water, is very short. In phanerogams, the sexual reproduction cycle is associated with flowers and no longer depends on the presence of water. Since the sequence algae - mosses - ferns - phanerogams repeats the evolutionary sequence of plants, we may observe a parallel between what happened during the course of evolution and the vegetational seriation we find in karstic caves. From the bottom of a cave upwards, we can ideally follow the evolutionary stages of plants, which colonized land by progressively freeing themselves from the presence of water. Cave flora. The close relationship of plants with light greatly influences their distribution underground. This is why, with the exception of some fungi and bacteria (traditionally included among flora, even though they are not vegetal species), we cannot speak of truly troglobitic flora, although some species definitely prefer certain areas of caves to others.

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South

North

EXTERNAL ZONE
IT

EA E H ORNB

L O W E R

0m

IV Y

1
THRESHOLD ZONE

2
AS

PLENIU M

-10 m
PO

LY P D I U M O

3
SUB-THRESHOLD ZONE

-20 m

MO

SSES

4
G

RE

EN ALGA

INTERNAL ZONE

-30 m
BL

UE ALGAE

TOTAL DARKNESS
FU N GI

L I M I T

O F FERNS

S U R V I V A L

F O R

F O L L O W I N G

G R O U P S :

PHANEROGAMS

MOSSES

GREEN ALGAE

BLUE ALGAE

Seriation of vegetation along walls of a karstic shaft

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In caves open to visitors and in artificial caves, lit by incandescent or neon bulbs, near the source of illumination we can see flora composed of external species like diatoms, blue and green algae, mosses and some ferns. The managers of some visitable caves have recently been using cold light sources, which do not allow these plant species to grow. Let us now briefly examine prokaryotic and plant organisms which may be found further inside natural caves. As regards bacteria, perhaps the most numerous but least known of the organisms which inhabit the subterranean world, studies have been carried out on some ferrobacteria which intervene in the deposition of iron in the form of hydroxide, and sulpho-bacteria which fix sulphur by oxidation of the sulphurated hydrogen produced by decomposing organic matter or other sulphur compounds present in thermo-mineral waters. Recent research on sedimentary formations has also confirmed the presence of bacteria specific to the subterranean environment. Lastly, blue algae, found in the entrance zones, are photosynthetic bacteria. In Italy also specimens of Geitleria calcarea have been found, a cave-dwelling blue alga present in some caves in central and southern parts of the country, which cannot survive if exposed to sunlight. Some authors consider some laboulbenials, parasitic fungi found on the integument of insects living in very damp environments, as cave-dwellers. This term is not really appropriate, since the laboulbenials which live on cavedwelling animals do not show evident morphological and adaptational differences with respect to those which are parasitic on external species. Italian caves contain the genera Rhachomyces and Laboulbenia, parasites of some carabid beetles; Troglomyces, parasitic on a diplopod; and Arthrorhynchus, which may be found on nycteribid dipterans living on bats. Some fungi, like basidiomycetes, the spores of which are easily windborne, are frequently found in caves on rotting wood. The small white cushions of the zygomycete Mucor mucedo may often be observed on bat guano. Green algae too may be found from the entrance zones, where they form large dark green patches, as far as the more internal parts of some caves, where a few species are found, together with some xanthophyceae. Little may be said of lichens: most of them live in external cave areas and entrance zones. The reason for their scarcity in these habitats is due to their need for light energy since they are symbionts between algae and fungi - and they prefer environments which are not too damp. Instead, far more numerous are bryophytes, mosses and liverworts. Together with green algae, from which they evolved, bryophytes are the plants most

