8382 faunistic affinities with southern France (e.g., among the isopods, Proasellusgr. cavaticus) and the Alps on one hand, and with Appennine and Tyrrhenianareas on the other. Biogeographically, therefore, this is a transition area whichhas little in common with the Alpine and pre-Alpine karst: we find richspecialized fauna and many endemic species belonging, for example, tovarious genera of isopods (Proasellus), diplopods (Glomeris, Spelaeoglomeris),carabids (Duvalius, Agostinia) and catopids (Parabathyscia).

• Appennines. The caves of the Appennine karst have troglobitic fauna whichis generally poorer and more monotonous than that in the Prealps, although insome regions this may be due to lack of research until now. These are areaswith endemic species (for aquatic fauna, in particular, the LigurianAppennines, Alburni mountains, Gran Sasso massif; and the Latium-Campania pre-Appennine area for terrestrial examples). But there are others,where endemic species are, with few exceptions, almost absent (Apuan Alps,Calabria). The faunal monotony between Liguria and Campania may be due tothe fact that the karstism in these areas is younger, and perhaps to the“peninsular effect” which definitely hindered colonization of the Appennines,but this is still a hypothesis. Novelties emerging from research conducted inthe last twenty years have not substantially changed this picture, althoughnew species of troglobites and eutroglophiles belonging to genera widespreadin Italy have been added, like pseudoscorpions (Neobisium), carabids(Duvalius) and leptodirines (Bathysciola). Aquatic fauna have revealedinteresting exceptions in the Gran Sasso and Alburni mountains, with thediscovery of probable marine relics of Miocene age (copepod crustaceans ofthe genus Pseudoectinosoma).Also of interest is recent research in the gypsum caves of the Emilian andBolognese Appennines: the terrestrial fauna is mainly composed of a feweutroglophiles, while aquatic fauna is rich in stygobites (copepods andamphipods), although none is exclusively cave-dwelling (that is, they are alsofound in the interstitial environment), signalling the fact that colonization ofthese caves is recent.Biospeleological research has also been carried out in the gypsum (Messinianevaporites) in Calabria (province of Crotone) and, here too, althoughinteresting data have come to light (for example, a new species of Niphargus),troglobites are infrequent. This may be due to a combination of unfavourablefactors, like the greater compactness of gypsum formations with respect tocarbonatic ones (and thus, a lower degree of fissuration) and the young ageand rapid evolution of pseudokarstic phenomena.

• Apulian karst. Very different from the Appennine karst, the Apulian version is

1 DINARIC KARST

2 ALPINE AND PREALPINE AREAS

3 LIGURIAN ALPS

4 APPENNINES

5 APULIAN KARST

6 SICILY

7 SARDINIA

TOMI

GE

TS

TN

VE

BO

FI

AN

AQ

CB

PG

ROMA

NA

BA

PZ

RCPA

CA

AO 12

3

4

5

7

6

Biogeographic provinces of karstic areas in Italy

8584 rich in endemic species and its cave-dwelling fauna is highly specialized. Ofthe thirty-odd troglobe species known, about twenty are exclusive to Apulia,some of which are genera or families in their own right. Aquatic fauna, stillrelatively poorly studied, contains exceptional endemic forms such as thesponge Higginsia ciccaresei, the mysids of the genera Spelaeomysis andStygiomysis, the decapod Typhlocaris salentina, and many highly specializedcopepods, isopods and amphipods. This is where we find some of the mostexceptional and enigmatic examples of Italian fauna, like the amphipodMetaingolfiella mirabilis. Among terrestrial troglobites are the isopod generaMurgeoniscus and Castellanethes and the carabid Italodytes stammeri, ofprobably ancient colonization, next to more recent ones like Aegonethes andPseudanapis.The richness of endemic, specialized species is due not only to the extent andantiquity of karstic phenomena, but also to the geological history of the area,considered by paleogeographers as part of the Aegeis or in any casebelonging to a microplate of different origin to that which gave rise to theremaining portion of the Italian peninsula.

• Sicily. Biogeographically composite, the cave-dwelling fauna of Sicily is stilllittle known, particularly its aquatic forms. Sicily contains both karstic and lavacaves. Various types of faunal research have been carried out in the volcaniccaves, but have not revealed cave-dwelling forms of particular interest, due tothe relative “youth” of such caves and to the structure of lava, which does notallow the creation of that indispensable system of crevices which troglobitesuse. Instead, interesting endemic forms are known in caves opening incarbonatic rocks: examples are the pseudoscorpion Roncus siculus, theamphipod Tyrrhenogammarus catacumbae, and isopods of the the genusSpelaeoniscus. Also endemic to the island are some species of trechinebeetles of the genus Duvalius.

• Sardinia. The karstic territory of Sardinia is a world in itself: it is rich in cave-dwelling species (more than 300 have been reported until now), many of themendemic, and it has faunal affinities with Provence and the Pyrenees on onehand and with Corsica and Tuscany on the other.This faunal originality derives from the paleo-geographic vicissitudes of theisland, which was once part, with the areas mentioned above, of a paleo-continent called Tyrrhenis. After its fragmentation, which started in theOligocene, Sardinia and Corsica became detached from the Provençal areaand migrated towards Italy with a slow, clockwise rotating movement. Likegreat rafts, they brought with them ancient elements of the Tyrrhenic fauna,called for this reason paleo-Tyrrhenic.

Classic examples are genera with clearcut affinities with the Pyrenées(Scotoniscus, Catalauniscus, Typhloblaniulus), Cataluña-Provence(Spelyngochthonius, Stygioglomeris, Oritoniscus), eastern Spain and theBalearic islands (Parablothrus, Syniulus, Speomolops, Ovobathysciola,Patriziella), southern France and the Tuscan coast (Stenasellus), next togenera with endemic species (Sardostalita, Sardaphaenops). There are alsomore widespread forms in the Italian peninsula, revealing more recent phasesof colonization.

Entrance to Grotta Zinzulusa, on Salento coast (Apulia)

Hydrozoa. Cave-dwelling hydrozoa areextremely rare. The siphons of somesprings and shafts may reveal species ofHydra, solitary polyps which lack themedusa-like generation typical of thisclass of cnidarians: little or nothing isknown of their ecology.The only hydrozoan considered to be atrue stygobite and cave-dweller isVelkovrhia enigmatica, widespread incaves in the Dinaric karst from Sloveniato Herzegovina. Its possible presence inItalian territory has not yet beenascertained.

Triclads. Triclads, or planarias, includemany species widespread in both sea

and fresh waters and in damp soil. Some genera are frequently found in theinterstitial environment and karsticsprings (Dugesia, Dendrocoelum,Polycelis). Stygobitic species are

87

Higginsia ciccaresei

Troglophile triclad of genus Dugesia

Taxonomy Fabio Stoch · Leonardo Latella · Luca Lapini

Porifers. Porifers, or sponges, areprimitive metazoa, mainly marine,although there are a few fresh-watermembers (spongillids). Some specieshave been reported in subterraneanwaters, generally as stygoxenes orstygophiles, exceptionally as stygobites.In Italy, although cave-dwelling spongeshave been found in some coastalgrottoes - Petrosia pulitzeri in the Gulf ofNaples, Sicily and the Adriatic,Myrmekioderma spelaea in Apulia, andPetrobiona incrustans in the Grotta delPresepe at Santa Maria di Leuca - theyare probably all eustygophile species.The most recent and exceptional findingis that of Higginsia ciccaresei, collectedby a diver from the Zinzulusa cave(Apulia), about 250 metres from theentrance and at a depth of 12 metres, intotal darkness. Its morphologicalcharacteristics and environment indicatethat it is a stygobitic species.

characterized not only bydepigmentation (also found in someabove-ground populations), reduction ordisappearance of sight organs, andslowness of the reproductive cycle andrespiratory processes, but also by a highnumber of chromosomes. A fewinfrequent cave-dwelling species areknown in Italy, including definitestygobities like Dendrocoelum collinii inthe Julian Prealps (also reporteddoubtfully in other parts of Italy),D. italicum in the Lombard Prealps,Atrioplanaria morisii near Cuneo,A. racovitzai in Sardinia and central-southern Italy, and Polycelis benazzii inLiguria.

Temnocephalids. This taxon, usuallyascribed to turbellarians, includes, inItalian caves, ectoparasitic species oftroglobitic crustaceans of the generaNiphargus (amphipods) and Troglocaris(decapods), which suck hemolymph.These tiny organisms (less than 2 mmlong) have tentacles and suckers withwhich they attach themselves to theirhost’s gills. In Italy, they have only beenreported in the phreatic waters of theGorizia and Trieste karst, where they aresometimes found in abundance onshrimps of Troglocaris. Three genera(Bubalocerus, Scutariella,Troglocaridicola) have been reported, butthe taxonomy of the entire group iscontroversial. Research has recently beenundertaken on the spermatozoa of theItalian species, with the aim ofreconstructing their phylogenetic affinities.

Nematodes. This is a highly diversifiedphylum (possibly including more than100,000 species), represented in

benthos, fresh and salt waters, and thesoil, or as species parasitic onvertebrates or other invertebrates. Due tothe sometimes very high number ofspecimens, even in caves, their roleappears to be essential to the functioningof Italian cave ecosystems, but nostudies have yet been carried out.Taxonomic difficulties in studying theseorganisms have prevented thepreparation of exhaustive lists for Italiancaves. An ongoing study in northern Italyhas identified many species found inpools and subterranean streams, generabeing Aphanolaimus, Dorylaimus,

Eudorylaimus, Paractinolaimus andTrypila. Our knowledge is still too sketchyto enable us to state definitely whethersome species may be consideredtroglobitic.

Nematomorphs. Nematomorphs makeup a small taxon similar to nematodes.Adults generally live in slow-flowingstreams, wells and drinking-places. Theyare filamentous in appearance, about 10cm long, and only about 1 mm thick.

Their filaments are sometimes so twistedthat they look like the Gordian Knot, fromwhich they derive one of their names(gordiaceans). The larvae areendoparasitic in various arthropods. Theyare occasionally found in cave streams innorthern Italy, but they are presumablystygoxene or perhaps stygophilespecies. No studies have been carriedout on Italian cave-dwellingnematomorphs.

Molluscs. Of the molluscs, only snailsand bivalves colonized undergroundenvironments and waters. Butaccumulations of debris and mud mixedwith the shells of many species maysometimes be found in caves. However,as these materials were carried downfrom the surface environment throughwater circulation and percolation, truetroglobitic species are in fact onlyrepresented by the genus Zospeum.Another ellobiid, belonging to the genusCarychium, which normally lives in otherhabitats in the external environment, likesvery damp litter and isolatedly appearsunderground, whereas many zonitids aretrue stygophiles. Subterranean watersare mainly colonized by hydrobiidprosobranchs; bivalves arerepresented by the single genusPisidium. Troglobitic molluscsalso underwenttransformations as a result ofliving underground: theshell is white andtransparent, and thediaphanous animal has noeye-spots. The shell in thevarious species of Zospeum(see drawing) is never morethan a couple of millimetres long.

These animals live in the dampest areas,or inside cracks, scraping the walls. Theshell may be very varied according towhere the specimens live and also withinthe same population.Interest in this species goes back toaround the mid-19th century. Since theirsize makes them difficult to identify,Pollonera, the famous researcher whowas among the first to study the species,advised the cave explorers of thosetimes to pass delicate brushes over thewalls of caves, holding boxes underneathto catch specimens as they fell, or tosearch for them in the sediments of riversflowing from karstic areas. In this way,many species and subspecies werediscovered and described in Italy, Austriaand the western Balkans, from Sloveniato Herzegovina. They have also recentlybeen found in the Pyrenées.The genus Zospeum was revised in the1970s for Slovenia and, similarly, ameticulous census has been made of allsites and findings in Italy, and aninterpretation of the various formsdescribed in the literature has beenproposed, from the Brescia area to theIsonzo basin. It appears that all the

westernmost populations ofZospeum, as far as the MonteGrappa massif, present a single

outer fold, whereas forms withtwo folds prevail to the east. Inthe latter specimens, the shellis also ribbed. It seems thatthe more westerly species

should be grouped asZospeum globosum andthe more easterly ones asZ. isselianum andZ. spelaeum, which is also

found in Slovenia. Instead,

88 89

Nematomorphs (gordiaceans)

Z. alpestre is almost exclusively found inSlovenian territory, although it hasrecently tentatively been identified insome caves in the Monte Musi chain(Julian Prealps). Among predatory troglophile molluscs,the genus Oxychilus is very widespread,although some species are morefrequent in some regions than in others.O. draparnaudi is almost ubiquitousthroughout Italy. Aegopis gemonensis isanother troglophile form frequently foundin the caves of the central-easternPrealps. Sometimes the genus Argna,with the species A. biplicata, typicallyfound below ground, appears in the cavehabitat.Among fresh and brackishwater snails, the hydrobiidfamily is the most numerousas regards number of generaand species. They are mainlyfound in karst springs orunderground and interstitialwaters. Some species,particularly those of thegenus Iglica, are even today

known only by the morphology of theirshell, because only already empty shellsreach the outside from springs andhyporheic waters. This indicates thatthese animals prefer the deepestsubterranean, poorly accessible habitats,sometimes far from the places in whichtheir shells are found. Hydrobiid shellsare very variable in aspect: high spiralforms prevail, but there are also generawith flattened spirals like Hauffenia,Hadziella (see drawings) or Islamia. Theyare always very small.Some stygophile species are found quiteextensively, like Graziana pupula in theEastern Alps, G. alpestris in the central-western sector, and Bythinella schmidtii,which ranges from the Alps to theAppennines, as far south as Campania.Most of the strictly stygobitic species aremore limited in their distribution, likemany species of Alzoniella,Pseudavenionia, Iglica, Paladilhiopsis,Phreatica and Hauffenia in the Alps;Avenionia, Fissura, Pezzolia and Islamiain the northern Appennines; andMoitessieria, Sardhoratia andSardopaladilhia in Sardinia.For some of these, like Istriana mirnae inthe flyschoid areas of Venezia Giulia andIstria, and Plagigeyeria stochi in theunderground parts of the Timavo, known

distribution is reduced to only afew locations.

Polychaetes. Polychaetes areessentially marine annelids and

some species colonizegrottoes along the coast.

Species suited to continentalfresh waters are very rare. In

Italy, the stygobitic speciesMarifugia cavatica is found in the

phreatic waters of caves in the Goriziaand Trieste Karst. This is a diaphanousworm, depigmented and eyeless, whichlives in a more or less calcified tube,sometimes more than 1 cm long. It formslarge colonies which coat the walls of thelarge galleries of the subterraneanTimavo near Trieste, and a richmicrofauna, composed of protozoa,gastropods, oligochaetes andcrustaceans inhabits the spaces betweenthe tubules or inside abandoned ones.M. cavatica ranges from the river Isonzoto the border with Albania. It has beenhypothesized as a very ancient relic ofmarine origin, presumably going back tothe Miocene.

Oligochaetes. These are annelids withlong bodies, composed of a successionof metamers without appendages, butbearing transversal rows of bristles, theform and arrangement of which is oftaxonomic value. The main species aredetritivorous or microphagous. Cavesfrequently contain some aquatic families(lumbriculids, naidids, tubificids) or onesof semi-aquatic or terrestrial type(enchytraeids, lumbricids).The terrestrial species, very common indamp soil and the deposits of organicmatter at the bottom of karstic shafts, aregenerally believed to be regular or simplesubterranean trogloxenes (e.g.,Allolobophora rosea, Octolasiumcyaneum, Dendrobaena cognettii). Someaquatic species, however, showingaffinities exclusively with marine generaor collected until now only insubterranean karstic waters, are to beconsidered true stygobites. Speciesknown until now only in caves are Habermonfalconensis, found at the Bocche del

Timavo in the Trieste Karst; Rhyacodrilusgasparoi from a cave in the JulianPrealps; R. dolcei from small pools in acave in the Trieste Karst; Tubifex pescei,frequent in phreatic waters in theMarches and Umbria and in a Sloveniancave; and Abyssidrilus cuspis, collectedfrom phreatic waters in Umbria andcaves in Liguria and Friuli-Venezia Giulia,a stygobitic species belonging to agenus which also has marine members.

Hirudineans. Hirudineans, or leeches,are not often found in Italiansubterranean waters, where they aremostly stygoxenic; only Dina krasensis,inhabiting springs on marly-sandstoneterrains in the most easterly part of theJulian Prealps and around Trieste,penetrates caves, where it is sometimesfound regularly and locally abundantly,and may be considered as a stygophile.Troglobitic species have been reported incaves in Herzegovina, Montenegro andTurkey, but not in Italy. The species inItalian caves do not suck blood, as onewould immediately think, but prey onother aquatic invertebrates.

Tardigrads. These small animals (rarelyexceeding 1 mm in length), aretransparent, and usually have four pairsof unjointed legs. There are marine andfresh water species, and terrestrial ones.They mainly frequent moss, where asmany as 2 million specimens have beenreported in one square metre. Tardigradsare active only in a thin sheet of water(e.g., after rain, or as dew); otherwise,they enter a stage of immobility(cryptobiosis) which may last for severalyears. Although data on cave-dwellingtardigrads are scarce, according to

90 91

Plagigeyeria stochi

ongoing research, the most commonspecies in this environment areMacrobiotus cf. richtersi and Diphasconnobilei, sometimes abundant in smallhollows in clay, but some have also beenfound in litter and moss.

Ostracods. These crustaceans areunmistakeable, with their bodiesenclosed in a generally ovoid, bean-shaped or trapezoidal bivalve carapace,sometimes ornated with tubercles orhollows. They are small (most stygobiticspecies do not exceed 1 mm in length),widespread in all fresh or salt waters,from small pools in tree-holes, topuddles, ponds and lakes, where theyare usually benthic or interstitial, veryrarely planktonic. Still very little is knownabout this group in Italian subterraneankarstic waters, although it is verywidespread in cave streams and siphons.Recent collections reveal richlydiversified fauna, with many blind,depigmented species new to science:among these, most belong to the vastcandonid family. One new species(Cypria cavernae) has recently been

described from the caves in the Goriziaand Trieste Karst, the only stygobiticspecies of the genus, common in theslow-moving waters of the phreatic zone.One very special species, whichdeserves to be mentioned for its life-style, is Sphaeromicola stammeri. Thiscommensal lives exclusively on variousabundant isopod stygobitic crustaceans,and is found clutching the appendagesof its host. Its distribution follows that ofits host species and it is thus found fromHerzegovina to Lombardy.Instead, S. sphaeromidicola lives on thelarge cave-dwelling cyrolanids of thegenus Sphaeromides, recently found alsoin Italy.

