ACTA CARSOLOGICA ISSN 0583-6050 © ZNANSTVENORAZISKOVALNI CENTER SAZU Uredniški odbor / Editorial Board Franco Cucchi, University of Trieste, Italy Jože Čar, University of Ljubljana, Slovenia Franci Gabrovšek, Karst Research Institute ZRC SAZU, Slovenia Ivan Gams, University of Ljubljana, Slovenia Matija Gogala, Slovenian Academy of Sciences and Arts, Slovenia Andrej Kranjc, Karst Research Institute ZRC SAZU, Slovenia Marcel Lalkovič, Te Slovak Muesum of Nature Protection and Speleology Jean Nicod, Emeritus Professor, Geographical Institute, Aix en Provence, France Mario Pleničar, University of Ljubljana, Slovenia Trevor R. Shaw, Karst Research Institute ZRC SAZU, Slovenia Tadej Slabe, Karst Research Institute ZRC SAZU, Slovenia Glavni in odgovorni urednik / Editor-in-Chief Andrej Kranjc Pomočnik urednika / Co-Editor Franci Gabrovšek Znanstveni svet / Advisory Board Ahmad Afrasibian, Philippe Audra, Ilona Bárány – Kevei, Pavel Bosák, Arrigo A. Cigna, David Drew, Wolfgang Dreybrodt, Derek Ford, Helen Goldie, Laszlo Kiraly, Alexander Klimchouk, Stein-Erik Lauritzen, Bogdan Onac, Armstrong Osborne, Arthur Palmer, Ugo Sauro, Boris Sket, Kazuko Urushibara-Yoshino. Naslov uredništva / Editor’s address: Inštitut za raziskovanje krasa ZRC SAZU - Karst Research Institute ZRC SAZU SI - 6230 Postojna, Titov trg 2, Slovenija Fax: +386 (0)5 700 19 99, e-mail: kranjc@zrc-sazu.si Spletni naslov / Web address: http://carsologica.zrc-sazu.si Distribucija in prodaja / Ordering address: Založba ZRC/ZRC Publishing Novi trg 2, P.O.Box 306, SI-1001 Ljubljana, Slovenia Fax: +386 (0)1 425 77 94, e-mail: zalozba@zrc-sazu.si, http://zalozba.zrc-sazu.si Sprejeto na seji uredniškega odbora 25. januarja 2007. Cover photo: Cover montage by Will Pearce. Images courtesy of Horton H. Hobbs III, John Mylroie, Arthur N. Palmer, and Ira D. Sasowsky.. Cena / Price Posamezni izvod / Single Issue Individual / Posameznik: 15 € Institutional / Institucija: 25 € Letna naročnina / Annual Subscription Individual / Posameznik: 25 € Institutional / Institucija: 40 € ACTA CARSOLOGICA 36/1 2007 SLOVENSKA AKADEMIJA ZNANOSTI IN UMETNOSTI ACADEMIA SCIENTIARUM ET ARTIUM SLOVENICA Razred za naravoslovne vede – Classis IV: Historia naturalis ZNANSTVENORAZISKOVALNI CENTER SAZU Inštitut za raziskovanje krasa – Institutum carsologicum LJUBLJANA 2007 LET / yEARS Inštitut za raziskovanje Krasa ZRC SAZU Karst Research Institute at ZRC SAZU ACTA CARSOLOGICA je vključena v / is included into: Current Geographical Contents / Ulrich's Periodicals Directory / COS GeoRef / BIOSIS Zoological Record. ACTA CARSOLOGICA izhaja s fnančno pomočjo / is published with the fnancial support of: Agencije za raziskovalno dejavnost RS / Slovenian Research Agency, Slovenske nacionalne komisije za UNESCO / Slovenian National Commission for UNESCO in / and Postojnska jama turizem d.d. CONTENTS VSEBINA PAPERS ČLANKI Franci GAbROvšEK 7 ON DENUDATION RATES IN KARST O hItROStI dENUdACIjE NA KRASU Arthur N. PALmER 15 VARIATION IN RATES OF KARST PROCESSES SPREmENLjIvOSt hItROStI KRAšKIh PROCESOv Wolfgang dREybROdt & douchko ROmANOv 25 TIME SCALES IN THE EVOLUTION OF SOLUTION POROSITy IN POROUS COASTAL CARBONATE AqUIFERS By MIxING CORROSION IN THE SALTwATER-FRESHwATER TRANSITION ZONE. ČASOvNO mERILO RAzvOjA POROzNOStI zARAdI KOROzIjE mEšANICE v mEjNEm ObmOČjU SLAdKOvOdNIh LEČ v mEdzRNSKO POROzNEm KARbONAtNEm ObALNEm vOdONOSNIKU Andrej mIhEvC 35 THE AGE OF KARST RELIEF IN wEST SLOVENIA StAROSt KRAšKEGA RELIEFA v zAhOdNI SLOvENIjI William b. WhItE 45 EVOLUTION AND AGE RELATIONS OF KARST LANDSCAPES RAzvOj IN StAROStNI OdNOSI KRAšKIh POKRAjIN Philippe AUdRA, Alfredo bINI, Franci GAbROvšEK, Philipp häUSELmANN, Fabien hObLéA, Pierre-yves jEANNIN, jurij KUNAvER, michel mONbARON, France šUštERšIČ, Paola tOGNINI, hubert tRImmEL & Andres WILdbERGER 53 CAVE AND KARST EVOLUTION IN THE ALPS AND THEIR RELATION TO PALEOCLIMATE AND PALEOTOPOGRAPHy RAzvOj jAm IN KRASA v ALPAh v LUČI PALEOKLImE IN PALEOtOPOGRAFIjE Leonardo LAtELLA & Ugo SAURO 69 ASPECTS OF THE EVOLUTION OF AN IMPORTANT GEO-ECOSySTEM IN THE LESSINIAN MOUNTAIN (VENETIAN PREALPS, ITALy) POGLEdI NA RAzvOj POmEmbNEGA GEO-EKOSIStEmA v GORAh LESSINI (bENEšKE PREdALPE, ItALIjA) Oana teodora mOLdOvAN & Géza RAjKA 77 HISTORICAL BIOGEOGRAPHy OF SUBTERRANEAN BEETLES – “PLATO’S CAVE” OR SCIENTIFIC EVIDENCE? zGOdOvINSKA bIOGEOGRAFIjA POdzEmELjSKIh hROšČEv – »PLAtONOvA jAmA« ALI zNANStvENI dOKAz? david C. CULvER & tanja PIPAN 87 wHAT DOES THE DISTRIBUTION OF STyGOBIOTIC COPEPODA (CRUSTACEA) TELL US ABOUT THEIR AGE? KAj NAm POvE RAzšIRjENOSt StIGObIONtSKIh CEPONOŽNIh RAKOv (CRUStACEA: COPEPOdA) O NjIhOvI StAROStI? Philipp häUSELmANN 93 HOw TO DATE NOTHING wITH COSMOGENIC NUCLIDES KAKO dAtIRAtI PRAzNINE S KOzmOGENImI NUKLIdI bojan OtONIČAR 101 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION (KRAS, MATARSKO PODOLJE AND ISTRIA; Sw SLOVENIA AND Nw CROATIA) zAKRASELA PERIFERNA IzbOKLINA POvEzANA z RAzvOjEm zGORNjEKREdNO-PALEOGENSKEGA PREdGORSKEGA bAzENA; KRAS, mAtARSKO POdOLjE IN IStRA (jz SLOvENIjA IN Sz hRvAšKA) Robert G. LOUCKS 121 A REVIEw OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS AND ASSOCIATED SUPRASTRATAL DEFORMATION mEdSEbOjNO zdRUŽENI PORUšENI PALEOKRAšKI jAmSKI SIStEmI IN dEFORmACIjE NAd NjImI LEŽEČIh PLAStI – PREGLEd R. Armstrong L. OSbORNE 133 THE wORLD’S OLDEST CAVES: - HOw DID THEy SURVIVE AND wHAT CAN THEy TELL US? NAjStAREjšE jAmE NA SvEtU: KAKO SO SE OhRANILE IN KAj NAm LAhKO POvEdO? Ira d. SASOWSKy 143 CLASTIC SEDIMENTS IN CAVES – IMPERFECT RECORDERS OF PROCESSES IN KARST KLAStIČNI SEdImENtI v jAmAh – NEPOPOLNI zAPIS KRAšKIh PROCESOv Ognjen bONACCI 151 ANALySIS OF LONG-TERM (1878-2004) MEAN ANNUAL DISCHARGES OF THE KARST SPRING FONTAINE DE VAUCLUSE (FRANCE) ANALIzA dOLGOČASOvNEGA (1878-2004) POvPREČNEGA LEtNEGA PREtOKA KRAšKEGA IzvIRA FONtAINE dE vAUCLUSE (FRANCIjA) Fred G. LUISzER 157 TIMING OF PASSAGE DEVELOPMENT AND SEDIMENTATION AT CAVE OF THE wINDS, MANITOU SPRINGS, COLORADO, USA ČASOvNO USKLAjEvANjE RAzvOjA jAmSKIh PROStOROv IN SEdImENtACIjA v jAmI CAvE OF thE WINdS, mANItOU SPRINGS, COLORAdO, zdA megan L. PORtER, Katharina dIttmAR & marcos PéREz-LOSAdA 173 HOw LONG DOES EVOLUTION OF THE TROGLOMORPHIC FORM TAKE? ESTIMATING DIVERGENCE TIMES IN ASTyANAx MExICANUS KAKO dOLGO tRAjA EvOLUCIjA tROGLOmORFNIh ObLIK? OCENjEvANjE dIvERGENČNIh ČASOv PRI AStyANAX mEXICANUS Peter tRONtELj, špela GORIČKI, Slavko POLAK, Rudi vEROvNIK, valerija zAKšEK & boris SKEt 183 AGE ESTIMATES FOR SOME SUBTERRANEAN TAxA AND LINEAGES IN THE DINARIC KARST OCENE StAROStI zA NEKAtERE POdzEmELjSKE tAKSONE IN ŽIvALSKE LINIjE NA dINARSKEm KRASU Eleonora tRAjANO 191 THE CHALLENGE OF ESTIMATING THE AGE OF SUBTERRANEAN LINEAGES: ExAMPLES FROM BRAZIL IzzIv OCENjEvANjA StAROStI POdzEmELjSKIh ŽIvALSKIh LINIj: PRImERI Iz bRAzILIjE Andreas WESSEL, Petra ERbE & hannelore hOCh 199 PATTERN AND PROCESS: EVOLUTION OF TROGLOMORPHy IN THE CAVE-PLANTHOPPERS OF AUSTRALIA AND HAwAI’I – PRELIMINARy OBSERVATIONS (INSECTA: HEMIPTERA: FULGOROMORPHA: CIxIIDAE) vzOREC IN PROCES: EvOLUCIjA tROGLOmORFNOStI PRI jAmSKIh mREŽEKRILNIh šKRŽAtKIh Iz AvStRALIjE IN hAvAjEv – PRELImINARNE UGOtOvItvE (INSECtA: hEmIPtERA: FULGOROmORPhA: CIXIIdAE) 207 ABSTRACTS FOREwORD Time occupies a curious place in science. In most of science, including karst science, “how” questions predominate. How are caves formed? How are caves eroded? How do animals survive in caves? How do animals come to lose their eyes and pigment in caves? But “when” questions have probably been asked from the very beginning of karst studies. Some of the fascination with time comes from the parent disciplines of biology and geology. Te diference between “catastrophism” and “uniformitarian-ism” in geology is also a question of diferences in time— rapid catastrophes versus slow, small changes. In biology, particularly in the late 19th and early 20th centuries, great controversies raged between the slow pace of evolution envisioned by Darwinians and the fast pace of evolution envisioned by neo-Lamarckians. questions of time have been especially fascinating in the karst sciences, probably because our senses tell us that caves (and cave animals) are very ancient. Caves are afer all the dwelling place in mythology of ancient creatures—dragons especially. Of course our senses (and our mythology) can be deceiving, and perhaps caves and cave animals are not as old as they seem to be. wide difer-ences of opinion have persisted about the ages of both caves and cave animals—estimates at present that range between less than a million years to up to 100 million years! Te time is ripe to examine time in karst. Te set of papers and abstracts in this volume is the result of a meeting, time in Karst, of karst scientists in Postojna, Slovenia, in March 2007. Jointly sponsored by the Karst Research Institute ZRCSAZU of Slovenia and the Karst waters Institute of the U.S.A., an international group of scientists came together to learn about and discuss time processes in karst from six perspectives: • Te age of karst landscapes, including caves and other karst landforms • Te biogeographic history of cave animals, especially as it relates to the present and past distributions of cave animals • methods of determining the age of caves, especially geophysical ones • Paleokarst and what it can tell us about age • Te sediment record • Te age of lineages of cave animals, especially using molecular clock techniques Both the Karst Research Institue ZRCSAZU and the Karst waters Institute have a history of promoting both international and interdisciplinary cooperation, and they are pleased to form a partnership in this international, interdisciplinary endeavour. Te participation of many researchers in early stages of their careers was made possible by project SMARTKARST of the Karst Research Institute, ZRC SAZU, funded by the Marie Curie programme, sponsored by the European Commission. Many members of the Karst Research Institute ZRCSAZU and the Karst waters Institute worked hard to make this meeting possible, including Drs. Daniel Fong, Franci Gabrovšek, Andrej Kranjc, Tanja Pipan, and Ira Sasowsky. tadej Slabe david C. Culver TIME in KARST, POSTOJNA 2007, 5 COBISS: 1.01 ON DENUDATION RATES IN KARST O HITROSTI DENUDACIJE NA KRASU Franci GABROVŠEK1 Abstract UDC 551.331.24:551.44 Franci Gabrovšek: On denudation rates in Karst Paper presents a simple mathematical model, which enables study of denudation rates in karst. A vertical fow of water which is uniformly infltrated at the surface is assumed. Denudation rate is calculated from the time needed to remove certain thickness of rock. Tis is done concretely on a limestone block dissected by a vertical array of fractures. It is shown that denudation rate increases with the thickness of removed layer and approaches an upper limit which is defned by the maximum denudation equations, which are based on assumption that all dissolution potential is projected into a surface lowering. Keywords: karst, denudation rate, limestone dissolution, mathematical model. Izvleček UDK 551.331.24:551.44 Franci Gabrovšek: O hitrosti denudacije na Krasu V prispevku predstavim enostaven matematični model s katerim raziskujem dinamiko zniževanja kraškega površja. Predpostavim enakomerno napajanje s površja in vertikalno pronicanje vode. Denudacijsko stopnjo izračunam iz časa, ki je potreben za odstranitev določene debeline kamninskega sloja. Konkretno to naredim na primeru apnenca v katerem se voda pretaka v sistemu vertikalnih razpok. Hitrost denudacije narašča z debelino odstranjene plasti in doseže zgornjo mejo, ki je določena z enačbami, ki temeljijo na predpostavki, da se celoten korozivni potencial vode manifestira v zniževanju površja. Ključne besede: kras, denudacijska stopnja, raztapljanje apnenca, matematični model. INTRODUCTION Uniform lowering or surface denudation is a dominant karstifcation process (Dreybrodt, 1988; Ford & williams, 1989; white, 1988). Te denudation rate is defned as the rate (LT-1) of lowering of a karst surface due to the dissolution of bedrock. A common approach used to estimate the denudation rate is based on the presumed equilibrium concentration (or hardness) and the amount of water which infltrates into the subsurface. It is summarized in the famous Corbel’s equation (Corbel, 1959): Dc (m/Ma) = (P-E)H 1000-p / Te infltrated water in mm/y is the diference between precipitation P and evapotranspiration E. H is the equilib- rium concentration (Hardness) in mg/L of dissolved rock, ? is the density of limestone in g/cm3, f denotes the portion of soluble mineral in the rock, which will be 1 in this paper. Te factor 1000 corrects for the mixture of units used in the equation. Tere are more general equations of this kind like that of white (1984, this issue). For a Limestone terrain in a temperate climate all these equations give denudation rate of the order of several tens of meters per million years. Similar results are obtained from fow and concentration measurements in rivers which drain a known catchment area. From the measured data the total rock volume removed from the area in a given time period can be calculated. Dividing the removed volume by the surface of the area and the time interval gives the denudation rate. 1 Karst Research Institute ZRC SAZU, Postojna, Slovenia, e-mail: gabrovsek@zrc-sazu.si Received/Prejeto: 01.02.2007 TIME in KARST, POSTOJNA 2007, 7–13 1 FRANCI GABROVŠEK Eq.1 implies that all dissolution capacity of water is used in the rock column, i.e. the solution at the exit of rock block is close to saturation. Among the many assumptions behind such estimations of the denudation rate I will address two of which at least one must be valid: 1. Most of the dissolution occurs close to the surface, i.e. within epikarst. 2. In the long term, the dissolution at depth is integrated into a surface denudation. It is the intention of this paper to theoretically validate “maximum denudation” approach. SURFACE LOwERING AND THE VOLUME OF DISSOLVED ROCK Dissolution of any rock is not instantaneous, but proceeds at some fnite rates. In conditions of difuse infltration through the karst surface and prevailing vertical fow, the concentration of dissolved rock in the infltrating water will normally increase with the depth as schematically shown by color intensity in Fig.1. Fig. 1. Section of a terrain with a uniform surface infltration of aggressive solution and prevailing vertical fow. Color intensity denotes that the concentration of dissolved rock increases with depth. Fig. 2 presents point at some depth z below the surface. Te volume ?V of rock dissolved per unit surface area S in time ?t between the surface and the point is given by AV/S = c(z)-q-&t /p 2 where c(z) is the concentration of dissolved rock [M/L3] at the depth z, q is the infltration rate at the surface [L3/ (L2T)] and ? is the density of the rock [M/L3]. Due to the surface lowering, the depth of the point is decreasing according to z(t) = z0 - D·t, where z0 is the Fig. 2: Idealized profle through the rock column at time t = 0 (lef) and t > 0 (right). Te depth of the point which is at z0 decreases in time due to the surface lowering. depth at t = 0 and D is the denudation rate (Fig. 2). Te volume of dissolved rock per surface area in time T above the point is then given by: T T AV/S (T) = ^ fc(z(t)) dt = - fc(z0 - D • t) dt 3 Po Po Introducing a new variable z=z0 - D·t into the right hand integral in Eq. 3 gives: AV W=w / cc-y^{\-s-1^1) 17 we will demonstrate the results on a characteristic data for a moderate climate with I=1000 mm/y and relatively bare karst area with ceq= 1mmol/l or H = 100 mg/L. For ? = 2.5 g/cm3. DC for this case is 40 m/Ma. we assume that the rain infltrates into a parallel set of fractures with spacing d = 1 m. Fig. 5a shows z0(TD) for four different saturation lengths arising from different infiltration intensities. yearly infiltration is 1000 mm/y for all curves. Therefore, the time period of dissolution is inversely proportional to the infiltration intensity. Dashed line shows the uniform lowering by D . we wee that all lines become practically parallel to maximum denudation line for zg > 2A2.The actual denudation rate becomes “maximal” when the removed thickness is larger than 2?2. This is about the depth where the concentration reaches 90% of saturation. The slope of the dotted lines presents the averaged denudation rates for curve with ?2 = 70 m. Fig. 5b shows the averaged rate for the same scenarios as Fig. 5a. Red dashed red curve clearly shows the fast approach of the actual rate to maximal for ?2 = 52.5 m. Another interesting conclusion can be made from Fig. 5a. Diferent saturation lengths ? can also arise from diferent fracture spacing (see Eq. 15 for qf) . If we imagine a region with high fracture density within a region of low fracture density, the frst will initially be denuded faster, but latter on both actual rates will become the same. Terefore the diference made at the onset will stay projected in the surface. Tis is shown by the double arrow between lines 3 and 4. TIME in KARST – 2007 11 FRANCI GABROVŠEK Fig. 5: a) Te time dependence of removed thickness for several infltration intensities. I=1000 mm/y, h = 100 mg/L, ? = 2.5 g/cm3, d = 100 cm, N = 2. dashed line show the “maximum denudation” rate which is 40 m/ma. dotted lines present the time averaged denudation rates (Eq.16). double arrow demonstrates the diference between the denuded thicknesses which is kept in time due to the initial rate diferences. b) dependence of average denudation rates on the removed thickness for the same scenarios as in Fig. 5a. dashed line presents the actual surface lowering for ?2 = 52.5 m. 12 TIME in KARST – 2007 ON DENUDATION RATES IN KARST CONCLUSION Denudation rate in a block with initially uniform porosity increases as the denudation proceeds and becomes maximum denudation (Eq.1), when the thickness of removed layer is about twice the typical saturation length. Initial diferences arising from diferent saturation lengths remain imprinted in the surface. If a soluble layer has a fnite thickness, the average denudation rate increases with the thickness, i.e. denudation is more efective on thick rock layers. Te presented results are based on many assumptions which might not be valid. Nevertheless, it gives some theoretical validation of maximum denudation formulae and suggest some mechanisms that can cause irregularities in karst surface. REFERENCES Appelo, C. A. J. & D. Postma, 1993: Geochemistry, ground-water and pollution. A.A. Balkema, xvi, 536 pp, Rotterdam; Brookfeld, VT. Bird, R. B., Stewart, w. E. & E.N. Lightfoot, 2002: transport phenomena. John wiley & Sons, Inc., xii, 895 p. pp, New york, Chichester. Buhmann, D. & w. Dreybrodt, 1985: Te kinetics of cal-cite dissolution and precipitation in geologically relevant situations of karst areas.1. Open system.-Chemical geology, 48, 189-211. Corbel, J., 1959: Vitesse de l’erosion.- Zeitschrif fur Geomorphologie, 3, 1-2. Dreybrodt, w. , Gabrovšek, F. & D. Romanov, 2005: Processes of speleogenesis: A modeling approach. Vol. 4, Carsologica, Založba ZRC, 375 pp, Ljubljana. Dreybrodt, w. , 1988: Processes in karst systems: physics, chemistry, and geology. Springer-Verlag, xii, 288 p. pp, Berlin; New york. Eisenlohr, L., Meteva, K., Gabrovšek, F. & w. Dreybrodt, 1999: Te inhibiting action of intrinsic impurities in natural calcium carbonate minerals to their dissolution kinetics in aqueous H2O-CO2 solutions.- Geo-chimica Et Cosmochimica Acta, 63, 989-1001. Ford, D.C. & P. williams, 1989: Karst geomorphology and hydrology. Unwin Hyman, 601 pp, London. Kaufmann, G. & w. Dreybrodt, 2007: Calcite dissolutio n kinetics in the system CaCO3-H2O-CaCO3 at high undersaturation.- Geochimica Et Cosmochimica Acta, In Press. white, w.B., 1984: Rate processes: chemical kinetics and karst landform development. In: La Fleur (Ed.): Groundwater as a geomorphic agent. Allen and Un-win, 227-248. white, w. B., 1988: Geomorphology and hydrology of karst terrains. Oxford University Press, ix, 464 pp, New york. TIME in KARST – 2007 13 COBISS: 1.01 VARIATION IN RATES OF KARST PROCESSES SPREMENLJIVOST HITROSTI KRAŠKIH PROCESOV Arthur N. PALMER1 Abstract UDC 551.44 Arthur N. Palmer: Variation in rates of Karst processes Te development of karst is not a linear process but instead takes place at irregular rates that typically include episodes of stagnation and even retrograde processes in which the evolution toward maturity is reversed. Te magnitude and nature of these irregularities difers with the length of time considered. Contemporary measurements in caves show fuctuations in dissolution rate with changes in season, discharge, and soil conditions. Dissolution is sometimes interrupted by intervals of mineral deposition. Observed dissolution rates can be extrapolated to obtain estimates of long-term growth of a solution feature. But this approach is fawed, because as the time scale increases, the rates are disrupted by climate changes, and by variations that are inherent within the evolutionary history of the karst feature (e.g., increased CO2 loss from caves as entrances develop). At time scales of 105-106 years, karst evolution can be interrupted or accelerated by widespread fuctuations in base level and surface river patterns. An example is the relation between karst and the development of the Ohio River valley in east-central U.S.A. At a scale of 106-108 years, tectonic and stratigraphic events cause long-term changes in the mechanism and style of karst development. For example, much of the karst in the Rocky Mountains of North America has experienced two phases of pre-burial Carboniferous karst, mineral accretion during deep burial from Permian to Cretaceous, extensive cave development during Paleocene-Eocene uplif, and stagnation and partial mineral deposition caused by late Tertiary aggradation. At such large time scales, it is difcult to determine rates of karst development precisely, if at all. Instead it is appropriate to divide the evolutionary history into discrete episodes that correlate with regional tectonic and stratigraphic events. Key words: Karst evolution, dissolution rates, retrograde processes, paleokarst. Izvleček UDK 551.44 Arthur N. Palmer: Spremenljivost hitrosti kraških procesov Razvoj krasa ni lineareni proces, pač pa poteka s spremenljivo hitrostjo, značilna so tudi obdobja stagnacije in obdobja, ko je razvoj obrnjen v smeri manj zrele faze. Velikost in narava sprememb sta odvisni tudi od časovnega merila v katerem jih opazujemo. Današnja merjenja v jamah kažejo, da je hitrost raztapljanja odvisna od letnega časa, pretoka in pogojev v prsti. Raztapljanje je občasno prekinjeno z obdobjem izločanja. Izmerjene hitrosti raztapljanja lahko ekstrapoliramo v času in na osnovi tega sklepamo o rasti določene korozijske oblike. Vendar bomo pri tem storili napako, saj merjenja ne vsebujejo dolgočasovnih sprememb. Te so lahko posledica različnih dejavnikov, kot so klimatske spremembe in spremembe, ki nastanejo zaradi samega razvoja krasa (npr. uhajanje CO2 zaradi odpiranja jamskih vhodov). V časovnem merilu 105-106 let razvoj krasa prekinjajo ali pospešujejo spremembe erozijske baze in spremembe površinskih vodotokov. Tak primer je povezava med razvojem krasa in doline reke Ohio v vzhodnem delu centralnih ZDA. V časovnem merilu 106-108 let tektonski in strati-grafski dogodki povzročajo dolgočasovne spremembe v razvoju krasu. Tak primer je kras v Skalnem gorovju v Severni Ameriki. Dvem fazam zakrasevanja v karbonu je sledil pokop in mineralna zapolnitev med permom in kredo. Temu je sledil obširen razvoj jam med paleocensko-eocenskim dvigom ter stagnacija in delna mineralna zapolnitev v poznoterciarni agradaciji. V tako velikem časovnem merilu je težko določiti hitrost razvoja krasa, če sploh. Primerneje je, da razvojno zgodovino razdelimo v obdobja, ki ustrezajo pomembnejšim regionalnim tektonskim in stratigrafskim dogajanjem. Ključne besede: razvoj krasa, hitrost raztapljanja, procesi nazadovanja, paleokras. 1 Department of Earth Sciences, State University of New york, Oneonta, Ny 13820-4015, U.S.A. e-mail: palmeran@oneonta.edu Received/Prejeto: 27.11.2006 TIME in KARST, POSTOJNA 2007, 15–24 ARTHUR N. PALMER INTRODUCTION In any discussion of the age of karst, one must consider the rates of the genetic processes and how they vary with time. Tese are infuenced by the length of time over which they have operated. Karst development undergoes large variations in rate and is commonly interrupted by periods of stagnation or even retrograde processes in One approach to interpreting karst history is to measure current rates of bedrock dissolution, for example by applying the mass balance, or by measuring rates of bedrock retreat with micrometers or standardized bedrock tablets. In the two following studies, empirical kinetic equations are applied. On the basis of prior dissolution experiments, feld measurements of water chemistry are used to estimate dissolution or accretion rates at specifc locations and times. Field example: eastern New York State Chemical measurements were made during 1985-1996 in streams of McFail’s Cave, New york (Fig. 1; Palmer, 1996). Suitable data-loggers were not available for use in this food-prone cave, so measurements were made randomly at every opportunity. Although statistically shaky Fig. 1: map of mcFail’s Cave, New york, showing location of sampling sites. which mass is accumulated instead of removed. Tis paper focuses on several feld examples that illustrate these processes and the difculty of quantifying them. Tese studies are still in progress and are used here only as points for discussion. compared to continuous or short-interval sampling, this approach allowed full chemical analyses. Te cave, in Silurian-Devonian limestones, consists of stream passages fed by dolines and ponors. Local soil PCO2 is 0.02-0.04 atm, but in this well-aerated cave the mean PCO2 of streams is only ~0.003 atm. Most measurements were made in the main passage and were correlated with discharge, but this location was not accessible during high fow. To provide broader coverage, additional measurements were made in similar passages with year-round accessibility. Chemical variations between sampling sites were negligible compared to variations with time. To allow extrapolation, the measurements were combined in a probability plot (Fig. 2), in which SI = log (IAP/K), IAP = (Ca2+)(CO32-), and K = calcite solubility product. Fig. 2: Probability plot of calcite saturation index in mcFail’s Cave for the period 1985-1996, where SI = log(IAP/K). data points are triangles; X = example of probability interval used in table 1. Although the passages involved are active canyons, the water is conspicuously supersaturated except during the highest 20-30% of fow. At low fow the calcite SI ofen exceeds +0.4 (~138% saturation). Calcite can precipitate SHORT-TERM VARIATIONS IN DISSOLUTION RATE 16 TIME in KARST – 2007 VARIATION IN RATES OF KARST PROCESSES at approximately SI > +0.2, so why does it not precipitate in the cave at those times? During a particularly dry summer (1995), a conspicuous calcite layer did accumulate on the canyon foors. Tis coating averaged 0.3 mm thick with inclusions of clay and quartz silt (Figs. 3 & 4). It was limited to surfaces that remained water-covered during lowest fow and formed a continuous layer in areas of steep gradient (supercritical fow) but only discontinuous patches in ponded water. Fig. 3: main stream of mcFail’s Cave during the summer of 1995, with calcite coating on foor of canyon. In mid-January, 1996, heavy rain fell on rapidly melting snow and produced a food with a return period of ~50 years. Te main cave entrance was covered by 5 m of water, and smaller inputs contained roaring waterfalls. Te calcite SI of the water entering the cave averaged -1.9 (cf. Fig. 2). Tis sample is not included in the statistics, as it was not random, but obtained purposely at the food peak, and it is not in the same class as the in-cave samples. However, it illustrates the high dissolutional capacity of extreme foodwater. Te rate of limestone removal can be estimated by S = 31.56 k (1 - C/Cs)n / ? cm/yr (Palmer, 1991), where S = rate of bedrock retreat, k = rate constant (mg-cm/L-sec), n = reaction order (di- mensionless), Cs = calcite saturation concentration, C = actual concentration of dissolved calcite, and ? = rock density (g/cm3). C/Cs is the saturation ratio, where 1.0 represents calcite saturation. From computer analysis, C/Cs ~ (IAP/K)0.35. For the cave conditions (mean PCO2 = 0.003 atm and T = 8°C), laboratory measurements by Plummer et al. (1978) show that k ~0.01 and n ~2.2 at C/Cs < 0.6, and k ~0.05 and n ~4 at C/Cs > 0.6 in open-system turbulent fow. Bedrock density is ~2.7 g/cm3 in this low-porosity rock. From chemical measurements during the winter and spring of 1996, it was predicted that the entire calcite coating of 1995 should have been removed by the time the cave became accessible in May. In fact, all but a few sheltered remnants of the calcite had been removed by then. Although mechanical abrasion may have aided the removal in places, the agreement between prediction and result is mild support for the validity of this approach. Fig. 2 includes a best-ft regression line through the chemical data. where this line extends below saturation, the probability scale was divided into 5% increments. From the mean SI in each increment, a net dissolution rate of 1.3 x 10-3 cm/yr was calculated for the period of study (Table 1). At that rate, the main cave stream would have deepened about 18 cm since the last glacial retreat in the region about 14,000 years ago. Tis is compatible with the presence of varved clays no more than a few centimeters above the lowest bedrock foors. Te clay was deposited when retreating glaciers blocked the local surface river, fooding the valley and neighboring caves. Probability range Mean C/Cs Mean S (cm/yr) Net annual entrenchment (cm) <0.05 ~0.52 ~0.017 ~8.5 x 10 4 0.05 - 0.10 0.65 0.0064 3.2 x 10 4 0.10 - 0.15 0.74 0.0019 9.5 x 105 0.15 - 0.20 0.88 8.8 x 105 4.4 x 106 0.20 - 0.25 0.89 7.4 x 105 3.7 x 106 0.25 - 0.30 0.95 2.1 x 106 1.1 x 107 TOTAL: 1.3 x 10-3 cm/yr 13 mm/1000 yrs tab. 1: Net dissolution rate in mcFail’s Cave canyons, 1985-1996,where the best-ft line in Fig. 2 falls below SI = 0. Entrenchment rates are calculated from the regression line, rather than from specifc data points, and provide only a rough approximation. At the estimated entrenchment rate, the 10 m depth of the main McFail’s canyon would have required more than 700,000 years to form. Tis rate seems low for an active canyon with a gradient of 1.2 degrees, but it is TIME in KARST – 2007 17 ARTHUR N. PALMER Fig. 4: Tin-section photomicrograph showing calcite crust on a limestone pebble from mcFail’s Cave (September, 1995). compatible with U/T speleothem dates. Related caves at the same elevation as McFail’s contain speleothems dated up to 277 ka (Dumont, 1995; Lauritzen & Mylroie, 2000; Mylroie & Mylroie, 2004). Some speleothems were located near the cave foors, so the passages themselves are far older. But the entrenchment rate during this period must have varied because of climate changes and burial beneath glacial ice for several tens of thousands of years. (Te only known glaciation in the area was wisconsin-an.) Te coarse bedload in parts of the cave also suggests mechanical abrasion during high fow. Te entrenchment rate has probably decreased with time. when entrances were blocked by glacial sediment, or had not yet enlarged enough to form open holes, escape of CO2 to the surface must have been severely lim- ited and the mean aggressiveness would have been higher than it is today. Also, calcareous glacial deposits cause low-fow inputs to be saturated with calcite before they even reach the cave. Te main canyon of the cave has an entirely vadose origin because it extends exactly down the local dip of the strata, except where it is defected by joints (Fig. 1). Terefore the canyon originated afer surface rivers had entrenched below its level (currently about 300 m above sea level). Although the age of the landscape is difcult to determine from the surface, data from the cave can provide helpful information. Mammoth Cave, Kentucky Meiman & Groves (1997), Anthony & Groves (1997), and Groves & Meiman (2005) conducted a similar study in the main river passage of Mammoth Cave, Kentucky. Tey made a high-frequency record of water levels in monitor wells, combined with periodic measurements of water chemistry. To calculate dissolution rates, they used the kinetic equation described above. Because of thick sediment, cave enlargement rates could not be estimated precisely. However, the authors determined that during the highest 5% of fow, 38% of the mass was removed (vs. about 65% in McFail’s). Te diference is probably due, at least partly, to the lack of entrances near the sampling sites in Mammoth Cave through which CO2 is lost, the higher carbonate content of soils in the New york karst, and the dominance of sinking-stream inputs to McFail’s Cave during severe foods. VARIATION IN KARST PROCESSES AT TIME SCALES OF 105–106 yEARS Te low-relief karst plateaus of Kentucky and Indiana, U.S.A., are developed on early Carboniferous carbonates and include extensive doline felds bordered by sinking streams. Tese include the Pennyroyal Plateau in Kentucky and the Mitchell Plain in Indiana. Tey are dissected to a maximum of 50-65 m by river valleys. Near rivers, inter-doline divides and residual fat areas lie 175-190 m above sea level, and up to a few tens of meters higher elsewhere. Although resistant beds form local fat areas, the overall surface is discordant to the strata. Te surface is mantled in many places by residual, colluvial, and alluvial sediment up to 30 m thick, the surface of which is concordant with the erosion surface on nearby bedrock. In the Mitchell Plain the deposits are attributed to a widespread Tertiary rise in base level (Palmer & Palmer, 1975). On the Pennyroyal, Ray (1996) calls this relatively fat surface the Green River Strath and attributes it to fu-vial processes. 