69

Fungi growing in absence of light on rotting organic matter

frequently found inside natural caves. Mosses are visible as far as the boundary of the external sector of the transition zone, where light intensity is reduced to one-thousandth that of the external zone. Liverworts may be found at the entrance of many caves, the most widespread in Italian caves being Lophocolea, Marchantia and Plagiochila. Mosses, more numerous than liverworts, very often grow in the external and transition zones. In our latitudes, the entrance zone may also contain various species belonging to the genera Homalothecium, Ctenidium and Plagiothecium, almost ubiquitous in these parts of caves. The transition zone contains Thamnobryum (Thamnium) alopecurum, the fronds of which become smaller and less ramified as they grow further inside the cave; Isopterygium depressum, observed in zones with luminosity reduced to one-2380th; various species of Fissidens and Mnium, and what many authors consider the cave moss par excellence: Schistostega pennata (= S. osmundacea). This particular bryophyte, which grows on rocks in non-calcareous caves, has special spherical cells acting as lenses, which allow it to concentrate the very small amount of light coming from the outside world on its chloroplasts - cells which work as reflectors, giving the tiny plant a luminescent aspect. Ferns (pteridophytes), water-loving plants, are represented underground by Asplenium trichomanes, which produces spores even in conditions of very little light, and sterile specimens of which may be found even far inside caves; Asplenium ruta-muraria, more external, and preferably found in caves with entrances opening to the south; and other common ferns like Adiantum capillus-veneris (maidenhair), Phyllitis Hepatica (Marchantia polymorpha)

70

Phyllitis scolopendrium

Polypodium interjectum

scolopendrium and polypods (Polypodium vulgare, P. interjectum, and others). Phanerogams, flowering plants, need quite a large amount of light for photosynthesis, and cannot penetrate very far inside caves. Among the very few higher green plants found inside the entrance zone are Geranium robertianum, which flowers in reduced light, and Mycelis muralis. Other phanerogams found here are Oxalis acetosella, Taraxacum officinale and Parietaria officinalis. The particular climatic conditions inside caves high humidity, constant temperature, little light have contributed towards making caves refuges for some species which are now no longer found outside but which were far more frequent in past ages when climatic conditions were more favourable. Examples are the subtropical fern Gymnogramme leptophylla, growing in some caves in central Alpine valleys, and Pinguicula alpina, found in the Alps around 2000 metres and collected in a ravine at 500 metres near the hills around Turin. Ecology and adaptation. and biotic components, and the food web. Every ecosystem, including the subterranean one, has two large components: biotic (associated with living things), and abiotic, i.e., environmental parameters (morphological or physico-chemical). Climatic factors were mentioned in the preceding section, and here only the influences of environmental parameters on the distribution and ecology of cave-dwelling organisms will be illustrated. Light. We have already seen the importance of light in the zoning of vegetation at cave and shafts entrances. Many cave-dwelling species have a

Abiotic

Asplenium trichomanes

Asplenium ruta-muraria

particular aversion towards light, often not tolerating direct exposure for long periods, whereas for other species light does not always represent a limiting factor. Already a century ago, for example, it was known that some stygobites (Monolistra, Niphargus) could regularly be collected in outside karst springs mouths, and that they had not always been transported there passively. True migrations by Niphargus were discovered, from the subterranean environment to the surface, for trophic purposes, i.e., to find safer and more abundant sources of nourishment, obviously mainly at night (but also by day in rivulets in the undergrowth). This behaviour has recently been documented in proteus. Temperature. Normally, as the temperature inside a cave corresponds to the annual mean value of the external temperature, it is a parameter closely connected with latitude and cave altitude. Internal shape, prevailing vertical or horizontal trends, and the presence of several entrances or watercourses also influence the internal temperature of a cave. The role of temperature as a limiting factor for cave-dwelling organisms has long been debated. It was noted that some animals with a high level of specialization, when kept in the laboratory, can tolerate temperature variations of 20-25C. For example, some species of Niphargus, although preferring temperatures between 8 and 14C, can tolerate variations between -0.5 and +24.5, and the development of juvenile specimens is regular between 5 and 18. Similar parameters have been found in some leptodirine beetles, which can only live between -5 and 25, with limits of activity between 0 and 20. Clearly, the thermal optimum varies in different cave-dwelling species, since they inhabit both cold high-altitude caves and hot caves in the Mediterranean area, and lethal limits are less ample in animals more closely associated with underground environments. It has been experimentally demonstrated that some grasshoppers of the species Troglophilus andreinii, living in caves around 16 with definitely troglophilous habits (not totally associated with the cave habitat), can support temperatures around 40. Humidity. In caves, humidity approaches the saturation point and this seems to be one of the most important limiting factors for terrestrial troglobites, many of which prefer high degrees of relative humidity (95-100%). Biospeleologists well know that the drier portions of caves where, for example, there may be air currents, are the poorest in troglobitic fauna. Relative humidity acts on general metabolism, respiration, and absorption of water through the cuticle. Although it seems obvious that stygobites are exclusively associated with water in the liquid state, crustaceans of the genus Niphargus are occasionally collected under stones or seen crossing damp soil, sometimes at quite a distance from water. So they can clearly tolerate its scarcity very well - a phenomenon which