Cladocerans. These are very commoncrustaceans found in fresh surfacewaters, where they populate seasonalpools, lakes and slow-flowingwatercourses; over a hundred speciesare known in Italy. They are frequentlyfound in the waters of Italian caves,generally as stygoxene forms. Theirscarce presence in subterranean watersis probably due to their feeding habits(many species are associated with thepresence of phytoplankton andsubmerged vegetation) and to theirdefinite preference for eutrophic waters.However, two stygobitic species haverecently been described (Alona stochi,A. sketi), blind, diaphanous, and small(0.5 mm). As the latter were found insprings in Slovenian caves only a fewhundred yards from the frontier with Italy,it is therefore probable that they also livethis side of the border.

Copepods. These are generally small(0.2-0.5 mm) crustaceans, widespread in

both marine and freshwaterenvironments. In Italian continentalwaters, they are essential components oflacustrine zooplankton, occupying bothflowing and standing waters, and oftenprevail in subterranean aquaticcommunities. In Italy, several hundredspecies exist and, of the ten ordersknown, only three (calanoids,harpacticoids, cyclopoids) havetroglobitic members widespread invadose and phreatic waters.• Calanoids: planktonic copepods, foundin lakes and slow-flowing watercourses.The only stygobitic and definitelytroglobitic species, Troglodiaptomussketi, lives in the Gorizia and Trieste Karstcaves in Italy, and also in Slovenia andCroatia. It is common in the siphons ofkarstic caves and shafts associated withthe underground course of the Timavo.• Cyclopoids: very widespread in surfacewaters, there are many stygobiticspecies, mainly benthic and interstitial,found throughout all Italian regions,although many must still be described indetail. The cyclopoids of Italian caves aredistinguished into three categories on the

basis of their ecology: mainly planktonic(e.g., Metacyclops, reported in the cavesof Venezia Giulia, Apulia and Sardinia);benthic, exclusive to the phreatic zone,or sometimes interstitial (e.g., stygobiticspecies of Eucyclops, Diacyclops andAcanthocyclops), and species inhabitingthe vadose zone, where they live in thenetwork of tiny crevices in carbonaticrocks (numerous species ofSpeocyclops).The most frequently found cyclopoids insubterranean waters belong to thelanguidoides group of the genusDiacyclops. This is a set of species,many of which have not yet beendescribed, populating both karstic areasand alluvial soils. Among the otherspecies of cyclopoids, north-eastern Italyhosts some endemics (such asMetacyclops gasparoi) or species whichshow a distribution restricted to theJulian-Slovenian (including the commonDiacyclops charon, Metacyclopspostojnae, Speocyclops infernus) orIllyrian-Balkan areas (e.g.,Acanthocyclops troglophilus and A.gordani). The Prealps and Alps generallyhost species more widespread in bothkarstic and non-karstic systems ofsouthern Europe (e.g., Eucyclopsgraeteri, Acanthocyclops kieferi,Graeteriella unisetigera); the cave-dwelling fauna of the Appennines ismuch less well known. Worthy of note isthe occurrence of Acanthocyclopsagamus, an exceptional species endemicto the Castelcivita cave (Alburni massif).The fauna of the Sicilian and Sardiniancaves is still poorly known.Some species are associated withanchialine caves, recently described inSardinia (but also reported in the

92 93

Troglobitic ostracods of genus Pseudocandona Troglobitic cyclopoids of genus Diacyclops

Balearic and Canary islands).It should not be forgotten that, next tostygobitic species, substygophile andstygoxene forms are widely found inItalian subterranean waters, where theymay be abundant (e.g., Eucyclopsserrulatus, Paracyclops fimbriatus,P. imminutus).• Harpacticoids: this order includes manybenthic and interstitial species, verycommon in caves. Stygophile speciesinclude members of Bryocamptus andAttheyella, although there are also someeustygophile (B. typhlops) and stygobiticforms (B. balcanicus, A. paranaphthalica).Stygobitic species also contain manyendemic forms in restricted karstic areas,belonging to Nitocrella, Elaphoidella,Lessinocamptus, Moraria, Morariopsis,Paramorariopsis, Ceuthonectes andParastenocaris. Here, many species haveonly recently been discovered and arestill being described, some being knownonly in a single cave: they generallyinhabit crevias in limestone and smallpools or percolation rivulets in cave. Thistype of habitat (which functions in acertain sense as as “island” inside acertain karstic system) and the isolationof the various carbonatic massifs askarstism progressed, presumablyfavoured speciation mechanisms whichgave rise to such a high number ofendemic species.Genera such as Elaphoidella andLessinocamptus probably colonizedsubterranean waters starting fromsurface waters or semi-terrestrialhabitats through “corridors” such as, forexample, hyporheic environments.Instead, members of Nitocrella andParapseudoleptomesochra presumablycolonized subterranean continental

waters starting from marine ancestors.Two different species belonging toPseudectinosoma, which until now hadonly contained a single Atlantic marine

species, were recently found in theinternal karstic systems of central-southern Italy (Gran Sasso massif,Abruzzo, Alburni massif, Campania). Thisfinding becomes even more important ifwe consider that the genus is unknownanywhere else in the Mediterraneanbasin and that these stygobitic,freshwater species may be therepresentatives of an ancient fauna of theMediterranean, which disappeared fromits original marine environment during theMessinian salinity crisis.

Isopods. This order of malacostracancrustaceans has reached an exceptionaldegree of adaptative radiation, with morethan 10,000 species covering terrestrial,fresh and salt water environments, oradapted to a parasitic life-style. Isopodspresumably colonized Italian cavesstarting from the sea (during the Mioceneand Pliocene regressions: cyrolanids,microparasellids, perhaps spheromatids)(see box for more details on thiscontroversial hypothesis), from surface

fresh waters (asellotes) or from soil andlitter (oniscoids). The adaptations andlife-styles of isopods, and theevolutionary history of their variousfamilies is so varied that each of themconstitutes a microcosm in its own right.• Spheromatids: the genus Monolistraoccurs in Italy, with many species incaves in the Prealpine belt from theItalian-Slovenian border to the Lake ofComo. Every species or subspecies is tobe viewed as endemic to a restrictedkarstic massif. M. (Microlistra)schottlaenderi, exclusive to some springsin the Isonzo Karst, and one species stillbeing described, found in the extremeeastern end of Venezia Giulia, are theItalian representatives of the subgenusMicrolistra, widespread in Slovenia andCroatia. All the species of this subgenushave large dorsal tubercles, sometimesextending to form long spines andinterpreted as efficient defencestructures, particularly when the animalcurls up to protect itself from predatorsor to allow itself to be transported bywater currents. Instead, M. (Monolistra)coeca julia is the only Italianrepresentative of the subgenusMonolistra and is endemic to the JulianPrealps. The remaining species andsubspecies in Italy belong to thesubgenus Typhlosphaeroma (or perhapsto several similar subgenera) and occurin the Trieste and Gorizia Karst, and inthe Carnic, Veneto (Lessini Hills, Brentamassif, Beric Hills) and Lombard Prealps.• Cirolanids: a family mainly containingmarine species, only discovered recentlyin Italian groundwaters.Typhlocirolana aff. moraguesi (identifiedby means of molecular biologytechniques as closely related to but

distinct from T. moraguesi from Majorca),found in artificial wells along the coastnear Siracusa in Sicily, presumablycolonized subterranean waters during themarine regressions of the Pliocene, asindicated by datings using moleculartechniques. Sphaeromides virei, found incaves in the Gorizia and Trieste Karst,Istria and Dalmatia, is a very largepredator, more than 3 cm long, which isat the apex of the food chains of simplesubterranean cave ecosystems: it maybe found along the coastline of Dalmatia(Illyrian distribution), and its manyendemic subspecies indicate that itcolonized the subterranean waters of theDinaric karst in very remote times.• Microparasellids: small (a few mm)isopods, generally interstitial. They wereoriginally marine species, whosedistribution recalls the ancient coastlinesof the Tertiary seas. Six Italian species ofthe genus Microcharon are known, butnone has as yet been collected in caves.• Asellids: a family including many fresh-water species, in both surface andunderground waters, with specializedlife-forms in both phreatic and caveenvironments. The family is found inEurasia, North Africa and northern andcentral America, but its affinities with

94 95

Aquatic isopods of genus Monolistra

Proasellus franciscoloi

96 97

Among the stygobites best known fromthe taxonomic and evolutionaryviewpoints are two genera of isopodcrustaceans (Monolistra andCoecosphaeroma), belonging to thefamily of spheromatids. Biospeleologicalinterest has focused on them for thefollowing important reasons:• their closest relatives are all, withoutexception, marine (Sphaeroma, whichgroups species very common in lagoonsand along pebbly beaches); nosimilar genera exist in freshsurface waters; • the geographicdistribution of the generafaithfully follows the limits ofTertiary seas, as alreadyhypothesized by severalresearch-workers ahundred years ago• their size andfrequency measured bymolecular analysis(electrophoresis) revealthat the origin ofMonolistra and of some ofits species can be datedvery precisely, andcoincides with the salinitycrisis of the Mediterraneanwhich led to its almost totaldrying-up during Messiniantimes (Upper Miocene).

All these facts mean thatmonolistrines are highly suitable“models” for formulating and testinghypotheses on the origin and evolution ofcave-dwelling organisms. However, heretoo, there are controversies, substantiallyas follows: the marine origin anddistribution of the group appear to becongruent with the “regression model”,

which presumes that these organismsare relics which adapted themselves tothe subterranean environment after themarine regression of the Miocene.Colonization thus followed a direct routefrom the sea to subterranean karsticwaters, probably independently for thetwo genera (Monolistra, of Mediterraneanorigin, and Coecosphaeroma, Atlantic).This theory has recently been criticized,in that it presumes that the animals were

“obliged” to colonizesubterranean waters as aresult of adverseenvironmental conditions,

i.e., increased salinity.Recent, still unpublisheddata, and the discovery ofgreat numbers of Tertiaryspheromatid fossils insurface streams andrivers, have given rise to

the following, moremodern hypothesis:• the marine species of

Sphaeroma freelycolonized fresh surfacewaters, occupyingavailable niches;• the Messinian salinity

crisis interrupted the geneflow with marinepopulations, giving rise to a

process of speciation;• karstification of the territory

favoured active colonization ofsubterranean waters, where empty

niches were available, together with lesscompetitive pressure, since insects didnot colonize groundwaters;• progressive karstification isolatedmany massifs and favoured processes ofallopatric speciation; currently, everyhydrographic basin has its own species

or subspecies of Monolistra;• climatic variations or other processesled to the disappearance of surface-livingspecies.Monolistrines have developedsometimes exceptional defencemechanisms, particularly in Monolistra,with long, sharp spines or tubercles; theirnatural enemies are mainly largeamphipods of Niphargus or olms.Another defence mechanism adopted by

these animals is their ability to curlthemselves up completely, thusprotecting their appendices and theunderside of their bodies from predators.Recent studies support thehypothesis that curling up is not onlya defence mechanism but also ameans of being transported morerapidly by water currents, the animalsbeing rolled along together with smallpebbles and gravel.

Biogeographic and evolutionary scenario of Monolistra species: theories compared Fabio Stoch

Present-day distribution of monolistrines, plotted on paleogeography of Messinian seas

MONOLISTRINES

MESSINIAN SEAS

EMERGED LAND

other marine isopod families give rise todoubts: subterranean waters weredefinitely colonized starting from surfacewaters. Populations of above-ground species have also beenfound in caves, depigmented, andwith reduced eyes(microphthalmic), denoting a stillongoing phase of activecolonization of the undergrounddomain. Almost all the stygobiticspecies are endemic to Italy andhave small areas of distribution.Within the ambit of the genusAsellus, Italy has A. cavernicolus,which lives in the basin of theTimavo in the province of Triesteand has recently been studied withmolecular techniques by aSlovenian team of researchers. It ispresumably a relic of pre-glacialcolonization of the Julian andSlovenian karstic areas by thesimilar surface-living species,A. aquaticus.The single speciesChthonasellus bodoni is the onlymember of the genus Chthonasellus,endemic to Italy (province of Cuneo). Ithas affinities only with the French genusGallasellus and with American species,revealing a perhaps very ancient origin.There are, instead, many morerepresentatives of Proasellus, with manystygobitic, cave-dwelling and interstitialspecies, throughout Italy; some of thesemust still be described.Among the most interesting cave-dwelling species are P. ligusticus, foundin Liguria and the Apuan Alps; P. pavaniin a cave near Brescia; P. cavaticus inwestern Piedmont, Liguria and France;and P. franciscoloi from caves in Liguria

and near Cuneo, together with very manyother species still not defined in detail,known until now under the collectivename of P. patrizii, exclusive to caves in

Sardinia. Also for the stygobiticspecies of Proasellus, which havesurface-living species in Italy,various hypotheses have beenformulated regarding their originas relics, i.e., derived from

populations which found refuge insubterranean waters as a result ofthe instability of surface waternetworks, or as active colonizers ofsubterranean waters.• Stenasellids: a widespread family,including exclusively very ancientstygobitic species. The number ofItalian species is not known exactly,since recent refinement of moleculartechniques is leading to extensiverevision of traditional systematics. InItaly, the family is limited to Tuscanyand Sardinia; the best-known

species is Stenasellus racovitzai(see drawing), found in Tuscancaves; the Sardinian fauna is

undergoing revision. Biochemicaltechniques have identified at least sixdifferent endemic species in Sardinia.Two of these (until now attributed to S.racovitzai) show affinities with Frenchspecies, and another two (S. nuragicus,S. assorgiai) with East European species.Lastly, two species, identified in theprovince of Nuoro, are similar to those ofSpain, revealing the complexpaleogeographic vicissitudes undergoneby Sardinia.• Microcerberids: present in Italy incontinental fresh waters with the singlestygobitic species Microcerberus ruffoi,from the phreatic water-table of the river

Adige, but never found in caves; otherspecies are of littoral marine type.• Oniscoids: terrestrial isopods havemany families with troglobiterepresentatives in Italy (styloniscids,trichoniscids, buddelundiellids,trachelipids, spelaeoniscids, armadillids).These organisms are not usually capableof dispersing easily and thus endemicspecies are frequent. Their distribution isof great interest for biogeographers,since it reveals many of thepaleogeographic events undergone bythe Italian peninsula. In north-westernItaly, for example, we find species whichalso populated the French AlpesMaritimes (sometimes troglophile in onepart of the area and troglobite inanother): typical of this area areTrichonicus voltai and Alpioniscusfeneriensis. Also of interest is theconcentration of endemic species ofBuddelundiella in the French AlpesMaritimes and Liguria. Proceeding from west to east along theAlpine arch, cases become morecomplex and taxonomy intricate:prevailing species belong to thesubgenus Dentigeroniscus of the genusAndroniscus, with a prevalence ofA. dentiger, presumably eutroglophile.Lastly, in Venezia Giulia, we find

troglobitic species with Illyrian affinities:Titanethes dahli, Androniscus stygius andAlpionisus (Illyrionethes) strasseri.The Appennines have a smaller numberof species (Miktoniscus patrizii in Latium,Trichoniscus callorii in the Monti Lepini),with the exception of Apulia, withinteresting endemic (Castellanethessanfilippoi, Murgeoniscus anellii) or trans-Adriatic species (Aegonethes cervinus).Sicily instead has a few endemicspecies, but they belong to genera whichmainly live above ground, likeArmadillium lagrecai. Lastly, Sardiniaoffers perhaps one of the mostinteresting faunal frameworks forbiogeographers, due to the occurrenceof paleo-Tyrrhenian species likeScotoniscus janas, Catalauniscushirundinella and C. puddui.