18 TIME in KARST – 2007 Caves are common in the karst plains and in adjacent sandstone-capped uplands. Mammoth Cave, Kentucky, is the best-known upland example. Its highest passages correlate with nearby low-relief areas of the Pennyroyal (Fig. 5), and passage patterns and gradients show that the Pennyroyal was the source of the cave water (Palmer, 1981). Tese passages are mostly large canyons flled partly or completely with stream sediment (Fig. 6). Dating of these sediments by cosmogenic radionuclides gives ages up to 4 Ma (Granger et al., 2001), but in areas bordering the Green River (the outlet for Mammoth Cave water), most samples date to ~2.2 Ma (see also Anthony & Granger, 2004, 2006). Tese passages record a history of slow Tertiary entrenchment interspersed with aggradation, and with a widespread rise in base level of more than 20 m at ~2.2 Ma. Te fragmentary sediment surfaces at the same elevation in the Pennyroyal must be correlative. Te cause of the widespread aggradation at VARIATION IN RATES OF KARST PROCESSES Fig.5: Location of mammoth Cave and surrounding landscapes. m = mitchell Plain, P = Pennyroyal Plateau, U = sandstone-capped uplands. X = pre-Pleistocene head of Ohio River. 1, 2, 3 = sequence of drainage from Appalachian mountains. 1 is probable but entirely hypothetical. 2 = late tertiary “teays River,” which is well known by its former valley, now flled with glacial sediment. 3 = course of the Ohio River since the early Pleistocene. Afer Palmer (1981); see also Granger et al., ( 2001) for explanation. Fig. 7: typical upper-level passage in mammoth Cave with detrital sediment fll. Tis is a former tourist trail that is no longer open to the public. Sediment once flled the passage almost half-way but later subsided into an underlying passage. Note banks of remaining sediment on the lef. Fig. 6: Simplifed cross section through the Pennyroyal Plateau and mammoth Cave, Kentucky (afer Palmer, 1981). 2.2 Ma is uncertain. It correlates roughly with the onset of widespread continental glaciation at higher latitudes, but it may relate more directly to a drying climate during the late Pliocene, which would have favored the accumulation of sediments in lowlands. Pleistocene continental glaciers extended southward as far as northern Kentucky and caused much rearrangement of surface drainage. Initial entrenchment below the uppermost passages in Mammoth Cave may have been triggered by the establishment of drainage from the Appalachian Mountains westward to the Mississippi River, to form the so-called “Teays River” (Fig. 7; see Granger et al., 2001). Later, the previously tiny Ohio River became one of the largest rivers on the continent when the Teays was diverted into it (Fig. 7). Tese shifs en- hanced the rate of river entrenchment into the sediment-mantled plains of carbonate rock. Subsurface karst drainage developed and the surfaces became “sinkhole plains.” Pleistocene cave passages formed at various levels as much as 60-70 m below the Tertiary passages. Again, caves provide clues to the interpretation of surface landscapes that cannot be discerned from surface observations alone. Could the karst plateaus have retained vestiges of their original fat surface for 2 Ma without signifcant lowering? Although dolines extend deeply into them, nearly fat remnants of the sediment-covered and resistant bedrock surfaces remain at approximately the same elevations as the sediment in the upper-level passages of Mammoth Cave, which suggests that parts of the original surface have survived with little or no lowering. what is the current karst denudation rate? Much of the Mammoth Cave area is drained by the Turnhole Spring basin, which has an area of 220 km2 (quinlan et al., 1983). In this basin, Hess (1974) measured a mean-annual Ca content of ~60 mg/L and Mg of ~7.5 mg/L (see also Hess & white, 1993). Tese measurements represent a mean dissolved load of ~0.044 cm3/L calcite and ~0.020 cm3/L dolomite (with the simplifying assumption that TIME in KARST – 2007 19 ARTHUR N. PALMER dolomite = Mg and calcite = Ca – Mg, in moles/L). Te annual precipitation is 1.26 m/yr, and about 2/3 of it lost to evapotranspiration, so a 220 km2 basin would have a mean runof of roughly 9 x 107 m3/yr. Te loss of carbonate rock is therefore about 6000 m3/yr. Roughly half of the basin consists of exposed carbonates, so the denudation rate on that half is about 5.5 cm/1000 years. Tis fg-ure corresponds to some of the lowest measured rates of carbonate denudation elsewhere (Ford & williams, 1989, p. 112–117). Transport of solids is neglected, as is subsurface dissolution. Te Mammoth Cave System represents a maximum porosity of about than 4%, even in areas of maximum passage density (Palmer, 1995). when the denudation rate is extrapolated to 2 million years, it indicates an overall lowering of the solu- ble Pennyroyal surface of roughly 100 m. Tis is impossible, because it exceeds the total relief between the original surface and the Green River. Tere is no doubt that most of the surface has been lowered (Fig. 6), but there were evidently long periods of stagnation, especially at the beginning, when large parts of the surface were mantled with thick sediment. Most of the denudation is in the form of doline growth. Gams (1965) points out that corrosion accelerates in dolines as they grow, because of enhanced CO2 production in their thickening soils. Apparently the rate of karst denudation is higher today than during the early Pleistocene. KARST DEVELOPMENT AT TIME SCALES OF 107–108 yEARS Karst that evolves throughout entire geologic periods borne et al. (2006) describe a similarly complex history or eras tends to do so in discontinuous steps in which in the Jenolan Caves of Australia. lengthy episodes of stagnation exceed those of active karst processes. For example, certain karst areas of the Rocky Mountains and Black Hills (western U.S.A.) have undergone at least 7 diferent stages over the past 350 my but were actively forming only about 20% of that time. Jewel and wind Caves in South Dakota are good examples (Fig. 8). with mapped lengths of 218 and 196 km, they are among the most complex caves in both pattern and diversity of geologic history. Each successive set of features was superposed on the previous ones, because each provided favorable sites for those that followed. Os- Fig. 8: Geologic setting of Wind Cave, South dakota. L = madison Limestone (early Carboniferous) underlain by thin Cambrian sandstone, S = late Carboniferous sandstone, Sh = mainly shale, K = Cretaceous sandstone, OS = Oligocene sediment (mainly siltstone, widely eroded). Te upper surface of the madison is irregular paleokarst. Wt = water table in lowest passage of Wind Cave. Te cave extends only a few meters below the water table. Arrows show dominant fow pattern of today. 20 TIME in KARST – 2007 Te major stages of karst development in the Black Hills are outlined below (Palmer & Palmer, 1989, 1995): 1. Early Carboniferous carbonates of the Madison Formation were deposited on a low-gradient continental shelf. Interbedded sulfates were included in the middle and upper Madison. 2. Brecciation and early voids formed by dissolution and reduction of sulfates, plus production of sulfuric acid (Fig. 9). Sulfate rocks were almost completely removed. 3. A mid-Carboniferous karst formed throughout much of western North America (Sando, 1988). Surface features included fssures and dolines up to 30 m deep. Caves concentrated at 20-50 m below the surface along former sulfate zones and intersect earlier breccias and caves (Fig. 10). Comparison with modern caves suggests some freshwater-saltwater mixing dissolution. 4. Te karst was buried by late Carboniferous detrital sediment, and most caves were completely flled. Te sedimentary burial continued through the Cretaceous to a depth of VARIATION IN RATES OF KARST PROCESSES Fig. 9: Early solution voids and brecciation related to early Carboniferous sulfate-carbonate interactions in jewel Cave, South dakota. Tese are exposed by collapse of wall of a later cave. height of photo is about 2 m. at least 2 km. Buried caves and vugs, as well as voids in the Carboniferous sediment, were lined by white scale-nohedral calcite about 1-2 cm thick (Fig. 11). Pre-burial Fig. 10: mid-Carboniferous paleokarst, bighorn mountains, Wyoming. Caves in clif were once flled with late Carboniferous sediment, but much of it has been removed by weathering and stream erosion. Fig. 11: top: Scalenohedral calcite coating of mesozoic age on walls of Carboniferous vug, Wind Cave (crystal length ~1.5 cm). bottom: Rhombohedral calcite coating of late tertiary age on weathered walls of an early tertiary passage, jewel Cave (maximum thickness of calcite = 15 cm). voids can be recognized by this distinctive coating. Along faults, surfaces were coated by euhedral quartz up to a 5 mm thick. 6. Te Black Hills and Rocky Mountains were uplifed by the Laramide orogeny (latest Cretaceous through Eocene; Fig. 8). Te climate was more humid than today’s, and the present topography above the caves was formed by the end of the Eocene. Enhanced groundwater fow enlarged earlier caves to their present form (Fig. 12). Teir layout shows evidence for mixing between shallow and deep water (Palmer and Palmer, 1989), although Bakalowicz et al. (1997) suggest a purely thermal origin. 7. Te caves drained and were exposed to subaerial weathering, which produced thick carbonate deposits in many passages. 8. Most of the Eocene landscape was buried by Oligocene sediments during a drying of the climate. Although much of this sediment has been removed by later TIME in KARST – 2007 21 ARTHUR N. PALMER Karst processes operate at rates that vary considerably with time, and the magnitude of that variation is generally greater as the developmental time span increases. At every time scale, the developmental history of karst (at least in the examples described here) includes episodes of stagnation and of retrograde development when material is deposited instead of removed. Modern measurements of the rates of karst processes can be extrapolated into the past, but this extrapolation becomes more suspect as the time span increases. Over the entire growth history of major cave systems (usually 106-107 years), many disruptions in rate are caused by changes in climate, base level, and river patterns. At time scales of 107-108 years, interpretation of evolutionary rates becomes difcult, and the history of karst is usually subdivided into discrete episodes, in the same manner as tectonic and sedimentary events. erosion, the Eocene landscape on the resistant Paleozoic-Mesozoic rocks has survived almost intact, as have the underlying caves, thanks to the present semi-arid climate. 9. Partial blockage of springs by Oligocene sediments caused a second phase of calcite coating (mainly rhombohedral) averaging 15 cm thick in Jewel Cave (Fig. 11) but thinner in wind Cave. Te earlier scalenohedral coating is still visible in pockets and vugs that were isolated from the cave development and exposed by later breakdown. In this sequence there is little information about developmental rates. Instead, the karst history is portrayed as a series of discrete episodes, which span a wide range of processes, groundwater conditions, tectonic relationships, and levels of diagenetic maturity of the host strata. All efects have overlapped, and in some caves it is possible to stand in a single spot and distinguish every phase of their history. Fig. 12: typical cave passage of Eocene age in Wind Cave, showing remnants of earlier breccia (b) and paleo-fll (P).height of photo is about 2 m. As a karst feature develops toward maturity, it tends to undergo inherent changes in developmental rate. For example, a cave may decrease in enlargement rate as entrances open and enlarge, allowing greater rates of CO2 loss. Rates of karst development may increase with time as dolines develop and enlarge, owing to greater exposure of soluble rock and accumulation of high-CO2 soils in depressions. It is impossible to interpret caves and karst without a solid understanding of their surrounding geology and physiography. But, despite uncertainties about their rates of development, karst features can provide more information about the surrounding landscape than vice versa. CONCLUSIONS 22 TIME in KARST – 2007 VARIATION IN RATES OF KARST PROCESSES REFERENCES Anthony, D.M. & D.E. Granger, 2004: A Late Tertiary origin for multilevel caves along the western escarpment of the Cumberland Plateau, Tennessee and Kentucky, established by cosmogenic 26Al and 10Be. - Journal of Cave and Karst Studies, 66, 2, 46-55. Anthony, D.M. & D.E. Granger, 2006: Five million years of Appalachian landscape evolution preserved in cave sediments. - In R.S. Harmon and C.M. wicks (eds.): Perspectives on karst geomorphology, hydrology, and geochemistry – A tribute volume to derek C. Ford and William b. White: Geological Society of America, Special Paper 404, 39-50. Anthony, D.M. & C.G. Groves, 1997: Preliminary investigations of seasonal changes in the geochemical evolution of the Logdson River, Mammoth Cave, Kentucky. - Proceedings of 6th Science Conference, 15-23, Mammoth Cave, Kentucky. Bakalowicz, M.J., D.C. Ford, T.E. Miller, A.N. Palmer & M.V. Palmer, 1987: Termal genesis of dissolution caves in the Black Hills, South Dakota. - Geological Society of America Bulletin, 99, 729-738. Dumont, K.A., 1995: Karst hydrology and geomorphology of the barrack zourie Cave System, Schoharie County, New york. - M.S. thesis, Mississippi State University, p. 71, Mississippi State, Mississippi. Gams, I., 1965: Types of accelerated corrosion. - In O. Štelcl (ed.): Problems of the speleological research. - International Congress of Speleology, 133–139, Brno, Czech. Granger, D.E., D. Fabel & A.N. Palmer, 2001: Pliocene-Pleistocene incision of the Green River, Kentucky, determined from radioactive decay of cosmogenic 26Al and 10Be in Mammoth Cave sediments. - Geological Society of America Bulletin, 113, 7, 825-836. Granger, D.E., J.w. Kirchner, & R.C. Finkel, 1997: quaternary downcutting rate of the New River, Virginia, measured from diferential decay of cosmogenic 26Al and 10Be in cave-deposited alluvium. - Geology, 25, 107–110. Groves, C. & J. Meiman, 2005: weathering, geomorphic work, and karst landscape evolution in the Cave City groundwater basin, Mammoth Cave, Kentucky. - Geomorphology, 67, 115-126. Hess, J.w., 1974: hydrochemical investigations of the central Kentucky karst aquifer system. - Ph.D. dissertation, Pennsylvania State University, p. 218, University Park, Pennsylvania. Hess, J.w. & w.B. white, 1993: Groundwater geochemistry of the carbonate aquifer, south-central Kentucky, U.S.A. - Applied Geochemistry, 8, 189-204. Lauritzen, S.-E. & J.E. Mylroie, 2000: Results of a speleo-them U/T dating reconnaissance from the Helderberg Plateau, New york. - Journal of Cave and Karst Studies, 62, 1, 20-26, Huntsville, Alabama. Meiman, J. & C. Groves, 1997: Magnitude/frequency analysis of cave passage development in the Central Kentucky Karst. - Proceedings of 6th Science Conference, 11-13, Mammoth Cave National Park, Kentucky. Mylroie, J.E. & J.R. Mylroie, 2004: Glaciated karst: How the Helderberg Plateau revised the geologic perception. - Northeastern Geology and Environmental Sciences, 26, 1-2, 82-92, Troy, New york. Osborne, R.A.L., H. Zwingmann, R.E. Pogson & D.M. Colchester, 2006: Carboniferous clay deposits from Jenolan Caves, New South wales: Implications for timing of speleogenesis and regional geology. - Australian Journal of Earth Science, 53, 377-406. Palmer, A.N., 1981: A geological guide to Mammoth Cave National Park. – Zephyrus Press, p. 210, Tean-eck, New Jersey. Palmer, A.N., 1991: Origin and morphology of limestone caves. - Geological Society of America Bulletin, 103, 1-21. Palmer, A.N.,1995: Geochemical models for the origin of macroscopic solution porosity in carbonate rocks. - In Budd, D.A., P.M. Harris, & A. Saller (eds.): Unconformities in carbonate strata: Teir recognition and the signifcance of associated porosity. - American Association of Petroleum Geologists, Memoir 63, 77–101. Palmer, A.N., 1996: Rates of limestone dissolution and calcite precipitation in cave streams of east-central New york State. - Abstracts of Northeastern Section meeting, Geological Society of America, 28, 3, 89. Palmer, A.N. & M.V. Palmer, 1989: Geologic history of the Black Hills caves, South Dakota. - National Speleological Society Bulletin, 51, 2, 72-99. Palmer, A.N. & M.V. Palmer, 1995: Te Kaskaskia paleo-karst of the Northern Rocky Mountains and Black Hills, northwestern U.S.A. - Carbonates and Evapo-rites, 10, 2, 148-160, Troy, New york. Palmer, M.V. & A.N. Palmer, 1975: Landform development in the Mitchell Plain of southern Indiana: Origin of a partially karsted plain. - Zeitschrif für Geomorphologie, 19, 1-39. TIME in KARST – 2007 23 ARTHUR N. PALMER Plummer, L.N., T.M.L. wigley, T.M.L. & D.L. Parkhurst, 1978: Te kinetics of calcite dissolution in CO2-wa-ter systems at 5° to 60° C and 0.0 to 1.0 atm CO2. - American Journal of Science, 278, 179-216. quinlan, J.F., R.O. Ewers, J.A. Ray, R.L. Powell & N.C. Krothe, 1983: Ground-water hydrology and geo-morphology of the Mammoth Cave Region, Kentucky, and of the Mitchell Plain, Indiana. - Indiana Geological Survey, Field trips in Midwestern geology, 2, 1-85, Bloomington, Indiana. Ray, J.A., 1996: Fluvial features of the karst-plain erosion surface in the Mammoth Cave region. - Proceedings of 5th Science Conference, 137-156, Mammoth Cave, Kentucky. Sando, w.J., 1988: Madison Limestone (Mississippian) paleokarst: A geologic synthesis. - In N.P. James and P. w. Choquette (eds.): Paleokarst: Springer-Verlag, 256-277, New york. 24 TIME in KARST – 2007 COBISS: 1.01 TIME SCALES IN THE EVOLUTION OF SOLUTION POROSITy IN POROUS COASTAL CARBONATE AqUIFERS By MIxING CORROSION IN THE SALTwATER-FRESHwATER TRANSITION ZONE. ČASOVNO MERILO RAZVOJA POROZNOSTI ZARADI KOROZIJE MEŠANICE V MEJNEM OBMOČJU SLADKOVODNIH LEČ V MEDZRNSKO POROZNEM KARBONATNEM OBALNEM VODONOSNIKU wolfgang DREyBRODT1 & Douchko ROMANOV2 Abstract UDC 556.3:552.54:539.217 Wolfgang Dreybrodt and Douchko Romanov: Time scales in the evolution of solution porosity in porous coastal carbonate aquifers by mixing corrosion in the saltwater-freshwater transition zone. Dissolution of calcium carbonate in the saltwater-freshwater mixing zone of coastal carbonate aquifers up to now has been treated by coupling geochemical equilibrium codes to a reactive-transport model. Te result is a complex nonlinear coupled set of diferential transport-advection equations, which need high computational eforts. However, if dissolution rates of calcite are sufciently fast, such that one can assume the solution to be in equilibrium with respect to calcite a highly simplifed modelling approach can be used. To calculate initial changes of porosity in the rock matrix one only needs to solve the advection-transport equation for salinity s in the freshwater lens and its transition zone below the island. Current codes on density driven fow such as SEAwAT can be used. To obtain the dissolution capacity of the mixed saltwater-freshwater solutions the calcium equilibrium concentration ceq(s) is obtained as a function of salinity by PHREEqC-2. Initial porosity changes can then be calculated by a simple analytical expression of the gradient of the spatial distribution s(x, y) of salinity, the distribution of fow fuxes q(x,y) and the second derivative of the calcium equilibrium concentration ceq(s) with respect to salinity s. Tis modelling approach is employed to porosity evolution in homogeneous and heterogeneous carbonate islands and coastal aquifers. Te geometrical patterns of porosity changes and the reasons of their origin will be discussed in detail. Te results reveal initial changes of porosity in the order of several 10-6 per year. Tis places the time scale of cavern evolution to orders from several tens of thousands to a hundred thousand years. Keywords: Calcite dissolution, mixing corrosion, saltwater-freshwater, mixing zone, coastal aquifer, evolution of porosity. Izvleček UDK 556.3:552.54:539.217 Wolfgang Dreybrodt and Douchko Romanov: Časovno merilo razvoja poroznosti zaradi korozije mešanice v mejnem območju sladkovodnih leč v medzrnsko poroznem karbonatnem obalnem vodonosniku Dosedanji modeli raztapljanja kalcijevega karbonata v območju mešanja sladke in slane vode temeljijo na združitvi geokemičnih ravnotežnih in reakcijsko transportnih modelov. Dobljeni sistem nelinearnih enačb zahteva veliko računske moči. Če je hitrost raztapljanja dovolj visoka in lahko predpostavimo, da je raztopina ves čas v ravnotežju glede na kalcit, rešimo problem z poenostavljenim modelskim pristopom. Začetno spreminjanje poroznosti v kamninski matriki določa advekcijsko tranportna enačbo, ki opisuje slanost v sladkovodni leči in prehodnem območju pod njo. Pri reševanju porabimo dostopne programske kode. Tokove nastale zaradi razlik v gostoti modeliramo s programom SEAwAT, topnost kalcita v mešanici sladke in slane vode v odvisnosti od slanosti pa izračunamo s programom PHREEqC-2. Začetno spreminjanje poroznosti lahko nato izračunamo z enostavnim analitičnim izrazom gradienta prostorske razporeditve slanosti s(x,y), razporeditve gostot toka q(x,y) in drugega odvoda ravnotežne koncentracije kalcija po slanosti. Tak modelski pristop uporabimo pri računanju razvoja poroznosti v homogenih in heterogenih karbonatnih otokih in obalnih vodonosnikih. Podrobno so prikazani vzroki in geometrijski vzorci spreminjanja poroznosti. Rezultati kažejo, da je začetna hitrost spremembe poroznosti reda velikosti 10-6 na leto. To postavi časovno merilo razvoja jam v območje nekaj deset tisoč do sto tisoč let. Ključne besede: Raztapljanje kalcita, korozija mešanice, območje mešanja sladke in slane vode, obalni vodonosnik, razvoj poroznosti. 1 Universitaet Bremen, FB1, Karst Processes Research Group, Bremen, Germany, e-mail: dreybrodt@ifp.uni-bremen.de 2 Freie Universitaet Berlin, Fachbereich Geowissenschafen, Berlin, Germany, e-mail: dromanov@zedat.fu-berlin.de Received/Prejeto: 21.12.2006 TIME in KARST, POSTOJNA 2007, 25–34 wOLFGANG DREyBRODT & DOUCHKO ROMANOV INTRODUCTION Carbonate islands consisting of porous rocks show typical karst features characterized by large dissolution chambers close to the coast, which have been created by mixing corrosion in the fresh-saltwater transition zone (Mylroie and Carew, 2000). Figure1 represents the basic concept. Due to meteoric precipitation a freshwater lens Fig. 1: Conceptual representation of a carbonate island from mylroie and Carew (2000). the chlorine concentration s, termed as chlorinity further on, of the mixture undersaturation or supersaturation may result. Figure 3 gives an example. It depicts the diference Fig. 3: Ace9(s) = ceg(s)-cmi[(j) as a function of chlorine concentration. Te curve extends from pure freshwater (right) to pure seawater (lef). builds up, foating on the denser saltwater (Vacher, 1988) Te transition from freshwater to seawater is not sharp. Depending on many factors, such as tidal pumping, periodicity of annual recharge, and the heterogeneity of the rock’s properties in the aquifer it exhibits a transition zone. Tis zone can range from a few meters to half the depth of the lens. In this zone mixing between saltwater and freshwater activates mixing corrosion, which creates large chambers. Tese are called fank-margin caves. Figure 2 shows such a cave with its typical solutional features on its ceiling. Fig. 2: Flank-margin cave. when seawater mixes with a solution of H2O-CO2-CaCO3 saturated with respect to CaCO3 the mixture is no longer in equilibrium with respect to calcite. Depending on hceq(s) = ceq(s)-cmix(s) of the calcium concentration c ix (s) of the mixture and that of its corresponding equilibrium concentration c (s) as a function of s. Tis is the amount of calcium, which can be dissolved or precipitated, when the mixed solution is in contact with carbonate rock. Te HO-CaCO-CO solution used to calculate this data is in 2 32 equilibrium with a partial pressure of CO2 of 0.01 atm at a temperature of 20°C. Te seawater also is at 20°C. Te data in Fig. 3 were obtained by use of the code PHREEqC-2 (Parkhurst and Apello, 1999 ). From Figure 3 it is evident that mixtures with low content of seawater, chlorinity s ? 0.3 mol/^, can dissolve calcite, whereas mixtures with higher chlorinity may precipitate calcite. Renewed aggressivity due to mixing therefore occurs only at the freshwater side of the mixing zone where chlorinity is low. If one assumes that dissolution of calcite proceeds sufciently fast the solution there will be saturated with respect to calcite. Dissolution of minerals under such conditions is termed a gradient reaction (Phillips, 1991). Here we use this as a novel instrument to explain the evolution of porosity in carbonate islands. Dissolution rates of limestone are sufciently fast, such that afer mixing between saltwater and freshwater we assume saturation with respect to calcite in the entire lens. Afer attaining equilibrium the local distribution of calcium concentration c (s(x,z)) becomes stationary and exhibits gradients. Necessarily advection and difusion must transport the dissolved limestone to the outfow of the aquifer. 26 TIME in KARST – 2007 TIME SCALES IN THE EVOLUTION OF SOLUTION POROSITy IN POROUS COASTAL CARBONATE AqUIFERS ... DISSOLUTION IN THE MIxING ZONE Te advection term: In Figure 4 we consider a volume element dxdydz at position (x,y,z), into which fow, with components qx and qz, enters perpendicular to dydz or dxdy. Te fux q is defned by the volume of fuid per time unit entering through a unit of surface area and is given in [cm3/(cm2s) = cms-1]. ^fi-esh + (c« »)¦ (5) Ac«? is the increase of equilibrium concentration as given in Figure 3, s is chlorinity of seawater. sea Te difusion term: Our mass balance so far, however, is incomplete because gradients of ceq cause transport by difusion. Te rate qD of mass transport by difu-sion is given by QD=-0-DV\cmbc+Ac ) (6) Fig. 4: mass balance for the advection term. Te component qx transports solution from the neighbouring elementary cell at position (x-dx, y,z) via the area dydz into the cell dxdydz. Tis solution has already attained equilibrium ceq(s(x-dx,y,z)) at position x-dx. when it enters into the volume element dxdydz it must dissolve or precipitate limestone to adjust its calcium concentration to equilibrium ceq(s(x,y,z)) at position x. On the other hand solution from the element dxdydz fows out into the neighbouring cell with fux qx(x,y,z). Mass conservation requires that the amount of limestone dissolved per time unit in the element dxdydz must be equal to the diference of mass transported into the cell and that transported out of it. From this one fnds (qx(x, z) ¦ Ceq(x, z) - qx(x -dx,z)- Ceq(x - dx, z))dydz fit (1) dxdydz An analogue equation exists for qz the amount of limestone dissolved by the fux component q entering via the surfaces (dx,dy). (q2 (x, z) • Ceq(x, z) - qz (x, z - dz) ¦ Ceq(x, z - dz))dxdy dxdydz Qz (2) Terefore Qadv =Qx+Qz=Q' gfodipeq (S(.X>Z)) + Ceq (S(X> Z) ' ^tvq (3) Because the fux q follows the Darcy law of incompressible fuids, div(q) =0. Qato-q-gradtfimu+te»,) (4) whereby we have replaced c^ = cmlx + Ace? ¦ cmlx is the calcium concentration resulting from the mixing of sea-water and freshwater and is a linear function of Cl-con-centration s. where D = qd/? + D is the m coefcient of dispersion. ? is the porosity of the rock and d its grain size. (Phillips, 1991). Dm is the constant of molecular diffusion (10-5cm s-1). Te total rate: Te total dissolution rate q is then given by qD+q d. Qui =Qgrad(cmi,)~ ^V2(cmjJ)+ qgrad(Aceq) - 0Z)V2(Ace?) (7) Due to the linearity of c with salinity s (eqn. 5) mix one fnds grad(c i) proportional to grad(s). Te distribution of salinity is governed by the ad-vection-difusion equation ds/dt = qgrads - (V)2.s = 0, (8) because the distribution s is stationary. From the linearity of s with cmix we have dc^,/dt=qgradcmlx-0D(V)2c^= 0, (9) Te total dissolution rate qtot is given by the master equation ßto(=?grarf(ACJ-0i5V2(ACe?) (10) Since ?ceq(s(x,z)) is a function of local distribution s(x,z) by diferentiating and using the chain rule, one fnds using equation 8 ßto,=-0(qd/0+Dm)-(Vs(x,z))2 ds2 (11) Tis master equation relates the amount of dissolved material per unit volume of the rock matrix [mol cm-3 s-1] TIME in KARST – 2007 27 wOLFGANG DREyBRODT & DOUCHKO ROMANOV to the gradient of salinity s, to the second derivative of ceq(s) = cmjx(s)+Aceq(s), and the fux q. d^c^/ds1 can be obtained by diferentiating twice the data set of Figure 3. Tis data set was obtained by using the program PHRE-EqC2 and calculating about 50 closely spaced points to avoid numerical errors, when diferentiating twice. Te result is shown in Figure 5. Fig. 5: Second derivative To obtain the initial distributions of fux q and chlorinity s in the lens of a carbonate island we have used SEAwAT by USGS (Guo and Langevin, 2002). Te modeling domain is shown in Figure 6. Te island is a strip of 1 km Fig. 6: modeling domain of a carbonate island. width. Porosity 0 and the hydraulic conductivity K are uniform (0=O.3O,K = l0m/day). Te transversal disper-sivity is aT = d = 0.01 cm, the longitudinal dispersivity is Te function |Aä(3c)| and the fow distribution |?(x)| can be obtained by the numerical hydrologic model SEA-wAT, as will be shown in the next sections. To calculate the initial change of porosity it is sufcient to obtain the fux and salinity distribution of an island without considering calcite dissolution, because the time to establish a stationary state of the lens is in the order of 100 years. It is a good approximation to assume that during this time the change of porosity is insignifcant. Equation. 11 can be written in terms of the change of porosity as — = —Qtat=-0D(ys(x,z))2-----ip— (1/s) (12) dt p ds p M = 100 g/mol is the molecular weight at CaCO3, r=2.7 g/cm3 is the density of compact CaCO3. q the mass of CaCO3 dissolved per time from a unit volume of the rock matrix is given in mol s-1 cm3. 90/dt is the amount of volume dissolved per time from a unit volume of the rock matrix (cm3s-1/cm3). By use of equation 12 it is now possible to construct a conceptual frame for the evolution of porosity. Tests of this approach on simple benchmark models have shown its reliability and have found agreement to experimental data (Romanov and Dreybrodt, 2006). al = 0.1 cm. Infltration is 3 . 10-3m/day =1.11m/year. Tis way the maximal depth of the lens is about 50 m below sea level. Te lower border of the domain reaches down to 70 m. At that boundary an impermeable layer imposes no-fow conditions. Te grid size in the domain is 1 m x 1 m in the part below sea level. In the part above sea level (2 m) the grid size is 0.2 m by 1 m. In its initial state when the island emerges out from the sea the entire aquifer is flled with seawater. when the island receives recharge from meteoric freshwater the lens builds up. A stable stationary lens is obtained afer about 30 years. Fig.7 shows the results of the model run. Figure 7a shows the freshwater lens (white), the transition zone and its distribution of Cl-concentration by a color code. From this distribution of chlorinity one can extract the scalar value Vs(jc) and d2ceq(s(x))/d s1. Figure 7b shows the chlorinity in units normalized to its maximum value along several horizontal sections as depicted in Figure 7a. Te lowest section at -68 m is entirely in saltwater with maximum Cl-concentration. Te section at -55 m extends through the almost horizontal base of INITIAL CHANGES OF POROSITy IN A HOMOGENEOUS ISLAND. 