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SOLAR ENERGY PRODUCERS SURFACE CONSUMERS

73

green plants
VEGETAL DETRITUS ANIMAL DETRITUS

animals

GUANO

ORGANIC MATTER DECOMPOSERS

bacteria and fungi

DETRITIVORES

CARNIVORES

Food web in cave environment

Different temperatures in a cave in summer (top) and winter (bottom)

is probably more highly developed in species populating the vadose and epiphreatic zones of caves, where water levels sometimes vary considerably. These species are known to be able to survive, sometimes for months, in small hollows in clay, where only a tiny amount of water is available for respiration. The possibility that these species can move outside water means that they are able to colonize new areas or search for food, but this field is unexplored and opens up fascinating possibilities for study. Biotic factors. Two categories of living creatures are distinguished in surface environments: autotrophs (including producers, essentially green plants), and heterotrophs (consumers, i.e., animals, but also fungi and many bacteria). In caves, as we have seen, autotrophs are rare, and their distribution is generally relegated to the first few metres from the cave entrance, where light penetrates sufficiently and allows at least the development of green and blue algae. In the true subterranean environment, autotrophs are completely absent and food chains are based only on nourishment from the outside. Food is often brought into caves in the form of organic debris only, so that at the base of food pyramids in caves we find the detritivores, primary consumers of decaying animal and vegetal matter. Debris of this kind is present in both soil and water, suspended or deposited on the bottom. Conversely, the higher

links of the food chain essentially contain predators. Caves thus have a simplified food network, since producers and all the phytophages are lacking, and the community is dominated by detritivores and carnivores. Recent studies have aimed at quantifying the role of biotic interactions (mainly predation and competition) in the structuring of simple underground communities. This role is supported by laboratory experiments and field investigations, but its importance in the natural environment is questionable and controversial. The old hypothesis according to which the presence of two or more congener species in the same cave is a very rare phenomenon and incompatible with the principle of competitive exclusion has been at least partially demolished. Recent research has demonstrated that this possibility depends on the structural complexity of the habitat and the existence of extensive interconnections with other caves: finding as many as five or six congener species (as in the genus Niphargus) in the same, perhaps only small, cave is an infrequent but not extremely rare fact. Very few field and laboratory observations have been carried out on more complex cave-dwelling communities and none of them in Italy. The field of competitive interactions in temperate caves is practically completely unexplored.

74

Ecological factors regulating biodiversity. As well as the history of a particular karstic area, biodiversity in caves depends on ecological factors which interact in a complex way. As well as the limiting effects of light, temperature, and humidity, some of the most important are briefly examined here. Heterogeneity of habitat. The structural complexity of the habitat, with consequent diversification of the environment (presence of cracks of various sizes and extents, water circulation) creates many spatial niches which may be occupied by various species and is thus directly proportional to the biodiversity of a cave. Recent research on microhabitats in large karstic systems, some of which has been carried out in north-east Italy, have demonstrated that the various micro-environments sometimes host completely different types of fauna and that the distribution of species in karstic areas is highly dishomogeneous. Area. The size of a karstic area is important, because larger areas obviously host a larger number of species, which may also contribute towards enriching local communities. That is, caves in large karstic areas may be richer in cavedwelling species of equal dimensions as in small ones. This fact also indicates how a cave may have free ecological niches, which are only occupied in certain geographic areas. Stability. Cave environments are generally believed to be stable, with small oscillations in environmental parameters, far smaller than on the surface. It was once believed that stable environments were richer in species, but then it was discovered that this is not so: environmental instability diversifies ecological niches and causes a clearcut succession of communities which differ during the course of the year. For this reason, unstable environments are often richer in species than stable ones, like caves. Productivity and availability of resources. Environments rich in resources generally host richer faunas, although the number of species tends to diminish after a certain threshold has been reached (in environments which are too rich in resources, i.e., eutrophic). Caves are generally resource-poor, although localized accumulations are not lacking - debris at the bottom of shafts, guano deposits, etc. - which may enrich the community. However, in these cases, there is often an increase in troglophiles and trogloxenes rather than in troglobites. Very poor (oligotrophic) caves are known to host poorer faunas; the availability of food therefore plays a fundamental role in regulating biodiversity. Adaptations in cave-dwelling organisms. The various adaptations made by cave-dwelling animals may be studied by analysing their adaptational