Amphipods. Like isopods, amphipodstoo are a rich order of mainly marine andfreshwater species, with a very smallnumber of subterrestrial ones.Amphipods colonized subterraneanwaters from the sea or from fresh surfacewaters, where they differentiated into avery high number of species, in somecases (e.g., Niphargus) showing splendidexamples of adaptative radiation. Italyhas almost a hundred species living insubterranean waters, most of them incaves, others in interstitial habitats inalluvial plains, in the interstices of marly-sandstone rocks, and even in cracks inigneous and metamorphic rocks.Here, we describe only the main familieswith stygobitic members.• Bogidiellids: mostly interstitial species,sometimes littoral (euryhaline or definitelymarine), occasionally collected ingrottoes (e.g., Bogidiella calicali from the

98

Titanethes dahli

99

Grotta del Fico on the island of SanPietro (off south-western Sardinia). • Gammarids: inhabiting mainlysuperficial fresh or marine waters, quitefrequent in caves, particularly inilluminated stretches of subterraneanstreams in karst springs and chasms, asstygoxenes (Gammarus,Echinogammarus). Specimens ofRhipidogammarus andTyrrhenogammarus have been found ingrottoes and coastal karst springs. Theonly definitely stygobitic Italian species,Ilvanella inexpectata, lives in phreatic andkarstic systems on the island of Elba andin Tuscany.• Hadziids: species of the genus Hadzia,presumably relics of the Tethys, are

mainly associated with interstitial andlittoral marine habitats. But there are alsocave-dwelling species, like Hadzia fragilisstochi, an endemic subspecies, recentlydiscovered in the Bocche del Timavo andin some caves in the Isonzo Karst: theyare definitely troglobites, with very long,delicate appendages, denotingspecialization to a habitat composed ofthe phreatic waters of karstic conduits.Other Italian species are Hadzia minuta,

found in caves in the Salento, andH. adriatica in Apulia.• Metacrangonictids: a family onlydiscovered very recently in Italy, itincludes phreatic and cave species of theMediterranean basin, Morocco andFuerteventura in the Canary Isles. In Italy,only Metacrangonyx ilvanus is found,endemic to Elba, where it is found in asingle shaft, in alluvial soil.• Niphargids: The systematics of thegenus Niphargus (more than 250 knownspecies, all - with very few exceptions -stygobites) is very intricate andcontroversial, and is perhaps one of themost problematic Italian groups from thetaxonomic viewpoint. Italy has more than60 species (but many others are currentlybeing described), with differences whichare often based on minute morphologicaldetails. The distribution of Niphargus isof southern European type, with aneastward extension as far as Iraq,whereas in the west it only marginallyaffects Spain; the genus is absent fromNorth Africa. The northern limit is clearlymarked by the maximum expansion ofthe Wurmian glaciers, with rare examplesof recolonization of glaciated areas bywidespread species. According to itsdistribution characteristics, the genus ispresumed to have colonized Europeanfreshwaters starting from the basins ofthe Paratethys in Tertiary times, althougha more ancient orgin has recently beenpostulated.The Alpine region has the greatestbiodiversity in Italy, and contains manyendemic species of Niphargus, referringto the groups stygius (East European)and longicaudatus (more southerly).Species of the stygius group may befound as far as central Italy along the

Appennines and in the islands of theTuscan archipelago, where speciessimilar to Niphargus speziae (Liguria) arefound. Southern Italy, Sicily and Sardiniaare much poorer in species, and thelongicaudatus group predominates.Another interesting group is the orcinusgroup, which includes large specieswidespread in southern Italy, the Balkansand the Middle East; in Italian territorysome species penetrate from theBalkans into the Trieste and Isonzo Karst.In Italian caves, there are yet otherrepresentatives of other groups, ofcomplex taxonomy, all widespread inEurope (groups aquilex, kochianus,bajuvaricus, puteanus), or endemicspecies of uncertain affinity (e.g.,Niphargus stefanellii, known in central-southern Italian caves with affinities onlywith one species inhabiting Istria andDalmatia).From an ecological viewpoint, Niphargusmembers occur in almost all caves whereminimal amounts of water are found.They play an important role within theframework of subterranean ecosystems,where they act as predators ordetritivores; they may be very large (up to4 cm in length) and are therefore at theapex of the food pyramids in these

habitats. They occupy all availableniches, from interstitial environments totiny clefts in carbonatic and non-carbonatic rocks, watercourses,subterranean pools and siphons, andlarge kastic conduits. Some species arevery resistant to water stress (e.g., if theirwater supplies dry up) and can survive inhollows in damp soil. This great variety ofmicro-habitats corresponds to aconsiderable variety in size (from 2 to 40mm) and body shape, ranging fromglobular forms, as in species withtendencies towards curling up, to longvermiform shapes in interstitial species.Their structure may be massive andstubby, as in species of the orcinusgroup, with large front legs of pincer type(gnathopods) or thin and long, with smallgnathopods, as in some species of thelongicaudatus group.• Pseudoniphargids: this family is notclosely connected with niphargids,although their name suggests it. But, likeniphargids, it has produced extraordinaryadaptive radiation in subterraneanwaters. For example, in most of Spain,Pseudoniphargus replaces Niphargus incaves, occupying the same ecologicalniches. In Italy, four species are known,of which the most common is P. adriaticus, found both in intersticesand in caves not far from the coast.• Salentinellids: the genus Salentinella iswidespread in regions along theMediterranean coastline, and ispresumed to be ancient (perhaps paleo-Mediterranean). Unknown marineancestors probably colonizedsubterranean waters, perhaps on severaloccasions from the Middle Mioceneonwards, as the distribution of the genusfollows ancient coastlines.

100 101

Niphargus gr. speziae

Niphargus julius

Pacific areas) reveals their considerableantiquity in continental waters and apresumably Tethyan origin.

Decapods. 170 stygobitic species ofdecapods are represented in thesubterranean waters of the world. In Italy,two cave-dwelling genera are known:Troglocaris (Isonzo and Trieste Karst) andTyphlocaris (Salento). Although Italy alsocontains three species of shrimps ofTroglocaris belonging to theanophthalmus group, they are “siblingspecies”, i.e., morphologicallyindistinguishable but clearly different onthe basis of molecular biology. Accordingto paleogeographic reconstructions andapproximate dating of the time ofdivergence by means of the molecularclock, it is believed that the three species

103The taxonomy of the Italian species ofSalentinella requires revision. Thecommonest species, S. angelieri,typically interstitial and frequent in

weakly brackish waters not far from thecoast, is also reported in caves inisolated karstic rocks, in typicallymountain environments. Presumably, inthese karstified areas there are distinctand endemic taxonomic units, alreadydescribed previously as true species.Instead, one definitely separate, endemicspecies, S. gracillima, inhabits phreaticcave waters in Apulia.• Ingolfiellids: small stygobiticamphipods, with long bodies, mainlyliving in interstices, in both salt and freshwater. Only one species is found in Italy,in fresh water: this is Ingolfiella(Tyrrhenidiella) cottarellii, only found in acave on the island of Tavolara (Sardinia). • Metaingolfiellids: Metaingolfiellamirabilis (see drawing) is one of the mostexceptional animals among Italian cave-dwelling fauna.The species has been collected onlyonce (but a large number of specimenswere caught) from a well in the Salento,pumping water from a deep karstic fault,and was described by Ruffo in 1969.Since then, despite muchbiospeleological research in the Apuliankarst, no other examples of this endemicspecies have been found, and it isconsidered extremely rare. This

crustacean is probably one of the oldestcomponents of all Italian fauna.

Bathynellacea. More than 200 speciesof these crustaceans are known, allstygobites. Some researchers believethat they form a group of very ancientorigin, perhaps already well diversified inthe Paleozoic. At that time, bathynellaceawere widespread in the littoral coastalwaters, lagooons and estuaries ofLaurasia, from which they colonizedGondwana in the Permo-Triassic, beforePangea split up, thereafter becomingwidespread in continental subterraneanwaters. This fascinating hypothesis, notyet supported by objective proof,indissolubly links taxonomic study ofthese organisms with the greatpaleogeographic events of our planet.In Italy, syncarid fauna has not beengreatly studied. The genera Bathynella,Anthrobathynella, Meridiobathynella,Sardobathynella and Hexabathynellahave been found, mainly containinginterstitial species. However, richpopulations of syncarids belonging toseveral species of Bathynella haverecently been found in cave pools in theLessini Hills, Julian Prealps and theGorizia and Trieste Karst, where theypresumably inhabit crevices in carbonaticrocks. Since this finding was due to newsampling methods in these specialenvironments, syncarids are probablymuch more widespread in Italian cavesthan hitherto believed.

Thermosbaenacea. In this case too, theorder is of very ancient origins, andincludes about thirty stygobitic speciesspread over five continents.The evolutionary history of these

102

Spelaeomysis bottazzii

Troglocaris gr. anophthalmus

Typhlocaris salentina

crustaceans is associated, as forbathynellacea, with the marine regressionof the Tethys, and followed itsvicissitudes. In Italy, the followingspecies are found in subterranean karsticwaters (rarely in interstitial waters):Limnosbaena finki, in the Gorizia andTrieste Karst (also known in Bosnia);Monodella stygicola, endemic to Apuliancaves and wells; Tethysbaena argentarii,endemic to a cave on Monte Argentario(Tuscany), and T. siracusae, endemic towells in Sicily.

Mysids. Mysids now number more than800 species, mainly living in marine orbrackish waters; only a few live in freshsubterranean waters.Most stygobitic species are found alongcoasts, proving their presumably recentorigin from marine ancestors. In Italy, twostygobitic species occur in caves andwells with phreatic waters in Apulia:Spelaeomysis bottazzii and Stygiomysishydruntina. Both species, which may livetogether, are endemic to Italy. If theiradaptation to subterranean waters, assuggested by recent electrophoreticresearch, is a relatively recent event(Pliocene), the wide geographicdistribution of these genera (embracingMediterranean, American and Indo-

differentiated between 700,000 and1,700,000 years ago, afterhydrogeological barriers formed betweenthe subterranean waters of the IsonzoKarst (associated with the basin of thatriver), the Trieste Karst (associated withthe basin of the Timavo) and small localbasins in the Istrian Karst, of which a tinytongue extends into Italy.These hydrogeological barriers partly nolonger exist, and it is for this reason that,in some localities, two of the threeTroglocaris species live together withoutcross-breeding. The other Italian species,Typhlocaris salentina, is endemic tocaves in Apulia; the other two knownspecies of Typhlocaris live insubterranean waters in Israel and Libya.

Mites. Mites are small or very smallarachnids, ranging in size from 0.1 mm to3 cm, mostly terrestrial, with some fresh-water species and a very few marine

ones. They are found in almost allenvironments, and may be predators,necrophages, phytophages, orcommensals of mammals and birds,consuming hairs, feathers, skin debris, orfatty secretions, or parasitic on otheranimals, both invertebrates andvertebrates.There are many terrestrial mites whichlive in caves, on guano or other organicmatter, on the soil, or in rock fissures. Asa few of them are associated with thishabitat, they are considered as cave-dwellers. Italian caves contain membersof various families, e.g., macrochelids(Geholaspis mandibularis); parasitids(Pergamasus quisquiliarum), widespreadthroughout Italy and its islands;polyaspinids (Uroseius sorrentinus), aspecies found in central-southern Italy;and ragidiids, with many speciesincluding Troglocheles strasseri,apparently associated with cave habitatsin north Italy, France, Switzerland, Austriaand ex-Jugoslavia. Then there areseveral species of the oribatid order,living on soil and guano.The position of a parasite living on bats,Ixodes vespertilionis (ixodid family) is ofgreat interest and has been the subjectof lengthy debate. Many authorsconsider it a troglobite, since it is oftenfound inside caves without its host and inall stages of development. Othersconsider it an occasional cave-dwellerassociated with bats.

Palpigrads. These small blind arachnids,with a long flagellum at the tip of theabdomen, are certainly the least knownof the cave-dwellers. The facts that allpalpigrads are colourless and lackingsight organs, that they live both in caves

and outside them, and that there are veryfew ecological data regarding this orderof arthropods, all mean that it is verydifficult to fit the species found in cavesinto one of the ecological categoriesnormally used for cave-dwellers.Palpigrads preferably eat spring-tails andother small arthropods which they find atthe base of stalagmites or under pebbleson the ground. Ten species have beenfound in Italian caves, all belonging to thegenus Eukoenenia. Of these, seven havebeen collected exclusively in the caveenvironment. Of particular interest are E.gasparoi in Venezia Giulia, E. brignolii inApulia, and E. patrizii and E. grafittii inSardinia, which have adopted specialstrategies suited for cave life.Other species are known in some cavesin the Alps.

Pseudoscorpions. Pseudoscorpions,small arachnids similar to true scorpionsbut lacking the famous tail with itspoisonous spine, are very wellrepresented in Italian caves and in thoseof many areas in Europe, Asia and theAmericas. They show sometimes highly

accentuated adaptations particularlysuited to life underground. They vary insize from 2 mm (above-ground forms,found under dead leaves in theundergrowth, under tree bark or deeplyburied pebbles) to 7-8 mm in the mosthighly specialized cave-dwelling forms.In caves, they are found under pebblesor on the ground, while they wait for theirprey, small invertebrates which theycatch with pincers on their pedipalps,which are particularly long in troglobiticspecies.Of the approximately 210 species knownuntil now in Italy, 124 are found in caves,and 80 of these are considered truecave-dwellers. The caves in whichpseudoscorpions have been foundnumber more than 440 and the best-known regions are Liguria, Veneto,Friuli-Venezia Giulia and Sardinia.Little or nothing is known about theiroccurrence in Val d’Aosta, Marches,

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Specialized mites of genus Rhagidia

Palpigrad of genus Eukoenenia

Neobisium lulense

Umbria or Molise.Italian cave-dwelling scorpions belong tothree separate families. One of these isthe chthonids, with the generaChthonius, Spelyngochthonius andTroglochthonius, of which the last twoare considered relics of a very ancientpopulation going back to thePre-Quaternary.Among the neobisiids is the genusNeobisium, found throughout thepeninsula with many ground-livingspecies, and a few species of thesubgenera Blothrus andOmmatoblothrus, respectively found incaves in the Veneto and Friuli-VeneziaGiulia and central-southern Italy; Roncus,generally less associated than the

previous one with central-southern Italiancaves, is represented by troglobiticspecies in the Alps and northernAppennines; Acanthocreagris, for which,for Italy, three troglobitic species arecurrently known in northern Italy andSardinia; and, lastly, Balcanoroncus, withB. boldorii in the Lombard andVeneto Prealps.Syarinids are represented byHadoblothrus, in Italian caves only inApulia by H. gigas, and Pseudoblothrus,with P. ellingseni in Piedmont caves: the

species of these last two genera, relics ofancient populations, have undergoneconsiderable modifications to suit themto the underground environment.

Phalangids (harvestmen). Arachnidsknown for their long legs, phalangids, orharvestmen, are easily confused withspiders. But more careful observers willnote that the abdomen of phalangids isnot attached to the rest of the body by athin peduncle, as in spiders, but acrossits whole width. Although these animalsare mainly active predators, in somecases they do not disdain carrion.There are many species of phalangidsliving in Italian caves, but only some areconsidered true troglobites or in any caseeutroglophiles. Among those showing theclosest links with the cave environment isthe travuniid, Buemarinoa patrizii, whichinhabits some caves in the province ofNuoro (Sardinia). In the same caves livesMitostoma patrizii which, together withM. anophthalmum in the LombardPrealps, are the troglobe representativesof the nemastomatid family. Anotherfamily, including various cave-dwellingspecies, is that of the ischyropsalidids towhich, among many others, belongIschyropsalis ravasinii (Veneto and Friuli),I. muellneri (Alps and Julian Prealps) andI. strandi (Monte Baldo and the LessiniMountains). There are also other speciesof this genus, not truly troglobitic, but inany case closely associated withunderground habitats, like I. adamii inAppennine caves, probably a recentcolonizer. All the cave-dwelling speciesof Ischyropsalis prefer environments withrather low temperatures. They normallyeat snails, which they capture with theirpowerful pincers.

Spiders. Spiders are the best-known ofthe large arachnid orders, which in Italycontains more than 1400 species, 200 ofwhich are found in caves, although onlysome of them are considered truecave-dwellers. Many surface-livingspecies are highly mobile, and easily(voluntarily or accidentally) enter the firststretches of caves, where they make upa quite large fraction of the trogloxenefauna.These formidable predators colonize allparts of the underground environment,from the entrance, where hydrophileand light-avoiding forms weavesometimes large webs on walls oramong pebbles on the ground, to themore internal parts, inhabited by morespecialized forms, which hunt byweaving webs or wandering in debris

or on calcitic concretions.Spiders normally have eight or six simpleeyes which, in cave-dwelling species,tend to be reduced or to disappearaltogether. However, this reduction is notrandom: the central eyes are the first todisappear; the lateral ones are onlyabsent in the more highly specializedforms.Parallel with the disappearance of eyes isthe development of trichobotria, longbristles which allow the animal to senseair movements and the surroundingenvironment.Inside caves, there are many areas inwhich spiders are found: cave-dwellingspecies often show close relationshipswith those living in moist habitats likeforest litter, in rocky areas with littlevegetation, or under stones in theground.The main families found in the caveenvironment are:• Dysderids: these are the most highlyspecialized troglobitic forms, allwandering predators. In Italy, they arerepresented by Stalita nocturna andS. taenaria (Venezia Giulia) and Sardostalitapatrizii (central-eastern Sardinia).• Leptonetids: members of this family arefive troglophile species of the genusLeptoneta (Liguria and Sardinia) and thetroglobitic L. baccettii (Elba), the onlyblind species. Paraleptoneta spinimana isa troglophile widespread along theTyrrhenian flank of Italy, Sicily andSardinia. Leptonetids capture their preyusing small webs strung betweenpebbles on the ground or in cracks incave walls.• Pholcids: small species with very longlegs: the only member associated partlywith caves is the troglophile Pholcus

106 107

Harvestman of genus Ischyropsalis

Nesticus eremita

Stalita taenaria

phalangioides, widespread in numerousItalian caves where it captures its prey byweaving irregular webs; it is asynanthropic species, and is also foundin cellars and other human habitations.• Tetragnathids: there are three cave-dwelling species belonging to the wallfauna: Metellina merianae, widespreadthroughout Italy; Meta menardi, largerthan the previous species and only foundon the Italian mainland, and M. bourneti,also large, and found on both mainlandand islands.• Linyphids: these small spiders weavehorizontal webs, are widespread in Italy,and have the highest number of cave-dwelling species. Many species ofTroglohyphantes live in Alpine caves andin one in the Ligurian Appennines. Thisgenus contains about 30 species, mainlynot particularly specialized, althoughsome are considered troglobites, likeT. konradi (Piedmont), T. bolognai andT. bonzanoi (Liguria), T. caporiaccoi,T. cavadinii and T. regalini (Lombardy),T. exul (Veneto) and T. juris andT. scientificus (Friuli). The same familyalso contains the genus Porrhomma, notparticularly associated with undergroundenvironments, of which the best-knownand most widespread member isP. convexum; Lepthyphantes, representedby troglophile species and found all overItaly; and Centromerus, composed ofsmall spiders, of which the only Italiantroglobitic member is the tinyC. cottarellii, known from a Ligurian cave.• Nesticids: seven species of this genusare known in Italy, all cave-dwelling. Intwo cases, these troglophiles are not veryspecialized: N. eremita, found inMediterranean Europe and very commonin Italian caves (except for Sardinia) and

N. cellulanus, very common in central-northern and western Europe, but foundin Italy only in the Alpine region. The otherfive species, found in small areas andcharacterized to various extents by areduction in eyes, are consideredtroglobites or specialized troglophiles.These are N. idriacus (Friuli, and in a fewlocations in the Veneto), N. morisii (foundin an artificial cave in Piedmont),N. menozzii (Liguria), N. speluncarum

(found in a small area between Liguria,Emilia and Tuscany) and N. sbordonii(a cave in Latium).• Agelenids: represented by the generaTegenaria, not particularly specialized, thecharacteristic funnel-shaped webs ofwhich are easily seen in caves, andHistopona, the only Italian specializedagelenid, H. palaeolithica (Liguria).• Lyocranids: the presence of Cybaeodesmolara, a microphthalmic speciesexclusive to Sicily, has been reported intwo caves on the island.