28 TIME in KARST – 2007 TIME SCALES IN THE EVOLUTION OF SOLUTION POROSITy IN POROUS COASTAL CARBONATE AqUIFERS ... Fig. 7: homogeneous island. a) Local distribution of chlorinity s(x). Te white region designates the freshwater lens. b) chlorinity along horizontal sections as indicated in a). the lens and shows a wide zone where the concentration raises to that of seawater. Te upper sections cut through the mixing zone and there the rise in concentration from freshwater to seawater becomes steeper. Te square of the gradient \Vs\ is shown by Figures 8a,b also normalized to its maximum value in Figure 8b. Figure 8a illustrates its local distribution, which exhibits large values only in the region of the transition zone. Te horizontal distribution along horizontal sections is depicted in Figure 8b. Te second derivative d2c (,s(x))/3 s2 obtained from the Cl-concentration in Figure 7a is given in Figure 9a. Its distribution is limited to that part of the transition zone with 0 < s < 0.03 mole/^. See Figure 5. Tis corresponds to a narrow fringe at the freshwater side of the transition zone with seawater content from zero up to about 4%. In any case creation of porosity is possible only in this restricted region. Figure 9b for completeness depicts some distributions of 32c (s(3c))/3 s2 along horizontal sections. To calculate the initial rate of change in porosity (conf. eqn 12) the Darcy fuxes q must be known. Tey are also obtained from the model run and shown in Fig 10. Te fux is low in the center of the island q^\ m/year), but increases by orders of magnitude when the fuid Fig. 8: homogeneous island. a) Local distribution of the square of gradients |Vs(x)| , b) square of gradients along horizontal sections as indicated in a). moves coastward, where it becomes about 0.2 m/day at the outfow. Te dispersion coefcient D = qd/O + Dm (conf. eqn. 11) depends on the fux q, but also on the coef-cient of molecular difusion Dm=10-5 cm2/s. For low fux q<10-4 cms-1 and particle diameters d?10-2 cm dispersion is dominated by molecular difusion. In the following scenarios we have used d=10-2 cm, a realistic value in porous limestone. Terefore in the range of fux, which can be read from Figure 10b the dispersion coefcient in the center of the island is D=10-5 cm2s-1. It increases by about 60% of this value at the coast. From the data given in Figures 7a, 8a, and 9a the initial porosity is obtained by use of eqn. 12. Figure 11 illustrates these results. Changes in porosity are restricted to a small fringe in the transition zone and are fairly even along it. Tey are in the order of 10-6 year-1. Tis is sufcient to create substantial porosity within 100,000 years. At the outfow fank margin caves can develop in 10,000 years. One has to keep in mind, however, that the approximation as a homogeneous island is a high idealization. Any disturbances, which increase the width of the transition zone, will reduce the gradients of chlorinity and therefore on more realistic settings the initial porosity changes accordingly. TIME in KARST – 2007 29 wOLFGANG DREyBRODT & DOUCHKO ROMANOV Fig. 9: homogeneous island. Local distribution of the second derivative —P, b) second derivative along horizontal sections ds2 as indicated in a). As we have stated already, the second derivative is restricted to narrow regions in the freshwater side of the transition zone. It exhibits signifcant values only at locations where the water contains between zero and 4% saltwater (see Figure 5). On the other hand the gradient in salinity is maximal at mixtures of about 50% seawater, because it arises from a difusive process. In the region of maximal gradients, however, the second derivative is small. Vice versa in the region of high values of the second derivative, the gradients of salinity are low. Tis is illustrated in Figure 12. Tis fgure is an overlay of the horizontal distributions (grads)2 in Figure 8b (red curves), the second derivative in Figure 9b (green curves), and the initial porosity change in Figure 11b (black curves). All curves are normalized to their individual maximum values. Terefore their values are not comparable in this fgure. what can be compared, are the locations. Evidently the curves for gradients and second derivative are well separated. Te curves of porosity change are proportional to the product of the square of the gradient and the second derivative. Porosity change displays high values Fig. 10: homogeneous island. a) Local distribution of fux b) fux along horizontal sections as indicated in a). in between their maxima but close to the region of high values of the second derivative. Figure 13 further illustrates this qualitatively. Te red region depicts the locations of the modeling domain where (grads)2 exhibits values val a 10"2 val^ valma is the maximal value. Te green region shows these locations for the second derivative and fnally the black region shows the locations of signifcant changes of porosity. Tese fndings agree with those of Sanford and Konikow (1989) who also found that changes in porosity are restricted to regions where waters contain between 0.5% and 3% of seawater. It should be noted here that any mechanism, which changes the sigmoid shape of the salinity distribution to a linear profle would enhance evolution of porosity dramatically. In this case salinity gradients become constant in the entire mixing zone and their value is at least one order of magnitude higher at the maximal value of the second derivative. One could speculate that tidal pumping and fuctuations of the water table due to seasonal changes of infltration could cause such linear mixing zones. Present observations in boreholes give some evidence for such transition zones. 30 TIME in KARST – 2007 TIME SCALES IN THE EVOLUTION OF SOLUTION POROSITy IN POROUS COASTAL CARBONATE AqUIFERS ... Fig. 11: homogeneous island. a) Local distribution of initial change of porosity dip/dt. b) d/ dt along horizontal sections as indicated in a). Fig. 12: homogeneous island. (grads)2 (red), ds2 (green), and) dtp/ dt (black) along horizontal sections of the island. Numbers on the sets of curves give the depth of the section. Fig. 13: homogeneous island. Regions of (grads)2 (red), of (green), and change of porosity (black). INITIAL CHANGES OF POROSITy IN A HETEROGENEOUS ISLAND A more realistic approach to nature can be taken by employing a geo-statistical distribution of hydraulic conductivities. Figure 14 shows such a distribution generated with the sofware of Chiang and Kinzelbach (1998). It covers conductivities of two orders of magnitude from about 380 m/day (red) down to 2 m/day (dark blue). Most of the aquifer is occupied by values between 10-200 m/day. Otherwise all previous boundary conditions are unchanged. Te fow feld is illustrated in Figure 15. Flux is unevenly distributed, because the heterogeneous distribution of conductivities distorts the pathways of fuid elements in comparison to the regular ones in a homogeneous island. Consequently the freshwater lens in Figure 16 shows a wide transition zone (compare to Figure 7a). Fig. 14: heterogeneous island. Statistical distribution of hydraulic conductivity in the modeling domain. TIME in KARST – 2007 31 wOLFGANG DREyBRODT & DOUCHKO ROMANOV Fig. 15: heterogeneous island. Local distribution of fux q. Fig. 16: heterogeneous island. Local distribution of chlorinity s(x). Te white region designates the freshwater lens. Fig. 17: heterogeneous island. Local distribution of |Vs(x)| . Te square of the gradient is limited to the seawa-ter side of the transition zone, as can be visualized from Figure 17. Te region of 0-4% mixtures extends far into Fig. 18: heterogeneous island. Local distribution of derivatives as2 Fig. 19: heterogeneous island. Local distribution of initial porosity change dl dt. Fig. 20: heterogeneous island. Regions of high values of (grads)2 o2 Ac (red), -----P (green), and change of porosity (black) in the modeling domain. the freshwater lens. Tis can be also visualized from the second derivatives as shown in Figure 18. Figure 19 illustrates the initial change of porosity, which exhibits high values of 3 . 10-6 year1 (red) at only a few locations close to the freshwater side of the transition 32 TIME in KARST – 2007 TIME SCALES IN THE EVOLUTION OF SOLUTION POROSITy IN POROUS COASTAL CARBONATE AqUIFERS ... zone. At some favorable locations (red and yellow) caves ure 21 depicts (grads)2 (red), d2c /ds2 (green), and d(p/dt may evolve there in several 10,000 to 100,000 years. (black) along selected horizontal sections. Tis is further illustrated by Figure 20, which shows In both fgures we fnd that the regions of (grads)2 the regions of high values for (grads)2 (red), d2c /ds2 (red), d2ceq /ds2 (green) are well separated and porosity (green), and 90/dt (black) in the modeling domain. Fig- develops in between. Due to the heterogeneity, however, the patterns become complex. Fig. 21: a) heterogeneous island. distributions of (grads)2 (red), ----^ (green), and porosity change (black) along selected horizontal sections. Number on the sets of curves give the depth of the section. INITIAL CHANGES OF POROSITy IN SALTwATER TONGUES. when impermeable strata underlay an island sufciently close to its surface the freshwater lens cannot extend below this layer and a saltwater tongue intrudes from the coastland inward until it reaches the impermeable layer. From thereon the freshwater lens is truncated by this layer. In this situation mixing of waters is restricted to the transition zone of the tongue and one expects high dissolution rates in this region. Fig. 22 shows the local distribution of chlorinity and the initial change of porosity using the statistical distribution in Figure 14 for the upper permeable part. Te mixing zone exhibits a structure similar to that of the heterogeneous island at the corresponding locations. Porosity changes at the outfow are low, but we fnd values up to 10-6 1/year land inward at various lo- Fig. 22: Coastal aquifer with heterogeneous conductivity down to 29 m as used in Fig. 14. Te strata below 29 m are impermeable (grey). Local distribution of Cl-concentration s(x) and initial porosity change d

780 ka (paleomag.) Tertiary weathered soils Upper Miocene? Audra 1996 Gr. Vallier Vercors 1500 / 200 m Tertiary, Lower Pleistocene glacial varves (paleomag.) Tertiary weathered soils yes Upper Miocene Audra & Rochette 1993 Réseau de la Dent de Crolles Chartreuse 1700 / 250 m > 400 ka (U/Th) Cretac. sandstones yes Upper Miocene? Audra 1994 Gr. Théophile Gdes Rousses 1900 / 1850 m 95 ka (U/Th) Middle Pleistocene Audra & Quinif 1997 Gr. de l’Adaouste Provence Stratigraphic correlation Miocene pebbles Artesian Tortonian Audra & al. 2002 Systeme du Granier Chartreuse 1500 / 1000 m > 1-1,5 Ma (234U / 238U equilibr., paleomag.) 1,8-5,3 Ma (cosmonucleides) Upper cretac. and oligo. limest. - Cretac. sandstones - weathered soils yes Upper Miocene? Hobléa 1999; Hobléa & Häuselmann 2007 Beatushöhle - Bärenschacht Siebenhengste 890 / 558 m > 350 ka (U/Th) Pleistocene Häuselmann 2002 Siebenhengste Siebenhengste 1900 / 558 m 4.4 Ma (cosmonucleides) Pliocene Häuselmann & Granger 2005 Jochloch Jungfrau 3470 m Lower Pleistocene? (palynology) practically no catchment today Lower Pleistocene Wildberger & Preiswerk 1997 Ofenloch Churfisten 655 / 419 m > 780 ka (paleomag.) Pliocene Müller 1995 Hölloch-Silberensystem Silberen 1650 / 640 m >350 ka (U/Th), <780 (paleomag) Lower Pleistocene? Battisti Paganella 1600 m > 1-1,5 Ma (234U / 238U equilibr.) Cherts from Eocene limestones yes Oligo-Miocene Conturines Dolomite 2775 m > 1-1,5 Ma (234U / 238U equilibr.) yes Oligo-Miocene Frisia & al. 1994 Capana Stoppani, Tacchi-Zelbio Pian del Tivano 900 / 200 m > 350 ka (U/Th) Boulders from glacial sinkholes yes Oligo-Miocene Tognini 1999, 2001 Gr. dell’Alpe Madrona Mte Bisbino 1000 / 200 m > 350 ka (U/Th) Miocene Tognini 1999, 2001 Covoli di Velo Ponte di Veia Mte Lessini 33-38 Ma (K/Ar) yes Eocene and Oligocene Rossi & Zorzin 1993 Gr. Masera Lario 200 / 361 m ? 2.6 to 7.2 Ma (cosmonucleides) Fluvial pebbles Pliocene or older Häuselmann unpub. Bini & Zuccoli 2004 Gr. On the Road Campo dei Fiori 805 / 300 m > 1-1,5 Ma (234U / 238U equilibr) Oligo-Miocene Uggeri 1992 Gr. Via col Vento Campo dei Fiori 1015 / 300 m > 350 ka (U/Th) Upper Plio. glacial sediments Oligo-Miocene Uggeri 1992 Gta. sopra Fontana Marella Campo dei Fiori 1040 / 300 m Middle Pleistocene (micro-fauna) Conglomerate with crystalline pebbles Ferralitic soils yes Oligo-Miocene Zanalda 1994 Ciota Ciara – Cuitarun caves Mte Fenara Large miocene f uvial pebbles yes Oligo-Miocene Fantoni & Fantoni 1991 Cosa Nostra-Bergerhöhle Tennengebirge 1600-1000 / 500 m > 780 ka (paleomag) yes Augensteine yes Miocene - Upper Pliocene Audra & al. 2002 Mammuthöhle Dachstein 1500-1300 / 500 m yes Augensteine yes Miocene Trimmel 1961, 1992; Frisch & al. 2002 Eisriesenwelt Tennengebirge 1500 / 600 m yes yes Lower Pliocene? Audra 1994 Feichtnerschacht Kitzsteinhorn 2000 / 1000 m 118 ka (U/Th) Pliocene? Audra 2001, Ciszewski & Recielski 2001 Poloska jama Mt Osojnica 750 / 500 m yes Crnelsko brezno Kanin 1400 / 400 m > 780 ka (paleomag) Glacial varves Audra 2000 Snezna jama Kamnik Alps 1600 / 600 m 1.8 to 3.6 or 5 Ma (paleomag) yes yes yes Miocene? Bosak & al. 2002 tab. 1: Synthesis of information about the quoted caves systems TIME in KARST – 2007 59 PHILIPPE AUDRA, ALFREDO BINI, FRANCI GABROVŠEK, PHILIPP HäUSELMANN, FABIEN HOBLéA, PIERRE-yVES JEANNIN, ... As a conclusion, a warm climate induces passage growth and speleothem deposition, whereas a cold climate generally tends to obstruct the lower passages by sediments. Glacial sediments covering older speleothems: cave systems may predate glaciations Some cave sediments correspond to very old glaciations, according to paleomagnetic measurements that show inverse polarity: Ofenloch/Churfrsten (Müller 1995), grotte Vallier/Vercors (Audra & Rochette 1993), Crnel-sko brezno/Kanin (Audra 2000). Tese sediments ofen overlie successions of alterites or massive fowstone deposits, which in turn prove the existence of a warm and humid climate, thus showing that the cave systems predate those glaciations. Some of the old speleothems are more or less intensely corroded by fowing water postdating their deposition. Cave development and glacial activity - Glacial abrasion at the surface and erosion in the va-dose zone. At the surface, the glacial activity is without doubt responsible for the abrasion of a variable amount of bedrock (50-250 m), which has surfaced old conduits that previously were deeply buried. Tis is manifested by wide open shafs, cut galleries and arches. During glacial melt, meltwater disappeared into distinct sectors. As soon as fractures were connected to preexisting conduits, they enlarged quickly and thus formed the “invasion vadose shafs” (Ford 1977), which can reach several hundred meters of depth: Granier, Silberen, Kanin (Ku-naver 1983, 1996). Te efectiveness of such meltwater is Some existing caves and karst features clearly correspond to a strongly diferent topography than today. Tey are therefore supposed to be older. In the following paragraphs, the position and morphology of caves are compared to today’s landscape. Ten cave sediment characteristics are presented and discussed. In a third part, links between caves and well-recognized paleotopographies are explained. All those indications are clear evidences for a high age of cave systems. CAVE SySTEMS VS. PRESENT TOPOGRAPHy Perched phreatic tubes Conduits with an elliptical morphology are sometimes perched considerably above the present base level (Tab. 1, mainly due to its velocity in the vertical cascades as well as their abrasive mineral load originating from bedrock and till material. - Some new cave systems appeared in the intra-Al-pine karst area due to glacial erosion. Tin limestone belts or marbles intercalated with metamorphic series were freed from their impervious cover by glacial erosion. Some caves are still in direct relationship with the peri-glacial fow, and act as swallowholes. Teir morphology refects the cascading waterfow and has a juvenile form: Perte du Grand Marchet/Vanoise, Sur Crap/Graubün-den (wildberger et al. 2001). At the Grotte Téophile/ Grandes Rousses, U/T datings evidenced that the cave was active at least along the two glacial-interglacial cycles that are marked by the sequence of passage-forming/fll-ing with gravel/sinter deposition (Audra & quinif 1997). Since cave development mainly occurred during inter-glacial, the efect of the glacier is only indirect, by eroding the impervious covers (Audra 2004). - Te lower phases of huge cave systems are indirectly generated by glacial valley-deepening. while the uppermost cave systems are ofen older than the glacia-tions (infra), the lower passages are ofen of quaternary age, since they are related to valleys evidently deepened by glaciers. In this respect, glaciers are indirectly responsible for the creation of new cave passages (Siebenhengste, Chartreuse, Vercors). Tis strongly contrasts with the South Alpine domain, where valleys were deepened during the Messinian event. Here, glaciations contributed merely to the inflling of the preexisting valleys. Tus, most of the South Alpine cave systems are thought to be older than the glaciations. 3rd column). Tey developed close to a paleo base level, long before today’s valley deepening. At the Siebenhengste, the highest phases even show a fow direction opposite to the present one. Caves intersected by current topography Old perched caves are ofen segmented by a subsequent lowering of the surface. Two situations are usually found in the feld: - Old phreatic caves at the surface of karst plateaus, which have been eroded by glacial abrasion (Grigna, Dolomites, Triglav, Kanin, Tennengebirge…) - Old phreatic caves along valley fanks, obviously cut by the lowering of the topography (Adda, Adige, Salzach, Isere): Pian del Tivano, Mt. Bisbino, Mt. Tremez- MORPHOLOGIC AND TOPOGRAPHIC EVIDENCES FOR A HIGH AGE OF CAVE SySTEMS 60 TIME in KARST – 2007 CAVE AND KARST EVOLUTION IN THE ALPS AND THEIR RELATION TO PALEOCLIMATE AND PALEOTOPOGRAPHy zo, Campo dei Fiori (Southern Alps), Paganella (Dolomites). Dimensions too large with respect to the present catchment and climate Te dimensions of some conduits are far too large compared to the present catchment area, thus proving that the older catchment areas had been much larger, but are now truncated by erosion (Eisriesenwelt/Tennengebirge (fg. 6); Antre de Vénus/Vercors; Snezna jama na Raduhi/ Kamnik Alps, caves at Pokljuka and Jelovica plateaus at Julian Alps, Siebenhengste, Pian del Tivano, Campo dei Fiori/Southern Alps). Spring location vs. present base level If the position of a spring is not due to a geologic perching above an impervious layer, it has to be close to base level (see part I). However, in some cases springs did not lowered down to today’s base level. In other cases springs are obviously located far below the base level. Tis can be explained by the following hypotheses: - Some springs are perched, because the valley incision is very recent and rapid (Pis del Pesio/Marguareis). - Others are presently submerged below the base level and hidden by alluvial fll or till (Emergence du Tour/Ara-vis; Campo dei Fiori). Tey were set into their place before the base level raised and they continue to function due to the high transmissivity of the sediment fll. A specialty is given when old vertical vadose caves are suddenly stopped by the present water table, proving that the horizontal drains are at much greater depth and completely drowned. Typical vadose morphologies (spe-leothems, karren) are known in some drowned conduits (Grotta Masera, Grotta di Fiumelatte/Lake of Como; Fontaine de Vaucluse/Provence). Here, the spring location is adapted to the present base level, but the caves are proof that the base level may, in some cases, also rise. Tis is especially true for areas afected by the Messinian crisis (Bini 1994; Audra & al. 2004). CAVE SEDIMENTS SHOwING EVIDENCE OF A REMOTE ORIGIN, DIFFERENT CLIMATE AND OLD AGE (tab. 1) Old fuvial material Te presence of some caves sediments is inexplicable with the present waterpaths. Big rounded pebbles found in caves perched high up on top of clifs mean that a valley bottom had to exist at this level. Aferwards, the valleys deepened so much that they are far below such perched massifs (Salzach/Salzburg Alps; Granier/Char-treuse). Ofen, gravels found in these caves have a petrog- raphy and mineralogy that is not found in the present rocks. Tey are issued either from caprock that has disappeared a long time ago (Fontana Marella, Campo dei Fiori) or from distant catchments, as proven by fuvial pebbles (Augensteine/Northern Limestone Alps in Austria), quartz sandstones (Slovenian Alps), fuvioglacial sediments (Lake of Como). Dating of fuvial pebbles by cosmogenic nuclides from the Grotta Masera (Como), yielded a probable age comprised between 2.6 to 7.2 Ma, showing a pliocene age, or maybe older (Häuselmann unpub.; Bini & Zuccoli 2004). In the Granier system, this method yields ages comprised between 1.8 to 5.3 Ma (Hobléa & Häuselmann 2007). Record of climatic changes in subterranean sediments Ofen, the analysis of the sediments evidences climate changes, with a change from biostatic conditions, marked by the rarity of allogenic sediments, towards rhexistatic conditions, with lots of allogenic sediments. Tese sediments come from the erosion of soils in a context of climate degradation and general cooling. Tey usually are interpreted to refect the climatic change in the Pliocene, before the onset of the glaciations. Such sediments are present in most of the old cave phases, which therefore should be older than the end of the Pliocene: Grotte Vallier/Vercors; Tennengebirge (Audra 1994, 1995), Campo dei Fiori (Bini et al. 1997), Monte Bisbino (Tognini 1999, 2001). In the Dachstein-Mammuthöhle, which dates back to the Tertiary and shows a phreatic tube perched 1000 m above the Traun valley, fowstones grown during the interglacials interfn-ger with a series of debris-fow conglomerates of glacial origin (Trimmel 1992). In the Grotta di Conturines/Do-lomites (2775 m ASL), the mean annual temperatures deduced from the 18O of speleothems were between 15 and 25°, which implies that speleothemes deposited in a warmer climate within the Tertiary, probably also at a lower altitude than it is found today (Frisia et al. 1994). Furthermore, in many caves, either conduits or fowstones have been deformed by late Alpine tectonic movements: Grotta Marelli, Grotta Frassino/Campo dei Fiori (Uggeri 1992; Bini et al. 1992, 1993). Dating results prove the antiquity of cave systems Te calculated age of old speleothems are regularly above the U/T limits (700 ka, even 1.5 Ma according to the 234U/238U equilibrium (Bini et al. 1997); Tab. 1). Te pale-omagnetic measurements ofen show inverse magnetism, sometimes with multiple inversion sequences, proving of a very old age of the cave sediments (Audra 1996, 2000; Audra & Rochette 1993; Audra et al. 2002b). Te use of the new cosmonucleide method to date old quartz sediments also confrms this trend and yield ages reaching TIME in KARST – 2007 61 PHILIPPE AUDRA, ALFREDO BINI, FRANCI GABROVŠEK, PHILIPP HäUSELMANN, FABIEN HOBLéA, PIERRE-yVES JEANNIN, ... back to about 5 Ma (see the details for Siebenhengste example in this volume). RELATIONS TO AN OLD TOPOGRAPHy Te geomorphologic approach, which uses external markers of old base levels (paleovalleys, paleoshelves with associated sediments) that are well dated, ofers precious possibilities for the dating of karst systems. Sadly, correlations are almost impossible up-to-date due to the scarcity of such information. In the northern fank of the Alps, the glaciations ofen caused the remnants of an old topography to disappear. Te southern Alps, less glaciated and better studied in this context, ofer more possibilities, also thanks to the presence of guiding events like the Messinian incision and the following Pliocene marine highstand. Old erosion surfaces Te identifcation of old erosion surfaces is a precious tool in geomorphology. Large surfaces ofen top the relief and cut across very old caves that are difcult to link to an old drainage system because of their fragmented character. Te cave systems developing below those high surfaces are more recent, such as the stacked surfaces in the Vercors, of Eocene, infra-Miocene and Pliocene age (Delannoy 1997). Shelves along slopes, created by lateral corrosion of the rim of ancient depressions, have the same signifcance as perched valley bottoms. In Vercors, Pliocene caves could be associated on them, such as the Antre de Vénus and the Grotte Vallier (Delannoy 1997). In the area of Varese (Lombardy), the Oligo-Miocene surface that cuts across limestone, porphyritic rocks and granites, is dissected by the late Miocene valleys that had PALEOKARST, A MILESTONE FOR OLD KARSTS Te study of paleokarsts is a separate domain. No cave system has survived in its integrality from the periods predating the Miocene. In the Northern Limestone Alps of Austria, the possibility that caves of the highest level (Ruinenhöhlen) may be relicts of an oligocene karstif-cation has been discussed (Frisch et al. 2002). However, Paleogene paleokarsts are frequent, as evidenced by natural or artifcial removal of their flling: - In Siebenhengste, upper Cretaceous paleotubes and fractures are found in Lower Cretaceous limestone, been deepened during the Messinian (Bini et al. 1978, 1994; Cita & Corselli 1990; Finckh 1978; Finckh et al. 1984). Morphological and sedimentological evidences of pre-pliocene paleovalleys A fuvial drainage pattern of Oligo-Miocene age, incised in the relief, predated the Alpine tectonic events of the late Miocene. Te drainage originated in the internal massifs, cut through the calcareous border chains, and ended in alluvial fans in the molasse basins. In the border chains, perched paleovalleys are found more than 1500 m above the present ones (Salzburg Alps), as well as fu-vial deposits coming from siliceous rocks (Augensteine/ Northern Calcareous Alps; siliceous sands/Julian Alps (Habic 1992)), sometimes buried in caves near the valley slopes (Grotta di Monte Fenera/Piemont, Grotta Fontana Marella/Campo dei Fiori). In the northern fank of the Alps, these valleys have been destroyed by the deepening of the hydrographic network, aided by the action of the glaciers. In the South, the old valleys have been deepened by the Messinian incision and flled by Pliocene sediments (Lake of Como/Adda, Varese, Tessin, Adige, Durance). As a consequence, the horizontal karstic drains that were linked to the old valleys had been truncated by slope recession, and are presently perched (Grotta Battisti/Paganella; Grotte Vallier/ Vercors; Pian del Tivano, Monte Bisbino (Tognini 2001); Campo dei Fiori (Uggeri 1992)). Te almost generally observed input of allogenic waters coming from impermeable rocks upstream, combined with a tropical humid climate with considerable foods, explains the giant dimensions of those caves. flled with Upper Cretaceous Sandstone (Häuselmann et al. 1999). - In many places, (Switzerland, Vercors, Chartreuse) vast pockets covering a karst relief and flling up some conduits can be observed. - In Southern Alps, upper Eocene and lower Oli-gocene sediments have been found into large cavities inflled by basaltic intrusions (Covoli di Velo, Ponte di Veia/Monte Lessini) Teir age could be determined by K/Ar datings (Rossi & Zorzin 1993). In several regions (Vercors and Chartreuse, Monte Lessini), karstifcation is more or less continuous from the Eocene onwards. However, the tectonic and paleo- AGE OF ALPINE KARSTIFICATION: FROM PALEOKARSTS TO RECENT MOUNTAIN DyNAMICS 62 TIME in KARST – 2007 CAVE AND KARST EVOLUTION IN THE ALPS AND THEIR RELATION TO PALEOCLIMATE AND PALEOTOPOGRAPHy geographic changes have only lef dispersed paleokarsts. Since the Miocene on, several massifs emerged from the molasse basins, thus allowing a karstifcation that continues today. ESTIMATION OF THE FIRST ExPOSURE ACCORDING TO MOLASSE PETROGRAPHy Te main phase of karstifcation begins when suitable rocks are exposed at land surface. Since the oldest remnants of karst are ofen eroded, it is possible to calibrate the beginning of the karstifcation by the foreland sediments (mainly the Molasse), which contain limestone pebbles eroded away at the surface. However, absence of evidence is not evidence of absence: sedimentary gaps are frequent, and a karst in biostatic conditions does not spread detritic elements towards the foreland. As a general rule, the Miocene molasse registered the beginning of the last big karstifcation phase, earlier in Italy, later in Switzerland: - Upper Oligocene-Lower Miocene (30 to 20 Ma) in the Southern Molasse, based on dated fuvial sediments located in paleovalleys (Gelati et al. 1988). - Lower Miocene (20 Ma) in the molasse south of Grenoble, corresponding to the erosion of the emerged anticlines of the Vercors and Chartreuse (Delannoy 1997). - Lower Miocene (20 Ma) in the Austrian Nord-Alpine molasse, corresponding to the erosion of the Augensteine cover, which is of Upper and Middle Oligocene age (Lemke 1984; Frisch et al. 2000). - Upper Freshwater Molasse in the Eastern Swiss basin (Hörnli fan, Middle Miocene 17-11 Ma) which contains pebbles of the frst erosion of Helvetic nappes (Siebenhengste, Silberen, Speck 1953; Bürgisser 1980). DATING THE yOUNGEST PHASES AND ExTRAPOLATION Te most generally applied dating method for cave sediments is U/T. It makes it possible to date speleothems. In best cases, it allows for going back to as far as 700 ka – dating only the sediment contained within the cave and not the cave itself. Te use of paleomagnetic dating makes it possible, in some scarce cases, to push back the datable range to 2.5 Ma. Te use of cosmogenic isotopes (Granger et al. 2001) is the only recent method that opens new possibilities, having a dating range between 300 ka and 5 Ma. Another solution consists in dating lower cave phases that are supposed to be younger, and in progressively going up the phases towards the oldest cave systems, until reaching the limits of the used methods. From the calculated rate of valley deepening, one can then extrapolate the age of the uppermost phases. Of course, such an approach can only give a general idea about the age. Te lowermost phases of the Siebenhengste cave system, St. Beatus Cave and Bärenschacht, have been dated by U/T. Te following ages have been obtained: Phase 558 (youngest) began at 39 ka (max. 114 ka) and is still active today; Phase 660 was active between 135 and 114 ka; Phase 700 was active between 180 and 135 ka; and Phase 760 started before 350 ka and ended at 235 ka (Fig. 4). Tese age values indicate a general valley incision rate of 0.5 to 0.8 mm/a, with a tendency to slow down as the age gets higher. Extrapolation indicated an age of about 2.6 Ma for the oldest cave systems, at 1850 m ASL. Absolute cosmogenic dating yielded an age of 4.4 Ma for the oldest sediment, contained in the second-highest cave phase at 1800 m, showing a slower entrenchment in the older phases (Häuselmann & Granger 2005; see also this volume). Dating of the cave systems at Hölloch/Sil-beren gave maximal rates of valley incision in the range of about 1.5 to 3.5 mm/a. RELATIVE UPLIFT RATES AND EROSION VOLUMES IN FORELAND SEDIMENTS Uplif rates are generally calculated for long periods of time, taking the average of variable rhythms and integrating vast parts of the area, without taking into account block tectonics which can difer considerably from one massif to the other. In the same range, the estimated volume of the foreland basins only gives a global approach. Such results only may give a general frame for a validation. Modeling the fssion-track measurements of the Swiss Central Alps (Reuss valley) give an average uplif of 0.55 mm/a (Kohl, oral comm. 2000) comparable to calculations of recent uplif (0.5 mm/a; Labhart 1992) and consistent with the rates inferred from dating in caves. Uplif is maximal in the central parts of the mountain chains, therefore the rocks are more deeply eroded in this area. As a consequence, the oldest caves had to have disappeared from the central zones, compared to the border chains where they are better preserved due to the slower erosion. TIME in KARST – 2007 63 PHILIPPE AUDRA, ALFREDO BINI, FRANCI GABROVŠEK, PHILIPP HäUSELMANN, FABIEN HOBLéA, PIERRE-yVES JEANNIN, ... CONCLUSION Te examples mentioned above are distributed throughout the Alpine belt. Terefore, the conclusions drawn here are valid for Alpine Caves at least, but they may be applied to other cave systems also. Te main following conclusions can be drawn from the above synthesis: - In contrast to some earlier views, caves are not directly linked to glaciations. On the contrary, there is evidence that during glaciations caves are mainly flled with sediments, while they are enlarged during the inter-glacials. Te main infuence of glaciers upon cave genesis is the deepening of the base level valley, thus inducing a new cave genesis phase to be formed. - U/T datings, coupled with paleomagnetism, inferred a Lower Pleistocene to Pliocene age for several cave sediments. Fossil or radiometric datings of solidifed cave flls (sandstone, volcanic rocks) gave ages reaching back to the Upper Cretaceous. It follows that caves are not inherent to the quaternary period, but are created whenever karstifable rocks are exposed to weathering. Due to later infll, however, most explorable caves range from Miocene to present age. - we have shown that caves are related to their spring, which is controlled by a base level that usually consists of a valley bottom. So, the study of caves gives very valuable information about valley deepening processes and therefore about landscape evolution. - Caves constitute real archives, where sediments are preserved despite the openness of the system. Te study of cave sediments gives information about paleo-climates. Moreover, the combination of cave morphology and datable sediments allow to reconstruct the timing of both paleoclimatic changes as well as landscape evolu- tion between the Tertiary and today. Diferential erosion rates and valley deepenings can be retraced. Information of this density and completeness has disappeared at the surface due to the erosion of the last glacial cycles and the present vegetation. - Correlations between well-dated cave systems can signifcantly contribute to the geodynamic understanding of the Alpine belt as a whole. Te location of most cave systems at the Alpine border chains is very lucky: since they are dependent on base level (in the foreland), recharge and topography (towards the central Alps). Tey inevitably registered changes in both domains. Caves are therefore not only a tool of local importance, but may have a wide regional/interregional signifcance. - Te dating method by cosmogenic nuclides was recently applied in some French, Italian and Swiss alpine cave systems which partially contain pre-glacial fuvial deposits. Te dated sediments yielded ages ranging between 0.18 and 5 Ma, which are consistent with other approaches. Advances in modern dating techniques (cosmogenic isotopes, U/Pb in speleothems) therefore open a huge feld of investigations that will very signif-cantly contribute to the reconstruction of paleoclimates and topography evolution along the last 5, possibly 15 to 20 Ma. - Te messinian event infuenced cave genesis over the whole southern and western sides of the Alps by overdeepening valleys. However, the subsequent base level rising fooded those deep systems creating huge deep phreatic aquifers and vauclusian springs (Audra et al. 2004). 