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Sun illuminates initial stretch of a natural cave

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responses, which are very often similar in sometimes very different animal groups. They represent interesting examples of evolutionary convergence, and are manifest on morphological, physiological and eco-ethological levels and in the genetic structure of populations. If we analyse the morphological adaptations of cave-dwellers, we first note that some structures clearly increase, whereas others are reduced or sometimes only rudimentary. For example, the lengthening of limbs and appendices is one of the most evident adaptations in cave-dwelling animals. The increased length of the antennae in troglobitic arthropods allows them to play an important tactile role in an environment where darkness is total; a similar role is played by the second legs of some cave-dwelling mites and phalangids. Antenna lengthening is also followed by development of the sense organs they contain (chemoceptors, hygroceptors, tactoceptors, etc.). Increased leg length allows the animal to insulate its body, with its lateral sense organs, from the soil and above all it increases its possibility of perceiving the surroundings. Long tactile bristles are also linked with perception of the environment in which the organism moves (water for some crustaceans, air for other terrestrial arthropods). A series of experiments on some leptodirine beetles has shown the importance of the development, in cave-dwelling animals, of antenna organs which perceive the degree of soil humidity and chemoreception. One of these in particular, called Hamanns organ, assesses the degree of humidity, thus allowing animals to avoid venturing into too dry zones, where they would not survive long. Another characteristic adaptation, associated with the need for dampness in troglobites, is the unusual development of the abdominal surface and wing covers which, mainly in some more specialized trechine and leptodirine beetles, take on a spherical aspect. This modification, called false physogastry, influences respiration and the need of these insects for damp air. And in effect, they have reduced respiratory apparatus (spiracles and tracheae), and respiration is direct between the tergal surface of the abdomen, reduced to a thin membrane, which is located in the respiratory chamber, formed of the convex elytra. In this way, the insect can store damp air under its wing covers and use it for breathing during its movements in drier areas. The most frequently observed physiological changes in cave-dwellers are reduced metabolism, reduced circadian rhythm, low fertility, and increased volume of eggs. Reduced metabolism, among other things, allows the animal to live on less food, which is generally not very abundant in temperate caves. Experiments comparing the consumption of oxygen in external and cavedwelling species of amphipods, isopods, spiders, fish and other groups have demonstrated that cave-dwelling animals consume less oxygen. Again

comparing organisms living above and below ground, the latter show reduced fertility, increased volume of eggs and yolk, and prolonged development of both embryo and adult phases. Another change linked to adaptations to cave life is the loss of circadian rhythm and seasonality, although the latter is not completely abandoned in all cave-dwellers and several troglobites Snail of genus Oxychilus preying on adult show seasonal behaviour associated butterfly (Scoliopteryx libatrix) with the slight climatic changes which occur in some caves or variations in the supply of food from the outside. Studies on American blind fish of the genus Amblyopsis show that brain structures respond to morphological changes and are subject to adaptational modifications. The size of the optic lobes in these fish is smaller in blind cavedwelling species than in ones living outside, and there is also an increase in the size of the telencephalon, brain stem, and semi-circular canals (movement receptors), otoliths (equilibrium receptors), and structures interpreting signals coming from tactoceptors and the lateral line. Population genetics also furnish information on adaptational strategies. This may seem obvious if we recall that genes are the main agents controlling morphological, physiological and behavioural characteristics. The many studies carried out by Sbordoni and co-workers on cave-dwelling species, in Italy and elsewhere, confirm the greater genetic variability of troglobitic species with respect to above-ground ones, and show how this variability tends to increase in troglobites which have long been accustomed to the cave environment with respect to more recent colonizers. The reasons for the differences are the reduced size of the population at the moment the gene flow with the outside is interrupted: the normal level of variability is then recovered as generations pass and the size of the population increases. Together with genetics, population ecology offers interesting information on adaptational strategies; parameters such as population dimensions, spatial distributions and age class structures can easily be measured in cave-dwelling populations. Many data are now available on the ecology of rhaphidophorid orthopterans and leptodirine beetles in Italian caves. Lastly, for about twenty years now, many studies have been carried out on the behaviour of troglobe animals (ethology), in particular on mating behaviour, the search for food, and inter- and intraspecific communications. These