Chilopods. Chilopods (or centipedes) areeasily recognizable by the number of legs

on each of the body segments: eachsegment has a single pair of legs (15 ormore in adults). Many centipedes areassociated with damp or lapidiculousenvironments, very similar to cavehabitats, which they often colonize.Italian caves have about 70 species ofcentipedes, most belonging to thelithobiid family with genera Lithobius,Eupolybothrus and Harpolithobius.Although these fast-moving predators arefrequent in underground environments,only very few of them have been foundonly inside natural caves, including:Lithobius scotophilus (Ligurian Alps andFrench Alpes Maritimes); L. doderoi(province of Nuoro, Sardinia) and L.electrinus (Tomba del Polacco cave in theBergamo Prealps, also found howeverabove ground in the Como area). Of allthe species present in Italy, only two aretrue troglobites: L. sbordonii, blind,depigmented, with very long antennae,living in some caves in eastern Sardinia,and Eupolybothrus obrovensis, inSlovenia and recently found also in theTrieste Karst. Among thescolopendromorphs there are therepresentatives of the cryptopid family, allblind but not generally troglobitic. Insidecaves, only a few species belonging toCryptops are found. Although they arequite common in Italian caves, they donot have particularly evident adaptationalfeatures.

Diplopods. Diplopods, or millipedes, aredistinguished immediately fromcentipedes because they have two pairsof legs on each trunk segment. Theseanimals are closely associated with dampenvironments, and are therefore frequentboth above ground and inside caves,

although it is not often easy to identify thespecies belonging to the subterraneanenvironment. Diplopods considered to betrue troglobites show the classicmorphological adaptations to cave life,being depigmented, blind, and with longantennae. Most of them eat decayingvegetal and occasionally animal remains.In caves, they may be seen near rotting

108 109

Lithobiid of genus Eupolybothrus

Polydesmus troglobius

Chersoiulus sphinx

Plectogona sanfilippoi bosseae

wood or leaves, under stones, or walkingon the dampest walls. About 150 speciesare known in Italy which, variously, haverelations with the cave environment.They are:• Glomerids and trachyspherids:with a reduced number ofsegments and the capacity to roll up intosmall balls (behaviour which may at firstbe confused with that of some oniscoidisopods); the most interesting genera inItaly are Spelaeoglomeris (Ligurian Alpsand Alpes Maritimes) and Trachysphaera.• Polydesmids: all blind, both cave- andsurface-living, with sometimes keeledrings, well represented in Italian caves byPolydesmus, Serradium,Mastigonodesmus, Sardodesmus andSchedoleiodesmus.• Craspedosomatids: a very widespreadfamily in Italy and Sardinia, with variousgenera and many species. They oftenhave lateral processes on each bodysegment and a pair of spinnerets forsecreting silk.• Callipods: again with spinnerets, thesecarnivores are present with the genusCallipus throughout Italy and Sardinia,Sardopus being endemic to the island.Despite their preference for subterraneanenvironments, these millipedes do nothave particular morphologicaladaptations to cave life.• Julids: present in Italian caves withvarious species belonging to Trogloiulus(northern Italy), Typhloiulus (some speciesin northern Italy and one in Latium) andSyniulus (endemic to Sardinia).

Spring-tails. This class is the oldestgroup of hexapods known. Spring-tailsare all small (1-2 mm long) and, althoughthey are perhaps the most numerous

arthropods found in caves, they have notbeen studied in depth. In spring-tails,depigmentation and the reduction ordisappearance of eyes are not exclusivecharacteristics of troglobitic species:however, in the latter, longer claws havebeen noted, probably to facilitatemovement on damp surfaces coveredwith a film of water, together with otheradaptations typical of troglobites (slowedmetabolism, reduced fertility, reducedcapacity for retaining water, greaterresistance to lack of food).About 50 species are known in Italy, allfound solely or with a certain frequency incaves, although the degree of preferencefor this environment has not beenestablished in most of them.These species belong to many families:hypogastrurids, such as Bonetogastruracavicola (Apulia) and B. subterranea, bothprobably recent colonizers of thesubterranean environment, and severalspecies of Mesogastrura; onychiuridswith several species of the genusOnychiurus; isotomids, to which belongsIsotomurus subterraneus, found in theTrento area; entomobryids, with morespecialized species, with longer legs andclaws, antennae, and the sense organsassociated with them. Among the most

interesting species are Pseudosinellaalpina, P. insubrica and P. concii, found inthe Alps from Piedmont to the Veneto.Of great interest is definitely thetomocerid Troglopedetes ruffoi, found inthe cave named l’Abisso, near CastroMarina in Apulia. Lastly, there is thearrhopalitid family, with some species ofthe genus Arrhopalites, and thesminthurid family to which belongs

Disparrhopalites patrizii found in caves onthe mainland and in Sicily.

Diplurans. This class of hexapodscontains about 800 species which mainlyprefer hot and temperate areas. They aremainly depigmented and have longantennae. Of the five Italian families, onlytwo contain species associated withcaves.• Campodeids: with long antennae, andtwo long multi-jointed cerci departingfrom the last abdominal segment. Of allthe areas of the world, central andsouthern Europe are those richest incave-dwelling campodeids. Italy hassome species of Campodea, mainly in

central-southern regions and Sardiniaand, in the Appennine caves,Plusiocampa. This genus contains thehighest number of cave-dwellingcampodeids (almost 40 species inEurope). Of particular interest is thepresence in Sardinia of the endemicPatrizicampa sardoa, a troglobite with ahigh degree of specialization.• Japygids: in the members of this family,the cerci are formed of a single joint andare transformed into a sort of pincer,which is used partly for defence butmainly for capturing prey - mites andother small arthropods. Less widespreadthan campodeids and associated withvery high humidity, japygids do not seemto be particularly suited to cave life,although they are quite frequently foundin caves. Known in Italy are Metajapyxmoroderi patrizianus (Sardinia), anendemic subspecies of a species knownonly in a few other caves in Spain, andM. peanoi, a cave-dweller of northernItaly, with particularly long antennae.

Orthopterans. Orthopterans, or hoppers,include crickets and grasshoppers. Theyare phytophagous insects (i.e., they eatvegetal matter), saprophages (eatingdecaying organic matter), or predators:the cave-dwelling species are generallysaprophagous, although some cases ofpredation and cannibalism have beenobserved. Other adaptations in cave-dwelling orthopterans are the classicreduction in pigmentation, long palps,and reduction or disappearance ofwings. In natural and artificial caves inItaly, orthopterans are represented by therhaphidophorid and gryllid families,containing more or less troglophilespecies.

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Campodeid dipluran

Spring-tail of genus Pseudosinella

• Rhaphidophorids: the generaDolichopoda and Troglophilus are foundin Italy. The former contains about tenspecies, including Dolichopoda ligustica(Liguria, Piedmont and Lombardy),D. laetitiae (central-norhern Italy),D. schiavazzii (Tuscan coast and Elba),D. geniculata (from Latium to Apulia) andD. calabra (Calabria). The SicilianD. palpata was described in 1776 by aGerman author, to whom severalspecimens from a cave near Siracusahad been sent, but it has never beenfound since then. Of interest is thefinding of a large population ofDolichopoda in a Veneto cave (Grottadella Poscola, on the Lessini Mountainsnear Vicenza), which appears to havebeen introduced accidentally by man.Three species of Troglophilus are foundin Italy: T. neglectus (Friuli-Venezia Giulia,Veneto, and Trentino-South Tyrol);T. cavicola (extending to Lombardy), andT. andreinii (only in Apulia, with twosubspecies, relics of an ancientpopulation which existed before theseparation of this region from theneighbouring Balkan peninsula).• Gryllids: species found in Italy areGryllomorpha dalmatina, widespread;Gryllomorphella uclensis, found inartificial and natural caves, damp areasand cellars; and Petaloptila andreinii,typically found in woodland litter but alsoin several Appennine caves (this speciesis dark brown in colour and has twosmall scaly wings). Acroneuroptilapuddui and A. sardoa are verycharacteristic orthopterans because,although being exclusivelycave-dwellers, they have conserved veryshort scaly wings but show markeddepigmentation.

The genus seems to be exclusive toSardinia.

Trichopterans (caddis flies). Theseinsects, winged when adult, are aquatic inthe larval stage. Adults are commonlyfound in caves as subtroglophiles,whereas the larvae, with their typical“larval cases”, are found in the entranceareas of caves with internal watercourses.In Italian caves, trichopterans are foundfrom February to November, with peaks inspring and summer, and may be a quiteimportant component of the wall fauna.About 15 species of stenophylacines arefound in Italy as subtroglophiles,including Stenophylax permistus, thelargest cave-dwelling stenophylacine andthe most widespread in caves onmainland and peninsular Italy andSardinia; S. vibex, quite rare in Italy,known in some caves in the central-northern Appennines; and S. mucronatus,S. crossotus and S. mitis.Passing to the genus Micropterna, wefind M. lateralis (northern Italy),M. nyctobia, which seems to prefer high-altitude caves between 1000 and 2500metres; M. sequax, M. malatesta,M. testacea and M. fissa, which is themost abundant trichopteran species in

Italy and for which mating has beenobserved from March to September.Mesophylax contains three cave-dwelling species: M. aspersus (fromnorthern Italy to Apulia, and the onlyspecies currently reported in Sicily);M. sardous (endemic to Sardinia); andM. impunctatus (a species reported froma few caves in northern Italy).

Lepidopterans (butterflies and moths).For many lepidopterans, caves are astable refuge in space and time. Some ofthem spend the winter on cave walls,others stay there in summer, and yetothers use caves during the day or night,and in bad weather.Although no species may truly beconsidered troglobite, many butterfliesare constantly found in caves. Somemicrolepidopterans, associated withguano, spend their entire lives in caveswhere bat droppings accumulate; theircaterpillars live on the guano, eatingfungi, the guano itself, and occasionallysmall invertebrates.Lepidopterans are an important source ofnourishment for many troglobitic animals,both as food, once dead, forsaprophages, and as prey for severaltypes of spider or for snails of the genus

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Gryllomorpha dalmatina

Dolichopoda calabra

Troglophilus neglectus

Stenophylax mitis

Scoliopteryx libatrix

Oxychilus, suited to eating arthropodssuch as butterflies of the genusScoliopteryx, on which active predationhas been observed several times.In Italy, about 15 species belonging tovarious families have been quitefrequently found in caves.Some of these are:• Psychids: Psyche casta, a small mothwhose females, like all those of thisfamily, have no wings and wait for themales inside the “larval cases” made ofvegetal fragments, in which they spendthe larval period; inside caves, it hasbeen observed only in Italy.• Tineids: mostly notorious for thedamage which some members of thefamily cause to clothes and other fabrics.Monopis rusticella is also known in somecaves in central-eastern Europe.• Acrolepids: represented in Italy byDigitivalva (Inuliphila) granitella, oftenfound in large numbers at caveentrances.• Alucitids: Alucita, with typically featherywings, has many subtroglophile speciesin several European countries.• Geometrids: Triphosa dubitata, whoseadults fly in summer and may be foundby day in summer months, or hibernatingduring the winter; and T. sabaudiata,found summering or wintering in Italianand all European caves.• Arctiids: the Appennine populations ofNudaria mundana, a species foundthroughout peninsular Italy, haverecently been studied. In some caves inthe Gran Sasso, they have been foundconstantly inside caves, in all phases oftheir life-cycle.• Noctuids: most lepidopterans found inItalian caves belong to this group:Rhyacia simulans and Apopestes

spectrum are widespread in all Italiancaves; Scoliopteryx libatrix passes thewinter in caves, only to come out duringspring; Autophila dilucida and Hypenaobsitalis are found in Italy, Slovenia andCroatia.• Nymphalids: among these butterflieswe quite often find Inachis io and Aglaisurticae, which may winter inside caves.

Dipterans. Well-known to be equippedwith a single pair of wings (the secondpair is very small and transformed intobalancing organs), the dipterans numberabout 100,000 known species, of whichabout a hundred are quite frequentlyfound in natural and artificial caves intemperate areas.In Italian caves, several species are moreor less closely associated with thesubterranean environment. The mostwidespread is Limonia nubeculosa(limonid family), subtroglophile, verycommon on the walls of caves near theentrance and more rarely further inside.

This species is mainly abundant duringsummer, when sometimes hundreds ofspecimens per square metre may becounted.In winter, on cave walls, we find species

of the culicid family, in particularthe females of Culex pipiens, thecommon mosquito.The most interesting, andperhaps only true troglobiteamong the Italian dipterans isAllopnyxia patrizii (sciarids). Inthis species, known only from acave near Rome, the male is very small(less than 1 mm long) with very smallwings (but equipped with balancingorgans and ocelli), whereas the female ismuch larger, about 2.5 millimetres long,without wings or balancing organs. Theocelli too, in the female, are absent, andthe abdomen is enormously developed,about three times longer than the rest ofthe body. Both sexes are depigmentedand their eyes are greatly reduced (8compound eyes in the male and 4 in thefemale). The larva of this species is stillunknown.In cold, high-altitude caves, wefrequently find tipulids (crane-flies) of thegenus Chionea, cryophilous, wingless,but apparently not particularly associatedwith caves, which they mainly use forrefuge. Cave-dwelling heleomyzidsinclude the so-called “guano fly”, Thelidaatricornis, a troglophile mainly found nearguano but also the excrement of othermammals and small carrion. It may befound in caves throughout the year invarious stages of its life-cycle. Phorids, in both natural and artificialItalian caves, are represented by somespecies of Triphleba, often near guano.Other families regularly or occasionallyfound in Italian caves are mycetophilids,chironomids, psychodids and phorids.Separate mention must be made ofnycteribiids (see drawing), ectoparasiticon bats, wingless, blind, with well-

developed legs and claws suitable forattaching the animal to its host. Only

females, before laying their eggs, andjuveniles in search of their first

host, leave the bat. Streblids (bat-flies) are also parasitic on bats, but

have wings and are less closelyassociated with their hosts.

Coleopterans (beetles).• Carabids: carabids are one of thelargest families of beetles, with more than40,000 species throughout the world,mainly predators (although theysometimes exploit other food sources).Hundreds of species of carabids havebeen described in various subterraneanenvironments. Their morphological andphysiological adaptations are often veryevident, from long legs and antennae,well-developed sensory bristles on wingcovers, and chemoceptors and olfactoryorgans, to the disappearance ofcircadian rhythms, slowed metabolism,and reduced number of eggs. Togetherwith the leptodirines, they represent thebest-studied and best-represented groupof Italian cave-dwelling fauna. In thevarious species of this family, all stagestowards the conquest of theunderground environment may beobserved: from species with typicalfeatures of surface-living animals,through various stages of adaptation,until the many troglobitic species, called“ultra-evolved”.Italian cave-dwelling carabids belong tofour tribes: clivinines, trechines,sphodrines and molopines. Among the clivinines is only onetroglobitic, Italodytes stammeri, endemicto Apulia, where it is found in the Murgiaand Salento areas.

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Limonia nubeculosa

Most numerous are definitely thetrechines, which form the majority of thecave-dwelling carabids, with elevengenera currently known in Italy, ofancient paleo-Aegeic or paleo-Mediterranean origin.Typhlotrechus bilimeki is a eutroglophilicspecies known in caves of the TriesteKarst and more commonly in Slovenia

and Croatia. Orotrechus includes morethan 30 species, of which at least 20 arepartly cave-dwelling and partly surface-living, found from the Verona LessiniMountains as far as Austria, Slovenia andCroatia. Most species of this genus areendemic to the eastern Italian Alps andPrealps, and many overlap.On the western Alps is the endemicgenus Doderotrechus, with threespecies: D. ghilianii and D. crissolensis,often found together, both above andbelow ground; and D. casalei, morestrictly cave-dwelling. Close affinitieswith Doderotrechus are evident inBoldoriella (central Alps), with a group ofspecies living above ground andinfrequently or never found inside caves,together with another more strictly cave-dwelling group, which also shows ahigher degree of morphologicalspecialization. Italaphaenops dimaioi,endemic to the Veronese LessiniMountains, found for the first time duringan exploration inside the Spluga dellaPreta, is the largest cave-dwellingtrechine in the world. Colonization ofcaves by this troglobitic may go back tobefore the last glaciations. Otherextremely interesting troglobitic are thethree currently known species ofLessinodytes, endemic to the VeroneseLessini Mountains and Brescian Prealpsand also systematically isolated fromother cave-dwelling trechines. Thedifficulty in finding specimens in thecaves in which they are known indicatesthat the preferred environment of theseanimals is not that of true caves, but ofthe surrounding superficial subterraneanenvironment. The most numerous ofItalian cave-dwelling trechine genera,Duvalius, is represented by more than 60

species, in several groups.This genus is widely distributed in theAlps, Appennines, Sicily and Sardinia.Some species seem to belong to a faunaof older origin and are more highlyspecialized and isolated geographically;others are more recent in origin.Agostinia launoi, the only species of thegenus in Italy, is endemic to MonteMarguareis in the Ligurian Alps, where itinhabits colder caves and lives togetherwith some species of Duvalius. Instead,the genus Anophthalmus includes manyspecies, both cave- and surface-dwelling, with various degrees ofadaptation to cave life. It is found inCroatia, Slovenia and north as far as theriver Drava: Italy has about ten species,also with various levels of specialization,found in the Julian Alps.Last on the list of trechines isSardaphaenops supramontanus, highlyspecialized, with Betic-Pyreneanaffinities. It is endemic to easternSardinia, where it lives in various caves inSupramonte di Orgosolo, Oliena and theUrzulei mountains.Most eutroglophile carabids belong to thesphodrine tribe; troglobitic sphodrines arenot known in Italian territory.This tribe contains Sphodropsis ghilianii,endemic to the western Alps, where it isknown in many caves and, in somehigh-altitude locations, also in surfaceenvironments and in marmot lairs.The genus Laemostenus also belongs tothe sphodrine tribe; it is subdivided intotwo subgenera (until some time ago,considered separate genera):Antisphodrus, inhabiting central-easternAlpine caves, and Actenipus, found incaves in the western Alps, central-southern Appennines and Sardinia.