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Strat., 100, 143-148 TIME in KARST – 2007 67 COBISS: 1.01 ASPECTS OF THE EVOLUTION OF AN IMPORTANT GEO-ECOSySTEM IN THE LESSINIAN MOUNTAIN (VENETIAN PREALPS, ITALy) POGLEDI NA RAZVOJ POMEMBNEGA GEO-EKOSISTEMA V GORAH LESSINI (BENEŠKE PREDALPE, ITALIJA) Leonardo LATELLA1 & Ugo SAURO2 Abstract UDC 551.442:574.4 (234.323.4) Leonardo Latella & Ugo Sauro: Aspects of the evolution of an important geo-ecosystem in the Lessinian Mountain (Venetian Prealps, Italy) Te Grotta dell’Arena (476 V/VR), located in the Lessinian Mountain, at the elevation of 1512 m a.s.l., is a very important underground karst system. Although it is only 74 m long, several of the geological, geomorphological and environmental features of the High Lessinian underground karst are present in this cave. Te Grotta dell’Arena shares some common geological and faunistic characters with other important and well known karst systems. Tis cave has also one of the highest number of troglobitic species in all Venetian Prealps and some of them possibly originated in the pre-quaternary. From the geological point of view the cave is the expression of a contact karst, where diferent limestone types come in contact both stratigraphically and along tectonic structures. Te Grotta dell’Arena is located at the stratigraphic contact between the “Calcari del Gruppo di San Vigilio” and the “Rosso Ammonitico” and it is very close to a fault plane putting in vertical contact the two above formations with the “Biancone”, a kind of limestone closely stratifed and densely fractured, very sensible to frost weathering. It is interesting to note the presence of a good number of species of Tertiary, or more generally pre-quaternary, originate in the Grotta dell’Arena. Tis presence is possibly related to the geology of caves. In this paper the diferent kinds of underground karst systems in the Grottta dell’Arena and Lessinian Mountain, are analyzed and the relation with the cave fauna distribution are taken in consideration. Key words: karst evolution, geomorphology, biospeleology, faunistic invasions, Venetian Prealps, Italy. Izvleček UDK 551.442:574.4 (234.323.4) Leonardo Latella & Ugo Sauro: Pogledi na razvoj pomembnega geo-ekosistema v gorah Lessini (Beneške Predalpe, Italija) Jama Grotta dell’Arena (476 V/VR) v gorah Lessini, 1512 m n.m., je zelo pomemben podzemeljski kraški sistem. Čeprav je dolga le 74 m, vsebuje geološke, geomorfološke in okoljske elemente, značilne za kraško podzemlje Visokih Lessini. Grot-ta dell’Arena ima nekaj geoloških in favnističnih značilnosti skupnih z drugimi pomembnimi in znanimi kraškimi sistemi. Jama je med tistimi z največjim številom troglobiontskih vrst v vseh Beneških Predalpah, od katerih nekatere verjetno izvirajo izpred kvartarja. Z geološkega vidika predstavlja jama kontaktni kras, kjer so vzdolž stratigrafskega in tektonskega stika različni apnenci. Grotta dell’Arena je na stratigrafskem stiku med apnenci “Calcari del Gruppo di San Vigilio” in “Rosso Ammonitico” in je zelo blizu prelomne ploskve, vzdolž katere se vertikalno stikata omenjeni formaciji s formacijo “Bianco-ne”, to je vrsta drobnoplastovitega in gosto prepokanega, slabo odpornega apnenca. Zanimiva je prisotnost precejšnjega števila terciarnih oziroma splošneje predkvartarnih vrst. To je verjetno v zvezi z jamsko geologijo. V prispevku so podrobneje obravnavane različne vrste podzemskih kraških sistemov v sami jami Grottta dell’Arena kot tudi v gorah Lessini in tudi njihovi odnosi z razporeditvijo jamskega živalstva. Ključne besede: razvoj krasa, geomorfologija, biospeleologija, invazija favne, Beneške Predalpe, Italija. 1 Museo Civico di Storia Naturale di Verona. Lungadige Porta Vittoria, 9, 37129 Verona, Italy. E-mail: leonardo.latella@comune.verona.it 2 Universita degli Studi di Padova, Dipartimento di Geografa. Via del Santo 26, 35123 Padova, Italy. E-mail: ugo.sauro@unipd.it Received/Prejeto: 21.12.2006 TIME in KARST, POSTOJNA 2007, 69–75 LEONARDO LATELLA & UGO SAURO INTRODUCTION Te Grotta dell’ Arena is registered with the number 476 in the Cadastre of the Caves of Veneto Region (the cave has been surveyed by A. Pasa in 1942, and GAS USV in 1972); the karst area is ML03 (Monti Lessini 03). Te cave is 74 m long with a diference in elevation of - 22 m. It is located in the Lessinian Mountain district of Bosco Chiesanuova, in Malga Bagorno area. G.C: 11° 06’ 02’’ E 45° 39’ 56” N, elevation 1512 m a.s.l. Te Grotta dell’Arena is a signifcant kind of underground karst system in Lessinian Mountain in fact: – it is a type of speleogenetic style in the morpho-dynamic context of the High Lessinians, – several of the geological, geomorphological and environmental features of the High Lessinian underground karst are present in this cave and played a signif-cant role in karst evolution, – some of the best known karst systems in the Les-sinian Mountain (Mietto & Sauro, 2000; Rossi & Sauro, 1977), such as the Abisso della Preta, the Covolo di Cam-posilvano, the Abisso del Giacinto, the Abisso dei Lesi, the Ponte di Veja, share some common characters with the Grotta dell’Arena, – from the biospeleological point of view, this cave has one of the highest number of troglobitic species in all Venetian Prealps, – several troglobitic species are endemic for the Grotta dell’Arena or the Lessinian Mountains and some of them possibly originated in the pre-quaternary. Te Grotta dell’Arena is a large chamber, roughly elliptical in plane section, with a main diameter of about 50 m. Te roof coincides mostly with bedding planes. Te southern part of the foor is characterized by a large, asymmetrical, funnel-shaped depression, a kind of subterranean doline developed in the collapse debris. Te chamber is situated a few meters below the topographical surface; it is connected to the surface through Gastropoda Opiliones Diplopoda Orthoptera Zospeum sp. Ischyropsalis strandi Lessinosoma paolettii Troglophilus sp. Anellida Copepoda Collembola Coleoptera Marionina n.sp. Speocyclops cfr. infernus Onychiurus hauseri Orotrechus vicentinus juccii Araneae Lessinocamptus caoduroi Pseudosinella concii Orotrechus pominii Troglohyphantes sp. Moraria n. sp. Sincarida Italaphaenops dimaioi Pseudoscorpiones Elaphoidella n. sp. Bathynella sp. Lessinodytes pivai Chthonius lessiniensis Isopoda Amphipoda Laemostenus schreibersi Neobisium torrei Androniscus degener Niphargus galvagnii similis Halberrria zorzii Balkanoroncus boldorii tab. 1: List of the cave-dwelling species in the Grotta dell’Arena. Fig. 1: Te collapse depression called Arena. 70 TIME in KARST – 2007 Fig. 2: Te large chamber in the Arena cave. In the foreground the debris blocks, in the background the inner “doline”. ASPECTS OF THE EVOLUTION OF AN IMPORTANT GEO-ECOSySTEM IN THE LESSINIAN MOUNTAIN some narrow passages which start from an open collapse depression located on a slope, which resembles a Roman theatre (i.e. an “Arena”, hence the name of the cave) (Fig.1, Fig. 2). Te depression is the result of the collapse of part of the subterranean room. To understand the signifcance of this cave it is necessary to: – delineate the geological, geomorphological, and, in general, environmental characteristics of this cave, THE ENVIRONM Te Grotta dell’Arena had been previously defned not as a distinct structure, but as a window on a subterranean space, that allows us to see only some features of a karst system (Castiglioni & Sauro, 2002). In fact, the subterranean environment is a much more complex system, mostly hidden to the human perception. From the geological point of view the cave is expression of a contact karst, where diferent limestone types come in contact both stratigraphically and along tectonic structures (Capello et al. 1954; Pasa, 1954; Sauro, 1973, 1974, 2001). In particular, the limestone formations present here are: – “Calcari del Gruppo di San Vigilio” of lower-middle Jurassic, about 60 m in depth, pure both oolitic and bio-sparitic/–ruditic, or reef limestones, relatively densely fractured, – “Rosso Ammonitico”, a condensed rock unit of middle- upper Jurassic age, about 30 m in depth, made up by nodular micritic limestone very resistant to erosion, crossed by widely spaced fractures, – “Biancone”, a chalk type unit, from the lower and middle Cretaceous, 100-200 meters in depth, made up by whitish marly limestone closely stratifed and densely fractured, very sensible to frost weathering. Te Scaglia Rossa formation of the upper Cretaceous, and the Eocene limestone, which lie above the Bi-ancone in the western and southern part of the plateau are not present in the studied area because they have been completely eroded. Below the “Calcari del Gruppo di San Vigilio” there is the formation “Calcari Grigi di Noriglio”, of lower Jurassic, which is about 300 m in depth and outcrops in the slopes of the main valleys, a kind of fuvio-karstic canyons. Te Grotta dell’Arena is located at the stratigraphic contact between the “Calcari del Gruppo di San Vigilio” and the “Rosso Ammonitico” and it is very close to a fault plane putting in vertical contact the two above formations with the Biancone (Fig. 3). Te cover rocks of the cave are made mostly by the massive beds of lower Rosso – reconstruct the framework of the spatial and temporal evolution of the High Lessini karst, – taking into account the climatic and environmental changes of the external environment surrounding the cave that occurred during the Pleistocene. – analyse the phyilogeographical and taxonomical afnities of the troglobitic elements of its fauna. CONTExT Ammonitico, whereas the inner cave is mostly developed inside the Calcari del Gruppo di San Vigilio. At the topographical surface, the line of the normal fault runs along Fig. 3: Sketches of the Arena cave system: I – Plan of the system; the grey corresponds to the biancone rock unit. II – vertical model of the karst system. Legend: 1) biancone formation, 2) Rosso Ammonitico, 3) Calcari del Gruppo di San vigilio and Calcari Grigi, 4) debris pipe in the cave, 5) bedding plane karst zone at the contact Rosso Ammonitico- Calcari del Gruppo di San vigilio, 6) fault plane karst zone, a) at the biancone side, b) at the Rosso Ammonitico side, 7) lateral fow inside and from the biancone aquifer, 8) vertical karst fow. TIME in KARST – 2007 71 LEONARDO LATELLA & UGO SAURO a small valley, a few meters to the east of the cave; the displacement of the fault is about 100 m. From the geomorphological point of view, Biancone is dissected by a network of dry valleys, whereas Rosso Ammonitico generates a rocky landscape with large fat karren separated by corridors, or rock cities of large blocks. From the hydrological viewpoint, the water circulates difusely inside the dense network of discontinuities SPATIAL AND TEMPORAL EVO It is easy to understand that the Grotta dell’Arena results of diferent spatial and temporal processes which occurred as a consequence of several predisposing factors. In fact, the cave is at the same time, an example of litho-logical contact karst, of intra-stratal karst, of fault zone karst and of a subterranean hydrological transition from a dispersed and sub-horizontal water fow to a more concentrated and sub-vertical one. Te Grotta dell’Arena system is fed by a lateral water fow coming from the Biancone aquifer and crossing the fault zone, facilitated by the westward dipping of the strata. Te speleogenesis of the cave has taken place in the lithological, tectonic and hydrological transition zone. Each cave we visit represents a moment of a long history, it is like the picture of a movie. Surely the present aspect of this cave and of its collapsed part are the result of relatively recent processes, occurred mostly during the middle and upper Pleistocene. But the karst system of which the cave is expression has surely begun to develop much earlier. Some caves, located in middle of the Lessinian plateau and in the Berici hills, are the result of the re-activation of old paleokarstic nets developed during the Paleo-gene (Rossi & Zorzin, 1989, 1991; Dal Molin et al. 2000); other caves with fllings from the early middle Pleistocene developed mostly during the lower Pleistocene. Te Grotta dell’Arena chamber seems to be related with the second group. Te fault to the east side of the cave is a paleotec-tonic feature of Jurassic age, reactivated during the Cretaceous and later by the Alpine orogenesis during the Paleogene and the Neogene. Te area where the cave is located probably emerged from the sea during the Oli-gocene, as the southern part of the Lessinian plateau. Te erosion of the Eocene rock unit occurred during late Pa-leogene and early Neogene. Te Scaglia Rossa formation was probably eroded during middle to late Neogene. At the beginning of the quaternary these two formations of the Biancone unit; the preferential fows is sub-parallel to the topographical surface and occurs mostly below the dry valley bottoms, but is also infuenced by the structural setting; vertical losses occur along the fault and fracture zones. In contrast, water circulation is more concentrated and mostly vertical in the Rosso Ammonitico. TION OF THE KARST SySTEM disappeared completely in the area (remnants of Scaglia are still present in the western High Lessinian). A model showing the sequence of landscapes developed in the diferent rocks by the erosion can be created, based on present-day landscapes of other parts of the Lessini Mountains, where the eroded geological formations are still present. Tus in the southwestern Lessinian Mountain (High Valpolicella) there is an active hydro-graphic network with gorges entrenched in both Eocene Limestones and in the Scaglia Rossa. Here, the early morphogenesis, afer the emersion and the uplif, has been mostly of the fuvial type, marked by the development of a network of valleys strongly controlled by the tectonic structure. So, a valley developed along the fault line. Following the incision of the Scaglia Rossa, the karst process begun to afect the fault zone. But, it is especially afer the erosion of the Scaglia Rossa that the aquifer hosted in the Biancone started to feed a new underground karst system located near to the fault zone of which the Grotta dell’Arena is the present day expression. From this simple model it is possible to infer that the evolution of the underground karst system started since Neogene, probably since middle- upper- Miocene. Te transition from the fuvial environment to the karst environment has been accompanied by the development of a fuviokarstic milieu in the Biancone. In this milieu, which is still present, there is not surface runof except during exceptional events, but there is a difuse circulation inside the rock, for some aspects similar to that occurring below the river beds, inside the alluvial deposits (Fig. 4). 72 TIME in KARST – 2007 ASPECTS OF THE EVOLUTION OF AN IMPORTANT GEO-ECOSySTEM IN THE LESSINIAN MOUNTAIN Fig. 4: Sketch of the morphological evolution of the alti Lessini according with the erosional stages reached by the relief (progressive erosion of the rock units). THE CLIMATE AND ENVIRONMENTAL CHANGES DURING THE PLEISTOCENE Te Lessinian Mountain plateau was afected by the climatic and environmental changes of the Pleistocene. In the cave area there is no evidence of past development of local glaciers (the nearest local glacier was more than 1 km to the northwest). However, traces of strong perigla-cial processes, such as remnants of small rock glaciers, nivation niches, etc. are present (Sauro, 2002). During the last würm sporadic permafrost was present in the area. Te material resulting from the collapse of the Are- na depression has been afected by cryoclastic processes, as shown by a large solifuction lobe located to the north side of the same hollow. Te climate and enviromental change occurred in the Pleistocene, afected the colonization of the subterranean environment by some actual troglobitic species and shaped the distribution of the species that colonized this environments before the Pleistocene. THE CAVE FAUNA AT PRESENT Te cave fauna of the Grotta dell’Arena is characterized ent times. Ancient elements of this fauna colonized the by the presence of high number of troglobitic and en- subterranean environments before the Pleistocene, and demic species (Caoduro & Rufo, 1998). Colonization of other species invaded the cave in diferent periods along the cave by the troglobitic elements occurred in difer- the quaternary. TIME in KARST – 2007 73 LEONARDO LATELLA & UGO SAURO Today, this cave has a high number of cave-adapted animals. Of the 43 taxa known for the cave, 24 could be considered eucavernicolous species (sensu Rufo, 1955: eutroglophiles+troglobites). Te specialization index Te subterranean karst of the alti Lessini is much more spatially developed than what is perceived by a speleologist. It consists not only in large pits and chambers but in a network of smaller cavities and fssures. In the two horizontal dimensions it is a kind of net, even if aniso-tropic, better developed along the fault zones and some bedding planes. In the vertical dimension the anisotropy is even greater, and the thickness overpasses one thousand of meters. In the time dimension, this karst network has evolved progressively, even with diferent speeds infu-enced by the changes of the morphostructural setting and of the external environment. Te karst morphogenesis occurred as result of the co-occurrence of various favourable conditions. Te hydro-geological condition of the alluvial deposits of the water courses of the early erosional stage, during middle Neogene, are no present here nowadays, but there are situation for some aspects similar both below the valley bottoms of the Biancone and in the difuse net of karst fssures developed inside this rock unit. Tis difuse aquifer is in contact with the more typical karst aquifer of the limestone of the Jurassic rock units. Likewise, some of the larger karst pockets developed in the Eocene limestone, may have had some connections with the karst cavities in the Scaglia, and, along the main fault structures or volcanic structures, also with the karst voids in the Jurassic rock units. we are grateful to Sandro Rufo for the helpful discussions and for the reading of the manuscript. we also thanks Augusto Vigna Taglianti for the informations re- (eutroglobites/ eucavernicolous), has a value of 0.91, this means that 91% of the cave species in the Grotta dell’Arena are troglobionts. Here, sudden and sharp changes of conditions of the underground environments have not occurred during the late Neogene and the Pleistocene. Even the abrupt climatic changes of the Pleistocene have had a limited infuence on the underground environments, according with the large thickness reached by it before the end of Neogene. It is interesting to note the presence of a good number of species of Terziary, or generally pre-quaternary, origin in the Grotta dell’Arena. Te most important relict species are: balkanoroncus boldorii (Beier, 1931), Lessino-camptus caoduroi Stoch, 1997, Italaphaenops dimaioi Ghi-dini, 1964 and Lessinodytes pivai Vigna Taglianti e Sciaky, 1988 (Casale & Vigna Taglianti, 1975; Vigna Taglianti & Sciaky, 1988; Gardini, 1991; Galassi pers. com.). Te presence and distribution of these species inside the caves of Lessinia (particularly the terrestrial species) has been usually related to certain environmental characteristic like temperature, humidity, air circulation etc. However, on the basis of the actual knowledge (Latella & Verdari, 2006), it appears that all these species are present in caves with a large range of temperatures, altitude and morphology. All these caves are developed inside, or in contact with, the Biancone or Scaglia (Cretaceous limestone) formations. It is likely that the geomorphology of the cave plays an important role not only in shaping the historical distribution, but also the actual presence, of cave animals in Lessinian area. garding Trechinae, Diana Galassi for the informations on Copepoda and Beatrice Sambugar for the Anellida. Tanks to Cristina Bruno for the linguistic review. FINAL REMARKS ACKNOwLEDGMENTS 74 TIME in KARST – 2007 ASPECTS OF THE EVOLUTION OF AN IMPORTANT GEO-ECOSySTEM IN THE LESSINIAN MOUNTAIN REFERENCES Caoduro, G. & Rufo, S., 1998: La Grotta dell’Arena, un biotopo di eccezionale interesse negli alti Lessini. La Lessinia ieri oggi domani: quaderno culturale 1998, 39-44. Casale, A. & Vigna TAglianti, A., 1976: Note su Itala-phaenops dimaioi Ghidini (Coleoptera, Carabide). Bollettino del Museo Civico di Storia Naturale di Verona, 2 (1975): 293-314. Capello, C.F., Nangeroni, G., Pasa, A., Lippi Boncampi, C., Antonelli, C. & Malesani, E., 1954: Les phéno-ménes karstiques et l’hydrologie souterraine dans certaines régions de l’ltalie. Assoc. Intern. Hydrol., vol. 37, n. 2, pp. 408-437, fgg. 5, Paris. Castiglioni, B. & Sauro, U., 2002: Paesaggi e geosistemi carsici: proposte metodologiche per una didattica dell’ambiente. In: Varotto M. & Zunica M. (a cura di) – Scritti in ricordo di Giovanna Brunetta. Dipar-timento di Geografa “G. Morandini”, Universita di Padova, 51-67. Dal Molin, L., Mietto, P. & Sauro, U., 2000: Considera-zioni sul paleocarsismo terziario dei Monti Berici: la Grotta della Guerra a Lumignano (Longare - Vicen-za). Natura Vicentina 4, 33-48 (ISSN 1591-3791). Gardini, G., 1991: Pseudoscorpioni cavernicoli del Vene-to (Arachnida). (Pseudoscorpioni d’Italia xIx). Bollettino del Museo Civico di Storia Naturale di Verona, 15 (1988): 167-214. Latella, L. & Verdari, N., 2006: Biodiversity and bioge-ography of Italian Alps and Prealps cave fauna. Abstracts 18th International Symposium of Biospeleol-ogy, Cluj-Napoca, Romania, 10-5 July 2006: 9-10. Mietto, P. & Sauro, U., (eds), 2000: Le Grotte del Veneto: paesaggi carsici e grotte del Veneto. Second edition, Regione del Veneto-La Grafca Editrice, 480 pp. Pasa, A., 1954: Carsismo ed idrografa carsica del Gruppo del Monte Baldo e dei Lessini Veronesi. C.N.R., Cen-tro Studi per la Geografa Fisica, Bologna, Ricerche sulla morfologia e idrografa carsica, n. 5, 150 pp. Rossi, G. & Sauro, U., 1977: L’Abisso di Lesi: analisi mor-fologica e ipotesi genetiche. Le Grotte d’Italia, (4) 6 1976): 73-100. Rossi, G. & Zorzin, R., 1989: Fenomeni paleocarsici nei Lessinian Mountain Centrali Veronesi. La Lessinia ieri oggi domani: quaderno culturale 1989, 47-54. Rossi, G. & Zorzin, R., 1991: Nuovi dati sui fenomeni pa-leocarsici dei Covoli di Velo (M.ti Lessini Verona). Atti xVI Congr. Naz. di Speleologia, Udine, 169-174. Rufo, S., 1955: Le attuali conoscenze sulla fauna caver-nicola della regione pugliese. Memorie di Biogeo-grafa adriatica, 3: 1-143. Sauro U., 2002: quando in Lessinia c’era il grande gelo. quaderno Culturale - La Lessinia ieri oggi domani - 2002, 85-94. Sauro, U., 1973: Il Paesaggio degli alti Lessini. Studio geo-morfologico. Museo Civ. di St. Nat. di Verona, Mem. f. s., 6, 161 pp. Sauro, U., 1974: Aspetti dell’evoluzione carsica legata a particolari condizioni litologiche e tettoniche negli Alti Lessini. Boll. Soc. Geol. It., 93, 945-969. Sauro, U., 2001: Aspects of contact karst in the Venetian Fore-Alps. Acta Carsologica, Ljubljana, 30(2), 89-102, 2001. Vigna Taglianti & Sciaky R., 1988: Il genere Lessinodytes Vigna Taglianti, 1982 (Coleoptera, Carabidae, Tre-chinae). Fragmenta Entomologica, 20 (2): 159-180. TIME in KARST – 2007 75 COBISS: 1.01 HISTORICAL BIOGEOGRAPHy OF SUBTERRANEAN BEETLES – “PLATO’S CAVE” OR SCIENTIFIC EVIDENCE? ZGODOVINSKA BIOGEOGRAFIJA PODZEMELJSKIH HROŠČEV – »PLATONOVA JAMA« ALI ZNANSTVENI DOKAZ? Oana Teodora MOLDOVAN1 & Géza RAJKA1 Abstract UDC 595.76:551.44 591.542 Oana Teodora Moldovan & Géza Rajka: Historical bioge-ography of subterranean beetles – “Plato’s cave” or scientifc evidence? Te last two decades were particularly prolifc in historical bio-geography because of new information introduced from other sciences, such as paleogeography, by the development of quantitative methods and by molecular phylogeny. Subterranean beetles represent an excellent object of study for historical bio-geography because they are the group with the best representation in the subterranean domain. In addition, species have reduced mobility, display diferent degrees of adaptations to life in caves and many specialists work on this group. Tree processes have shaped the present distribution of the tribe Leptodirini (Coleoptera Cholevinae) in the world: dispersal, vicariance, and extinction. Terefore, three successive stages can be established in the space-time evolution of Leptodirini: (1) dispersal from a center of origin in the present area(s); (2) dispersal, extinction and vicariance among the present area(s); and (3) colonization and speciation in the subterranean domain. Te Romanian Leptodirini, especially those from western Carpathians is examined with respect to these processes. Teir pattern of distribution in diferent massifs and at diferent altitudes is discussed, with possible explanations from a historical biogeo-graphic point of view. Key words: Historical biogeography, cave beetles, Leptodirini, Romania. Izvleček UDK 595.76:551.44 591.542 Oana Teodora Moldovan & Géza Rajka: Zgodovinska biogeo-grafja podzemeljskih hroščev – jama »Platonova« ali znanstveni dokaz? Zadnji dve desetletji sta bili za historično biogeografjo še posebej bogati, predvsem zaradi številnih novih informacij in dognanj paleogeografje, razvoja kvantitativnih metod ter molekularne flogenije. Podzemeljski hrošči so odličen model za proučevanje historične biogeografje, saj spadajo v tisto skupino organizmov, ki je v podzemlju najpogosteje zastopana. Hrošči iz podzemlja imajo zmanjšano mobilnost, razvili so številne načine prilagoditev na življenje v tem habitatu. S to skupino organizmov se ukvarjajo številni raziskovalci po svetu. Na trenutno razširjenost vrst rodu Leptodirini (Coleoptera Cholevinae) so vplivali trije procesi: disperzija, vikarianca in izumiranje. V prostorsko-časovnem razvoju Leptodirinov se lahko pojavijo tri zaporedne faze: (1) razširjanje iz izvornega mesta na sedanje/a poročje/a, (2) razširjanje, izumrtje in vi-karianca med sedanjimi območji, ter (3) kolonizacija in spe-ciacija v podzemeljskih habitatih. S tega vidika smo proučevali Leptodirine iz Romunije, s poudarkom na vrstah iz zahodnih Karpatov. V prispevku je opisan vzorec razširjenosti hroščev v različnih gorskih predelih ter na različnih nadmorskih višinah. V razpravo smo z vidika historične biogeografje vključili tudi verjetno interpretacijo. Ključne besede: zgodovinska biogeografja, speleobiologija, Leptodirini, Romunija. 1 Institutul de Speologie “Emil Racovitza”, Cluj Department, Clinicilor 5, P.O.Box 58, 400006 Cluj-Napoca, Romania; e-mail: oanamol@hasdeu.ubbcluj.ro Received/Prejeto: 04.12.2006 TIME in KARST, POSTOJNA 2007, 77–86 OANA TEODORA MOLDOVAN & GéZA RAJKA INTRODUCTION Te Greek philosopher, Plato (428-348 BC), in his book, Te Republic, tells about Socrates teaching lessons of wisdom. One of these is about human beings kept in a cave with one source of artifcial light coming from behind. Te idea of the allegory is that we might have a wrong perception about what is reality, or, that most people live in a world of ignorance because they rely only on their narrow experiences and rely on their own truths. Another possible interpretation of Plato’s allegory is that we might be wrong if we consider concepts and perceived objects on the same level. Historical biogeog-raphy is a science based on concepts and suppositions and there is no direct evidence available in the attempt to build credible scenarios about past and present animal distributions. However, development of this science on circumstantial evidence ensures better understanding of the objects under study. Biogeography studies geographic distribution of organisms. Tis simple defnition describes an extremely complex science. Geology, geography and various branches of biology defne a discipline that is continuously developing. Te Swiss botanist de Candolle (1820) was the frst to speak about ecological and historical bio-geography as separate branches of biogeography. Tey difer mainly in what concerns spatial and temporal scales. Historical biogeography reports on evolutionary processes over millions of years, mostly on a global scale (Crisci 2001). Pleistocene glaciations are sometimes collectively considered a separate or intermediary branch between historical and ecological biogeography. Te last two decades were particularly prolifc in papers on historical biogeography due largely to new information introduced from other sciences, such as paleogeography, by the development of quantitative methods (Morrone & Crisci 1995) and by the development of molecular phy-logeny. Morrone & Crisci (1995) and Crisci (2001) defne the biogeographic processes that modify the spatial distribution of taxa and recognize nine basic approaches to historical biogeography: (1) Identifcation of the centers of origin, or the existence of “Eden” where diferent lineages of all living beings moved from by dispersal to the present areas; (2) Panbiogeography, which plots the distribution of diferent taxa on maps, connecting their distribution areas together with lines; (3) Phylogenetic biogeography and (4) Cladistic biogeography, both assuming correspondence between taxonomic relationships and area relationships; (5) Parsimony analysis of endemicity that classifes areas by their shared taxa; (6) Event-based methods; (7) Phylogeography; (8) Ancestral areas; and (9) Experimental biogeography. Te evolution of subterranean animals is a process that can be presumed but not directly proven. Te origin, migration and colonization of the subterranean realm can be explained by a multitude of arguments and indirect evidences which support or falsify the proposed hypotheses. Te role of historical biogeography is to explain the way subterranean animals gain their present distribution, using available data from biology and other sciences. Trough this process, we can gain a new insight into the mechanisms of colonization that have afected some of the extreme areas or habitats which exist in the subterranean domain. Chronologically, the history of a taxonomic group (like the beetles), or of a phyletic lineage must begin with its origin. To understand present distribution patterns and why some areas were colonized and others were not, we must frst establish temporal and spatial reference points. SUBTERRANEAN COLEOPTERA ExAMPLES IN HISTORICAL BIOGEOGRAPHy Tere are several reasons why subterranean beetles represent an excellent object of study for historical biogeog-raphy: 1. Tey are the best represented group in the subterranean karst environment or domain, with many species inhabiting caves and the mesovoid shallow substratum (also called MSS, see Juberthie et al. 1980); 2. Most species are terrestrial and therefore have reduced mobility; and while they are not limited to limestone/karst areas, most taxa inhabit caves; 3. A group whose representatives that display different degrees of adaptations to life in caves. Some of the lines have endogean and hypogean species, of which the last is more or less adapted to subterranean environment. 