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The proteus, or olm, the only troglobitic vertebrate among Italian fauna

researches too provide information, in some cases still to be interpreted, on the strategies devised by animals colonizing the subterranean environment. Origin, evolution and distribution of troglobites Relics or active colonizers? Much has been written and recently debated in scientific conferences and journals, on the origin of cave-dwelling fauna. There are substantially two main theories: one, developed by many classic authors and summarized in 1991 by Botosaneanu and Holsinger, is based on the concepts of relics and refuge; the other, expressed in definitive form by Rouch and Danielopol in 1987, speaks of active colonization. Most models explaining the origin of troglobitic and stygobitic organisms start from the idea that the subterranean environment is one of refuge. This concept is based on study of temperate-zone caves, which acted as shelters into which the ancestors of present-day troglobites adventured to avoid the climatic vicissitudes of the Pleistocene, with its alternating periods of glaciation and drought. The richness of troglobites in temperate zones does contrast with the faunal poverty of tropical caves, where clearly glaciation was not so much of a problem. This interpretation in particular sums up the views of two important authors, Jeannel (in the years from 1923 to 1956) and Vandel (1964). Jeannel, a famous French expert on cave-dwelling beetles, believed that troglobites belonged to phylogenetically ancient, highly specialized lineages, which found refuge in the subsoil during the worst periods of drought, when they would not have been able to survive on the surface. Vandel too, in a wellknown treatise on biospeleology, expressed the view that cave-dwellers derive from senescent phylogenetic lines, incapable of migrating or coping with climatic changes and thus obliged to take refuge underground. For these

authors, cave-dwellers are relics of now extinct surface-dwelling species and are therefore true living fossils, the title of one of Jeannels books. The origin of aquatic fauna has also been modelled using the concept of refuge, like the evolutionary model associated with marine regression of the Dutch author Stock. Stock believed that, during periods of retreat by the sea, some organisms were stranded in subterranean coastal waters. However, these models have recently been confuted by the researchers Rouch and Danielopol, who have made many criticisms of these hypotheses which are fascinating but, it must be said, to a certain extent naive. Firstly, recent research in tropical areas (Africa, Asia, Brazil) has revealed the existence of highly specialized troglobites in the tropics too. Secondly, the refuge model does not explain why, today too, colonization of the subterranean environment still continues, whith populations of partially depigmented animals with reduced eyes, in the absence of unfavourable climatic conditions. Last but not least, there is the objection that the refuge model implies an external cause for the process of colonization, a sort of thrust towards the subsoil which does not need to be invoked by any scientific requirement. Rouch and Danielopol have replaced the refuge model with that of active colonization, which presents a single, general scenario for subsoil colonization, independent of climatic events, and puts the subterranean habitat on the same footing as others, denying that it is special or hostile. Here, we may speak of true adaptative radiations of some taxa, which give rise to numerical explosions in the number of species in the subsoil (Stochs theory of adaptative zone). For example, there are hundreds of amphipod species of the genus Niphargus, in comparison with the few surface-living amphipod species which inhabit the same geographic areas. Together with adaptative radiations in the subsoil, which evolved to occupy free niches, we may observe repeated and ongoing phenomena of colonization and invasion of the subsoil (theory of multiple colonizations) by surface-living species. Isolation and speciation. Colonization of the underground world is followed by speciation, but it is not a direct, compulsory consequence. In order for speciation to occur, the karstic massif must be isolated, with interruption of the gene flow towards the surface or adjacent karstic massifs. The presence of many troglophiles and stygophiles indicates that this process only occurs in some cases. Speciation in caves is always allopatric, i.e., due to isolation; we have no evidence of speciation events of any other type. Speciation in cave-dwellers has been studied in many taxa, sometimes by

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MARINE BENTHIC SPECIES PHASE 1: VERTICAL TRANSACTION (ACTIVE) MARINE INTERSTITIAL SPECIES