The molopines belong to severalground-living species but there is also atroglobe of particular importance:Speomolops sardous, endemic toSardinia, where it inhabits a few cavesin the province of Nuoro. Its systematicaffinities with other genera of theBalearic islands and Spain set it in thatcategory of animals which reveal thelinks between Sardinian and westernTyrrhenian fauna.• Cholevids: this beetle family probablyoriginated in the Jurassic (or evenearlier), inhabiting the forests whichcovered the southern part of the onlycontinent then existing, Pangea. Thesplitting up of Pangea and the laterdispersal of the continental blocks

which resulted, led to the present-dayarrangement of emerged land whichpartly explains the current distribution ofanimals which originally lived in thesingle ancient continent. There are eightsubfamilies of these coleopterans, mostof which live in the soil. Some of themare associated with varioussubterranean environments, from truecaves to the network of cracksinaccessible to man, to more superficialnetworks and artificial caves. All thecholevids have feeding habits of

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Orotrechus muellerianus

Italaphaenops dimaioi

Duvalius apuanus

Sphodropsis ghilianii

saprobiotic type, eating decayingorganic matter. Together with thecarabids, they show the mostextraordinary and deeply studiedexamples of adaptation to undergroundlife. The most evident aspect isregression of the eyes, already veryadvanced in subterranean species buttaken to extremes in troglobites, inwhich the total disappearance ofganglionic optic centres has beenobserved in some cases. The absenceof body pigment and membranouswings and the lengthening of legs arealso well-known forms of adaptation,found in these insects too. Severalleptodirine colevids and trechinecarabids also have a particularlyrounded form of the wing covers, called“false physogastry”, an adaptationassociated with the need to maintain ahigh degree of humidity for respirationthrough the dorsal surface of theabdomen (see section on troglobitieadaptations).Many studies on the ecology andethology of leptodirines have revealedadaptational strategies, includingreduction in number of eggs,disappearance of circadian rhythms,slowed metabolism, and the exploitationof special aggregation pheromones inresponse to the scarcity of food andparticular living conditions insideterrestrial caves.Of the eight subfamilies into which thecholevids are divided, only two arefound in Italy, and not all of them havethe same preference for subterraneanhabitats. The nemadine subfamily, infact, although including some cave-dwelling species found in Anatolia,Afghanistan, Pakistan and Arizona, is

not represented in Italian caves by anyspecies associated with this particularenvironment. The same may be said ofthe ptomophagines, of which manyeutroglophile and troglobitic species areknown in various parts of the world, butthey are only very rarely found insideItalian caves and never with speciesexclusive to them. The same discourseapplies to the anemadines. There arealso some groups which, although notstrictly cave-dwelling, are frequentlyfound in Italian caves. They includeCatops speluncarum (Sardinia), threespecies belonging to Choleva(peninsular Italy), and Apocatopsmonguzzii (northern Italy).Lastly, the leptodirines represent abouthalf the cholevid species known in theworld, and are those most closely linkedwith the underground environment.Among them we find the most extremeexamples of adaptational strategies,both as regards single organisms andentire populations.In Ligurian caves, there are manytroglobitic and ground-living species ofthe genus Parabathyscia,representatives of which are also foundin the French Alpes Maritimes, Corsicaand England, as well as in Piedmont

and peninsular Italy (some parts ofTuscany, Latium, Campania andCalabria) where, however, they are notfound inside caves but only above onthe ground. In Piedmont, on the easternslopes of the Alpine arch, two speciesof Dellabeffaella are found: D. olmii,living in the superficial subterraneanenvironment of the Cottian Alps, andD. roccai, in caves in the Graian Alps.Three other species are found in thecaves and superficial undergroundenvironment of Piedmont: Canavesiellalanai, C. casalei and Archeoboldoriadoderoana. In Lombardy, there are:Insubriella, with a single species(I. paradoxa) of the superficialsubterranean environment near Brescia;many species of Pseudoboldoria; threetroglobite and one ground-living speciesof Viallia near Bergamo; and manyspecies of Boldoria, many of which aretroglobites, extending their habitat as faras Monte Baldo in the Veneto.In the eastern Alps, the boundary ofwhich with the western Alps is clearlymarked, as regards leptodirines, by theAdige valley, the situation is definitelymore complex and variegated. Manygenera with varying degrees ofspecialization and differing phylogeneticaffinities occur: examples areMonguzziella grottoloi, belonging to thephylogenetic series of Boldoria, the onlycomponent of the eastern Alps showingaffinities with the western genera; andNeobathyscia, which includes severalspecies concentrated mainly in theLessini caves in the Veneto, which havean interesting distribution depending onaltitude between 500 and 1100 metres.Two species of Lessiniella are found inthe province of Vicenza, and two

belonging to the troglobitic genusCansiliella live respectively in a cave onthe Cansiglio and in one on MonteCiaurlec (Friuli), mainly found walking onthe thin film of water which coversconcretions. In caves in the TridentinePrealps, Lessini Hills and the Altopianodei Sette Comuni (Veneto), generally inhigh altitude caves, are five knownspecies of Halbherria. Species ofOrostygia are found in caves in theVeneto and Friuli Prealps - mainlytroglobites which, in particularenvironmental conditions of high stablehumidity, e.g., in the Cansiglio forest,are also found above ground. Of interestalso is Oryotus, in the Prealps from theVeneto to Slovenia: in the eastern areasof this territory they are also found inhigh caves with ice.Of the three species and as many as 15subspecies belonging to the genusAphaobius, only one reaches Italy,A. milleri forojulensis in Friuli; the othersare distributed in Slovenia and Croatia.This genus is closely related with thetwo preceding ones and with Cansiliella.The Italian genus of leptodirines whichshows the greatest degree ofmorphological modification in adaptingto the subterranean environment is

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Leptodirine beetles mating Leptodirus hohenwarthi reticulatus

undoubtedly Leptodirus.This genus, with only one species, isfound in a cave in the Trieste Karst,subspecies L. hohenwarthi reticulatus,but occurs more extensively in Sloveniaand Croatia. As already mentioned, thiswas the first insect to be discoveredinside a cave.As well as the already-mentionedParabathyscia, species of Bathysciolaare found in the Appennines, Piedmont,Liguria, the Veneto and Sicily. They are,however, mainly ground-living speciesand only a few are troglobites:interesting examples of cave-dwellers,like Bathysciola sisernica, B. sbordoniiand B. delayi, are found in thePreappennines in Latium andCampania.Other interesting examples of troglobiticleptodirines are found in Sardinia: theendemic genera Ovobathysciola andPatriziella, and Speonomus, alsocommon in the Pyrenees. Thedistribution of this last genus is anindication of the paleogeographicvicissitudes of the island.• Staphylinids (rove beetles): A family ofbeetles easily recognizable by their longbodies and very small wing covers, ithas few representatives in the caveenvironment. Troglophile species areAtheta spelaea and Quediusmesomelinus, typical of the guanobeassociation; Lathrobium alzonai isendemic to the caves of the Berici Hills.Today, also ascribed to the staphylinidsare the pselaphids (as subfamilypselaphines), small (less than 2.5 mm),avoiding light and hydrophile. Outsidecaves they are found in ant-hills, termitenests, litter, decaying vegetal matter androtten wood. They are all predators,

mainly eating spring-tails. Italy hasabout ten species dwelling in or nearcaves, whereas others are only found incaves occasionally. The definitely cave-dwelling species belong to the generaTychobythinus (species in caves inpeninsular Italy and Sardinia); Bryaxis (insome north-eastern Italian caves);Machaerites (with M. ravasinii in theTrieste Karst and Slovenia, one of themost highly specialized forms of cavelife in Italian fauna); Bythoxenus (withB. italicus reported only from one cavein the province of Udine); andGasparobythus in the Trieste Karst.• Histerids: these beetles, mainly roundor oval in shape, live in dung, carrion,under the bark of dead or decayingtrees, rotten wood, decaying fungi, andin other decomposing materials. Theyare also found in birds’ nests and thelairs of mammals, and are associatedwith ants of various species. They preyon other arthropods, in both larval andadult stages.Only a few histerids have colonized thecave environment. Five troglobiticspecies are currently known in Italy:Spelaeabraeus agazzii (with threesubspecies), S. giorgii, S. infidus andS. tormenei, only found in some cavesin north-eastern Italy), and Sardulusspelaeus (endemic to a cave in theprovince of Nuoro, Sardinia). All thesespecies show total regression of eyes,absence of wings, fusion of wingcovers, and partial lengthening of legs.Occasionally in caves but morefrequently near the entrance, otherhisterids may be found, mainly speciesof Gnathoncus which, however, do notpresent any of the morphologicaladaptations typical of cave life and

cannot therefore be considered astroglobites.• Curculionids (weevils): easilyrecognizable thanks to the greatlengthening of the head, which givesthem a snout-like aspect. Weevils arephytophages: many are highlyspecialized for this type of food, to theextent that they are associated solelywith some parts of specific plants. It isprecisely this feeding characteristicwhich has prevented them fromcolonizing caves on a grand scale. Thespecies most frequently found in cavesare those which eat roots and wood,i.e., more associated with the soil andthe underground environment.Most of these species belong to thegenus Otiorhynchus (subgeneraTroglorhynchus and Lixorhynchus) andare frequent in (or exclusive to) caves incentral-northern Italy. However, southernItaly does have O. (L.) monteleonii in theCampo Braca well and a secondspecies, again of Lixorhynchus, stillundescribed, in a cave in the MontiAlburni. In Sardinia, O. (L.) doderoi hasbeen found in the cave of Sos Turrittas(Golfo Aranci), although it may becollected in great numbers by sieving inthe Mediterranean maquis.

Other genera represented in Italiancaves are Absoloniella, with A. reitteri(two caves in the Trieste Karst),Amaurorhinus caoduroi and Pselactuscaoduroi (two caves in Apulia) andPararaymondyonimus mingazzini(Emilia Romagna).

Amphibians. Excluding trogloxenespecies, which sometimes (likeSalamandra salamandra, the spottedsalamander) successfully breed instreams in caves, the species belongingto the plethodontids and proteids areconsidered to be cave-dwellers.• Plethodontids: this is a family ofcaudates without lungs, with a very

particular distribution. America has 250species; in Europe there are at leastseven relict species found in Italy(Ligurian Alps, central-northernAppennines, Sardinia) and south-eastern France. The species ofSpeleomantes (cave salamanders) areone of the most important naturalisticfeatures of Italy. They live in cliffhabitats, very variable but allcharacterized by little direct exposure tothe sun and high relative humidity, andare always associated with interstices inrocks or underground caves, from which

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Curculionids of genus Otiorhynchus mating

Speleomantes

they seldom move far. Although duringthe night and on the wettest days theymay sometimes be found even above-ground, it is easier to observe theminside karst caves, where they are verynumerous even on vertical, subvertical,or jutting rocky surfaces. On thesedamp surfaces, often exposed toconstantly dripping water, cavesalamanders move slowly, exploitingboth the air-water interface and theadesive capacity of their webbed feet.These animals are quite opportunistic asregards food. Even in total darkness,they capture invertebrates of varioustaxonomic groups, using their sticky,mushroom-shaped tongues which arerapidly projected outside their mouths.In Italian caves, the following speciesare commonly recognized:Speleomantes ambrosii (provinces of LaSpezia and Massa Carrara, less so inthe north-western parts, where the

following species are found): S. strinatii(south-western Piedmont, Liguria,provinces of Savona, Genoa andImperia); S. italicus (Garfagnana, ApuanAlps, provinces of Reggio Emilia, Lucca,Pesaro and Pescara, introduced into thearea near Siena); S. flavus (province ofNuoro); S. genei (Sulcis-Iglesiente,province of Cagliari, Sardinia);

S. imperialis (Nuoro, Oristano andCagliari); S. supramontis (Oliena, Nuoro).Cave-dwelling salamanders areprotected by the Bern Convention andare included in the Habitat Directive ofthe European Community. Their limiteddistribution, sometimes circumscribedto a few systems of underground caves,makes them vulnerable, since in theseconditions they cannot survive radicaltransformations of their habitat. For thisreason, some species are also includedin the Red List of the IUCN (InternationalUnion for the Conservation of Nature).• Proteids: the only Europeanrepresentative of this family, and theonly troglobitic amphibian of Italianfauna, is Proteus anguinus, the proteus,or olm. The other genus of the family,Necturus, is American and lives on thesurface in rivers and lakes. The olm iscertainly one of the cave-dwellers best-known to the public. The history of itsdiscovery goes back as far as 1689, theyear in which Valvasor reported seeing a“pétit dragon” in the caves of Postumia(now in Slovenia).The first exact scientific description wasgiven by Laurenti in 1768.Olms have an eel-shaped body and theeyes, in adults, are atrophied andhidden under the skin; the front feethave three toes and the back ones two.The colour is whitish, or slightly pink,but exposure to sunlight induces weak,temporary pigmentation. Among thecharacteristics which make the olm anexceptional species among Italian faunais undoubtedly the phenomenon ofneoteny. Neotenic animals are thosewhich reach sexual maturity andreproduce although still retainingmorphological features typical of larvae.

The phenomenon is frequent in somecaudates, like newts, which sometimesmaintain the external gills typical of thelarval stage even when they are fullygrown. However, although in newtsmetamorphosis may be induced bynatural stimuli or in the laboratory, in theolm neoteny is obligatory andirreversible.The reproduction of the olm has longbeen a mystery. Once, there wereopposing theories which led to acontroversy as to whether it wasviviparous or oviparous. In fact, the olmis oviparous: in the natural state, eggsare laid one at a time over a period ofabout one month, numbering from 20 to80. Normally, they are attached to eachother on rocks and stones, and thelarvae are born at ambient temperatureafter 13-20 weeks. Surprisingly, thelarvae, greyish in colour, possess clearlydistinct eyes until the age of about 2months. Measuring 16-22 mm at birth,the larvae grow slowly and until the ageof 3 months are nourished exclusivelyby their yolk-sacs; only later do theybecome predators. In nature,reproduction rarely occurs before thetenth year of age (11-14 in males, 15-18in females). Adult olms are predators

and mainly eat aquatic oligochaetes andcrustaceans; in Italy too, cases of olmsmoving to the surface to prey on smallarthropods have been reported.The origin of the olm and the antiquity ofits colonization of subterranean watersare often debated. Fossil remains (ofHylaeobatrachus croyi) found atBernissart in Belgium, together withthose of iguanodonts, going back to theLower Cretaceous, indicate thatproteids lived in surface waters in thosetimes. The most accredited theory in thepast was that the above-groundpopulations of proteids persisted inEuropean superficial waters at least untilthe mid-Miocene, when the areas wherethe olm currently lives were in a pre-karstic phase and there was still anextensive surface water network. Withprogressive karstism in the Pliocene, theolm presumably actively colonized thesubterranean waters of the Dinarickarst. However, this theory was recentlyconfuted by Slovenian researchers, afterthe discovery of a new subspecies(Proteus anguinus parkelj, described in1994). Specimens with a flatter muzzle,eyes, and pigmentation, were found inthe Bela Krajina karst, and geneticanalysis demonstrated that they are verysimilar to the troglobitic species of thesame area, so that a much more recentcolonization of the underground domainwas postulated. However, recent studieshave shown that there may in fact bemore than one species of olm in theDinaric area (in 1850, as many as sevensubspecies, according to the number ofvertebrae, were described), thus againraising the possibility of pre-Quaternarycolonization of underground waters. Butthe genetic distance between the

122 123

Speleomantes strinatii

Proteus anguinus

populations of Bela Krajina and those ofPostumia date the divergence to theUpper Miocene, i.e., the old hypothesisis sustained.The olm is found from the river Isonzo(province of Gorizia) as far asMontenegro, and also in the hinterlandof Slovenia, Croatia and Bosnia. In Italy,it is known in many caves in the Isonzoand Trieste Karst where, althoughthreatened by recent pollution anddisturbance of its habitat, it is,fortunately, still common. There is alsoan isolated colony in the Grotta Paroliniat Oliero (Valstagna, province ofVicenza). It is known, however, withcertainty that this population derivesfrom the introduction, in 1850, of someSlovenian specimens, whichacclimatized well. Due to its exceptionalnature and the restricted area itoccupies in the present-day territory ofthe European Community, the olm isprotected by various international normsand, in particular, is on Annexe II of theHabitat Directive and the IUCN Red List.

Bats. Although bats are not alwaysclosely associated with caves, thecollective image attributes to them aspecial link with underground caverns.There are almost one thousand species,all greatly varying in aspect, size andhabits. As well as being able to fly bymeans of the wide flaps of skin spreadbetween the phalanges of the “hand”,hind legs and tail, the most spectacularfeature of bats is their capacity for echolocation. Although many other mammalsuse ultrasound for communication, tolocate prey, or to perceive thetopography of their surroundings(whales, shrews, etc.), in insect-eating

microchiropterans this capacity hasbeen brought to a point of exceptionalrefinement, since bats use it constantly,even during their most rapid aerialacrobatics. In a whole series ofactivities, they emit rapid sequences ofultrasonic clicks and use the returnechoes both to locate their prey in flight,to capture it, and to orient themselves inthe absence of light and thus avoidobstacles. For this reason, they cannothunt during rain, and generally takerefuge in their hiding-places a fewminutes before the onset of anydownpour.European bats generally have somecapacity for thermoregulation. Thiscondition, known as imperfecthomeothermy, is quite widespreadamong mammals in temperate climates(hedgehogs, dormice, marmots, a fewcarnivores, etc.) and allows them toconserve metabolic energy both duringperiods of rest during the day and inwinter, when the environment isparticularly lacking in resources. In thisperiod, the animals sleep inside ahibernacle with constant temperature

and humidity, in which they may alsomate. The places chosen for hibernationare usually cavities in cliffs or trees,cracks in ceilings or badly fitting doorsand windows, or abandoned buildings,and differ according to species.Although some bats (e.g., barbastels)like cold well-ventilated cavities, theygenerally choose places which aredeep, damp and preferably constant intemperature even during winter. Insidethese hibernation cells, gregarious batsmay gather in impressive numbers.Dwarf bats (Pipistrellus pipistrellus), forexample, gather in very crowdedhibernacles, sometimes hundreds orthousands of them, all heaped one ontop of the other. In one Roumaniancave, one of these colonies wasestimated to number 100,000 bats.However, their hibernation is far frombeing continuous. If they are disturbed,or if the temperature falls drastically, thebats wake and actively seek betterplaces. These winter movements maybe very spectacular, sometimesinvolving hundreds of animals in searchof new locations in which to spend

the winter.The reproductive cycle of bats is alsovery peculiar. The mating period beginswhen the mature females scatter fromthe nursery, between the end of Augustand autumn. Mating may last all winterand include most of the spring, but itoften occurs inside the hibernacles(Myotis). However, in various species ofthe genus Nyctalus and Pipistrellus, themales actively defend true harems of 2-5 females, which group together at thebeginning of the mating season and aredefended by the same male for severalyears running. Births occur around mid-June, when the pregnant females havealready grouped into large nurseries,sometimes numbering hundreds orthousands of pregnant or nursingmothers and small numbers of males.Knowledge about Italian bats is stillscanty. In Italy, there is no specifictradition of studies on bats, although inthe late 1980s the attention ofresearchers did focus on these animals.In Italy, there are about 30 species ofbats, only some of which are closelyassociated with underground caves

124 125

Plecotus auritus Myotis myotis Myotis myotis hibernating

(horseshoe-nosed bats, miniopterans,barbastels, and several species ofMyotis). Some also frequently usevarious types of buildings or cavities intrees at different moments of their life-cycle. Tracing the boundaries betweentroglophiles, anthropophiles and forest-loving bats is thus really very difficult,mainly because buildings are truereplacements for cliff habitats, whichwould otherwise be quite scarce formost of the anthropophile species. Togive only one example, the greatbiological success of Kuhl’s bat(Pipistrellus kuhlii) is due to its capacityto use both urban and suburbanbuildings, which are very common,where crowded bat nurseries may reachgreat densities.During a cave excursion, it is easy tosee signs of bats. In many cases, theseare small heaps of droppings under theirfavourite resting-places. It is not alwayseasy to actually see them, eitherbecause they hide inside tiny cracks orbecause the droppings may be old andare only a souvenir of their past activity.Large accumulations of guano areinvariably the result of late spring orsummer reproduction. Outside theirshort reproductive period, the samecavities are not used and are deserted.Inside Italian caves in summer, it is quite

easy to locate the nurseries of varioustroglophile species.The most common are:Miniopterus schreibersii (colonies ofthousands of bats, hanging by all fourfeet, often two or three layers deep, andoften associated with various species ofMyotis or Rhinolophus; Myotis myotisand Myotis blythii (dozens of animalsclose together, hanging by their hindlegs, sometimes inside buildings);Myotis daubentonii (dozens of batsclose together, hanging by their hindlegs, sometimes in buildings, nearwater); Rhinolophus hipposideros,R. ferrumequinum, and R. euryale(dozens of specimens hanging by theirhind legs, neatly folded in theirwing-flaps, and not in contact witheach other).In winter months, it is easier to see thevarious hibernating species. They arenot necessarily definitely troglophiles.Apart from the great bats, or noctules(gen. Nyctalus), most European batsmay hibernate underground. In order to reproduce or hibernate, batsneed peace and quiet. Even duringmonitoring and measurements, humansshould not impose their presence formore than five minutes consecutively.