4. Tey are a well known group, with many specialists studying various aspects of their biology, including taxonomy, adaptations, behavior and molecular phylogeny. Two families encompass most of the world’s subterranean beetles, the Trechinae (predator Carabidae) and the Cholevinae (detritivorous and saprophagous Leiodi-dae). Our example is from one of the best represented subterranean tribe of Cholevinae, the Leptodirini (for- 78 TIME in KARST – 2007 HISTORICAL BIOGEOGRAPHy OF SUBTERRANEAN BEETLES – “PLATO’S CAVE” OR SCIENTIFIC EVIDENCE? mer Bathysciini). According to ecological biogeography, Leptodirini are presently distributed in caves, MSS, and other dark and humid habitats, such as litter and moss, mostly in the Palearctic region. For historical biogeography the life of the tribe Lep-todirini begins in the Paleozoic. Te present distribution of Leptodirini can only be explained through wegener’s theory of continent drif and the dispersal, vicariance and extinction processes. Dispersal was the main concept in biogeography before wegener and it explains the area of a population by the mechanisms of migration and crossing over geographical barriers. Extinction means the death of local populations, species or even supraspecifc taxa, and its role in biogeography has not always been recognized. Vicariance represents the splitting of an ancestral population into several subpopulations, which will later evolve into species through isolation. Tese three processes have shaped the present distribution of Leptodiri-ni, and three successive stages can be established in the space-time evolution of this group: I. Dispersal from a center of origin in the present area(s); II. Dispersal, vicariance, and extinction among the present area(s); III. Colonization and speciation in the subterranean domain. Fig. 1: historical biogeography of ancestors of Leptodirini, which migrated from Gondwana (a) together with continental microplates (b), up to Eurasia (c), from there dispersed west to the mediterranean region (d) (modifed from Giachino et al. 1998): areas covered by Leptodirini are represented by grey ellipses. I. According to Giachino & Vailati (1998) the ancestral family of Oricatopidae inhabited the southern part of the Gondwana supercontinent (Fig. 1). Descending from this family, ancestors of Leptodirini and other tribes migrated at the end of Paleozoic to the what is now the south of Eurasia on the microplates that broke of from Gondwana. Tus, Eurasia was colonized by the ancestors of Leptodirini 120–150 Ma ago. More recently, 50-65 Ma ago, the group dispersed from northeast, through the central south of Asia, up to eastern Eurasia and then toward the west, along the Mediterranean basin. Epigean individuals successively migrated at the surface, and they were probably pre-adapted to low, constant temperatures and high humidity. Jeannel & Leleup (1952) provide excellent examples for preadaptated ancestors of pselaphid beetles, afer studying high altitude (2000-2900 m) beetles on Mount Kivu (Congo). At this level, the species are exclusively humic inhabitants, deepened at few centimeters in humus, where proper conditions, such as constant temperature (10°C) and high humidity, are fulflled. Tey also described a species with similar adaptations to those inhabiting caves, and also found deeper, under the humus. II. Te second phase of evolution of the group probably happened before the Miocene, and possibly in the late Oligocene. Te dispersal of beetles was from Asia, along the Miocene Alpine chain, whose remains are the Cantabric chain, Pyrenees, Central French Massif, Alps, Dinarides, Balkans, Pindus chain, Peloponnesus chain, and Pontic Alps. Aferwards, some species colonized the Apennines, Jura, Carpathians, Rodops, Taurus, Caucasus and Mediterranean inlands (Giachino et al. 1998). Due to major geological and geographical transformations of the landscape, extinction and vicari-ance alternated during the next periods. A large and continuous distribution area of epigean and probably endogean ancestors of Leptodirini that migrated from east was then fragmented, even before the colonization of the subterranean domain. New paleogeographic data about the evolution of the Paratethys from Late Eocene to Pliocene has been recently published (Steininger & Rögl 1985, Popov et al. 2004), and it appears that paleoconfguration of the Paratethys shaped the distribution of Leptodirini in Europe. TIME in KARST – 2007 79 OANA TEODORA MOLDOVAN & GéZA RAJKA III. Te third stage is represented by colonization and speciation in the subterranean domain. Two scenarios were proposed for explaining the mechanisms of underground colonization by epigean and endogean representatives of various groups of fauna (Holsinger 2000, 2005): (1) Te climatic-relict model, in which preadapted or adapted animals were “forced” or constrained by climatic fuctuations to fnd refuge in caves. Te best documented are the Pleistocene glacial-interglacial periods. Eventually, surface ancestors of these successful colonizers went extinct. Tis model not only fts temperate climate regions, but also any region that has previously sustained drastic climatic changes. (2) Te adaptive-shif model is mostly applied to lava tubes and tropical karst regions. Proposed by How-arth (1981), it explains the active colonization by pre-adapted ancestors and exploitation of new and empty niches to avoid competition. In this model, adaptation to the new environment does not depends on physical isolation from surface ancestors, as is necessaryin the previous model. In our opinion, colonization of the subterranean domain is an active process, as is the case of empty niche colonization anywhere on Earth. Climatic changes have made important contributions in shaping the distribution areas and breaking of gene fow with surface relatives. Climatic changes can also interrupt energetic fow between surface and subterranean environments, leading to extinction of populations on one or both sides. Tis apparently happened in parts of the world directly affected by or covered by Pleistocene glaciers. Bellés (1991) enumerates three reasons for cave colonization: 1. Survival, when external stress determines animals to fnd refugee in caves; 2. Opportunism or colonization of an empty space; 3. Convenience or escaping surface competition which uses the same trophic resources. Peck (1980), Vailati (1988) and Juberthie (1988) proposed scenarios for cave colonization by beetles in the family Leiodidae from North America and Europe. Juberthie’s model uses complex data from studies on the ecology, ethology, genetics, tectonics, paleoclimate and geology of species from the Speonomus delarouzeei complex (Leiodidae: Cholevinae: Leptodirini). Tese species inhabit the MSS and caves on Mount Canigou in the French Pyrenees. Te history of this complex begins with the frst glacial period (2.3 Ma), when high altitude species separated from those at low altitude in the Mediterranean climate. Te two species, S. brucki and respectively S. delarouzeei are characterized by different mating behavior, by reproductive isolation and by genetic distance. Tere are also diferences in fecundity and egg development speed at diferent temperatures. Tus, depression of temperature during glaciation selected cold resistant genotypes at 1000 m altitude, where annual mean temperature in caves is today 8-10°C, while S. delarouzeei remained at low altitude at temperatures of 14°C. During glacial periods, the forest belt displaced several hundred meters downslope on Mount Canigou and was replaced by steppe vegetation. Cracks, voids, MSS and even caves formed during these periods, while flling happened during interglacial periods. At altitudes between 500 and 1000 m two other species, S. emiliae and S. charlottae, inhabit subterranean habitats. S. emil-iae lives in the MSS at 720 m altitude and is presumed to be the ancestor of S. brucki, and probably populated caves and MSS at diferent altitudes. S. emiliae migrated together with the forests during glacial periods through cracks and MSS down to the present altitude. Te isolation of this species and S. brucki, which formed a population at high altitude, happened by inflling of the MSS and related cracks and crevices. Similar mechanism acted in a previous period for separating S. charlottae from ancestors of S. brucki. By comparing cuticular hydrocarbons (pheromones that act at short distance or by contact) of species of the S. delarouzeei complex, Moldovan (1997) and Moldovan et al. (2003) obtained a diferent composition in the mountain species S. brucki and the Mediterranean species, S. delarouzeei: shorter chains in the frst one and longer in the second. S. emiliae, at an intermediate altitude, displays a mixture of short and long molecules in the cu-ticular hydrocarbon cocktail. Te result can be explained by temperature infuence on cuticular hydrocarbon composition. A small variation in temperature can change hydrocarbon composition even from the frst generation (Toolson et al. 1990). For subterranean beetles, adaptation to a new climate can rapidly change the pheromone composition, thus representing an important mechanism of isolation that acts prior to mating. Climate changed the distribution of populations on the slope of Canigou Mountain through migrations of ancestral populations. Consequently, composition in cuticular hydrocarbons changed and preceded genetic mutations. Hydrocarbon changes become stable if climate is maintained over long periods of time, eventually causing isolation of populations and genetic mutations. Terefore, speciation of subterranean inhabitants can occur without the existence of physical barriers as proposed in Juberthie’s model. 80 TIME in KARST – 2007 HISTORICAL BIOGEOGRAPHy OF SUBTERRANEAN BEETLES – “PLATO’S CAVE” OR SCIENTIFIC EVIDENCE? REFERENCE MARKS IN HISTORICAL BIOGEOGRAPHy OF ROMANIAN LEPTODIRINI with its geographic position in eastern Europe, Romania is very rich in subterranean fauna for a non-Mediterranean country. Tis is due to the special features defned as follows (Moldovan et al. 2005): 1. Te geographic position of the country, with climatic and fauna infuences from various regions; 2. Te reduced total surface of limestone; karst areas are distributed along the Carpathians and in Dobrogea, covering only 2% of the surface of Romania (Onac & Cocean 1996); 3. Te high number of caves/surface units; even if the covered surface is so small, the speleological potential is high, with almost 12,000 caves discovered prior to 1989 (Goran 1989); 4. Te distribution of caves at low altitude, with 27% of karstic rocks at altitudes below 500 m and 47% up to 1000 m a.s.l. (Bleahu & Rusu 1965); 5. Te patchy distribution of limestone, with small outcrops scattered especially in western and southern regions. From an ecological point of view Romanian karst forms small continental islands between non-karstic rocks, which act as natural barriers to migration of subterranean organisms. Each area represents an island to its inhabitants, which in turn leads to isolation and promotes evolution and formation of new species. Te discovery of the MSS has added new insights into the availability of subterranean habitats, but it can explain only short distance migrations between geographically close areas. Fig. 2: Genera of Leptodirini distributed in the Western and Southern Carpathians of Romania. Te above-mentioned features explain the fragmentation of the initial distribution area of surface ancestors of cave animals and speciation processes. It also explains the high number of genera and species for a country at 45° northern latitude. Other countries at the same latitude are much poorer in species (e.g., Austria - 2 species, Switzerland – 1 species), even though their limestone areas in these countries are larger. Romanian Leptodirini is represented by 8 genera and 6 subgenera with 50 species and 46 subspecies, composed of 1 epigean, 10 endogean and 85 strictly caver-nicolous taxa. Concerning the distribution of subterranean beetles, the Romanian Carpathians can be divided in three geographical units: western, Southern and Eastern Carpathians. Te last unit has few caves and no representatives of Leptodirini. More than half of the karst surface of the country belongs to the western unit (the Apuseni Mountains). Tis region also has the highest degree of speciation with 63 taxa in the genera drimeotus, Pholeuon and Protopholeuon. Tese species inhabit caves and the MSS. More genera but fewer species are found in the Southern Carpathians: the epigean monospecifc mehadiella, and the 34 taxa of subterranean banatiola, Sophrochaeta, Closania and tismanella (Fig. 2). Te origin of Romanian Leptodirini is strictly linked to dispersal of ancestral lineages that inhabited the Alpine Miocene chain and to a Paratethys evolution. Jean-nel (1924, 1931) and Decu & Negrea (1969) suggested a Dinaric origin for the Romanian Leptodirini, based on morphological similarities, and especially on features of the male geni-talia. Tis theory explains the colonization of the Apuseni Mountains through the Bohemian massif, and direct colonization of the Southern Carpathians from the Dinarides. Diferences between the two phy-letic lineages (western drimeotus and Southern Sophrochaeta) justi-fed this hypothesis, which is also supported by new paleogeographic data (Steininger & Rögl 1985, Popov et al. 2004). when these diferent waves of colonization occurred is questionable, because direct Dina-rides-Carpathians connection is either very old or too recent. Migration of Asian ancestors of Cholevinae was not possible until the Upper Oligocene-Early Miocene, when a Dinarian-Pelagonian- TIME in KARST – 2007 81 OANA TEODORA MOLDOVAN & GéZA RAJKA Anatolian landmass was formed, and linked to the rest of Europe by the recurring Slovenian corridor. Tis was the frst connection between the Dinarides and Carpathians and lasted until Lower Badenian (16 Ma) when the Central Paratethys was fooded. It provided the possibility of populating the Southern Carpathians by Dinaric lineages, which was also at the time of Carpathian system development (Fig. 3). In Upper Ottnangian (17-18 Ma), the basin of the Paratethys was profoundly altered and a connection between the Alps and the Carpathians was established. Te frst Alps-Bohemian Massif-Carpathians connection provided conditions for the Apuseni Mountains colonization through the Bohemian massif. In Lower Badenian (15-16 Ma) a major transgression temporarily interrupted the connection between the Carpathians and the Bohemian Massif. Later, in Middle Badenian (14-15 Ma), the connection was defnitively established. A connection between the Dinarides and the Carpathians was also established during the Messinian crisis (5-6 Ma) but cannot explain the processes of Southern Carpathian colonization and speciation. Subterranean beetles are less mobile, even if they can migrate over relatively short distances through the non-calcareous MSS. Supposing that epigean and edaphobiont forms migrated and colonized this region, it is improbable to admit that adaptation to deep subterranean habitats and speciation could have occurred in such a short period of time. In conclusion, the Dinaric origin of subterranean Romanian beetles can be explained by a frst migration wave of a Dinaride lineage over the Southern Carpathians and a later one through the Bohemian massif of a lineage that colonized the Apuseni Mountains. Both lineages are morphologically linked to Dinaric species, with some peculiar features in the drimeotus western lineage. An additional migration from the southwest during the Messinian crisis also could have been possible but only for species less adapted to caves. Te next step in the evolution of the Apuseni beetles was subterranean domain colonization. Te processes and mechanisms that could have driven the subterranean colonization were presented in the previous section. we also analyzed spatial distribution of species and subspecies of the Drimeotus phyletic lineage to obtain information which can be corroborated with available taxonomic and molecular data. Romanian Leptodirini ofer good material for studying speciation and vicari-ance processes, because it involves insular distribution, which is diferent from the large, continuous limestone surfaces such as those of the Dinarides and the Pyrenees, where speciation has occurred in the absence of geographical barriers. Te Apuseni Mountains are inhabited by three genera (Protopholeuon, Pholeuon and drimeo-tus), belonging to the drimeotus phyletic lineage (Fig. 4). Fig. 3: Evolution of the Paratethys in Upper Ottnangian (a), middle badenian (b) and Lower badenian (c) (Cluj is located in North-Western Romania) (simplifed afer Rögl & Steininger 1984): grey – marine realms, dark grey – evaporitic basins, light grey – important areas with fuvio-terrestrial sedimentation and/or lignite formation, white – continental realms, ? - basins narrowed post-sedimentation by tectonic processes. Te lineage is monophyletic (Bucur 2003) and suggests a common ancestor, probably epigean. Protopholeuon, which is monospecifc, inhabits the Metaliferi Mountains, while the other two genera have larger distribution. Most species of Pholeuon and drimeotus occur in the Pa-durea Craiului and the Bihor mountains. Pholeuon also 82 TIME in KARST – 2007 HISTORICAL BIOGEOGRAPHy OF SUBTERRANEAN BEETLES – “PLATO’S CAVE” OR SCIENTIFIC EVIDENCE? has sub-genera in the Codru-Moma unit, while drimeo-tus is in the Metaliferi and the Trascau Mountains. Each mountain is inhabited by a diferent sub-genus. It is possible that future searches will enlarge the distribution of drimeotus also in Codru-Moma. (Te frst and last specimen of Drimeotus in the Metaliferi was found as recently as 2001 in a cave well known for Protopholeuon and this was afer several trapping episodes where Protopholeuon/ drimeotus ratio was 150/1.) Fig. 4: distribution of Leptodirini in the Western Carpathians (Apuseni mountains): ¦k - drimeotus, ¦ - Pholeuon, ¦ - Protopholeuon, grey areas - karst. Comparing the largest geographical units in the Apuseni, Padurea Craiului and Bihor, the number of species and subspecies of less adapted drimeotus and the more adapted Pholeuon is diferent between genera and between units. drimeotus (20 species and 12 subspecies) has higher specifc diversity in both units and lower sub-specifc diversity than Pholeuon (6 species and 20 subspecies). Tis can be explained by the diference in adaptation and diferent histories of the two genera. Pholeuon, very adapted, is less mobile and very few individuals were found in MSS or under rocks. Preadapted ancestors of Pholeuon colonized deep subterranean environment in the entire area and since then small modifcations have occurred. Lacking competition, infra-generic differentiation of Pholeuon is slow. drimeotus, less adapted and relatively mobile between limestone areas, is under stronger climatic and biologically stronger selective pressures, which explains larger distribution areas and higher specifc diversity. Tere are also diferences between the Padurea Craiului and the Bihor drimeotus, with more than two-thirds of the species in the frst mountains. In Padurea Craiului the climate is warmer and less humid, with caves at lower altitude than in Bihor. Terefore, migration and gene fow between populations inhabiting diferent limestone areas or caves are limited and specia-tion is stronger. Te more humid and colder Bihor Mountains allows superfcial migrations and gene interchange between geographically close populations. Te altitudinal distribution of Leptodirini was frst discussed by Jeannel (1952), who mentioned the presence of the same genus and even same species at the surface at 1500 m altitude, under the rocks at 1000 m, and strictly cavernicolous at 500 m. A sound analysis of the distribution published by Decu (1980) emphasizes the lack of correlation between body size and altitude. we found diferent results, given in Table 1. For drimeotus, correlation between altitude and body length or relative length of antennae is negative for Padurea Craiului samples, and positive for Bihor samples. An approach that links geography to morphology is the kriging method, which uses a topographic representation of the data sets. Using the Golden Sofware Surfer 8 we obtained the maps in Fig. 5. As one can see, the vectors defne centers of origin at about 500-700 m altitude. Ancestors of the drimeotus lineage probably inhabited superfcial habitats between 500 and 1000 m altitude. Colonizing caves at lower or higher altitude induced body increase, a process explained by lack of competition and/or decrease of temperature. Body growth, as a result of cave colonization, is not only known for subterranean beetles. Te t test showed no signifcant diference in body length between populations of Padurea Craiului and Bihor. Te same test on the relative length of antennae gives shorter antennae in Bihor than in the Padurea Craiului samples. Tis can be due to the fact that the higher Bihor Mountains are inhabited by populations less confned to cave, compared to the Padurea Craiului. Pholeuon has the same tendencies as drimeotus, with a negative correlation of body length and altitude in Padurea Craiului and TIME in KARST – 2007 83 OANA TEODORA MOLDOVAN & GéZA RAJKA a positive correlation in Bihor, and the antennae length decrease with altitude. tab. 1: Linear regression (y) and coefcient of determination (R2) at drimeotus (13 populations) and Pholeuon (6 populations) from the Padurea Craiului and the bihor mountains (Western Carpathians): bL body length, AL/bL antenna/body lengths ratio. Altitude (m) 0-499 500-999 > 1000 Drimeotus BL y = -0.0003X + 4.4511 R2 = 0.0096 y = 0.0004x + 4.1025 R2 = 0.2337 y = 0.0007x + 3.5404 R2 = 0.3301 AL/BL y = -8E-05X + 0.5969 R2 = 0.1077 y = 3E-05x + 0.5327 R2 = 0.0195 y = 0.0001x + 0.4088 R2 = 0.2449 Pholeuon BL y = -0.0001X+ 3.9152 R2 = 0.0239 y = 0.0031x + 2.142 R2 = 0.7849 y = 1E-04x + 4.6179 R2 = 0.0150 AL/BL y = -1E-04x + 0.8619 R2 = 0.2445 y = -0.0002X + 0.9593 R2 = 0.7847 y = -0.0001x + 0.9044 R2 = 0.2529 Mantels test is a regression in which variables are matrices summarizing pairwise similarities among sample locations. Geographic distance was used as a predictor variable, and morphological features of populations belonging to drimeotus and Pholeuon, from the Padurea Craiului Mountains, as dependant distance matrices. For ? = 0.01, the test gives strong correlation for drimeotus samples (Fig. 6) and no correlation for Pholeuon. For the moment, we cannot explain this result, although presumptions can be formulated. Fig. 5: Antennae relative length (a) and altitude (b) variations in geographical coordinates for 15 populations of drimeotus s. str. (red dots) of Padurea Craiului mountains, in 3d and vectorial overlayed representations. Mantel’s test (xLSTAT 2006.5 sofware) was also used to correlate geographic distance with morphological features, such as body and relative antennae lengths. Fig. 6: histogram representing results of the mantel test on representatives of drimeotus s. str. from Padurea Craiului. 84 TIME in KARST – 2007 HISTORICAL BIOGEOGRAPHy OF SUBTERRANEAN BEETLES – “PLATO’S CAVE” OR SCIENTIFIC EVIDENCE? CONCLUSIONS Plato’s allegory still raises questions in historical bioge-ography of cave beetles, but new scientifc acquisitions will also enhance the chances for a more objective view in explaining the history of one group or another and what shaped their present distribution. we have only presented few results and further research must take into consideration more populations and the surface of the populated area, as in insular studies. In caves, as well as on oceanic islands, the number of endemics can be related to the size of the area. Studies of Barr (1985) and Culver et al. (1973) applied the insular theory of McArthur & wilson to karst areas, which have the characteristics of continental islands separated by non-calcareous “seas”. Tus, the number of subterranean species is strongly correlated with the surface of the limestone. Te recent interest in biogeographic studies has resulted from conservation necessities, especially in the last we are grateful to John Holsinger and David Culver for valuable suggestions, our colleagues Gheorghe Racovita and Tudor Tamas for useful discussions, and Andrej Mi- Barr Jr., T.C., 1985: Pattern and process in speciation of trechine beetles in eastern North America (Coleop-tera: Carabidae: Trechinae). In: Ball, G.E. (ed.) Phy-logeny and zoogeography of beetles and ants, Junk, Dordrecht, 350-407. Bellés, x., 1991: Survival, opportunism and convenience in the processes of cave colonization by terrestrial faunas. In: Ros, J.D. & N. Prat (eds.) homage to Ramon margalef; or, Why there is such pleasure in studying nature. – Oecol. Aquat. 10 325-335. Bleahu, M. & T. Rusu, 1965: Carstul din Romania. O scurta privire de ansamblu. – Lucr. Inst. Speol. “E. Racovita” 4 59-73. Bucur, R., Kosuch, J. & A. Seitz, 2003: Molecular phylo-genetic relationships of Romanian cave Leptodiri-nae (Coleoptera: Cholevidae). – Atti. Mus. Civ. Stor. Nat. Trieste 50 231-265. decade. For example, the concept of habitat fragmentation became one of the priority themes of conservation researches. Te concept is considered ambiguous, and empirical studies demonstrate a wide variety of direct and indirect efects, even with opposing implications (Haila 2002). Te efects of habitat fragmentation are considered extremely dangerous for species and population preservation, and are ofen mentioned when establishing protection areas for rare and vulnerable species. From a biospeleological point of view, habitat fragmentation represents one of the main mechanisms that enhanced speciation processes in reduced areas (at least for terrestrial fauna). Tis idea, diferent from the conservationist concern, can be extremely useful in solving protection problems. Unfortunately, none of the main contributions in conservation biology mentions caves and cave fauna as examples of survival in small, fragmented areas. hevc, Liviu Buzila and Alexandru Imbroane for advice and maps. de Candolle, A. P. , 1820: Géographie botanique. In: dic-tionnaire des Sciences Naturelles F.G. Levrault, Strasbourg, 18 359-422. Crisci, J.V., 2001: Te voice of historical biogeography. – J. Biogeogr. 28 157-168. Culver, D., Holsinger, J. R. & R. Baroody, 1973: Toward a predictive cave biogeography: the Greenbrier valley as a case study. – Evolution 27, 4 689-695. Decu, V. , 1980: Analyse de la repartition selon l’altitude des coléopteres cavernicoles Bathysciinae et Trechi-nae des Carpates de Roumanie.-Mém. Biospéol. 7 99-118. Decu, V. & S. Negrea, 1969: Aperçu zoogéographique sur la faune cavernicole terrestre de Roumanie. – Acta Zool. Cracov. 14, 20 471-546. Giachino, P. M., Decu, V. & C. Juberthie, 1998: Coleop-tera Cholevidae. In: Juberthie, C. & V. Decu (eds.) Encyclopaedia biospeologica, Fabbro Saint-Girons, France, 1083-1122. ACKNOwLEDGEMENTS REFERENCES TIME in KARST – 2007 85 OANA TEODORA MOLDOVAN & GéZA RAJKA Giachino, P.M. & D. Vailati, 1993: Revisione degli Ane-madinae Hatch, 1928 (Coleoptera Cholevidae).– Mus. Civ. Stor. Nat. Brescia, Monografa di “Natura Bresciana” 18 1-314. Goran, C., 1989: La spéléogramme de la Roumanie.-Mis-cell. Speol. Rom. 1: 335-346. Haila, y., 2002: A conceptual genealogy of fragmentation research: from island biogeography to landscape ecology.-Ecol. Appl. 12 312-334. Holsinger, J.R., 2000: Ecological derivation, colonization, and speciation. In: wilkens, H., Culver, D.C. & w.F. Humphreys (eds.) Subterranean ecosystems. Ecosystems of the world 30, Elsevier, Amsterdam, 399-415. Holsinger, J. R., 2005: Vicariance and dispersalist bio-geography. In: Culver, D. C. & white w. B. (eds.) Encyclopedia of Caves, Elsevier Academic Press, 591-599. Howarth, F.G., 1981: Non-relictual terrestrial troglobites in the tropic Hawaiian caves. Proc. 8th Internat. Congr. Speleol., Bowling Green, Kentucky (USA), 2 539-541. Jeannel, R., 1924: Monographie des Bathysciinae.–Arch. Zool. Exp. Gén. 63 1-436. Jeannel, R., 1931: Origine et évolution de la faune cav-ernicole du Bihar et des Carpathes du Banat.-Arch. Zool. Ital., Atti xI Congr. Internaz. Zool. Padova, 1930. 16 47-60. Jeannel, R. & N. Leleup, 1952: L’évolution souterraine dans la region méditerranéenne et sur les Mon-tagnes du Kivu.-Notes Biospéologiques 7 7-13. Juberthie, C., Delay, B. & M. Bouillon, 1980: Sur l’existence d’un milieu souterrain superfciel en zone non cal-caire.-C. R. Acad. Sc. Fr. 290 49-52. Juberthie, C., 1988: Paleoenvironment and speciation in the cave beetle complex Speonomus delarouzeei (Coleoptera, Bathysciinae).-Int. J. Speleol. 17 31-50. Moldovan, O., 1997: Reconnaissance des sexes et isole-ment reproductif chez les coléopteres bathysciinae souterrains: approche taxonomique, biochimique et expérimentale, PhD thesis, «Paul Sabatier» University, Toulouse (France), 130 pp. Moldovan, O.T., Juberthie, C., Jallon, J.-M. & H. Alves, 2000: Importance of cuticular hydrocarbons for speciation in the Speonomus delarouzeei complex (Coleoptera: Cholevidae: Leptodirinae).-Evolution & Adaptation 6 110-115. Moldovan, O.T., Iepure, S. & A. Persoiu, 2005: Biodiversity and protection of Romanian karst areas: the example of interstitial fauna. In: Stevanović, Z. & P. Milanović (eds.), Water resources and environmental problems in karst. Proc. Internat. Conf. & Field Semin., Belgrade & Kotor (Serbia & Montenegro), 13-19 September 2005, p. 831-836, Belgrade. Morrone, J.J. & J.V. Crisci, 1995: Historical biogeography: Introduction to methods.-Annu. Rev. Ecol. Syst. 26 373-401. Onac, B.P. & P. Cocean, 1996: Une vue global sur le karst roumain.-Kras i Speleologia 8, 17 105-112. Peck, S.B., 1980: Climatic change and the evolution of cave invertebrates in the Grand Canyon, Arizona.– Nat. Speleo. Soc. Bull. 42 53-60. Popov, S.V., Rögl, F., Rozanov, A.y., Steininger, F.F., Shcherba, I.G. & M. Kovac, 2004: Lithological-Pa-leogeographic maps of Paratethys Late Eocene to Pliocene. Courier Forschungsinstitut Senckenburg, Band 250, Frankfurt am Main, 46 pp., maps 1-10. Rögl, F. & F.F. Steininger, 1984: Neogene Paratethys, Mediterranean and Indo-pacifc seaways. Implications for the paleobiogeography of marine and terrestrial biotas. In: Brenchley, P. (ed.) Fossils and Climate, John wiley & Sons Ltd. 171-200. Steininger, F.F. & F. Rögl, 1985: Paleogeography and pal-inspastic reconstruction of the Neogene of the Mediterranean and Paratethys. In: Dixon, J.E. & A.H.F. Robertson (eds.), Te Geological Evolution of the Eastern Mediterranean, Special Publication of the Geological Society, No. 17, Blackwell Scientifc Publications, Oxford, pp. 659-668. Toolson, E.C., Markow, T.A., Jackson, L.J. & R.w. Howard, 1990: Epicuticular hydrocarbon composition of wild and laboratory-reared drosophila mojavensis Patterson and Crow (Diptera: Drospophilidae).-Annls. Ent. Soc. Am. 83 1165-1176. Vailati, D., 1988: Studi sui Bathysciinae delle Prealpi centro-occidentali. Revisione sistematica, ecologia, biogeografa della “seria fletica di boldoria” (Co-leoptera Catopidae).–Mus. Civ. Stor. Nat. Brescia, Monografa di “Natura Bresciana” 11 1-331. 