PHASE 1: HORIZONTAL TRANSACTION (ACTIVE)

PHASE 2: VERTICAL TRANSACTION (ACTIVE)

SPECIES LIVING IN SUPERFICIAL WATERS

PHASE 2: HORIZONTAL TRANSACTION (ACTIVE, RARELY PASSIVE)

SPECIES LIVING IN SUBTERRANEAN WATERS

Colonization of subterranean fresh waters

molecular biology, but a simple glance at the taxonomy of the groups which include cave-dwellers and their distribution reveals the incredible success of some of them in originating new species and in adapting themselves to the subterranean environment in such a highly specialized way. The consequence of isolation is the formation of many endemic species, i.e., with distribution limited to a restricted geographic area, sometimes a single karstic massif, sometimes only a single cave. This tendency towards endemicity means that cave-dwellers are some of the most important components of Italian fauna, and their importance is even sanctioned by law, since one of the criteria of the Habitat Directive of the European Community when choosing species and habitats to be protected is precisely that of their endemicity. Biogeography of Italian cave-dwelling fauna. To subdivide Italian karstic territory into areas which have a certain homogeneity in their troglobitic fauna is definitely an arduous task. The fragmentation of the genera into so many endemic troglobitic species and subspecies means that, in reality, every karstic massif may be considered something unique, a cradle of species which do not exist in any other place on Earth. However, if we examine relationships among the species and their distributions and correlate them with the paleogeographic vicissitudes which modelled Italy, we see that some karstic areas are well characterized from a faunistic viewpoint. Let us examine this briefly. Dinaric karst. Definitely one of the areas best and longest known to speleologists and biospeleologists, the Dinaric karst is located in north-east

Italy in the Trieste and Isonzo Karst, the so-called classic karst. Covering not more than 200 sq.km., it lies between the river Isonzo, the sea, and the border between Italy and Slovenia, and its fauna certainly has affinities with that of the karst in Slovenia, Istria and Dalmatia. In it prevail endemic or Illyrian and Illyrian-Balkan species, and the Isonzo is often viewed as the western limit of their geographic distribution. Among the fauna which (with very rare exceptions) have not moved beyond the Isonzo are entire genera of gastropods (Plagygeyeria), crustaceans (Sphaeromides), spiders (Stalita), beetles (Leptodirus) and the well-known proteus, or olm (Proteus anguinus). Alps and Prealps. The Prealps are definitely one of the areas best explored by biospeleologists, who know very well that every Prealpine massif (from Piedmont as far as Slovenia) is a tiny world rich in endemic species. Typical terrestrial fauna include carabids (ground beetles) of the genera Orotrechus and Anophthalmus, next to highly specialized endemic genera like Italaphaenops and Lessinodytes. In aquatic fauna, typical isopods here are those of Monolistra, the many endemic species of amphipods of Niphargus, and copepods of the genus Lessinocamptus. These are all species with affinities with the Balkans or Eastern Europe, which colonized the area in a definitely pre-glacial period. The Alpine karst in a strict sense - that is, caves located in the internal massifs of the Alps, often at high altitudes - is still imperfectly explored. It was impoverished as a result of the Quaternary glaciations and thus some biospeleologists believe it is of no particular interest. This is not true, since recent research on terrestrial fauna has revealed endemic species of carabids (Anophthalmus), leptodirines (Pseudoboldoria) and phalangids (Ischyropsalis). Studies on aquatic fauna have also shown a fauna of considerable biogeographic interest, since this is the only area in which we find some species of amphipod crustaceans of the genus Niphargus (such as N. strouhali, exclusive to high karst areas, sometimes exceeding 2000 metres a.s.l.), next to species of both northern provenance and others similar to those populating the Prealps. Although some forms found extensively north of the Alps (like the copepod Eucyclops graeteri or the amphipod Niphargus foreli), typical of cold climates, colonized the area after the retreat of the large Quaternary glaciers, most of the species may have recolonized Alpine caves from marginal pre-Alpine areas or moved from deep phreatic faults or refuge massifs (various species of Elaphoidella, Lessinocamptus, Speocyclops, Niphargus): in the last case, the species are always endemic. Ligurian Alps. The karst in the extreme western part of the Alps shows

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