126 127

Rhinolophus hipposideros

Bat colony

129128

The only European vertebrate perenniallyassociated with water in subterraneancaves is, as we have seen, the typicalsubspecies of the olm. In Italy and insome French caves, there are somecaudate amphibians closely associatedwith the cool, dampenvironment typical ofcaves, cave salamanders(Speleomantes), but theyare not rigorouslyassociated with thedarkness of caves and,at nightfall, maysometimes be foundon the surface,where they frequentwell-fissured cliffhabitats, cool,shady and damp,preferably withdripping water.Bats toofrequentundergroundcaves, but onlytemporarily,either to sleepduring the day inthe breedingseason, or (forsome species) tohibernate.Some strigiforms(tawny owls),columbiforms (wild pigeon),rodents (dormice, fieldmice, black rats)and carnivores (wild cats, foxes,badgers, bears, pine-martens) are alsoseen near the entrances to large caves,but in this case their presence istemporary, almost always connectedwith their resting periods during the day,or else reproduction. All these species

are defined as troglophiles and theiractivity inside caves, although frequent,is never exclusive. Signs of theirpresence are easy to see during a visit toa cave - if not the actual animal, thentraces of its activity or its remains.

Excrement may reveal recentunderground activity, and in

certain cases, it issurprising to finddroppings in placessometimes very farfrom the entrance tothe cave (fieldmice,black rats, pine-martens). Theexplanation here isvery often that thecaves have tinyopenings, sometimesunknown or very farfrom those officiallyknown to man andentered by

speleologists, and theanimals habitually use

them.However, although clear

signs of predatory activityby troglophiles (bones

of small mammalsregurgitated by owls,

dormouse droppings,remains of fox’s prey)

may be found at the entrance to a cave,the presence of Lataste’s frogs,European frogs, green frogs, newts orspotted salamanders - all surface-livinganimals - must be explained in someother way. Typically, these animals areinvoluntarily carried into caves duringtheir larval or adult stages, and are allmainly found in caves after extraordinaryevents, such as sudden floods,

accidental falls into subvertical shafts,etc.. Sometimes even snakesremain imprisoned inunderground cavities, butthey are generally therein a spontaneoussearch forsomewhere tospend the winter.Caves may alsocontain bones ofuncertain origin andattribution, They areoften fossilized orsubfossilized remains,but may often appear veryrecent. When they are true fossilremains, they have a particularly highspecific weight. Their internal structure isoften completely mineralized, so that ifthey are exposed to a naked flame theydo not burn or emit the smell typical ofburning fresh or subfossil bones. Incertain cases, they are the remains ofpast underground activities of nowextinct troglophile species (e.g., the cavebear, Ursus spelaeus). But not allaccumulations of bones found insidecaves are there as a result of their use bypast troglophile species, intrusive orliving in clefts. For example, IsabellePathou, a French researcher who hasattempted to establish the origin of theremains of subfossil bones of marmotsin some Pyrenean caves, reached theconclusion that they were the remains ofmeals by ancient populations of hunter-gatherers. By carefully examining thesurfaces of these bones, she foundscratches, cuts and non-parallelabrasions, certainly made by primitivebut sharp stone instruments. The mostfrequent origin of bone deposits near theentrances of most caves is

Vertebrates occasionally found in caves Luca Lapini

anthropogenous.In many cases, they

are recent accumulationsof the remains of domestic

animals, mostly dogs, but also sheep,goats, cattle and pigs, which werethrown into shafts and subvertical holesin the ground. In some areas in theTrieste and Gorizia Karst, but also inSardinia, Sicily and the central-southernAppennines, this habit was commonamong rural populations, who were thusable to get rid of sometimes largeamounts of waste quickly. This rapidwaste elimination system was (and stillis) used in Prealpine areas by huntersand poachers, and in caves in the JulianPrealps and the Karst the remains ofchamois, roe deer and boar are quiteoften found. The skeleton of a jackal(Canis aureus) was recently found in asmall shaft near Doberdò del Lago(Gorizia), where it had been illegally killedand hidden by poachers. It lay on top ofa large number of skeletons of dogs, thesilent witnesses of a long tradition ofusing that particular shaft for thepurpose. In some places, these shafts(notoriously called foibe in Italian) werealso exploited in wartime. It was asimple matter to throw unwantedpersons or prisoners awaiting execution,alive or dead, down them.

Tawny owl

Skull of cave bear

131

■ Exploitation of caves for tourist, speleological and conservation purposes

The extent and distribution ofunderground tourism. Caves andunderground environments in generalare entered by two types of visitors:speleologists and tourists. The 800large accessible caves in the world arevisited by about 170 million peopleevery year, and play a definite socio-economic role in providing earnings tomore than 10 million people.In Italy, subterranean tourism in naturaland artificial caves involves about 2.5million people every year, and is aconsiderable source of revenue and of

potential naturalistic interest which cannot be neglected.Most of these people (about 1.5 million per year) visit about 70 “show caves”and consequent environmental problems only concern a few dozen caves:about 0.3% of currently known natural caves (there are more than 33,000 inItaly). Show caves are large-scale infrastructures where income from “cavemanagement” must co-exist with the proper conservation of the very placewhich produces that revenue.Parallel with this influx of visitors, there are very many enthusiasts ofspeleological excursions who visit all the remotest corners of the “non-tourist”caves. Including those who follow courses and visit caves as tourists, it isestimated that about 12,000 speleologists organize excursions every year: ifthis activity were homogenously distributed over the 33,000 known caves, therelative environmental damage would be negligible. But this is not so: there arecaves all over Italy which have been explored by so many thousands of peoplethat they have sometimes been seriously and irreversibly damaged.

Protection and conservation of theunderground environmentMAURO CHIESI · LUCA LAPINI · FABIO STOCH

Grotta Gigante, near Trieste (Venezia Giulia), one of the most popular tourist caves in Italy

UFFIZI MUSEUM, FLORENCE (21%)

POMPEI (23%)

COLOSSEUM (27%)

CAVES OPEN TO TOURISTS (29%)

Extent of cave tourism in Italy in 1998

133132 Speleologists and show caves. Withthe growth of interest in speleology,and in view of the delicate equilibriumof karstic environments, the first signsof excessive exploitation began tobecome apparent - precisely in thoseplaces visited not only for their beautyand integrity, but also for theirvulnerability and the peculiarly fragileenvironmental mechanisms whichcreated them. Thus, parallel with thegrowth of environmental awareness ofspeleology, there arose the need toplace restrictions on “direct

consumption” (the caves themselves, and removal of concretions) and theunconscious exploitation (the impact of humans) of karstic territories. Itbecame apparent that there is a “tolerability threshold” even as regards thenumber of visitors - whether they are tourists or speleologists or occasionalvisitors - to a single cave or just part of one.Speleologists, despite great technical difficulties, may spend hours insidecaves, emerging with the impression that, during their sojourn underground,they have touched the centre of the Earth. Conversely, the more easilyaccessible parts of certain relatively shallow caves are suitable for touristvisits: pathways can be laid out, galleries excavated, and lighting of all kindsinstalled, to enable the cave to be exploited as a place in which, as in no other,a set of fantastically shaped concretions may be seen and admired. Over theyears, techniques have been refined, courses in speleology are now offered inplaces where, a few decades ago, they had never been heard of. No materialor waste of any kind must be abandoned in caves, just as it must not beabandoned on paths on the surface. Caves have turned out to be much largerthan previously believed, but also much less capable of tolerating the impactof those who wish to explore them.Attractive “show caves” have also shown themselves to be very fragile andvulnerable: they deteriorate under the impact of millions of visitors. Theparallel pathways - speleology in itself, and tourist exploitation of thesubterranean world - have begun to converge and even meet: now we knowthat caves must be protected both from careless visitors and from ignorantspeleologists.Those speleologists who have an aversion towards “show caves” are

concerned about the damage which tourists cause and about theunconscious mass-tourist concept of “consumption” and thus of theirreversible destruction of natural beauties - the object of the original touristattraction itself - and known precisely thanks to the work of exploration anddocumentation provided by speleologists.All visits to underground environments, even sporadic, cause alterations to theecosystem. They may be temporary, with a general increase in energy to thesystem, or permanent, like that abominable habit of ignorant persons whobreak off concretions and take them away as souvenirs. Pollution anddestruction are caused by ignorance or under-estimation of the damageresulting from certain actions.Only by studying the effects of mass exploitation of show caves can we beginto understand what, and to what extent, damage may be caused to the fragilesubterranean world.

The “energy level” of a cave. The concept of a cave’s energy level (developedby Heaton in 1986) is an essential parameter which allows us to predict,approximately, the relative “weight” of the influence of man on the subterraneanenvironment. There are three different levels, decreasing in degree:1. high-energy caves: periodically involved in large-scale events like floods;2. medium-energy caves: energy is supplied by small watercourses, wind, andanimals;3. low-energy caves: the flow of energy is practically reduced only to thedripping of water.

Obviously, people can enter the cavesof the first category without causingproblems: the periodic natural suppliesof energy are capable of cancelling anychanges caused by the presence ofvisitors. Instead, medium-energycaves, rich in concretions, may beseriously disturbed by great numbersof visitors, when the consequentenergy supply becomes comparablewith the energy balance of the caveitself. Lastly, low-energy caves shouldnot be visited: the very presence ofhumans can irreversibly disturb theirgeneral equilibrium.

Caves at Monsummano Terme (Tuscany) usedfor thermal spa purposes

San Giovanni d’Antro: church at entrance tofortified cave with spring

135134 The concept of the “receptive capacity” of caves. From the viewpoint of theflow of energy or mass, most caves are almost isolated environments: with theaim of keeping environmental disturbance under an acceptable threshold levelof reversibility, their basic environmental parameters must be changed as littleas possible. The “receptive capacity” of a cave may be defined as: “... themaximum number of visitors acceptable over a certain period of time and indefined conditions, which does not imply any permanent change in anysignificant parameter”. (A. Cigna). This definition may be applied to allunderground environments. Recognizing that environmental parametersundergo natural oscillations in time, attention must be paid to those variationswhich, due to visitors, move too far away from these natural trends. In thisway, we can determine which “critical” parameters must constantly bechecked with especial care. This is why preliminary monitoring of any projectfor tourist purposes is indispensable. We must measure the natural referencevalues and establish the natural dynamics (i.e., the energy) of the system,which is exclusive and characteristic to every cave or undergroundenvironment.Putting the “speleologist” and the “tourist” in the same boat is not anexaggeration. From the viewpoint of the cave ecosystem there is nosubstantial difference, either as regards the scale of the problem (in our case,a single cavity) or the analysis of possible factors of deterioration. On the scaleof a single cave, the quality of damage which may be produced is basically thesame: damage increases according to the numbers of people who passthrough the various “sensitive” areas of the cave. Its adaptation for touristpurposes therefore, mainly concerns the quantity rather than the quality of thedamage - obviously, except for caves in which tourist adaptation correspondsto actual infrastructures inside them (pathways, removal of “obstacles”,widening, opening of new entrances, etc.) and external ones (construction ofaccess roads, car-parks, etc.), which are gross and irreversible. Possibleanswers to these problems are in fact identical, and all concern:- prior assessment of environmental conditions preceding explorations or visits;- minimization of anthropic impact;- code of ethical and deontological behaviour, and need for surveillance.

■ Environmental degradation and water pollution

Today, karst aquifers represent about 40% of our sources of water supply (fordrinking and other uses) in the Mediterranean basin, and they always tend tobe increasingly exploited, in spite of the fact that these sources of supply areMain causes of degradation in natural caves

contamination:wastecarbonbatteriesclothingdyes

physical damage:breaking off of concretionspeople treading everywhereexplosionsvandalismwidening of narrow points

removals (thefts):concretionsarcheological depositsmineralsfauna

graffiti:carbide from lampspaintsscratches and cutsin rock surfaces

quarrying and mineexplosives

caves “eaten away” by quarrying and mining caves filled with:solid urban wasteinert aggregatecementindustrial wastefarming and food wasteshydrocarbonssewageremains of dead animals

pollution byfarming waste

pollution byindustrial waste

illegal dumping of waste

H U M A N C A U S E S · S P E L E O L O G I S T S D E T E R I O R A T I O N

S E R I O U S D A M A G E T O C AV E S

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

D E S T R U C T I O N O F C AV E S

I N D U S T R Y T O T A L D E S T R U C T I O N

under the surface. This is the case of all aquifers in gypsum which, forexample, makes up about one-third of the thirsty Sicilian provinces ofAgrigento, Enna and Caltanissetta.Lastly, mention must be made of the “tamping” effect in karstic aquifers incases of flood. It may act directly both favourably, by diluting a pollutant (ifclean waters are “jettisoned” from the network of cracks in the aquifer). Viceversa, it may act very negatively (in the case of the large-scale removal ofpollutants trapped in siphons and bodies of waters in the vadose zone), sinceconcentrated waves of polluting substances are returned to the surface.Safeguarding the quality of subterranean waters also means conserving theintegrity of its delicate animal community, both to protect that animal life, andfor practical reasons, since organisms play an essential role in recyclingorganic matter and thus in purifying waters. When water quality is referred tothe integrity of its populations, we speak of biological quality.Methods for assessing the biological quality of surface waters have beenroutinely used to assess their degree of pollution for about 20 years. Theadvantages of these methods, based on study of benthic macroinvertebrates

137136 poorly known, poorly studied, and even more poorly protected. Their peculiarcharacteristics mean that they are generally very vulnerable to pollution andgreatly exposed to the risk of contamination, both accidental and permanent.Some of the main sources of pollution and contamination are waste productsfrom industry, farms and agriculture, as well as deliberate dumping of solidurban and industrial waste and, last but not least, permanent or accidentalpollution due to emissions of fumes from traffic.The dynamic characteristics of karstic aquifers are such that they can do verylittle to combat the spread of polluting substances: they have fast-flowingwaters and very little capacity for self-depuration.From the surface to the surface again, the course of a polluting substance isas follows:- the pollutant enters the system;- it migrates to an unsaturated (vadose) zone;- its propagates in the saturated (phreatic) zone;- it is sent back to the outside world.If the karst cover (where present) can reduce the effects of pollution, in thecourse from the vadose zone to the phreatic zone it is flow velocity whichdetermines self-purifying capacities directly, influencing sedimentation andoxygenation and thus the overall biological conditions (bacterial and antibioticactions, closure of the biological food cycle by predatory organisms). Thesephenomena, essential in self-depuration of surface waters, are far less activeand efficacious in the non-saturated karst zone.In the phreatic zone, in the absence of oxygenation, purifying power ispractically limited to possible dilution from non-polluting sources.Lastly, the return of the pollutant depends on the overall hydrogeologicalfeatures: if the aquifer runs mainly in conduits, its response is extremely rapidand concentrated. In systems of networks of cracks, the polluting load arrivesslowly and is diluted, often to a considerable extent, over time. It is interestingto note how the time factor acts differently according to type of pollutant: it isimportant in the case of pathogenic bacteria but has no influence in othercases, e.g., chlorides.Simplifying again, extremely unfavourable factors are the absence of vegetalcover, the presence of active vertical shafts, and the outflow of rapidly-movingwaters.Polluting load being equal, the size and shape of the aquifer directly influenceits degree of vulnerability: springs which drain waters from very large areaswith very thick non-saturated areas (karstic massifs) are less vulnerable thanquite thin aquifers with phreatic networks located only a few dozen metres

Degradation induced in underground environments as a result of visits by humans

SPELEOLOGISTS AND TOURISTS

CONTAMINATION

PHYSICAL DAMAGE

REMOVALS (THEFTS)

GRAFFITI

IN ADDITION: ALL DAMAGE DUE

TO ADAPTATIONS TO CAVE FOR

TOURIST PURPOSES - VERY

OFTEN IRREVERSIBLE

DAMAGE IS PRODUCED

ACCORDING TO:

1. QUANTITY

2. QUALITY

3. NUMBER OF PEOPLE IN CAVE

(CAVE TOURISM)

4. TIME SPENT IN CAVE

ENVIRONMENTAL RE-EQUILIBRATION

LIGHTING

EXCAVATIONS

CAT-WALKS, WALK-WAYS

SMALL-GAUGE TRAINS

SUPPORTS

TECHNOLOGICAL PLANT

CLEANING SYSTEMS

CONTAMINATION

FURTHER GRAFFITI

139138 (bottom-living organisms larger than 1 mm), are known to be better thanchemical or microbiological methods. Aquatic organisms, being sensitive toeven slight degrees of pollution and living all their lives in a wateryenvironment, record variations in water quality with considerable precision andmay successfully be used even when chemical analysis does not supplyadequate results (e.g., in cases of intermittent or episodic polluting events).The use of bio-markers in the study of underground water quality is morecomplex, due to a series of problems which are difficult to solve:- lower biodiversity (insects being almost completely missing) means thatidentification of organisms must be made at species level and not at that ofgenus or family, as happens in surface waters; in addition, meiofauna (i.e.,organisms between 0.3 and 1 mm in size) must be examined;- the need for special techniques for sampling, study and identification ofspecimens is often time-consuming and based only on specialized literature.This last point is further complicated by two more factors:- we have insufficient knowledge of the taxonomy and distribution ofspecies living in Italian underground waters, in which species new to scienceor to Italy are discovered every year;- endemic species tend to form rapidly, so that the fauna may be different ineach single karstic massif.In spite of these technical problems, French, Dutch and American studieshave shown the efficiency of this method in the ecological surveillance ofaquifers, and techniques suitable for interstitial waters have been developed.For karstic waters, the problem of sampling is more complex and techniquesare still being perfected. In Italy, they are rarely applied to cave and karstspring waters. In general, in cases of slight organic pollution, it has been notedthat the number of stygobites, which are the most sensitive organisms,decreases in favour of stygophiles and stygoxenes, which cover a largerecological valency, until we reach threshold situations in which stygobitesdisappear completely, to give way to dense populations of stygoxenes whichtake advantage of the accumulations of decaying organic matter. Thisphenomenon was recently observed in a study on the fauna of the Lessini Hillswhere, in a cave with a high degree of organic pollution (Spurga di Peri), onlyordinary stygoxene species of tubificid oligochaetes, copepods and ostracodswere found, accompanied by cladocerans and dytiscid beetles which areinfrequent in subterranean waters. This encouraging result means that thepopulations of underground karstic waters allows us to assess the state ofbiological integrity of their ecosystems, and reveals the possibility of creatingindices of water quality in the very near future.