86 TIME in KARST – 2007 COBISS: 1.01 wHAT DOES THE DISTRIBUTION OF STyGOBIOTIC COPEPODA (CRUSTACEA) TELL US ABOUT THEIR AGE? KAJ NAM POVE RAZŠIRJENOST STIGOBIONTSKIH CEPONOŽNIH RAKOV (CRUSTACEA: COPEPODA) O NJIHOVI STAROSTI? David C. CULVER1 & Tanja PIPAN2 Abstract UDC 595.3-15 591.5:595.3 David C. Culver & Tanja Pipan: What Does the Distribution of Stygobiotic Copepoda (Crustacea) Tell Us About Teir Age? Geographic distribution of stygobionts is ofen used to estimate age of a group by assuming vicariant speciation with little or no subsequent dispersal. we investigated the utility of using distributional data for Slovenian stygobiotic copepods by assuming that dispersal is a way to measure age of a species. we list some species of Copepoda that, on the basis of their range and frequency of occupancy within their range, should be older. Body size is not predictor either of range or frequency of occupancy. Key words: Speleobiology, Copepoda, stygobionts, dispersal biogeography. Izvleček UDK 595.3-15 591.5:595.3 David C. Culver & Tanja Pipan: Kaj nam pove razširjenost stigobiontskih ceponožnih rakov (Crustacea: Copepoda) o njihovi starosti? Ob predpostavki, da je nastajanje novih vrst posledica vikari-ance, brez naknadne disperzije, se za ocenjevanje starosti živalskih skupin pogosto uporablja geografska razširjenost sti-gobiontov. Ob domnevi, da je disperzija merilo za določanje starosti vrst, smo proučevali primernost podatkov o razširjenosti stigobiontskih kopepodov v Sloveniji. Na osnovi analize obsega naselitve in pogostosti naseljevanja znotraj območja smo v prispevku priložili seznam nekaterih vrst ceponožnih rakov, ki naj bi bile evolucijsko starejše. Telesna velikost ne določa obsega naselitve in pogostosti pojavljanja. Ključne besede: speleobiologija, Copepoda, stigobionti, dis-perzijska biogeografja. INTRODUCTION Te distribution of stygobionts has ofen been used to infer the age of a fauna. Te general procedure has been to assume that little or no migration has occurred, and that the extant distribution represents the site of original colonization and isolation in subterranean habitats. Te vicariance biogeographic view, now dominant in modern biogeography (e.g., Crisci, Katinas, and Posadas 2003) largely supplanted the old idea of centers of origin with species dispersing out from this central place (Matthew 1915). Given the reduced opportunities for dispersal of subterranean animals, it is not surprising that there have been a number of studies that show a correspondence between ancient shorelines and current distributions, especially Tethyan and Paratethyan distributions (Culver and Pipan in press). In some cases, it has been possible to match distributions to historical events and to obtain support from molecular clock data (see Verovnik, Sket, and Trontelj 2004). However, not all subterranean dis- 1 Department of Biology, American University, 4400 Massachusetts Ave., Nw, washington D.C., U.S.A.; e-mail: dculver@american.edu 2 Karst Research Institute ZRC-SAZU, Titov trg 2, SI-6230 Postojna, Slovenia; e-mail: pipan@zrc-sazu.si Received/Prejeto: 27.11.2006 TIME in KARST, POSTOJNA 2007, 87–91 DAVID C. CULVER & TANJA PIPAN tributions can be explained solely by vicariance. A particularly interesting example is the cirolanid isopod An-trolana lira Bowman. In general, subterranean cirolanids are found near to marine shores (Botosaneanu, Bruce, and Notenboom 1986), suggesting a marine ancestor with vicariant isolation. But, A. lira is found in caves in the Appalachian Valley of Virginia and more than 200 km from not only the present ocean shore, but from any ocean dating back at least to the Paleozoic (Holsinger, Hubbard, and Bowman 1994). In this contribution, we take a dispersalist rather a vicariance view of subterranean biogeography. we consider a model of colonization and isolation as follows. A species colonizes and is isolated in a subterranean site. As adaptation occurs, the species occupies more sites in the vicinity of the colonization. Tat is, the frequency of occupancy of subterranean sites increases. In the next stage, the species expands its range, with a high occupan- cy of suitable sites in its range. Finally, as other species also evolve, the original species may be out-competed or it may become specialized in response to competition. In this scenario, it will then occupy a lower frequency of sites within its range. Tus, we can rank the ages of species in increasing age as follows: 1. Species with small ranges and occupying few (sometimes only one) sites 2. Species with small ranges but occupying a high frequency of sites within its range 3. Species with large ranges and occupying a high frequency of sites within its range 4. Species with large ranges and occupying a low frequency of sites within its range. we examine this hypothesis using distributional data of subterranean copepods from Slovenia (see Pipan 2005), and make assess the utility of this approach. METHODS AND MATERIALS From information in Pipan (2005) and Culver, Pipan, and Schneider (in press) we generated list of stygobiotic copepods from seven Slovenian caves, with information on ranges, frequency of occupancy of well-sampled caves, and average body size. Ranges were categorized into three groups: 1. Slovenian endemics 2. Balkan endemics 3. European endemics 4. Cosmopolitan species To measure frequency of occupancy, we used data from Pipan (2005), which was intensive enough that it is likely that most species were found (Pipan and Cul- ver in press). Body sizes were taken from original species descriptions and direct measurement by T P. Data were available for 37 species. Analysis was done by grouping ranges into two categories (Small—categories 1 and 2 and Large—categories 3 and 4), frequency of occupancy into two categories (Low—1 to 3 caves and High—4 to 7 caves), and size into two categories (Small—less than the overall mean of 0.61 and large—greater than or equal to the overall mean of 0.61). Te resulting 2x2 contingency tables were analyzed for independence using Fisher’s Exact Test in JMPTM (Sall, Creighton, and Lehman 2005). RESULTS Available data for the 37 species of stybobiotic copepods are shown in Table 1. In Table 2, all species are categorized into four groups based on range and occupancy. Tere was no signifcant diference between observed and expected although there was a small excess of species with large ranges that were also found in a high frequency of caves. Tose species hypothesized to be the oldest (large ranges, low occupancy) were: • Acanthocyclops kieferi • Acanthocyclops venustus stammeri • diacyclops clandestinus • dicyclops languidoides • Elaphoidella elaphoides • morariopsis scotenophila Te group hypothesized to be the next oldest are those with large ranges and high occupancy: • Elaphoidella jeanneli • bryocamptus balcanicus • Acanthocyclops venustus • Parastenocaris nolli alpina 88 TIME in KARST – 2007 wHAT DOES THE DISTRIBUTION OF STyGOBIOTIC COPEPODA (CRUSTACEA) TELL US ABOUT THEIR AGE? tab. 1: Stygobiotic copepod species found in seven well-sampled caves in Slovenia. See Pipan (2005) and Culver et al. (in press). Species Name/Taxonomic Authority Mean Body Size No. Caves Occupied Range Acanthocyclops kieferi (Chappuis, 1925) 0.73 2 3 Acanthocyclops venustus (Norman & Scott, 1906) 1.07 1 3 Acanthocyclops venustus stammeri (Kiefer, 1930) 1.07 5 3 Bryocamptus balcanicus (Kiefer 1933) 0.40 4 3 Bryocamptus borus Karanovic & Bobic, 1998 1 2 Bryocamptus pyrenaicus (Chappuis, 1923) 0.80 7 3 Bryocamptus sp. 2 1 cf. Stygepactophanes sp. 0.35 3 1 Diacyclops charon (Kiefer, 1931) 1.00 7 2 Diacyclops clandestinus (Kiefer, 1926) 0.40 3 4 Diacyclops hypogeus (Kiefer, 1930) 0.50 2 1 Diacyclops languidoides (Lilljeborg, 1901) 0.80 3 4 Diacyclops slovenicus (Petkovski, 1954) 0.68 3 1 Echinocamptus georgevitchi (Chappuis, 1924) 0.70 1 2 Elaphoidella cvetkae Petkovski, 1983 0.75 4 2 Elaphoidella elaphoides (Chappuis, 1924) 0.60 1 3 Elaphoidella franci Petkovski, 1983 0.64 1 1 Elaphoidella jeannelli Chappuis, 1928 0.60 4 3 Elaphoidellakarstica Dussart & Defaye (1990) 1 1 Elaphoidella sp. A 2 1 Elaphoidella sp. B 2 1 Elaphoidella stammeri Chappuis, 1936 0.62 4 1 Maraenobiotus cf. brucei 0.60 1 1 Metacyclops postojnae Brancelj, 1990 >0.61 1 2 Moraria sp. A 2 1 Mor aria sp. B 1 1 Moraria stankovitchi Chappuis, 1924 0.55 2 2 Morariopsis dumonti Brancelj, 2000 0.39 2 1 Morariopsis scotenophila (Kiefer 1930) 0.49 3 4 Nitocrella sp. 0.50 2 1 Parastenocaris cf. andreji 0.40 2 1 Parastenocaris nolli alpina (Kiefer, 1938) 0.42 5 3 Parastenocaris sp. A 0.40 2 1 Parastenocaris sp. B 0.40 4 1 Parastenocaris sp. C 0.40 2 1 Speocyclops infernus (Kiefer 1930) 0.47 6 2 Troglodiaptomus sketi Petkovski, 1978 0.88 3 2 we investigated whether there was a body size bias pan and Culver 2006). In any case, there was no relation-for occupancy or range size. Smaller copepods might be ship between frequency of occupancy and body size and able to disperse more easily but they may also be more no relationship between range and body size (Table 3). subject to the vagaries of water movement in epikarst (Pi- TIME in KARST – 2007 89 DAVID C. CULVER & TANJA PIPAN tab. 2: Number of species of stygobiotic copepods in categories of large and small range and high and low frequency of site occupancy. Numbers in parentheses are the expected numbers. Observed and expected numbers do not signifcantly difer (p=0.21, Fisher’s Exact test). High Occupancy Low Occupancy Large Range 4 (2.4) 5 (6.6) Small Range 6 (7.6) 22 (20.4) tab. 3: Number of species of stygobiotic copepods in categories of high and low frequency of site occupancy (A), large and small range (b) and body size. Numbers in parentheses are the expected numbers. Neither association was statistically signifcant (p=0.71 for A, p=0.71 for b, Fisher’s Exact test). A. High Occupancy Low Occupancy Large Body Size 5 (4.3) 8 (8.7) Small Body Size 5 (6.7) 12 (11.3) B. Large Range Small Range Large Body Size 5 (4.5) 8 (8.5) Small Body Size 5 (5.5) 11 (10.5) we have created a list of copepod species that, according to the hypothesis outlined in the introduction, should be older than other stygobiotic copepod species discussed in this study. Unfortunately, we know of no detailed phy-logeny that would allow for such a comparison but we think that it would make for a very interesting study to do so. what is known about copepod phylogeny is that the Cyclopoida seem to be a more recent group than the Harpacticoida, accoding to the phylogeny of Huys and Boxshall (1991). Te fact that cyclopoids are over-represented among species with large ranges (Table 4) contradicts the hypothesis put forward. Of course, just because Te authors were supported by funds from the Center for Subterranean Biodiversity of the Karst waters Insti- Finally, we investigated the taxonomic position of the putative older species, i.e., those with larger ranges. Of the ten species listed above, fve are cyclopoids and fve are harpacticoids. Tere is an excess of large ranged cyclopoids but the diference was only signifcant at p~0.10 (Table 4). Acanthocyclops is especially noteworthy. All three stygobiotic species (A. kieferi, A. venustus, and A. venustus stammeri) had large ranges. In contrast none of the three species of Moraria (m. stankovitchi, sp. A, and sp. b) have large ranges. Te lone calanoid species (troglodiaptomus sketi) also has a small range. tab. 4: Relationship between range and taxonomic group (Cyclopoida vs. harpacticoida). Expected numbers are given in parentheses. Te relationship was marginally signifcant (p~0.10, Fisher’s Exact test). Large Range Small Range Cyclopoida 5 (2.8) 5 (7.2) Harpacticoida 5 (7.2) 21 (18.8) cyclopoids as a group are younger does not mean that the species are all younger than harpacticoids. Alternatively, it may be that harpacticoids are in general being outcom-peted by cyclopoids, and this has resulted, not only in reduction in occupancy frequency, but also in range contraction. we think that examination of the kinds of distribution patterns (range size and occupancy) discussed here will yield interesting results. Tis analysis would enrich phylogeography studies as well as provide additional hypotheses about the origin and evolution of subterranean groups. tute and the Ministry of Higher Education, Science, and Technology of the Republic of Slovenia. DISCUSSION ACKNOwLEDGEMENTS 90 TIME in KARST – 2007 wHAT DOES THE DISTRIBUTION OF STyGOBIOTIC COPEPODA (CRUSTACEA) TELL US ABOUT THEIR AGE? REFERENCES Botosaneanu, L., N. Bruce, & J. Notenboom., 1986: Isopoda: Cirolanidae, pp. 412-421, in L. Botosanea-nu [ed.] Stygofauna mundi. E.J. Brill, Leiden, Te Netherlands. Crisci, J.V., L. Katinas, & P. Posadas., 2003: historical bio-geography. An Introduction. p. 250, Harvard Univ. Press, Cambridge. Culver, D.C., T. Pipan., & K. Schneider., in press: Vicari-ance, dispersal, and scale in the aquatic subterranean fauna of karst regions. Freshwater biology Culver, D.C., & T. Pipan., in press: Subterranean ecosystems. In S.A. Levin [ed.] Encyclopedia of biodiversity, second edition. Elsevier, Amsterdam. Holsinger, J. R., D. A. Hubbard, Jr , & T. E. Bowman., 1994: Biogeographic and ecological implicationd of newly discovered populations of the stygobiont iso-pod crustacean Antrolana lira Bowman (Cirolani-dae). journal of Natural history 28, 1047-1058. Huys, R., and G. Boxshall., 1991: Copepod evolution. p. 468, Te Ray Society, London, Matthew, w.D., 1915: Climate and evolution. Annals of the New york Academy of Sciences 24, 171-318. Pipan, T., 2005: Epikarst – a Promising habitat. 100 p. Karst Researach Institute at ZRC-SAZU, ZRC Publishing, Postojna. Pipan, T., & D.C. Culver., 2006: Copepod distribution as an indicator of epikarst system connectivity. hydro-geology journal Pipan, T., & D.C. Culver., in press: Regional species richness in an obligate subterranean dwelling fauna— epikarst. journal of biogeography. Sall, J., L. Creighton, & A. Lehman., 2005: jmP Start Statistics. Brook/Cole—Tomson Learning, Belmont, California. Verovnik, R., B. Sket, & P. Trontelj., 2004: Phylogeog-raphy of subterranean and surface populations of water lice Asellus aquaticus (Crustacea: Isopoda). molecular Ecology 13, 1519-1532. TIME in KARST – 2007 91 COBISS: 1.01 HOw TO DATE NOTHING wITH COSMOGENIC NUCLIDES KAKO DATIRATI PRAZNINE S KOZMOGENIMI NUKLIDI Philipp HäUSELMANN1 Abstract UDC 539.16:552.5(494) Philipp Häuselmann: How to date nothing with cosmogenic nuclides A cave is a natural void in the rock. Terefore, a cave in itself cannot be dated, and one has to resort to datable sediments to get ideas about the age of the void itself. Te problem then is that it is never very certain that the obtained age really is coincident with the true age of the cave. Here, we present the use of a method which couples sedimentary and morphologic information to get a relative chronology of events. Datings within this relative chronology can be used for assessing ages of forms, processes, and sediments, and the obtained dates also fx some milestones within the chronology, which then can be used to retrace, among other things, paleoclimatic variations. For many cave systems, the dating limits of the most widely used U/T method on speleothems are too low (350 to max. 700 ka) to get ages that inform us about the age of the cave. Te recent use of cosmogenic nuclides on quartz-containing sediment permits to push the datable range back to 5 Ma. while the theoretical background is explained elsewhere (Granger, this volume), we concentrate on the Siebenhengste example (Switzerland). Key words: relative chronology, cosmogenic nuclides, cave dating methodology, Siebenhengste. Izvleček UDK 539.16:552.5(494) Philipp Häuselmann: Kako datirati praznine s kozmogenimi nuklidi Jame predstavljajo praznino v kamninski masi in jim kot takim ne moremo določiti starost. Zato z datiranjem jamskih sedimen-tov sklepamo tudi o starosti jame, pri čemer seveda ne moremo trditi, da je dobljena starost tudi prava starost jame. V članku predstavimo metodo pri kateri z združitvijo sedimentarnih in morfoloških izsledkov sklepamo o relativni kronologiji dogodkov. Datiranje v oviru relativne kronologije lahko uporabimo za določevanje starosti različnih oblik, procesov in sedimentov. Dobljene rezultate pa lahko uporabimo kot pomembne mejnike v kronologiji, npr. pri intepretaciji klimatskih sprememb. Veliko jam je starejšiih od zgornje meje starosti (350 do 700 ka), ki jo lahko določimo z uran-torijevo metodo, ki je zelo razširjena. V zadnjem času se zato uveljavlja metoda datacije s kozmogenimi nuklidi, ki omogoča datiranje dogodkov do starosti 5 Ma. Ker je teoretično ozadje te metode predstavljeno drugje (npr. Granger v tej številki), se tu omejimo le na uporabo metode v jamskem sistemu Siebenhengste (Švica). Ključne besede: relativna kronologija, kozmogeni nuklidi, metodika datiranja jam, Siebenhengste. INTRODUCTION For many cave scientists, it might not be evident that a while the sediments found within the cave give variable cave does not exist - only the surrounding rock gives existence to the void called cave. Terefore, a cave cannot be dated by conventional methods (Sasowsky 1998), but one has to use datable sediments. In karstic caves, the age of the surrounding rock gives a maximal age of the cave, ages from today (in the case of still active speleothems) up to the last stages of speleogenesis (in the case of spe-cifc sand deposits dated by cosmogenic nuclides) and therefore to the age of nothing itself. 1 Swiss Institute of Speleology and Karst studies SISKA, c.p. 818, 2301 La Chaux-de-Fonds, Switzerland, Fax 0041 32 913 3555, e-mail: praezis@speleo.ch Received/Prejeto: 11.12.2006 TIME in KARST, POSTOJNA 2007, 93–100 PHILIPP HäUSELMANN Tis paper contains two parts. In the frst part, the concept of relative chronology is explained. Te link between morphology and sediment succession leads to a relative chronology of erosional and depositional events. Any dating of sediment with the purpose of studying the age of nothing basically requires such a relative chronology, which places the obtained data into a timeframe. In the second part, the dating of sandy cave sediments with cosmogenic nuclides is briefy presented. Es- THE CONCEPT OF RE INTRODUCTION Geologists and other scientists are usually aware of the laws of stratigraphy, which say that a younger sediment overlies an older one. Tese laws are the base of a relative chronology. Tis chronology is normally used to assess the correctness of an obtained age - the numerical value has to be concordant with stratigraphy, or the dated age may not be correct. Most of the time, this principle is used with stalagmites, where the obtained ages must be older at the base and younger at the top (e.g. Spötl et al. 2002). Morphological indications, on the other hand, also give chronological information. A keyhole passage informs us that a phreatic phase was followed by a vadose one. Successions of speleogenetic phases are found in many cave systems. while some of them indicate base level rises (Audra et al., 2004), most of them indicate a downcutting of the regional base level (uplif, valley deepening, e.g. Ford & williams 1989; Rossi, Cortel & Arcenegui 1997). Tis in itself is also a chronological information: the oldest cave passages are on top, the youngest ones near the present baselevel. Te difculty now is to connect the sediments of several, basically independent, sedimentary profles and to link them with the morphological succession of the cave passages. Tus, the sedimentary profles are not independent from each other, and a relative chronology of erosional and depositional events over the whole cave can be made. ExAMPLE Figure 1 shows a real situation encountered in St. Beatus Cave (Switzerland): To the right side is a typical keyhole passage which proves that a phreatic initiation of the ellipse on top was followed by a canyon incision. In the middle part of the fgure, the meander gradually disappears and is replaced by a more or less elliptic passage that continues towards pecially when dealing with sands, a relative chronology is very important to date only meaningful sediments. Te theoretical background is only very briefy presented, and the reader is referred to Granger (this volume) for more thorough information. Te Siebenhengste example, the use of the relative chronology, and the obtained results are presented in more detail. TIVE CHRONOLOGy the lef side of the fgure. we see therefore a transition of a vadose feature into a phreatic one, and thus an old water level. In the profle to the right, we observe fowstone deposition that was truncated by the river incising the meander. Terefore, the fowstone predates the canyon, but postdates the initial genesis of the elliptic passage to the right. Te meander changes into an elliptic passage, thus the two forms are contemporaneous. Consequently, the older fowstone disappears in the area of this transition. within all the passages, silts were deposited. Tey are younger than the meander incision, and younger than the passage to the lef, and prove of an inundation of the whole cave. Stalagmites grow on the silts and are partially still active. Tis example can be written as a table (Tab. 1). -------------------------------------- Phreatic genesis of top ellipse -------------------------------------- Water level lowering Deposition of fowstone Erosion of fowstone Erosion of meander -------------------------------------- Water level lowering Silt deposition Stalagmite growth -------------------------------------- tab. 1: Chronology of erosional and depositional events (Fig. 1) Tis table is a frst relative chronology that links the sediments and the morphology of the cave.For practical reasons, the table presenting the chronology of events in a large cave system is not rewritten with each sedimentary succession found. Instead, the single sedimentary sequence is coupled with morphology, and is written as a column in the table. Te next sedimentary sequence, again coupled with morphology, is written as another column. Tus, the above example would then look like Table 2. 94 TIME in KARST – 2007 HOw TO DATE NOTHING wITH COSMOGENIC NUCLIDES Sequence at left Sequence at right Phreatic genesis Phreatic genesis Phreatic genesis Water level lowering Silt deposition Stalagmite growth Phreatic genesis of top ellipse Water level lowering Deposition of fowstone Erosion of fowstone Erosion of meander Silt deposition Stalagmite growth tab. 2: Chronological table with columnar writing of Fig. 1 ExPANSION If we continue up- and downstream of that profle, we fnd several other morphological indications and sedimentary successions, each of them having a link with our initial profle - until we encounter the next paleo-water-level and thus the next morphological change. Tere, the links have to be established again. Te table thus slowly grows and gets more complete. Of course, the example presented above is an ideal case. Ofen, the passages lack some information, thus making it difcult to establish an unambiguous chron-Table 3: A more complicated example from St. Beatus Cave Lower passage Phreatic genesis Water level lowering Pebble deposition Speleothem Erosion Sand deposition Speleothem Sand deposition Silt deposition Pebble deposition Sand deposition Silt deposition Erosion Speleothem Erosion Silt deposition Speleothem Erosion Silt deposition ? ? Upper passage Phreatic genesis Water level lowering Speleothem Silt deposition Speleothem Silt deposition Speleothem Silt deposition Erosion Speleothem Silt deposition ? ological table. Table 3 give an example: here, the upper passage lacks incision of a canyon. Terefore, it is not clear whether the sediments found in the upper passage were all deposited while the lower passage was still in its initial genesis, or whether the sediments can be partly correlated. In this case, a relative correlation of the sediments by observation only is not possible: some absolute dates have to be obtained. Of course, these ages have to be in stratigraphic order of both the sediment succession and the morphologic indications. Te above example had been dated by U/T on speleothems. Te resulting table is presented in Table 4. Here, the speleothems with roughly the same age have been grouped together. Ten, laminated silt deposits that are thought to be a product of glacial damming (Bini, Tognini & Zuccoli 1998; Audra et al., this volume), are parallelized, inferring that the whole cave was fooded in such conditions. Of course, some uncertainties still persist. Table 4: The more complicated example, dated and expanded Lower passage Phreatic genesis Water level lowering Pebble deposition Speleothem Erosion Sand deposition Speleothem (235 ka) Sand deposition Silt deposition Pebble deposition Sand deposition Silt deposition Erosion Speleothem (180 ka) Erosion Silt deposition Speleothem (91 ka) Erosion Silt deposition Upper passage Phreatic genesis Water level lowering Speleothem (>350 ka) Silt deposition Speleothem (337 ka) Silt deposition Speleothem (114 ka) Silt deposition Erosion Speleothem (99 ka) Silt deposition tab. 3: A more complicated example from St. beatus Cave tab. 4: Te more complicated example, dated and expanded TIME in KARST – 2007 95 PHILIPP HäUSELMANN Fig. 1: Schematic section through a part of St. beatus Cave (Switzerland), showing the relationship between sediments and morphology. wHy A RELATIVE CHRONOLOGy? parallelized all the sedimentary sequences, it is possible Te huge advantage of such a table of relative chronology to make a synthetic and dated sediment profle of the is that it ofers more control on the correct stratigraphic whole cave, which can then be used to get information order than single sections, in ideal cases also the cave on climatic variations and the presence or absence of gla-genesis can be dated, and last but not least, when having ciers damming the cave’s exit (Häuselmann 2002). DATING wITH COSMOGENIC NUCLIDES INTRODUCTION Cosmogenic nuclides are generated by the interaction of cosmic rays (mainly protons, neutrons, and muons) with atoms in the Earth’s atmosphere and lithosphere. Te production rate of cosmogenic isotopes depends on the intensity of the cosmic rays, which is subject to change. Te atmosphere then absorbs most of the primary rays and thus causes production rates to depend on elevation. Finally, the geometry of the sample location (and eventual snow or soil cover) also has its efects. Te radioactive nuclides most widely used for dating purposes are 10Be and 26Al produced in quartz. THE PRINCIPLE AND POSSIBILITIES OF BURIAL DATING Burial dating of cave sediments is a relatively new technique that indicates the time sediment has been underground (Granger, Fabel & Palmer 2001). It relays on the radioactive decay of the nuclides that were previously accumulated when the sediment was exposed at the surface. whereas the intensity of the cosmic rays may vary with time, the ratio of produced 10Be to 26Al remains always approximately 1:7. Te 10Be/26Al ratio can thus be calculated from the production rates and radioactive decay. If a sample that contains 10Be and 26Al is washed underground to sufcient depth to be shielded from further radiation, the nuclide concentrations diminish. Since 26Al has a half-life of 720 ka, opposed to the one of 10Be of 1.34 Ma, the ratio of 1:7 is gradually lowered. Measurement of that ratio therefore gives a direct indication of the time the sample remained underground. Of course, several prerequisites have to be fulflled in order to get a burial age: - First of all, the sediment must contain quartz that was irradiated sufciently prior to burial. Te grain size should be minimally fne sand (otherwise the cleaning process also eliminates the quartz), but may reach pebble size without problem. - Ten, burial should ideally be 20-30 m below the surface to be sufciently shielded from radiation. - In order to make a measurement meaningful, the stratigraphic relationship of the sampled sand with the passage and other sediments should be clearly established - the relative chronology is needed. Burial dating has a range from about 100’000 years up to 5 Ma. Afer that time, the amount of remaining isotopes is usually too small to be measured accurately (Granger & Muzikar 2001). It is one of only a few radio-metric methods that date lower quarternary and Pliocene deposits. It is of great interest for cave dating, frst because many old caves were created in the Pliocene or even earlier, and second because caves are very efective at shielding the sediment from further cosmic ray bombardment. As with other cave-dating methods, burial dating may also be used to date the age of the passage, 96 TIME in KARST – 2007 HOw TO DATE NOTHING wITH COSMOGENIC NUCLIDES Caves of the region Siebenhengste - Hohgant Perspective view from 370g, as of January, 2002 Only the most important passages shown Spring for 1440 - 558 Aare valley Fitzlischacht I 0.5 1 1.5 km mi* Siebenhengste IA201IISHPT Spring for 1950-1505 Eriz Hohgant "~760 558 St. Beatus Caves Bärenschacht Haglätsch by HRH & toporobot Fig. 2: Projection (370 degrees) of the Siebenhengste caves with the speleogenetic phases. Stars indicate sampling places for cosmogenic dates. From häuselmann & Granger (2005), modifed. thus indicating valley deepening rates and evolution of the surface outside the cave. THE SIEBENHENGSTE ExAMPLE we used burial dating to date the old passages of the Siebenhengste cave system in Switzerland. Te Siebenhengste region is situated in the north-western part of the Alps, adjacent to the molasse basin. From Lake Tun, the mountain range extends to the Schrattenfuh, 20 km away. Te cave region is one of the longest and deepest worldwide, with the Réseau Siebenhengste-Hohgant Fig. 3: Plot of ages (vertical) versus altitude (horizontal). having 154 km length and -1340 m depth. Te caves comprise 14 diferent speleogenetic phases, which can be related to paleo-valley bottoms (Jeannin, Bitterli & Häuselmann 2000). Te highest and oldest fve phases (at presumed spring elevations of >1900, 1800, 1720, 1585, and 1505 m a.s.l.) had their springs in the Eriz valley (Fig. 2). Te next phase, at 1440 m, shows a change in fow direction of 180°. Te spring was then located in the area of Lake Tun. Te infuence of today’s Aare valley (the site of Lake Tun today) therefore became predominant. All subsequent springs (at 1145, 1050, 890, 805, 760, 700, 660, and 558 m a.s.l.) drained towards the Aare valley. In the area between Lake Tun and Hohgant, a total of 23 sites were selected for sampling (see Fig. 2: stars indicate sites). Selection was made on the basis of a relative chronology, and care has been given to ensure that either the oldest possible sediment, or a series in stratigraphic order, was sampled. Due to the limited amount of time in which sampling could be done, the relative chronology is incomplete (Tab. 5), although the main events were retraced. 21 samples were analysed (Häuselmann & Granger 2005). Te results show a great diversity of ages, ranging from 118 ka up to TIME in KARST – 2007 97 Faustloch Bätterich PHILIPP HäUSELMANN bold = morphologic event (a 0 denotes phreatic genesis, a v vadose enlargement), italic = dated event A201 ShP low SHP up Haglätsch A2TR A2CHU A2NS RBL L18 Faustloch Beatus Age Interpretation 18000 18000 18000 SHP 7 Sediment Erosion Erosion SHP2 SHP 3 Silt 18000 4.39 2.35 17200 Flowst. Flowst. Silt lake Silt Erosion Erosion Erosion Flowst. 17200 17200 Sand Erosion A201 SHP5 Sand Silt 1.9-1.84 Sand Silt 15850 15850 15850 15850 15850 Erosion Parag.? Flowst. Flowst. Silt Flowst. Erosion lake SHP1 Flowst. SHP6 HGLP Flowst. Flowst. Erosion Erosion Canyon HGLS Paragen. epiphreatic L18 Silt 1.54-1.60 1.04-1.09 (.93?) SHP4 HGLT A2TR A2CHU A2NS 1505/14400 Flowst. RBL2 Erosion RBL1 10500 1505/14400 0.78-0.80 (.93?) 0.63 10500 1050V 8900 Flowst. flooding FSTL 8050 7600 0.47 BG23 0.23 BG1 0.18 BG20 0.16 8900 8050 7600 tab. 5: Relative chronology of events around the Siebenhengste 1500 1000 500 1 2 3 4 6 Burial age (10 years) 4.4 Ma (Tab. 6). Te surface sample (MwA) has a burial age of 106 ± 176 ka. Tus, the value is indistinguishable from zero, and we may assume that the sample was never buried. Te sample from St. Beatus Cave (BG1) has an age of 182 ± 122 ka. Its true value, bracketed by U/T Fig. 4: Rate of valley lowering in the Siebenhengste. Only maximum and minimum ages are displayed; however the valley deepening rates as well as the knickpoint at ~800 ka are easily visible. 98 TIME in KARST – 2007 5 A relative chronology of events, albeit incomplete, coupled with burial age dating by cosmogenic nuclides, permitted to obtain a continuous history of valley incision in the Alps. Such data cannot be obtained in the same precision with other methods or at the surface. Te results presented here are the frst cosmogenic dates for an Alpine cave system in a glacially infuenced area. Te results indicate an onset of karstifcation in the Siebenhengste before 4.4 Ma, that is in the Pliocene or even earlier. Together with U/T dates obtained earlier (Häuselmann 2002), the history of the Siebenhengste cave system and HOw TO DATE NOTHING wITH COSMOGENIC NUCLIDES ages, should be between 160 and 235 ka, which is again the case. Tese values indicate that the method yields young ages where expected. A difculty for dating with cosmogenic nuclides is mobility of the sediment. For instance, recent sand can be transported into a fossilized cave by a food and then be deposited. Our results show that this process happens: for any speleogenetic phase, there is a range of ages observed (Fig. 3). However, Fig. 3 also indicates that the re-mobilization and re-deposition of old sediments is rarely observable: if this would be the case, we would expect a random distribution of ages throughout the phases. However, the maximum age decreases with the next lower phase. we can thus construct a gradual valley lowering with time which is represented in Fig. 4. we see a knickpoint in the line connecting the ages: this knickpoint occurs at around 800 ka and 1500 m. Tis point refects a dramatic increase in valley deepening rate and coincides with the change in fow direction from Eriz to the Aare valley. tab. 6: Results of dating. its surrounding environment can be traced back over a huge time span. Te construction of a complete relative chronology is very time-consuming, but can be extremely rewarding given the information one can extract from the cave. If speleogenetic phases, which are related to the overall geomorphic evolution of an area, can be expanded by such relative chronologies as well as absolute dates, the rate, duration, and extent of valley deepenings can be assessed, and a paleoclimatic history can be drawn as well. TIME in KARST – 2007 99 PHILIPP HäUSELMANN REFERENCES Audra, Ph., L. Mocochain, H. Camus, E. Gilli, G. Clauzon & J.-y. Bigot, 2004: Te efent of the Messinian Deep Stage on karst development around the Mediterranean Sea. Examples from Southern France. - Geo-dinamica Acta, 17, 6, 27-38. Bini, A., P. Tognini, & L. Zuccoli, 1998: Rapport entre karst et glaciers durant les glaciations dans les val-lées préalpines du Sud des Alpes. - Karstologia, 32, 2, 7-26. Ford, D.C. & P. williams, 1989: Karst geomorphology and hydrology. Chapman & Hall, London, 601 p. Granger, D.E, D. Fabel & A.N. Palmer, 2001: Pliocene-Pleistocene incision of the Green River, Kentucky, determined from radioactive decay of cosmogenic 26Al and 10Be in Mammoth Cave sediments. - GSA Bulletin, 113, 7, 825-836. Granger, D.E. & P.F. Muzikar, 2001: Dating sediment burial with in-situ produced cosmogenic nuclides: theory, techniques, and limitations. - Earth and Planetary Science Letters, 188, 269-281. Häuselmann, Ph., 2002: Cave genesis and its relationship to surface processes: Investigations in the Siebenhengste region (bE, Switzerland). - PhD thesis, Université de Fribourg, 168 p. Häuselmann, Ph. & D.E. Granger, 2005: Dating of caves by cosmogenic nuclides: Method, possibilities, and the Siebenhengste example (Switzerland). - Acta Carsologica, 34, 1, 43-50. Jeannin, P.-y., T. Bitterli, T. & Ph. Häuselmann, 2000: Genesis of a large cave system: the case study of the North of Lake Tun system (Canton Bern, Switzerland). In: A. Klimchouk, D. C. Ford, A. N. Palmer, & w. Dreybrodt (Eds.), Speleogenesis: Evolution of Karst Aquifers, pp. 338-347. Rossi, C., A. Cortel & R. Arcenegui, 1997: Multiple pa-leo-water tables in Agujas Cave System (Sierra de Penalabra, Cantabrian Mountains, N Spain): Criteria for recognition and model for vertical evolution. - Proceedings 12th Int. Congress of Speleology, La Chaux-de-Fonds, Switzerland, 1, 183-187. Sasowsky, I.D., 1998: Determining the age of what is not there. - Science, 279, 1874. Spötl, C., M. Unterwurzacher, A. Mangini & F.J. Long-stafe, 2002: Carbonate speleothems in the dry, inneralpine Vinschgau valley, northernmost Italy: witnesses of changes in climate and hydrology since the last glacial maximum. - Journal of Sedimentary Research, 72, 6, 793-808 100 TIME in KARST – 2007 COBISS: 1.01 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION (KRAS, MATARSKO PODOLJE AND ISTRIA; Sw SLOVENIA AND Nw CROATIA) ZAKRASELA PERIFERNA IZBOKLINA POVEZANA Z RAZVOJEM ZGORNJEKREDNO-PALEOGENSKEGA PREDGORSKEGA BAZENA; KRAS, MATARSKO PODOLJE IN ISTRA (JZ SLOVENIJA IN SZ HRVAŠKA) Bojan OTONIČAR1 Abstract UDC 551.44.551.7(497.4-14) 552.541.551.7(497.4-14) Bojan Otoničar: Upper Cretaceous to Paleogene forbulge unconformity associated with foreland basin evolution (Kras, Matarsko Podolje and Istria; SW Slovenia and NW Croatia) A regional unconformity separates the Cretaceous passive margin shallow-marine carbonate sequence of Adriatic Carbonate Platform from the Upper Cretaceous and/or Paleogene shallow-marine sequences of synorogenic carbonate platform in southwestern Slovenia and Istria (a part of southwestern Slovenia and northwestern Croatia). Te unconformity is expressed by irregular paleokarstic surface, locally marked by bauxite deposits. Distinctive subsurface paleokarstic features occur below the surface (e.g. flled phreatic caves, spongework horizons…). Te age of the limestones that immediately underlie the unconformity and the extent of the chronostratigraphic gap in southwestern Slovenia and Istria systematically increase from northeast towards southwest, while the age of the overlying limestones decreases in this direction. Similarly, the deposits of synorogenic carbonate platform, pelagic marls and fysch (i.e. underflled trinity), deposits typical of underflled peripheral foreland basin, are also diachronous over the area and had been advancing from northeast towards southwest from Campan-ian to Eocene. Systematic trends of isochrones of the carbonate rocks that immediately under- and overlie the paleokarstic surface, and consequently, of the extent of the chronostratigraphic gap can be explained mainly by the evolution and topography of peripheral foreland bulge (the forebulge). Te advancing fexural foreland profle was the result of vertical loading of the foreland lithospheric plate (Adria microplate) by the evolving orogenic wedge. Because of syn- and post-orogenic tectonic processes, and time discrepancy between adjacent foreland basin deposits and tectonic (“orogenic”) phases it is difcult to defne the exact tectonic phase responsible for the evolution of the foreland complex. According to position and migration of the subaerially exposed forebulge, distribution of the foreland Izvleček UDK 551.44.551.7(497.4-14) 552.541.551.7(497.4-14) Bojan Otoničar: Zakrasela periferna izboklina povezana z razvojem zgornjekredno-paleogenskega predgorskega bazena; Kras, Matarsko podolje in Istra (JZ Slovenija in SZ Hrvaška) V jugozahodni Sloveniji in Istri so kredna karbonatna zaporedja Jadranske karbonatne platforme pasivnega obrobja Jadranske mikroplošče ločena z regionalno diskordanco od zgornjekrednih in paleogenskih karbonatnih zaporedij sino-rogene karbonatne platforme. Razgibano paleokraško površje, ki diskordanco označuje, je lokalno prekrito z boksitom. Pod površjem se pojavljajo različne podpovršinske paleokraške oblike, med drugim večje zapolnjene freatične jame in diskretni horizonti drobnih prepletajočih se kanalčov. Starost apnencev neposredno pod paleokraškim površjem in obseg stratigrafske vrzeli v jugozahodni Sloveniji in Istri sistematično naraščata od severovzhoda proti jugozahodu, nasprotno pa starost apnencev, ki paleokraško površje pokrivajo v tej smeri upada. Preko obravnavanega območja so med campanijem in eocenom od severovzhoda proti jugozahodu napredovala tudi sedimentna zaporedja sinorogenih karbonatnih platform (karbonatne kamnine Kraške grupe) ter pelagičnih laporjev in fiša, ki predstavljajo sedimente podhranjenega predgorskega bazena. Sistematične trende izohron karbonatnih kamnin, ki ležijo neposredno pod in nad paleokraškim površjem in posledično razpona stratigrafske vrzeli lahko v veliki meri razložimo z evolucijo in topografjo periferne predgorske izbokline. Napredujoči feksurni predgorski profl je nastal zaradi vertikalne obremenitve predgorske litosferske plošče (Jadranske mikroplošče) z nastajajočim orogenim klinom. Zaradi sočasnih in postorogenih tektonskih procesov ter časovnega neskladja med sedimenti sosednjih predgorskih bazenov in med različnimi tektonskimi (»orogenimi«) fazami tega dela zahodne Tetide v kredi in paleogenu, je opredelitev tektonske faze, ki je neposredno odgovorna za evolucijo obravnavanega predgorja otežena. Glede na položaj in migracijo periferne iz- 1 Karst Research Institute ZRC SAZU, Titov trg 2, Si-6230 Postojna, Slovenia, e-mail: otonicar@zrc-sazu.si Received/Prejeto: 01.02.2006 TIME in KARST, POSTOJNA 2007, 101–120 BOJAN OTONIČAR related macrofacies and orientation of tectonic structures, especially of Dinaric nappes, and Dinaric mountain chain I suggest that the foreland basin complex in western Slovenia and Istria was formed during mesoalpine (“Dinaric”) tectonic phase due to oblique collision between Austroalpine terrane/Tisia micro-plate and Adria microplate when probably also a segmentation of the foreland plate (Adria microplate) occurred. Key words: forebulge unconformity, paleokarst, chronostrati-graphic gap, fysch, Adriatic Carbonate Platform, synorogenic carbonate platform, foreland basin, Adria microplate, Dinaric orogene, Cretaceous, Paleogene, Sw Slovenia, Istria. Plate tectonics theory had a crucial impact on our understanding of sedimentary basins, and consequently, of carbonate sedimentary systems. Plate tectonics determines not only the gross architecture (dimension and shape) and lithological/structural characteristics of carbonate platforms (Bosellini, 1989), but also their evolution and the longevity. Tose characteristics are largely defned by specifc geotectonic setting in which certain carbonate platform begin to grow. Carbonate platform(s), which colonize certain area through longer or shorter period of geologic history, constantly change its/their position in relation to the equator and plate boundaries and pass through diferent phases of the wilson cycle. Te sedimentary and diagenetic character of the carbonate platform(s) constantly change(s) during this journey and at a stretch, the platform evolution may be stoped. In this case, the area formerly inhabited by the carbonate platform may fall under conditions which are not favourable for considerable carbonate production. In one scenario it may immediately afer the deposition or later in the geologic history be uplifed, subaerially exposed and karstifed. Similarly as the plate tectonics governs the sedimentary evolution of the carbonate platforms, it may also determins their diagenetic evolution, including karsti-fcation. Te gross architecture, lithological/structural caharacteristics, and the evolution and the longevity of the uplifed area with subaerially exposed carbonate platform are mainly dependent on its geotectonic position regard to plate boundaries, former geodynamics and consequently topography of the area, especially of the carbonate platform. Although important for the appearance of the karstic landscape, the efects of other variables, such as climate and ground water level, may be just superimposed on the geotectonically predisposed framework. bokline, razporeditev makrofaciesov podhranjenega predgor-skega bazena ter usmerjenost tektonskih struktur, predvsem Dinarskih pokrovov, in Dinarskega gorstva v celoti domnevam, da je nastal predgorski sistem v zahodni Sloveniji in Istri med mezoalpidsko (»Dinarsko«) tektonsko fazo, kot posledica bočne kolizije med Avstroalpidskim terranom in/ali Tisa mikroploščo ter Jadransko mikroploščo, pri čemer je verjetno prišlo tudi do segmentacije Jadranske mikroplošče. Ključne besede: diskordanca, paleokras, kronostratigrafska vrzel, fiš, Jadranska karbonatna platforma, periferna predgorska izboklina, sinorogena karbonatna platforma, predgorski bazen, Jadranska mikroplošča, Dinarski orogen, kreda, paleogen, jugozahodna Slovenija, Istra. Each karstic landscape carries its specifc geotecton-ic signature which can be read from and explained with specifc evolution of karstic features and a karst system as a whole. In addition, studies of sedimentary successions of rocks that under- and overlie the (paleo-) karstic surface and that of the adjacent sedimentary basins as well as the general geologic conditions of the region may signifcantly improve our knowledge on geodynamics of the uplifed area. Te paper documents an example of paleokarst that occurred during the uplif of the Adriatic Carbonate Platform (sensu Vlahović et al., 2005) in the distant foreland region of the evolving collision related orogenic belt between the Adria microplate (sensu Stampfi et al., 1998) and the Austroalpine terrane and/or Tisia micro-plate (sensu Neugebauer et al., 2001) in the Late Cretaceous and the Early Paleogene. Te study is based on 36 geological profles from the karstic regions of southwestern Slovenia, both Slovenian and Croatian part of Istria peninsula and the area between Trieste bay and Italian-Slovenian border in northeastern Italy (Figs. 1, 2). To get a whole picture of conditions that dominated the region during the emersion period, I expend the area of interest to syno-rogenic carbonate platform that onlap the paleokarstic surface and to siliciclastic fysch regions of afore mentioned areas and the adjacent regions of western Slovenia and northeastern Italy (along the border between Italy and Slovenia). Te aim of this work is to show the causes of the uplif and subaerial exposure of the northwestern part of the Cretaceous Adriatic Carbonate Platform in Late Cretaceous and Early Paleogene. Te data presented here were provided mainly from the studies of paleogeograph-ic and topographic extent of the emersion, stratigraphy of the carbonate successions that immediately under- and INTRODUCTION 102 TIME in KARST – 2007 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION Fig. 1: Geographical position and simplifed geological map of the western Slovenia and Istria showing major structural elements (modifed from Placer, 1999). overlain the paleokarstic surface, stratigraphy and sedi- time of the uplif is correlated with events on the adjacent mentology of the onlapping synorogenic carbonate plat- plate boundaries of the western Tethian domain (tradi-form and adjacent deeper marine basin as well as from tional “orogenic phases”) and global eustatic curve. regional geotectonic and general geologic situation. Te GEOLOGy OF THE AREA Te geology of southwestern Slovenia and Istria has been carbonate successions of diferent Cretaceous formations studied from the late 19th century on. Since that time also from shallow-marine limestones of the Upper Creta-a regional unconformity which separates shallow-marine ceous/Lower Paleogene Liburnia Formation or Eocene TIME in KARST – 2007 103 BOJAN OTONIČAR Fig. 2: Simplifed lithostratigraphic columns of Cretaceous to Eocene successions in southwestern Slovenia and Istria (NW Croatia and SW Slovenia), with one column from NW Italy. Authors of original geological columns are listed below: 1) šribar (1995); Rižnar (1997), 2) drobne (1977, 1979), 3) drobne et al. (1988, 1996); šribar (1995); jurkovšek et al. (1996), 4) jurkovšek et al. (1996), 5) drobne (1981); jurkovšek et al. (1996), 6) jurkovšek et al. (1997), 7) hamrla (1959); drobne (1977, 1979); Pavlovec et al. (1991), 8) hamrla (1959, 1960); jurkovšek et al. (1996), 9) brazzatti et al. (1996), 10) hamrla (1960); drobne et al. (1991); jurkovšek et al. (1996), 11) hamrla (1959); buser & Lukacs (1979); delvalle & buser (1990); jurkovšek et al. (1997); this study, 12) drobne (1977); delvalle & buser (1990), 13) delvalle & buser (1990); šribar (1995); buser & Radoičić (1987), 14) šikić et al. (1972); drobne (1977), 15) drobne (1977); this study, 16) drobne (1977, 1981); hottinger & drobne (1980); drobne & Pavlovec (1979); drobne et al. (1991); turnšek & drobne (1998); this study, 17) drobne (1977), 18) šikič et al. (1972); drobne (1977), 19) biondić et al. (1995), 20) šikić et al. (1972); drobne (1977), 21) šikić et al. (1968); drobne (1977), 22) Pleničar et al. (1969); drobne (1977); Gabrić et al. (1995), 23) Pleničar et al. (1969); drobne (1977), 24) hamrla (1959); Pleničar et al. (1973); drobne (1977); velić & vlahović (1994); matičec et al. (1996), 25) šikić et al. (1968); drobne (1977); hottinger & drobne (1980); drobne et al. (1991), 26) matičec et al. (1996), 27) tarlao et al. (1995), 28) buser & Lukacs (1972); drobne (1977); hottinger & drobne (1980); matičec et al. (1996), 29) Polšak & šikić (1973); drobne (1977), 30) drobne et al. (1991), 31) - 34) matičec et al. (1996), 35) šikić et al. (1968); magaš (1973); šikić et al. (1973); šikić & Polšak (1973); höttinger & drobne (1980); Otoničar et al. (2003), 36) Polšak (1970); drobne (1977); matičec et al. (1996). 104 TIME in KARST – 2007 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION Alveolina-Nummulites Limestone has been known. Te Liburnia Formation, Alveolina-Nummulites Limestone and intermediate Trstelj Beds represent the Kras Group (Košir, 2003) (Fig. 3), which corresponds to the lower unit of the underflled peripheral foreland basin stratigraphy (i.e. the lower unit of the “underflled trinity” of Sinclair, 1997). Tus the unconformity represents a megasequence boundary and typically separates the underlying passive margin carbonate succession from the overlying deposits of the synorogenic carbonate platform at periphery of the foreland basin (Košir & Otoničar, 2001). Te synorogen-ic carbonate platform was fnally buried by prograding hemipelagic marls (i.e. the middle unit of the “underflled trinity” of Sinclair, 1997) and deep-water clastics (fysch) (i.e. the upper unit of the “underflled trinity” of Sinclair, 1997) (Fig. 3). Because the name of the carbonatre platform that overlie the unconformity has not been defned yet, I will use in this paper only the general geodynamic term – i.e. the synorogenic carbonate platform. Fig. 3: Generalized stratigraphic column of Upper Cretaceous-Eocene succession in the Kras (Karst) and matarsko podolje regions, SW Slovenia, showing major lithostratigraphic units (modifed from Košir, 2004). Te unconformity is expressed by an irregular paleokarstic surface, locally marked by bauxite depos- its. Although the unconformity has been repeatedly mentioned, no systematic study of paleokrast has been performed. Relatively numerous papers on biostratig-raphy, especially on the carbonate successions of the Kras Group, have been published (see list of references attached to Fig. 2), yet not more than few attempts on explanation of the sedimentology of the paleokarstic deposits and onlapping beds have been done (Otoničar, 1997; Debeljak et al., 1999; Durn et al., 2003). Only occasionally, the geotectonic conditions under which the paleokarst (uplif) evolved have been briefy mentioned (Košir & Otoničar, 2001; Otoničar & Košir, 2001; Durn et al., 2003). Tectonically, the discussed area corresponds to three macrotectonic units, the Southern Alps, the Ex- Fig. 4: Illustrative geological map showing distribution of fysch deposits and major structural elements in western Slovenia. Te map is based mainly on data from basic geological maps of yugoslavia, 1:100.000, sheets beljak & Ponteba (jurkovšek, 1986), Udine-tolmin & videm (Udine) (buser, 1986), Kranj (Grad & Ferjančič, 1974), Gorica (buser et al., 1968), Postojna (buser et al., 1967), trst (Pleničar et al., 1969) and Ilirska bistrica (šikić et al., 1972). Copyright: Geološki zavod Slovenije (Geological survey of Slovenia), 2002 – All rights reserved. TIME in KARST – 2007 105 BOJAN OTONIČAR Fig. 5: A) Paleogeographical map showing major geotectonic units at Santonian-Campanian boundary in western tethys and central Atlantic (modifed from Neugebauer et al., 2001). b) Geotectonic and paleogeographic units of Adria microplate and adjacent areas. ternal Dinarides and the Dinaric foreland (Placer, 1999) (Fig. 1). while the fysch-related sediments can be followed across the all three units (Fig. 4), the unconformity and the overlying carbonate successions of the Kras Group correspond to the most external thrust unit of the Dinaric fold and thrust belt – the northwestern External Dinarides in southwestern Slovenia, Italian part of the Kras plateau and northeastern Istria, and to more stable foreland domain of the Dinaric mountain belt in other parts of Istria (Figs. 1, 2). Te nappe structure of northwestern part of the External Dinarides comprises fve successively lower and younger thrust units from northeast to southwest: Trnovo Nappe, Hrušica Nappe, Snežnik Trust Sheet, Komen Trust Sheet and Kras Trust Edge (Placer, 1981, 1999, 2002) (Fig. 1). Te External Dinarides and the Dinaric foreland correspond to the northwestern part of the Cretaceous Adriatic Carbonate Platform and the Upper Cretaceous-Eocene synorogenic carbonate platform which occupied northeastern part of the Adria microplate s.s. (Fig. 5). In the Cretaceous the area of present day Southern Alps was a part of deeper marine realm which comprised the Slovenian Basin formed in the Middle Triassic (Cousin, 1981; Buser, 1989) and the area of former Julian Carbonate Platform which was drowned in the Lower and Middle Jurassic (Cousin, 1981; Buser, 1989). Te geologic and paleogeographic situation started to change severely in the Late Cretaceous (see below). It is important to note, that the described region is recently confned from the north side by the Periadriatic fault zone, from the west by the deposits of the Southern Alpine Molasse Basin and from the south and southwest by the Adriatic Sea and its sediments (Fig. 1). To understand the mechanisms that governed the uplif and emersion, regional geotectonic conditions of the wider area of the Late Cretaceous-Early Paleogene western Tethys were taken into consideration. During the Mesozoic, the area between Eurasia and Gondwana or the western part of the extensive Tethys bay of the Pangea was occupied by more or less uniform Adria microplate surrounded by smaller tectonic units or terranes (Fig. 5). with regard to major geotectonic events, the extent and shape of Adria microplate was 106 TIME in KARST – 2007 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION changing constantly through the geologic history. Te results of these events (e.g. tecto-sedimentary successions or cycles) could be correlated between geographically and geologically distant parts of the Adria microplate. Afer substantial Permian to Middle Triassic and Triassic/Jurassic extensional tectonics, the Adria domain became encircled by oceanic bays and dissected by numerous deepwater basins and drowned carbonate platforms (Fig. 5). It is considered that since Early Jurassic the Adriatic Carbonate Platform had been isolated by deeper marine realms (Vlahović et al., 2005). At the Middle/Late Jurassic boundary compression-al tectonic regime prevailed over the peri-Adriatic region. It was caused by the beginning of closure (subduction) of the adjacent oceanic bays of the western Tethys. During the Late Jurassic and Cretaceous gentle broad-scale positive and negative lithospheric defections periodically occurred on the Adriatic Carbonate Platform. Te defections were expressed by coexistence of karstic areas and somewhat deeper marine intra-platform basins (Tišljar et al., 1995; 1998; Vlahović et al., 2005). Distinctive defections correspond to period of ophiolite emplacement [e.g. the Late Jurassic/Early Cretaceous obduction of ophiolite suite of the Dinaric Tethys on the E margin of the Adria microplate (Pamić et al., 1998; 2000)] and distant collisions [e.g. the mid-Cretaceous Eoalpine orogenesis in the Pelso/Austroalpine/Tisia domain (Faupl & wagreich, 2000; Neugebauer et al., 2001)]. Topographic disunity over the platform gave rise to irregular facies distribution and thickness of carbonate successions of diferent parts of the platform. In the investigated area both surface and subsurface pale-okarstic features occur. In places the paleokarstic surface is denoted by surface karst forms like karrens, dolines and depressions of decimetric amplitude (Fig. 6a). Pedogenic features and enlarged root-related channels characterize the upper part of the vadose zone, the epikarst. Vadose channels, shafs and pits penetrate up to a few tens of meters bellow the paleokarstic surface, where they may merge with originally horizontally oriented phreatic cavities. Te latter comprise characteristics of caves forming in fresh/brackish water lenses. At least some of them may be defned as fank margin caves (Fig. 6b, 6c). In extensive outcrops, the remains of such caves can be followed as much as few hundreds of meters along strike. In one case a breccia body which was defned as paleokrastic cave related deposit (Otoničar et al., 2003), is so extensive that was used even as mappable unit for Basic geologic map of Signifcant interruptions of carbonate successions are also related to global eustatic oscillations and/or oceanic anoxic events, but they are mainly superimposed on tectonically induced changes of relative sea-level. Tus before the beginning of the uplif of northern part of the Adriatic Carbonate Platform in the Late Cretaceous and the synchronous onset of fysch sedimentation in the area north and north-eastern of the platform, the whole region was already topographically distinctly heterogeneous. Flysch started to deposit in deeper marine basin with partly inherited bathymetry from former deeper marine domain of Slovenian Basin and drowned Julian Carbonate Platform (Fig. 5). Deeper marine realms with more or less uninterrupted sedimentation had still encircled the carbonate platform from its western and southwestern side (Vlahović et al., 2005) (darker grey area on Fig. 5). Later tectonic activity which shortened the area and displaced diferent parts of the region, prevent more accurate interpretation of geotectonic conditions at those time. Namely, except the substantial shortening of the region due to diferent “thrusting” phases of Alpine oro-geney, the area north from the Periadriatic Fault Zone was displaced for at least 100 km eastward during the Miocene (Ratschbacher et al., 1991; Frisch et al., 1998; Vrabec & Fodor, 2006), in some estimates up to 500 km (Haas et al., 1995). It should be noted that western Istria (i.e. Dinaric Foreland on Figure 1) experienced signif-cant counterclockwise rotation most likely between the end of Miocene and the earliest Pliocene (Márton et al., 1995; Márton, 2006). yugoslavia 1:100.000 (see Magaš, 1965). Te cavities are usually irregular and elongated in shape, and could be up to few tens of meters long and up few meters high (Fig. 6b). Depending on locality, the phreatic cavities were found in diferent positions regarding to the paleokrastic surface, the lowest one some 75 meters below it. Te cavities had been subsequently partly reshaped and entirely flled with sediments and fowstones in the upper part of the phreatic, epiphreatic and vadose zones (Figs. 6b, 6c). Similarly, the vadose channels and voids are also flled by sediments and fowstones, but they usually difer from these of phreatic cavities in higher content of noncar-bonate material, lower ?13C values of carbonate material and more distinctive pedogenic modifcation. Te denudation had frequently exposed flled paleokarstic subsurface cavities on the paleokarstic surface, where they may be identifed only by the remains of their fll (Otoničar PALEO KARST TIME in KARST – 2007 107 BOJAN OTONIČAR et al., 2003) (Fig. 6c). Te internal sediments and fow-stones ofen occur as grains in deposits that cover the paleokarstic surface or fll subsurface paleokarstic cavities of diferent generations. Paleokarstic surface with its depressions as well as subsurface channels and voids are ofen covered and flled by bauxite deposits which were locally exploited (Fig. 6d) (Gabrić et al., 1995). Certain limestone lithofacies of immediate cover of the unconformity are commonly locally confned, sug- Fig. 6: A) Paleokarstic surface is locally denoted by small scale depressions (motorway road-cut at Kozina village, SW Slovenia). Note colour contrast between Upper Cretaceous shallow marine limestone of the Lipica Formation and dark grey palustrine limestone of the Liburnia Formation. hammer for scale is about 30 cm high. b) horizontally oriented cave of irregular shape largely flled with reddish-stained calcareous mudstone/siltstone (Podgrad, matarsko Podolje, SW Slovenia). Te maximal height of the cave is approximately 4 meters. Te cave deposits are artifcially marked by reddish transparent colour on the photograph. C) breccia body represents a part of flled roofess paleokarstic phreatic cave at Koromačno in Istria, NW Croatia. (1,8 m tall geologist for scale in the upper right corner) d) Excavated paleokarstic cavity (vadose shaf?) originally flled with bauxite (minjera, Istria, NW Croatia). 