■ Norms for protection and discipline in speleological practice

For a country in which more than one-third of the territory is karstified andmore than 40% of water resources for drinking purposes comes from karsticaquifers, the current overall norms for local and regional protection of thesespecial and essential resources are not simply inadequate - they areunacceptable.Little or nothing concrete has been done to safeguard Italian karst areas - neitherto prevent the consequences of the impact of public and private works on bothsuperficial and subterranean karstic sites, nor to remedy those consequences, ifpossible. Every time pollution of a karstic aquifer has to be investigated, thepossible ways and means of acting are uncertain. For example:- Italian law no. 1089 of 1939 (recently substituted by legislative decree no.490 of October 29 1999), which safeguards assets of artistic, historical,archeological, ethnological and paleontological interest, is only applicable tothose caves (quite numerous) which contain assets of those types: i.e., itprotects the “contents” but not the “container” (the karst landscape, thesurface area, the cave itself, and the aquifer);- law no. 1497 of 1939 (recently substituted by legislative decree no. 490 ofOctober 29 1999) precludes the possibility of protecting superficial karsticareas which do not have peculiar landscape or geological characteristics -whereas we know that a landscape without these “peculiarities” may be ofessential importance in draining surface waters in covered karst. But even incases of constraint under this law, it is only the external aspect of the cave(and only that), for which any modification must be authorized by the localauthorities (Sopraintendenza). If the law is not observed, penal regulationsapply (Art. 734 of the Italian Penal Code, destruction of or damage to classifiednatural beauties), involving fines from 1 to 6 thousand euro; - law no. 319 of 1976, dealing with water pollution, aimed at disciplining andregulating waste dumping in water, soil and subsoil, but even in its laterarticles and assignments to local government, it never constituted a validresponse to the peculiar needs of karstic aquifers and their protection. Thereare innumerable cases of dumping of sewage, apparently lawful, which dailyaccumulates inside karstic caves and which, every so often, simply returnsconcentrated pollutants with the next outflow. And what can be said of thewaste dumps, controlled and uncontrolled, scattered over the plains ofApulia? A perfect example of an aquifer in very rapid and irreversibledegradation, partly due to excessive pumping of water, has given rise topermanent invasion by seawater over enormous areas - and that water cannot

141140 even be used for agricultural irrigation. In this framework, therefore, it is notsurprising that, in order to protect these natural resources, frequent appealsquote Royal Decree no. 1016 of 1939 (a happy year, evidently for legislationprotecting the beauties of Italy, but now too far removed from present-dayreality). The decree also contains norms regulating hunting, in which Art. 38covers special protection for bats. Again, the legislative decree of December12 1923 concerns hydrogeological constraints, and royal decree no. 1443 ofthe 1927 laws covers mining.In Italy, there is no national law which sanctions, by safeguarding theirintegrity, the peculiarities of karstic environments containing aquifers providingwater for drinking and agricultural purposes. This situation has requiredmeasures on the part of various regional authorities - sometimes well-definedbut in themselves poorly organized and sometimes, for this reason, ineffective.The very regional laws “favouring speleological activities”, already operationalin some regions since 1972, do not safeguard the karstic environment. Theonly practical impetus resulting from these legislative instruments has beenthe Catasto delle Grotte, the Cave Registry, an essential and dynamicinstrument for proper knowledge of the territory.Laws for national parks and others deriving from legislation providing forregional and provincial territorial landscape plans, again due to the lack of anational law safeguarding the karstic environment, would represent a uniqueopportunity to make up for lost time and delays, and would fill many gaps. Butthere are very few cases in which this has happened - all linked to thefortuitous but combative presence of speleologists among those responsiblefor regional planning. Further progress was made with legislative decree no.152 of May 11 1999, concerning measures for protecting waters frompollution, including subterranean waters. However, this law too, like itsprecedessors, considers waters according to their use by humans (for drinkingpurposes, agricultural or industrial use) and not according to the interest whichtheir environment may hold.

■ The Habitat Directive and the subterranean environment

Directive 92/43/CEE of the European Community, dated May 21 1992,regarding conservation of natural and semi-natural habitats and wild flora andfauna, is called the “Habitat Directive”. It has recently been integrated (byDirective 97/62/CE of October 27 1997, updated to reflect technical andscientific progress with respect to the original directive), and finds itsapplicational regulation in Italy in DPR no. 357 of September 8 1997. TheseHall with concretions in Grotta Mitica (Julian Prealps, Friuli)

powerful instruments aim at guaranteeing the conservation of natural habitatsand of the karstic environment, together with its flora and fauna.The Directive has as its aim the safeguarding of habitats and species of interestto the European Community, and supplies clearcut criteria of selection,according to the presence of rare, endemic, vulnerable or at-risk species.Although information on the state of risk is scanty, rarity and endemicity arecriteria which may be applied perfectly to troglobite and stygobite species, andtherefore many may be included in any forthcoming normative updating amongthe species of Community interest. Not only this: the Directive already supplieslists of habitats and species of interest to the European Community.Of essential importance is Annexe I of the Habitat Directive (Annexe A of DPR357), which lists the types of natural habitats of Community interest, conservationof which requires the designation of special conservation areas. Art. 8 of the97/62/CE Directive indicates “Rocky habitats and caves”, and includes:- 8310 (65 of the preceding norms): caves not yet exploited for tourist purposes;- 8320: lava fields and natural cavities;- 8330: submerged or semi-submerged sea grottoes.Clearly, member states are specifically required to safeguard these environments.But the caves included in sites of Community or national interest (Natura 2000network and Bioitaly sites) are limited in numbers, with the exception of those insome autonomous regions or provinces (e.g., the Trentino) which have listedmany sites to be protected. The weak point of decisions taken until now hascertainly been that of protecting mainly single caves and not entire karstic areasor water networks; but the protection programme has now been started andcannot be stopped. As regards fauna, Annexe II (Annexe B in DPR 357) listsanimal and plant species of Community interest, the conservation of whichrequires the designation of special areas, and Annexe IV (Annexe D in DPR 357)(animal and vegetal species of Community interest which require strictprotection). Among the vertebrates we find all the bats frequenting Italian caves(and particularly all species of the genus Rhinolophus in Annexe II, the words “Allspecies” of microchiropterans in Annexe IV) and, among the amphibians, cavesalamanders (Speleomantes) and the olm (Proteus anguinus). The list ofinvertebrates, in which there are practically no troglobites, is highly deficient.Clearly, caves with bats, newts and olms can be proposed as sites to beprotected according to the Habitat Directive.Art. 8 of DPR 357 states that, for species in Annexe D (i.e., all bats, newts andolms), it is forbidden to capture or kill specimens of these species in the naturalenvironment; to disturb them, particularly during all stages of their breeding, orduring hibernation, wintering or migration; to damage or destroy their breeding or

143resting areas (i.e., all caves in which these species may do no more than rest).Their possession, transport, exchange or marketing in the natural environment isalso forbidden, except for specimens legally collected before the coming into

force of the DPR. Both speleologistsand biospeleologists must be clearlyaware of these restrictions.

■ Strategies for protecting bats

The equilibrium of predators with lowreproductive rates is fragile. If for somereason their numbers are critical, theymay be threatened by even slightsituations of disturbance, which inoptimal conditions could more easilybe well tolerated or even ignored. Batsenter this category and are currently

viewed as among the vertebrates with the highest risk of extinction in manyindustrialized countries. Bats are particularly delicate animals, partly becauseof their imperfect homeothermy, i.e., they are very sensitive to even slightclimatic and seasonal oscillations. For example, prolonged rainfall during thebreeding season of various species may lead to very high neonatal mortality(miniopterans, minor horseshoe-nosed bats) and their distribution may even beinfluenced by recent changes in the climate, at least partly caused by man.Although information on the current situation of European bat populations isstill scanty, there are various anecdotal data referring to the past, whichhypothesize that bat populations have been in a critical state for a long time.These data are often discontinuous but common to various central Europeancountries (Great Britain, France, the Low Countries, Switzerland, Germany),and appear to agree on the general trend towards reduced numbers of mostlarge bat colonies which have been monitored for longest. Although previouscomparative data are available on limited local situations and few species, thebats most at risk appear to be the major and minor horseshoe-nosed bats(central Europe), major vespertilionids (Germany), barbastels and dwarf bats.The situation is not homogeneous, since signs of recovery are noted for somespecies in some areas (Daubenton’s bat), but the relative density of batpopulations is declining everywhere.The causes for this are still not completely clear, but may be grouped intocategories, all associated with recent developments in man’s activities.

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145144 In Italy, all species are also protected by law no. 157 of 1992, which forbidsthem to be killed or disturbed. Various ways of reducing the impact of man’sactivities on bat populations have been suggested, but in many cases they aresimply proposals or declarations of intent. In view of the great variability ofdirect and indirect possiblities of conflict between the logistic and economicneeds of man and those of these delicate animals, few of these measures arereally effective or sustainable.If they were properly applied, they would influence the very models ofdevelopment of Western societies, by drastically restricting the use ofchemicals in agriculture, imposing different criteria on buildings, and providingfor proper forestry management. Even the distribution of nesting-boxes in

certain areas is merely a palliative, since they are onlyused by a few species and are in any case very

limited, mainly as supports to researchprogrammes. However, as regards bats

breeding in caves, the most practical andfunctional protective measure against

direct disturbance by man is to denyphysical access to the most frequently

visited caves (see drawing).

■ The ethics of speleology

Mountains belong to everybody. Speleologists, who are beginning to knowthem inside too, have some rights but also many more obligations. Everyconscientious speleologist is obliged to prevent and correct any damage tocaves and to the water resources which they contain.Some recommended ethical and deontological rules are shown below.

Prevention of damage to the external environment- Request authorization for visits whenever possible, scrupulously respectingthe arrangements made by owners or managers of natural parks and reserves;- Always inform others about the particular dynamics and vulnerability ofkarstic aquifers, mentioning their special capacity for accumulating pollutants,their incapacity for self-depuration, and the possibilities of mass(concentrated) pollutants which have already accumulated, in order to avoidpossible contamination of drinking waters;- Avoid publicizing the discovery of a new cave before the necessaryprotective measures have been taken: this will avoid attracting simply the

The origin of this decline in bat numbers is probably mainly due to the transitionbetween an agricultural economy of traditional type, and its more modernversion, which aims at improving crop yields. This has led to extensive use ofbiocides, to reduce on one hand losses due to fungi and insects, and, on theother, to limit competition by weeds.Reduction in the quantity of insects to prey upon, and the accumulation ofpesticides in the fatty tissues of bats, which are very small predators, haveaffected both forest-dwelling and troglophile species.Accumulation of pesticides in subcutaneous fat in bats has not been studied indepth, but it is certainly known to cause great damage. It is revealed in periodsof greatest energy demand, when absorption of fats by the organism is intense,thus interfering with reproduction, suckling, and waking from hibernation.This situation of generalized crisis is aggravated by a whole series of more orless serious circumstances of disturbance, always associated with exploitationof the territory by man. For example, the continuous rejuvenation of forestalcenoses as a result of various types of reafforestation, and the tree healthmeasures and applications associated with it, represent a double-edgedweapon: on one hand, the number of holes in trees available to the more forest-dwelling species (various species of noctules and vespertilionids) has fallen; onthe other, the disturbance of subterranean caves for tourist purposes preventstroglophiles from gathering in them, both during the breeding season andduring hibernation (horseshoe-nosed bats, miniopterans, various species ofvespertilionids).Even anthropophile bats have suffered, due to recent widespread changes inbuilding criteria, but the great adaptative responses of some species and thenumber of buildings available appear to have limited the damage in somecases (see the previously mentioned case of Kuhl’s bat). In spite of this, thespecies most closely associated with old buildings constructed partially inwood (major vespertilionids, long-eared bats, some horseshoe-nosed bats)have abandoned them, partly due to their demolition and partly due to modernchemical treatments applied to the wood of the buildings still available.Most European bats are specifically protected by international treaties (e.g.,Bern and Bonn Conventions) and, as we have seen, all species are listed in theHabitat Directive.Many of them are on the Red Lists of the IUCN (International Union for theConservation of Nature), in various levels ranging between LR (low risk) to VUA2c (exposed to serious risk of extinction due to a 20% reduction predictedover the next 10 years or in the next three generations, due to a decline inoccupied surface area and/or quality of the habitat).

147146 curious (inexperienced people who may also risk accidents) and illegalcollectors of concretions, mineralizations, fauna or anything else, who wouldprobably irreparably damage the integrity of the new cave;- Place great importance on ecological education supplied during courses onspeleology and on proper training of guides taking visitors on caveexcursions;- Give “underground tourists” all useful information on proper understandingof the importance of conserving subterranean assets and karstic aquifers.

Preventing damage to the subterranean environment- Avoid too many visits, even though the most delicate caves are the oneswhich are most attractive. If possible, discourage visits by large groups,reserving information on the less delicate caves to speleology courses;- Always avoid walking about too much all over caves, even when they arevery large: a simple ribbon of red cotton on the ground means that everyonecan walk along the same track and limit damage elsewhere.

■ Strategies for choosing karstic caves and areas for faunal protection

The urgent need to safeguard at-risk karstic caves and their surroundingsposes serious problems regarding the criteria to be used to select cavesbelieved to have “priority”, out of the more than 33,000 Italian cavesregistered until now.Obviously, improvisation or other people’s opinions are of little value, butscientifically rigorous methods must be used to quantify the value of eachkarstic area, so that it may be protected properly.As regards fauna, objective methods are required to identify the “hot spots” ofbiodiversity, rarity and endemicity. In particular, it has been shown that thepoints of major interest are those with a high rate of endemicity, which onlypartially coincides with the points of major biodiversity.Modern statistical techniques and GIS programmes (Geographic InformationSystems, computerized cartography) now available to researchers areextremely powerful tools, and allow experts to select, map and attributeparameters for territorial conservation and to identify priority sites. Butalthough they are powerful and sophisticated, they cannot be based on aweak, incomplete data-bank. To apply any kind of statistics, properknowledge of the territory is required - knowledge that can only be acquired byspeleologists and biospeleologists who must collaborate closely if their aimsare to be achieved.

■ Why protect caves?

This volume has described all the most salient characteristics of the Italiankarstic environment, and given a brief overview of the zoological groups withcave-dwelling members, their origin, distribution and ecology. A detailedexamination has also been made of the problem of strategies to protect caves,legislative deficiencies, and also progress made in the last few years. Butalthough speleologists are thoroughly aware that caves and karstic areasdeserve immediate safeguarding, the reasons why these environments are ofsuch extraordinary importance must be clearly illustrated to those who havenever set foot underground.There are three main reasons why Italian caves must be protected and whysuch protection can no longer be postponed.

Intrinsic value of caves. First, caves inherently contain the same scientificand cultural values which make them the patrimony of all mankind: theirgeology, mineralogy, hydrogeology, landscape, archeology, and paleontology,and the fact that they are habitats for rare and sometimes unique species inthe worldwide faunal panorama.

Economic and social value. Caves also have economic and social value,both as regards tourism (which, as we have seen, provides considerablerevenue) and recreation, and because they supply drinking-water to very largenumbers of people.

Ethic-aesthetic value. There is a third, more profound, aspect, rooted in thevery essence of the human spirit, too often neglected in plans for protectionand exploitation of the natural patrimony.This aspect deals with feelings: we appreciate caves and the strangecreatures which inhabit them for what they are, without the need for scientificor economic support.We consider them as natural monuments which, like architectural ones, areimportant inasmuch as they exist and, for this reason, they deserve to besafeguarded.Science and economics can supply indispensable indicators for planningchoices and orienting efforts at protection, but our emotions and the sense ofmarvel which many of us feel when we enter a cave, listening to the sound ofits waters and observing the strange, white, blind creatures which live andmove in total darkness - these things alone justify the need for safeguarding.

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Study of subterranean karstic phenomena is often viewed as a specializedsubject, suitable only for universities or the last years of high school. This isnot in fact so: the approach to peculiar and richly fascinating evironments, likedolinas and caves, may intimidate some children but at the same time attractthem, and may represent a very useful instrument for training the young toknow and understand the land they live in. Interest in caves, also as regardsprehistory and archeology, immediately create that multidisciplinary link whichallows us to study an environment from many points of view and to enrich theminds of our students. It was for these reasons that here, at the end of thisvolume, we present two teaching suggestions for all those who are interestedin introducing others to the fascinations of speleology.

■ Topography of a dolina

• Aims of a topographic exercise: to stimulate knowledge of one’s ownterritory; to develop capacities for observation, analysis and abstract thought;to learn to interpret symbols used on maps.

• Field work: excursions in the field, to compare expectations and reality.

• Scholastic level: last years of junior school and middle school (children of 7-8 years upwards); the depth of teaching is directly linked with the children’slevel of preparation (see box on dolinas, page 24).

STUDY PHASES

1. Study topographic maps of karstic areas, preferably Technical Maps, scale1:5,000 (or 1:25,000) and identify the most interesting areas where bothsuperficial forms of karstism (dolina, polje, uvala, etc.) are widespread andmapped, and underground ones (entrances to caves); names of villages andtowns are particularly useful for a multidisciplinary approach to mapping.2. Once a particularly significant map area has been identified, supplystudents with scale copies; a few other instruments (rulers, goniometers,squared graph paper, pencils) are sufficient for the exercise.3. Organize individual work by each student on the toponyms referring tomorphology: various symbols which represent water networks, reliefs,

Suggestions for teachingMARGHERITA SOLARI

Grotta di Punta Galera (Palinuro, Campania)

151150 escarpments, vegetation, villages and towns, communications networks,paths, etc..4. Observe and compare various cartographic representations of superficialkarstic forms: for example, dolinas may be shown as concentric level curves,wedges placed in a circle, or simply altimetric references.5. Exercises: identify dolina perimeters, represented by concentric contourlines and definition of gradients; deduce unspecified altitudes with the help oflevel curves; similarly, deduce altitudes of valley bottoms, if unspecified.6. Identify the maximum diameter of a dolina, i.e., the segment joining the twomost distant points of the perimeter, and the minimum diameter perpendicularto it; calculate the direction of the diameter with respect to compass north,using a goniometer.7. Draw the profile of a dolina: draw the maximum diameter of the dolina and,using graph paper, mark on it the distances of the points intersecting withcontour lines; mark main contours (every 25 m) for large dolinas andsecondary curves (every 5 m) for smaller ones (or even 1-m contour lines) forminor forms.8. Sketch of the dolina: make a drawing and complete it with symbols forvegetation; intepret the key and extrapolate information on the “landscape”;prepare proper symbols and represent vegetation indicating species (trees,vineyards, open-field crops, etc.) and density; note possible diversities in thevegetal cover on the four flanks of the dolina.9. Additional exercise: analyse and compare the representation of a smallrelief; draw its profile; examine problems of cartographic representation.

FURTHER WORK

10. Last field excursion: verification of expectations, comparison between reallandscape and that deduced from topographic map; exercises in orientation:identification of cardinal points and maximum diameter of landform; ifpossible, descent into the dolina in groups, identifying points which served fordrawing its profile (intersection of diameter with contour lines) andreconstruction of contour lines shown on topographic map by means ofribbon or cord; guided observations on natural environment, with particularemphasis on vegetation on flanks and bottom and surrounding territory.

■ A tour of a cave

• Aims of visiting a natural cave: to go further into the themes covered byteaching; to develop capacities for exploration, orientation and observation,

Plan and cross-section of a large dolina in Trieste Karst, showing changes in vegetation according tomicroclimate: from left to right (i.e., from south to north), cool-loving, water-loving and heat-lovingwoodland

153152 essential for fixing acquired knowledge firmly in the memory; to completetaught material with direct experience and to strengthen knowledge by clearlydefining experienced situations; to combine amusement and study (“businesswith pleasure”) in a multidisciplinary research. Before beginning to organizethis type of activity, check if any students suffer (or may suffer) fromclaustrophobia: in this case, only try to create motivations for overcomingslight fears, but do not force any student.

• Level: junior school, middle school (from 7-8 years of age), althoughinitiatives of this type, suitably modified and prepared, are possible with evenyounger children.According to aims and students’ capacities, various levels of excursions maybe organized, according to their past experience of guided tours:- visit to a tourist cave (wearing gym shoes or trainers, and windjackets),focusing on general observations of the underground environment, which maybe anthropized and degraded in this case;- visit to a natural cave presenting slight difficulties;- full-scale visit, with special equipment.For the choice of which cave to explore and the behaviour to adopt, seesection on the ethics of speleology.

• Essential help: preferably, contact a speleological association capable ofsupplying both logistic support and, if one of its members is specialized inteaching speleology, ask for specific useful information.Come to an arrangement with the speleologists on which cave to visit,ascertain any difficulties in its itinerary, according to the needs of the group,and plan for an adequate number of support staff (in general, oneaccompanying person for every 3-4 students is reommended for a visit ofmedium difficulty).

• Equipment: for a cave of medium difficulty, all visitors must be properlyequipped with: helmets with lamps, rubber boots or stout shoes, woollen orinsulated clothing. If the environment is muddy, a change of clothes can be leftat the entrance to the cave.

PRELIMINARY ACTIVITIES

1. General theoretical introduction to the phenomenon of karstism and theimportance of the subterranean environment, to create expectations andstimulate curiosity and interest.2. Choice of cave, suited to students’ capacities.3. Pre-visit check-up by organizer and support staff.4. Bibliographic and cartographic study, in class, to identify the cave.

155154 5. Class debate to organize material necessary for the excursion, choice ofroute to reach the cave, setting-up of working groups assigned with varioustasks (general preparation, survey charts, preparation of notes on caveitinerary, etc.). Information on correct behaviour to be adopted in caves: followstaff indications, do not leave waste of any kind, do not touch (OR BREAKOFF) concretions, and so on.6. Preparation of excursion equipment by groups.

EXCURSION

7. Check route: the group with this task guides everyone to the entrance to thecave.8. Notes on the itinerary, for later preparation of reports: type of path, altitude,orientation, vegetation present, time taken to reach cave, geomorphology, anyforms of superficial karstism:9. When the entrance to the cave has been found, take notes onmorphological, physical and biotic data, e.g.:- shape and orientation of entrance, possible presence of water indicating thatthe cave is active (cave or spring);- variations in temperature, light, vegetation, presence of material (branches orleaves) from the exterior;- possible signs of degradation: presence of waste, carbide residues fromlamps, state of conservation of concretions, etc..10. After entering the cave, note variations in amount of light perceived, andcheck temperature variations with a thermometer.11. Take notes on the itinerary, observing significant elements helpingorientation: forks, watercourses, etc.. Try out initial preparation of a planimetricrelief with compass and cord marked off in metres, at least along one stretchof the itinerary.12. Guided observations on:- environment: possible presence of draughts indicating secondary entrancesor large open spaces inside the cave; sound made by flowing water, drippingwater, or streams; presence of humidity (small clouds of water vapour afterbreathing out);- vegetation: observations on seriation (point out interesting parallels withevolution); abundant vegetation at the entrance, gradually becoming less withless light (e.g., Parietaria, Lunaria annua, etc.);- cave morphology: the cross-section is mainly determined by water flowing inconduits and by tectonic factors in galleries; observation of the ground,possible fractures, faults, landslides and collapses; various types of

concretions (stalactites and stalagmites, solution pans, columns, draperies,etc.) from viewpoints of both morphological and mineralogical composition(generally, but not always, calcitic); possible relationships betweencharacteristic forms of concretions and tectonic structures; in the first fewyards, note presence of rocks and landslides; ceiling vaults of galleries andhalls (visitors always tend to look at the ground and walls, forgetting that theceiling is in fact the first part of any cave to form);- fauna: presence of guano, indicating bat colonies; animals living on the walls:orthopterans, dipterans, lepidopterans, spiders; search for possibletroglobites and observation of their typical colours.13. Observation of one’s own movements: how does one have to move in acave? (crawling, climbing, etc.)14. Observation of the simplicity and fragility of the subterranean ecosystem,importance of respecting the environment, and on one’s capacity toappreciate it without removing anything from it or damaging it; recovery of allwaste.15. Moment for relax, during lunch interval: perhaps (if possible), allow alllights to be turned off and sit in absolute silence, trying to experience thenatural environment without adding one’s own outside impressions to it;sharing impressions and feelings.16. Consultation and interpretation of notes taken during the visit, in order tofind the way back to the entrance.17. Sharing of impressions once outside the cave again, particularly asregards sensations of light and smells in the outside environment.

CONCLUSIVE ACTIVITIES IN CLASS

18. “Reprocessing” in class: preparation of maps, scientific reports on the cave,preparation of essays on personal impressions of the new experience, etc..

CONTINUATION OF WORK

19. The set of observations made during a visit to a natural cave takes onfurther significance when re-interpreted in comparison with a superficialenvironment.The diversities between the two focus on the particular fragility of thesubterranean environment and on the need to protect it. To complete theexperience and go deeper into these topics, the excursion may later becombined with a visit to a museum with a section on karstism, or asubterranean biology laboratory (found throughout Italy: Trieste, VittorioVeneto, Verona, Brindisi, Rome, and many others).

157

VARIOUS AUTHORS, 1978 - Manuale di Speleologia. Longanesi, Milan.An ample, complete treatise prepared by the Italian Speleological Society, devoted to allaspects of speleology. Although somewhat out-of-date as regards some specific parts(e.g., description of karstic areas), it is still extremely valid for general topics.

VARIOUS AUTHORS, 1982 - Biogeografia delle caverne italiane. Lavori della Società Italianadi Biogeografia, new series, vol. 7 (1978).The most complete collection of original scientific articles on the cave-dwelling fauna ofsome Italian regions, with particular attention to biogeographic aspects.

VARIOUS AUTHORS, 1989 - Problemi di inquinamento e salvaguardia delle aree carsiche(ed. M. Chiesi), SSI, CAI, Nuova Editrice Apulia.A volume entirely devoted to describing the mechanisms of how aquifers and both thesuperficial and deep karstic landscape are compromised by pollution, with informationon ways and means of providing both regional and local protection.

BADINO G., 1998 - Fisica del clima sotterraneo. Memorie dell’Istituto Italiano diSpeleologia, 2nd series, 7, Bologna.An ample work giving a detailed description of the physics of climates underground: themost complete work ever published on cave climatology.

BOTOSANEANU L. (ed.), 1986 - Stygofauna Mundi. E.J. Brill/Dr. W. Backhuys, Leiden.An exhaustive and rigorous treatment of all taxa found in subterranean waters in theworld. A short diagnosis and a list of all species known until 1985, with their worldwidedistribution, is given for each animal group.

CASTIGLIONI G.B., 1979 - Geomorfologia. Utet, Turin.In this valuable work, one lengthy chapter (by U. Sauro) is devoted to karsticmorphology. The illustrations are excellent, and were also a source of inspiration forsome drawings in the present volume.

CHIESI M., FERRINI G. & BADINO G., 1999 - L’impatto dell’uomo sull’ambiente di grotta.Quaderni Didattici, no. 5, Società Speleologica Italiana, Bologna.A teaching manual, for use in introductory courses to speleology, published by theItalian Speleological Society, summarizing the main environmental problems caused byhumans entering caves.

COLLIGNON B., 1992 - Il manuale di Speleologia. Zanichelli, Bologna.A work covering the various aspects of scientific speleology, with particular attention tokarstism, although biospeleology and other topics connected with study of naturalcaves are also discussed. Richly illustrated and completed by a vast bibliography, theItalian translation of this volume contains many explicit references to Italy.

Select bibliography

Subterranean glacier in a cave in Mount Canin (Julian Alps, Friuli)

158 FILECCIA A., 1996 - Speleologia subacquea. Vallardi, Milan.A manual devoted to this particular aspect of speleological activity, complete withdescriptions of the areas containing the most interesting grottoes in Italy and the mostinteresting cave siphons.

FORNASARI L., VIOLANI C. & ZAVA B., 1997 - I Chirotteri italiani. Epos, Palermo.An attractive volume, useful for teaching those who are interested in the world of bats,without going into excessive detail.

JUBERTHIE C., DECU V. (eds.), 1994 - Encyclopaedia Biospeleologica. Société deBiospéléologie, Moulis (C.N.R.S.), Academia Roumaine, vol. I, Bucharest.The first volume in an extensive series devoted to biospeleology. Vol. I deals withnumerous invertebrate taxa on a worldwide scale, and examines troglobitic fauna inAmerica and Europe. One chapter is devoted to the fauna of Italian caves.

JUBERTHIE C., DECU V. (eds.) 1998 - Encyclopaedia Biospeleologica. Société deBiospéléologie, Moulis (C.N.R.S.), Academia Roumaine, vol. II, Bucharest.In this second volume, ample space is devoted not only to some minor taxonomicgroups, but also to cave-dwelling insects (particularly beetles), vertebrates and flora.Vol. 3 is in preparation.

LANZA B., 1983 - Anfibi, rettili. Guide per il riconoscimento delle specie animali delleacque interne italiane. 27. CNR, Rome.This work, a mix between scientific literature and clearly presented, simpler information,contains good descriptions of olm, cave salamanders, and other species of amphibiansfound in Italian caves.

MINELLI A., RUFFO S., LA POSTA S., 1993 - Checklist delle specie della fauna italiana(Checklist of species of Italian fauna). Calderini, Bologna.Lists all known species of Italian fauna, allowing correct, standardized nomenclature tobe used. Series composed of 110 sections bound in 24 booklets.

TOSCHI A. & LANZA B., 1959 - Fauna d’Italia. IV. Mammalia (Generalità, Insectivora,Chiroptera). Calderini, Bologna.In this volume on the fauna of Italy, a very large part (pp. 187-473) is devoted to Italianbats, with explicit references to cave-dwelling species. Do not be deceived by the dateof publication: this work is still today an excellent scientific reference work for all thosewho want to learn about bats.

VIANELLI L., 2000 - I fiumi della notte. Bollati Boringhieri, Turin.A valuable work, also from the photographic point of view, examining the variouscharacteristics of underground waters and their essential role in the development ofkarstism. Some of the most important subterranean rivers and karstic aquifers in Italyare described.

Much more information may be obtained from the numerous journals devoted tospeleology published in Italy. Examples are the publications of the Società SpeleologicaItaliana (“Speleologia” and “Le Grotte d’Italia”), the journals “Atti e Memorie dellaCommissione Grotte E. Boegan”, “Mondo Sotterraneo” and “International Journal ofSpeleology” (sections on biology and earth sciences), and many bulletins published bylocal speleological groups.

Glossary

> Allopatria: condition of complete separation of the areas in which one or more specieslive; allopatric speciation is the differentiation of new species due to the isolation ofsome populations of a common progenitor.> Benthic organisms: organisms living in water, always in very close relationship with thebottom substrate.> Dissolution: a chemical process giving rise to the solubilization of limestone by waterenriched in carbon dioxide.> Endemic: an adjective indicating a species exclusive to a certain relatively small geo-graphic territory; caves are among the environments richest in endemic species.> Biotic interactions: general interactions among all kinds of living organisms (e.g., com-petition, predation, etc.).> Phreatic zone: the permanently water-saturated area in a karstic massif, in which hori-zontal flow of subterranean waters prevails.> Gour: French term, used to indicate a pool.> MMS: acronym used by French authors to indicate the soil layer mainly resulting fromdismantling of underlying rocks (Milieu Souterrain Superficiel); it is considered to be anintermediate habitat when a cave environment is colonized.> Plankton: organisms living in suspension in waters.> Speleothemes: various types of deposits in a cave (concretions, etc.).> Speciation: evolutionary process leading to the differentiation of new species.> Stygobites: organisms exclusive to subterranean waters (karstic or otherwise), in whi-ch they spend their entire life-cycle; they always present special adaptations to under-ground life.> Stygophiles: organisms found in subterranean waters (karstic or otherwise), wherethey may reproduce, but they are not exclusive to these environments, since they maycomplete their life-cycle partly in superficial waters.> Stygoxenes: organisms living in superficial waters but also occasionally found in sub-terranean waters, as a result of passive transport or because they seek special condi-tions in some phases of their life-cycle.> Troglobites: organisms exclusive to caves, where they spend their entire life-cycle;they always present special adaptations to underground life.> Troglophiles: organisms regularly found in caves, where they may reproduce, but theyare not exclusive to these environments; in some cases (eutroglophile organisms) theymay present partial adaptations to underground life.> Trogloxenes: surface-living organisms occasionally or accidentally found in subterra-nean environments; in some cases, they frequent caves during particular phases of theirlife-cycle or in some seasons (regular trogloxenes).> Vadose zone: that part of a karstic massif in which vertical flow of waters prevails(unsaturated zone), with dripping and percolation of various intensity.

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Special thanks go to Paolo Forti for his usefuladvice and critical reading of the part devotedto karstism.

Many speleologists, to whom go our thanks,collaborated on the preparation of chaptersfocusing on various regions (Marco Bani, EzioBurri, Gian Domenico Cella, Paolo Giuliani,Pino Guidi, Carmine Marotta, Paolo Mietto,Mauro Muccedda, Paolo Zambotto).

The authors would like to thank all thespecialists who provided them with informationand who applied their specific competences torevising and integrating various sections in thisvolume:Maria Manuela Giovannelli, Marco Bodon andWilly de Mattia (molluscs);Beatrice Sambugar (oligochaetes);Diana Galassi (copepods);Roberto Argano (isopods);Sandro Ruffo (malacostracans);Fulvio Gasparo (palpigrads, spiders);Claudio Chemini (harvestmen);Giulio Gardini (pseudoscorpions);Marzio Zapparoli (chilopods);Alessandro Minelli (chilopods, diplopods);Pietro Paolo Fanciulli (spring-tails);Paolo Fontana (orthopterans);Carlo Morandini (lepidopterans);Achille Casale and Augusto Vigna Taglianti(carabid beetles);Stefano Zoia (leptodirine beetles);Adriano Zanetti (rove beetles);Giorgio Sabella and Roberto Poggi(pselaphid beetles);Fabio Penati (histerid beetles);Giuseppe Osella (curculionid beetles).Achille Casale also kindly revised the sectionon the biogeography of cave-dwelling fauna,and Valerio Sbordoni made many valuablesugggestions.Many thanks are to to Grahm S. Proudlove foruseful suggestions on English terminology.The authors are exclusively responsible fortextual contents, and for any errors andomissions.

This volume was published with funds from theItalian Ministry of the Environment andTerritorial Protection.

Printed byArti Grafiche Friulane, Tavagnacco, Udine, Italy.