108 TIME in KARST – 2007 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION gesting highly irregular topography of the karstic surface before the beginning of transgression. In places it is clear that the incipient transgression involved gradual increase of groundwater table and, eventually, ponds or “blue holes” were formed in karstic depressions (Durn et al., 2003). In the Kozina site (southwestern Slovenia) during the “blue hole” stage of the transgression, a paleokarstic pit was flled by coarse grained breccia with vertebrate remains, mainly dinosaurian and crocodilian bone fragments and teeth (Debeljak et al., 1999, 2002). Generally, the cover sequence (i.e. the Liburnian Formation of Maastrichtian and early Paleogene age) is characterized by restricted, marginal marine and palustrine lithofacies, which frequently show pedogenic modifcations. EVOLUTION OF THE PERIPHERAL BULGE (THE FOREBULGE) Besides the research on paleokarst related phenomena, the study of sedimentary successions of the host rock in which the paleokarstic features occur and those that overlie the paleokarstic surface is of crucial importance to understand the uplif of substantial part of the Adriatic Carbonate Platform above the sea-level in the Late Cretaceous and Paleogene. To explain the mechanisms that govern the uplif, regional and global geotectonic and eu-static conditions were taken into consideration, too. STRATIGRAPHy Te age of the limestones that immediately underlie the unconformity and the extent of the chronostratigraphic gap in southwestern Slovenia and Istria systematically increase from northeast towards southwest (Figs. 2, 7a, 7b), while the age of the overlying limestones decrease in this direction (Figs. 2, 7c). In western part of Istria the orientation of the isochrones is slightly diferent and VN. »im a i If ^^+jSijf/ / / / J^ I h Fig. 7: A) Isochrones of carbonate rocks that immediately underlie the unconformity. b) Isochrones of the extent of the chronostratigraphic gap. C) Isochrones of carbonate rocks that immediately overlie the unconformity. Isochrones in all fgures are in ma. major structural elements of the area (see Fig. 1) and positions of the geological profles used in the research (see Fig. 2) are also shown in the fgures. TIME in KARST – 2007 109 BOJAN OTONIČAR shows a dome-like topography of the forebulge. Te isochrones represent a statistic result acquired by kriging in Surfer programme version 8.00 (© Golden Sofware, Inc.). Te data were provided from 36 geological profles from the karstic regions of southwestern Slovenia, both Slovenian and Croatian part of Istria peninsula and the area between Trieste bay and Italian-Slovenian border in northeastern Italy (Figure 2 and red dots on Figures 7a, 7b, 7c). Te youngest rocks below the unconformity belong to mid-Campanian and occur in the central and northeastern part of the Kras (Karst) plateau (the Komen thrust sheet) (Fig. 1) (Jurkovšek et al., 1996) and close to Postojna (the Snežnik thrust sheet) (Fig. 1) (Šribar, 1995; Rižnar, 1997) in southeastern Slovenia, while the oldest one, Valanginian and Hauterivian in age, crop out in the western part of Istria (Matičec et al., 1996) (Figs. 2, 7a). Te beds that cover the unconformity correspond to diferent ages, litofacies, members and formations. As mentioned afore, the age trend of the immediate cover is opposite to that of the footwall. In this case the oldest rocks occur in southwestern Slovenia and belong to the youngest stage of the Late Cretaceous - the Maastrich-tian. Towards southwest, progressively younger deposits onlap the paleokarstic surface (Figs. 2, 7c). However, the youngest strata that onlap the unconformity don’t ft exactly with the oldest one immediately below it. with regard to described situation, the chronostratigraphic gap increases considerably from few Ma on the Kras plateau (southwestern Slovenia) to more than 80 Ma in western Istria (Figs. 2, 7b). Te lithofacies of the lower part of the cover sequence (Te Liburnian formation) frequently show features typical of subaerial exposure surfaces, including calcrete, pseudomicrokarst, brecciated horizons and karstic surfaces. Locally, the lowermost subaerial exposure surface of the Liburnija Formation, which shows karstic topography of decimetric amplitude, and the main paleokarstic surface form a composite unconformity. Sporadically, thin coal beds and seams occur in the lower part of the sequence. Although the stratigraphy of the Kras Group, “Transitional Beds” and Flysch (Fig. 3) shows overall deepening of the basin, prominent subaerial exposure surfaces also occur in carbonate successions of Trstelj Beds and Alveolina-Nummulites Limestone (Košir & Otoničar, 1997; Košir, 2003). Much thicker successions of paralic sediments with more frequent unconformities and marsh related sediments occur in southwestern Slovenia and northeastern Istria in comparison with other parts of Istria, yet local variation can be signifcant (Figs. 2, 8). In western Istria, where the chronostratigraphic gap is the most extensive, the foraminiferal limestones frequently lie directly on the paleokarstic surface (Matičec at al., 1996). Te thickness of the Kras Group generally decreases from northeast toward southwest, although also in this case signifcant deviations may occur (Figs. 2, 8). Te point where the unconformity pinch-out towards the foreland basin occurs somewhere between the northeastern part of the Kras plateau on the Komen Trust Sheet and some 10 km (approximately 25 km in original position – see Placer, 1999) distant Mt. Nanos on the Hrušica nappe (Fig. 1). From this point on towards the foreland basin, the uplif of the forebulge didn’t take place because the area was so close to the orogene that experienced only a subsidence. Here, the sedimentary succession of the Adriatic Carbonate Platform gradually passes into progressively deeper-marine carbonate succession of synorogenic carbonate platform. Namely, on the Mt. Nanos at Campanian-Maastrichtian boundary, the deepening of the shallow marine carbonate platform without any evidence of preceding emersion is documented (Šribar, 1995). Further towards the northeast, in the Julian Alps (the eastern part of the Southern Calcareous Alps) and in the most northern part of recent Dinaric mountain belt in western Slovenia and northeastern Italy (the Trnovo Nappe), the turbiditic siliciclastic sediments (fysch) started to deposit in Campanian and Maastrichtian over the rocks of diferent lithology, age and origin (Pavšič, 1994). Flysch ofen overlies deeper marine pelagic marls of “scaglia” type and alodapic carbonates, which were receiving the material from Adriatic Carbonate Platform. It is important to note that in this part of western Slovenia deep-marine basin existed before fysch or above mentioned deeper marine pelagic marls started to deposit. However, the oldest pelagic marls (pre-fysch deposits) which overlie the Upper Cretaceous shallow marine carbonates of the northeastern margin of the Adriatic Carbonate Platform also belong to Maastrichtian. Similar as I stated for chronostratigraphic gap, the pelagic marls and fysch deposits are also diachronous over the area. From northeast toward southwest, successively younger strata onlap the pre-foreland basin deposits (Fig. 4). Te successions of pelagic marls and especially si-liciclastic fysch were periodically interrupted by deposition of calcarenitic and calcruditic beds/megabeds, locally even of olistostrome character. Tose beds were supplied by turbiditic currents from the fault-related escarpments of distorted and seismically active marginal areas of former Adriatic Carbonate Platform (Skaberne, 1987; Tunis & Venturini, 1987) and later also from outer parts of synorogenic carbonate platforms (distally steepened ramps?) (Fig. 9). Te synorogenic carbonate and siliciclastic deposits of other parts of External Dinarides (e.g. Dalmatia) are 110 TIME in KARST – 2007 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION Fig. 8: Lithostratigraphic columns for three adjacent sites in matarsko Podolje and mt. Slavnik (SW Slovenia). Note signifcant variations in thickness of lithostratigraphic units and in time span of stratigraphic gap. younger than these described here. Tey started to deposit not before Eocene (Marjanac & Ćosović, 2000) and probably represent deposits of diferent foreland system or at least of diferent segment of the one described here. DISCUSSION Systematic trends expressed by isochrones showing the age of the carbonate rocks that immediately under- and overlie the paleokarstic surface (Figs. 7a, 7c), and consequently, the extent of the chronostratigraphic gap (Fig. 7b), can be explained mainly by the evolution and topography of peripheral foreland bulge (the forebulge) (Fig. 9). when the foreland continental lithospheric plate is vertically loaded by the fold and thrust belt, it responds with fexure. In front of the evolving orogen an asymmetric foreland basin is formed; the deepest part of the basin (the foredeep) is located adjacent to the orogenic wedge (Fig. 9). Because of the isostatic rebound on vertical loading of the lithosphere, the opposite side of the basin (opposite to the orogenic wedge) is instantaneously upwarped and the bulge with subtle relief is formed, the peripheral bulge or the forebulge. Te bulge is especially well expressed in early, fysch stage of the foreland basin evolution (Crampton & Allen, 1995). while the wavelength of the defection is approximately the same for both, foreland basin and peripheral bulge, the amplitude of the basin subsidence is typically much greater as the uplif of the bulge (Crampton & Allen, 1995; Miall, 1995). If the conditions are suitable, synorogenic carbonate platforms with distinctive ramp topography may colonise the gentle slope of the forebulge toward the fore-deep (Dorobek, 1995). Signifcantly, as the whole complex of the orogenic wedge advances forelandward, the fexural profle produced by the orogenic wedge advances with it. Topography of the forebulge is controlled by numerous factors, among which the rigidity of the foreland lithospheric plate and the rate of emplacement of the load are the most important (Allen & Allen, 1992; Dorobek, 1995; Miall, 1995). An expected maximal height of the forebulge above the sea level (if the foreland plate is at or close to sea-level prior to fexural loading) would be in the range of up to a few tens to few hundreds meters (Crampton & TIME in KARST – 2007 111 BOJAN OTONIČAR Fig. 9: Schematic block diagram of foreland basin complex showing the position of the orogenic wedge, foredeep and forebulge with distribution of macrofacies belts before plate convergence ended (modifed from bradley & Kidd, 1991). Allen, 1995; Miall, 1995). According to topography of the forebulge, the rate of erosion (see white, 2000) and the style of migration of the orogenic wedge, the area of maximal denudation should occur in the central part of the region, which is over-passed by the bulge (Crampton & Allen, 1995). In addition, non-fexural deformations (e.g. reactivation of pre-existing heterogeneities, enhanced defections because of horizontal in-plane stresses…) and inherited topography may signifcantly infuence the evolution and topography of the forebulge (Allen & Allen, 1992; Dorobek, 1995; Miall, 1995; Crampton & Allen, 1995). On Mt. Nanos (Hrušica Nappe; Fig. 1) shallow water rudist limestone of the Adriatic Carbonate Platform gradually passes over limestone with orbitoidiform larger foraminifera into pelagic marls without any emersion at the base of the deepening sequence – the erosional gap reduces to conformity. Te age span of this transition falls within a period of the shortest documented chronostrati-graphic gap between the northeastern part of former Adriatic Carbonate Platform and the overlying synogen-ic carbonate platform (Fig. 2), which extends from mid-Campanian to Late Maastrichtian. Maastrichtian in age are also the oldest pelagic marls which in places directly overlie the Upper Cretaceous shallow water carbonates of former northeastern margin of the Adriatic Carbonate Platform. Although the oldest turbiditic siliciclastic fysch was deposited in a basin with inherited deeper marine bathymetry (former Slovenian Basin) its Campan-ian and Maastrichtian age could be correlated with other incipient foreland related deposits and phenomena. with regard to these criteria and trends of unconformity related isochrones elsewhere (Figs. 2, 7a, 7b, 7c), I suggest that northern part of the Adriatic Carbonate Platform had thrived more or less prosperously till the end of Campanian, when an initial uplif of the forebulge occurred. Te carbonate sediments that had originally been deposited till that time, and are now missing in carbonate successions immediately below the unconformity, had been erased during the paleokarstic period by the karstic denudation processes. According to topography of the forebulge and advancing nature of the foreland geodynamic complex as a whole, the most extensive denudation is expected in the central area over which the forebulge migrates. Te western part of Istria, where the chronostratigraphic gap is the largest and the beds immediately below the unconformity are the oldest (Fifs. 2, 7a, 7b), most probably corresponds to this zone. However, in an ideal conceptual/mathematical model of the forebulge unconformity, the amount of erosion should remain more or less constant over vast area in the central part of the region over-passed by the bulge, and decreases on its distal slope towards back-bulge basin (Crampton & Allen, 1995). Instead, in western Istria the isochrones of the beds underlying the unconformity show distinctive condensation compared to situation in northeastern Istria and southwestern Slovenia (Fig. 7a). I suggest that this is not the result of rapid increase of the amount of footwall eroded but rather of denudation of primarily much thinner Cretaceous carbonate successions in western Istria. Namely, in this part of Istria the carbonate successions are relatively thin (Matičec et al., 1996), partly because of repeating emersions throughout the Cretaceous (Velić et al., 1989) and partly because of reduced accommodation space of Cretaceous shallow marine environments. Evidence of considerable Late Jurassic and Cretaceous land areas in the vicinity of western Istria (probably ofshore form its recent west coast), came also from dinosaur record (footprints and bones) (Della Vecchia et al., 2000; Mauko & Florjančič, 2003; Mezga et al., 2003) and distribution of sedimentary fa-cies of the adjacent peritidal to deeper marine environments of intraplatform basins (Tišljar et al., 1995; 1998). why was the area of western Istria beeing preferentially uplifed during the Cretaceous is still questionable, but the reasons for defections should be searched at adjacent plate boundaries where their reorganisation and difer-ent collision-related events and processes (see Faupl & wagreich, 2000; Neugebauer et al., 2001) produced hori- 112 TIME in KARST – 2007 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION zontal in-plane stresses that may be transmitted many hundreds of kilometers inboard of actual collision (Zei-gler et al., 1995). It is also possible that the central zone of the foreb-ulge and the slope towards back-bulge basin in their fnal position occurred ofshore of recent Istrian west coast. However, we should be aware that the Late Cretaceous Adriatic Carbonate Platform was surrounded from the western side by deeper marine interplatform basins (Vlahović et al., 2005) what might considerably afected the appearance of the forebulge and the back-bulge area. Although the “abnormal thickness” of denuded stratigraphy in western Istria is mainly the result of previous sedimentary history, some uncertainties may also arise from diferential uplif/subsidence of certain parts of the forebulge. Evidence for diferential subsidence along reactivated ancient tectonic structures is for example well documented in carbonate successions of the Kras Group, where the thickness of chrono- and lithostratigraphic units may vary considerably over short distances (Figs. 2, 8). In conclusion I suggest that the denudation exposed the oldest carbonate rocks in the western Istria partly because of specifc evolution (migration) and topography of the forebulge and partly because of primarily thinner carbonate successions in this part of Istria compared to more northeastern parts of the investigated area. Te rate of transgression over the paleokarstic surface is expressed by the isochrones of the strata that onlap the unconformity (Fig. 7c). while the large scale diachro-nism of the onlapping strata shown in Figure 7c is the result of specifc large-scale topography and migration of the forebulge as a whole, local smaller scale spatial differences in the onlap pattern (not observable in Figure 7c) are due to shorter oscillations of relative-sea level and deposition over topographically irregular paleokarstic surface (e.g. dolines, shafs… – a “blue hole phase” of the transgression). Te pattern of the isochrones shown in Figure 7c suggests that the transgression during its earlier stages (southwestern Slovenia and northeastern Istria) was slower compared to its later stages (western Istria). Although subsequent tectonic deformations, such as tectonic shortening, faulting and rotation, substantially afected the area, the rate of the onlap in southwestern Slovenia and northeastern Istria is estimated to about 2-3 km/Ma while in southwestern Istria to about 4-5 km/Ma. we should be aware that some apparent anomalies, especially at terminations of the isochrones may be the result not only of later tectonic deformations of the area but also of limited number of data points which are not uniformly distributed, spatially confned area of the investigation along the strike of the forebulge and defectiveness of statistic method (kriging) used. Slightly diferent orienta- tion of the isochrones in western part of Istria compared to those in southwestern Slovenia and northeastern Istria (Figs. 7a, 7b, 7c) may also be the result of diferent syde-positional or synorogenic orientation of the prevailing stresses (see Marinčić & Matičec, 1991; Matičec et al., 1996) during the Cretaceous and Paleogene and later counterclockwise rotation of the area (see Márton et al., 1995 amd Márton, 2006). In spite of all that, the reasons for diferent stratigraphic pinch-out rate are many sided and may arise from diferential rheologic and structural characteristics of the foreland plate itself, events at collision zone and adjacent plate boundaries, sublithospheric processes and external reasons like eustatic sea-level oscillations and climate changes. In our case it is difcult to determine the exact reason for the increasing rate of the onlap in Lower Eocene, not only because diferent processes may lead to the same result, but also because they can act simultaneously. Long term sea-level fall (i.e. second-order cycle of Haq et al., 1987) may for example slow-down the onlap rate and vice-versa long term sea-level rise may increase the onlap rate. If we observe the eustatic curve for the Cretaceous and Paleogene (Haq et al., 1988) we can notice that the rate of the onlap is in relatively good agreement with mid-Campanian to Late Paleocene second-order fall and Early Eocene rise of the sea-level. However, the foreland basin should progressively widen and pinch-out migration rate would increase also if, for example, the orogenic wedge loaded a progressively stronger elastic lithosphere (Allen & Allen, 1992). Although not all local variations of relative sea-level oscillations and so the onlap rate could be identifed from isochrones in the Figure 7c, they could be observed in the feld. Namely, the subaerial exposure surfaces that periodically interrupt the carbonate sedimentation of the Liburnia Formation refect relative sea-level falls. Short term falls (i.e. third-order cycles of Haq et al., 1987), which were documented in Late Maastrichtian, Late Pa-leocene and Early Eocene (Haq et al., 1988), could cause these unconformities. On the other hand, a few other processes may infu-ence the rate of the onlap. Te forebulge should increase in height and migrate toward the orogenic wedge over time if the foreland lithosphere behaves viscoelastical-ly even when the load is unchanging (Tankard, 1986). However, estimations for time constants of the viscous relaxation of stresses are longer than actual amount of time available for the forebulge migration (Allen & Allen, 1992; Dorobek, 1995). Variation in onlap rate may refect also changes in sediment supply, or within the orogenic wedge, such as the formation of a new thrust complex (Crampton & Allen, 1995) or transition from passive to active thrusting phase. An increase in com- TIME in KARST – 2007 113 BOJAN OTONIČAR pressive in-plane stress produced during convergence also might enhance uplif of the forebulge and causing shoreline regression along its fank (Allen & Allen, 1992; Dorobek, 1995). Evidence of short term sea-level oscillations could also be recognized from the specifc evolution of the pa-leokarst, especially phreatic caves. If the majority of lenticular caves with irregular walls and discrete horizons of spongework or swisse-cheese like vugs on young carbonate islands originated at/in fresh/brackish water lenses (see Mylroie & Carew, 1995), then in our case the major part of the cavities had been emplaced in the vadose zone prior to submergence and burial. Namely, the caves are frequently completely flled with deposits originated in vadose zone, like fowstone and bauxite, or they had been opened to the paleokarstic surface by complete denudation of the roof (i.e. roofess caves of Mihevc, 2001). If the water-level is stagnant and the forebulge migrates, than in the conceptual sense only those phreatic cavities developed below that forebulge fank that facing towards back-bulge basin should be uplifed in the vadose zone before subsidence. On the contrary, phreatic caves developed below the fank facing the foreland basin and the advancing orogenic wedge should sufer nothing but subsidence and subsequent burial. Teoretically it is possible that because of the advancing character of the forebulge, caves formed in diferent sides of the forebulge may occur in the same karstic profle. Phreatic cavities developed below the fank facing towards the back-bulge region should be uplifed and modifed in the vadose zone. Subsequently, afer the crest of the forebulge migrates over the back-bulge fank, the “back-bulge” phre-atic caves should re-immerge into phreatic zone, but this time below the fank facing towards the foreland basin. It is important to note that frequently observed multiphase modifcations of originally phreatic caves could also be the result of the same causes of relative sea-level oscillations that govern the onlap character of the beds that overlie the unconformity (e.g. relaxation of the viscoelas-tic bulge, formation of a new thrust complex, increase of horizontal in-plane stress, eustatic sea-level fall…). Te carbonate platform was subsequently re-established and fnally buried by prograding deeper-marine clastics (pelagic marls and fysch) of the migrating foreland basin (Fig. 9). As it has been already discussed, shallow-water carbonate successions that cover the unconformity may yield a considerable amount of information about relative sea-level oscillations and geodynamics of the forebulge. Paralic/shallow-marine successions with frequent unconformities and palustrine deposits of the Liburnia Formation (Fig. 3) are usually much thicker in southwestern Slovenia and northeastern Istria than in central and western Istria (Fig. 2). Tere the paleokarstic surface is frequently directly overlain by foraminiferal limestones (Matičec at al., 1996). Te general trend of thickness and the rate of transition from shallow to deep marine environments (drowning) (Fig. 2) are in good agreement with the rate of the onlap (Fig. 7c) and should be the result of the same processes that caused the diferentiations in the onlap pattern. I suggest that the anomalies in thickness and facies distribution that could be in places quite distinctive may arise from reactivation of inherited geological structures due to the approaching orogenic wedge. It has been discussed already, that the orogenic phases could be recognised from structural and stratigraphic data even in areas that are located at some distance from the source of tectonic activity at plate boundaries (e.g. collision and orogenesis). Because of later tectonic deformations it is sometimes difcult to defne the exact tectonic phase which afects the area and the actual source of tectonic activity. In our case, the structural and stratigraphic data indicate the evolution of migrating synorogenic foreland basin complex, which should be the result of collision processes and the evolution of the advancing orogenic wedge (see e.g. Allen & Allen, 1992; Crampton & Allen, 1995; Miall, 1995). At frst sight it seems normal to link the foreland complex to tectonic phase that generated structures by mainly NE-Sw compression (mesoalpine phase of some authors; see Doglioni & Bosellini, 1987) and gave rise to Dinaric mountain belt during its fnal stages. However, the Dinaric orogenic belt of which fnal uplif occurred during the Oligocene-Miocene (Vlahović et al., 2005) is supposed to be the result of collision between Tisia and Adria microplates with onset of collision during the Eocene (Pamić et al., 1998; Pamić, 2002), what is also the age of the oldest synorogenic deposits of the “coastal” part of the External Dinarides (Marjanac & Ćosović, 2000). On the contrary, although the nappe structures of western Slovenia and Late Cretaceous – Pa-leogene compressional deformations of northeastern Italy indicate NE-Sw or ENE-wSw compression, and so “Dinaric” orientation of prevailing regional stress, the oldest foreland basin deposits in these regions are much older than those of other parts of the External Dinarides and belong to the latest stages of Late Cretaceous (Pavšič, 1994; Doglioni, 1987; Doglioni & Bosellini, 1987). As it is shown on Figure 4 the age distribution of fysch deposits indicates the advancing nature of foreland basin from northeast towards southwest what is in accordance with “Dinaric” orientation of the prevailing regional stress. while south of Zagreb-Zemplen fault line, the remnants of oceanic lithosphere (i.e. ophiolite melange) as well as subduction and collision related rocks of Internal Dinarides (i.e. the Sava-Vardar zone by Pamić et 114 TIME in KARST – 2007 UPPER CRETACEOUS TO PALEOGENE FORBULGE UNCONFORMITy ASSOCIATED wITH FORELAND BASIN EVOLUTION al, 1998), which could be linked to closing processes of the Vardar Ocean and collision between Tisia and Adria (Pamić, 2000) are widespread, north of Zagreb-Zemplen line no such rock has been found so far. It seems possible that in central Slovenia, in prolongation of the Sava-Var-dar zone, such rocks have been buried by Tertiary sediments and Southern Alpine nappes. In addition, on the NNE side the nappe structure of western Slovenia was cut from its “root zone” by Periadriatic fault. Te “root zone” should be displaced for at least 100 km eastward during the Miocene (Ratschbacher et al., 1991; Frisch et al., 1998; Vrabec & Fodor, 2006). Although, the structural and sedimentary features of eoalpine tectonic phase which culminated in mid-Cretaceous orogeny in the Austroalpine domain (Faupl & wagreich, 2000) and also afected the central and western part of the Italian Southern Alps (Doglioni, 1987; Doglioni & Bosellini, 1987) mostly pre-date the foreland related features and sediments described here, it should In spite of all structural and depositional heterogeneities and subsequent tectonic deformation of the area the paleokarstic unconformity marked by distinctive surface and subsurface paleokarstic features exhibits characteristics typical of a forebulge unconformity: 1) From northeast towards southwest the unconformity cuts progressively older units which are onlapped by progressively younger shallow water carbonates; the chronostratigraphic gap progressively increases. 2) Deepening upward sequences of synorogenic ramp-like carbonate systems overlie the unconformity. In marginal parts of the former Adriatic Carbonate Platform towards the foreland basin, a deepening upward sequence is documented also without intermediate unconformity – here the sequence is conformable because the orogenic wedge was so close that the area experienced only subsidence and forbulge uplif had no taken place. 3) Te foreland basin with siliciclastic turbiditic fysch deposits was developing synchronously with the forebulge and synorogenic carbonate platforms. It was also advancing synchronously in the same direction as they were forebulge and synorogenic carbonate platforms. Te stratigraphy overlying the unconformity (i.e. underflled trinity) representing subsidence in under-flled peripheral foreland basin. 4) Evidence of contemporary seismic activity arises from periodic carbonate resediments (megabeds, olistostromes) fnd in siliciclastic fysch successions. be noted that in Istria Tertiary tectonic cycle (from Eocene on) display distinctively diferent orientation of the prevailing stress than Mesozoic one (Marinčić & Matičec, 1991; Matičec et al., 1996). In conclusion, the foreland basin complex in western Slovenia and Istria was probably formed during me-soalpine (“Dinaric”) tectonic phase, although some infu-ences of eoalpine tectonic phase could be important in earlier stages of its evolution. Te time discrepancy and also the exact orientation of prevailing regional stress are probably the result of oblique collision between Adria and Tisia microplates (and/or Austroalpine terrane?) and/or segmentation of the foreland plate (see Ricci-Luc-chi, 1986; Allen & Allen, 1992). Oligocene to recent tectonic events especially in Dinarides and Apennines, and conter-clockwise rotation of Adria importantly modifed the area formerly occupied by the forebulge, but this is already beyond the scope of this paper. Tey were supplied by turbiditic currents from the fault related escarpments of the forebulge slope (reactivated ancient faults). Besides fexural upwarping because of the isostatic rebound on vertical loading of the foreland lithosphere, other smaller scale fexural and non-fexural deformations signifcantly infuenced the evolution and appearance of the forebulge (incuding its diagenesis and karstifcation), lithofacies distribution and thickness of the carbonate successions above the unconformity. At least some infuence of eustatic sea-level oscillations cannot be excluded. 5) Te subaerially exposed area and the facies belts of progressive forelandward advancing shallow-marine, pelagic, and turbiditic depositional environments ahead of the orogenic front are roughly parallel to the Dinaric mountain chain. However, the Dinaric foreland-related system supposedly began to evolve during the Eocene when Tisia and Adria microplates began to collide what is much later comparing to Late Cretaceous onset of foreland basin evolution and forebulge uplif in western Slovenia and Istria. In Istria the orientation of the prevailing regional stress during Cretaceous tectonic cycle difers signifcantly from Eocene one. I suggest that the foreland basin complex in western Slovenia and Istria was probably formed during mesoalpine (“Dinaric”) tectonic phase, due to oblique collision of Adria and Ti-sia microplates (and/or Austroalpine terrane?) and segmentation of the foreland plate. TIME in KARST – 2007 115 BOJAN OTONIČAR ACKNOwLEDGMENTS Most of this work was performed as part of doctoral dissertation at University of Ljubljana. I am grateful to David Allen, P.A. & Allen, J.R., 1992: Basin analysis - principles & applications.- Blackwell science, p. 451, London. Biondić, B., Braun, K., Vlahović, I., Mlinar, Ž., Andrić, M., Balen, I., Pollak, D., Fuček, L., Oštrić, N., Prtol-jan, B., Šaban, B., Blagus, Z., Dukarić, F., Buljan, R. & Biondić, R., 1995: Inženjerskogeološki model željezničkog tunela Ćićarija.- In: I. Vlahovič, I. Velić & M. 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Grenerczy, J. weber, S. Stein & D. Medak (Eds.), Te Adria microplate: GPS geodesy, tectonics and hazards. NATO Science Series, I V, Earth and Environmental Sciences. Springer, 151-168, Dordrecht. white, w.B., 2000: Dissolution of limestone from feld observations.- In: A.B. Klimchouk, D.C. Ford, A.N. Palmer & w. Dreybrodt (Eds). Speleogenesis - Evolution of karst aquifers. National speleological society, 149-155, Huntsville. Ziegler, P.A., Cloetingh, S. & Van wees, J.D., 1995: Dynamics of intra-plate compressional deformation: Te Alpine foreland and other examples.- Tectono-physics, 252, 1-4, 7-59. 120 TIME in KARST – 2007 COBISS: 1.01 A REVIEw OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS AND ASSOCIATED SUPRASTRATAL DEFORMATION MEDSEBOJNO ZDRUŽENI PORUŠENI PALEOKRAŠKI JAMSKI SISTEMI IN DEFORMACIJE NAD NJIMI LEŽEČIH PLASTI – PREGLED Robert G. LOUCKS1 Abstract UDC 551.44 Robert G. Loucks: A Review of Coalesced, Collapsed-Paleo-cave Systems and Associated Suprastratal Deformation Coalesced, collapsed-paleocave systems and associated supra-stratal deformation appear to be prominent diagenetic/struc-tural features in carbonate sections at/near composite unconformities. Te basic architecture of the system can be divided into two sections. Te lower karsted section, where high-density cave formation took place, is preserved as massive breccias commonly displaying a rectilinear pattern in map view. Te overlying suprastratal deformation section is characterized by large, circular to linear sag structures containing faults and fractures. Regional distribution of coalesced, collapsed-cave systems commonly appears as large-scale (hundreds to thousands of square kilometers in area), rectilinear patterns with areas of concentrated, coalesced breccias separated by relatively undisturbed host rock. Tis pattern may refect development of the paleocave system along fracture swarms. Collapsed-paleocave systems are large, complex features that show broad-scale organization. Te complete paleocave system may need seismic data or large, mountain-scale outcrops to de-fne their architecture and distribution. Key Words: Paleocaves, Paleokarst, karst, suprastratal deformation, cave systems. Izvleček UDK 551.44 Robert G. Loucks: Medsebojno združeni porušeni paleokraški jamski sistemi in deformacije nad njimi ležečih plasti – pregled Medsebojno združeni porušeni paleokraški jamski sistemi in deformacije nad njimi ležečih plasti predstavljajo izrazite dia-genetsko/strukturne oblike karbonatnih zaporedij v bližini sestavljenih geoloških nezveznosti. Osnovno zgradbo posameznega sistema lahko razdelimo na dva dela. Spodnji zakraseli del, kjer je gostota jam velika, je ohranjen v obliki masivnih breč, ki pogosto kažejo v tlorisu vzorec sestavljen iz ravnih odsekov. Za deformirane plasti, ki prekrivajo porušene jamske sisteme, so značilne velike skledaste do škatlaste uleknine, ki jih sekajo prelomi in razpoke. Regionalno gradijo združeni paleokraški jamski sistemi tega tipa vzorec velikega merila (zajemajo območja velika stotine do tisoče kvadratnih kilometrov), sestavljen iz ravnih odsekov in vključuje območja zgoščenih združenih brečastih teles, ločenih z relativno neprizadeto prikamnino. Tak vzorec lahko kaže na razvoj paleokraškega jamskega sistema vzdolž razpok-linskih con. Porušeni paleokraški jamski sistemi predstavljajo velike kompleksne pojave, ki odražajo organiziranost velikega merila. Za opredelitev zgradbe in razprostranjenosti popolnega paleokraškega jamskega sistema teh dimenzij potrebujemo podatke seizmičnih raziskav ali izdanke dimenzij gorovja. Ključne besede: pelokraški jamski sistemi, paleokras, deformacije, jamski sistemi. 1 Bureau of Economic Geology , John A. and Katherine G. Jackson School of Geosciences, Te University of Texas at Austin, University Station Box x, Austin, Texas 78713-8924 U.S.A., Fax: 512-471-0140 , email: bob.loucks@beg.utexas.edu Received/Prejeto: 27.11.2006 TIME in KARST, POSTOJNA 2007, 121–132 ROBERT G. LOUCKS INTRODUCTION At several composite unconformities in the stratigraphic record, carbonate sections display extensive karsting that leads to multiple development of cave systems (Esteban, 1991). Tese cave systems underwent extensive collapse and mechanical compaction with burial. Deformation of the overlying strata is associated with burial collapse of the cave system. Te efects of this suprastratal deformation can be noted 700+ m up section above the karsted interval. Tis review will describe the evolution of cave systems during burial and what the characteristics of the cave systems are at diferent stages of burial. Also the characteristics of suprastratal deformation will be described. Paleocave systems have been investigated by several authors including Lucia (1968, 1995, 1996), Loucks and Anderson (1980, 1985), Kerans (1988, 1989, 1990), wilson et al. (1991) wright et al. (1991), Candelaria and Reed (1992), Loucks and Handford (1992), Lucia et al. (1992), Kerans et al. (1994), Hammes et al. (1996), Maz-zullo and Chilingarian (1996), McMechan et al. (1998), Loucks (1999, 2001, 2003), Loucks et al. (2000, 2004), Loucks and Mescher (2001), McMechan et al. (2002), and Combs et al. (2003). Te review will mainly synthesize material from these studies. CLASSIFICATIONS OF CAVE PRODUCTS AND FACIES Loucks (1999) and Loucks and Mescher (2001) produced classifcations of cave products and cave facies. Loucks (1999) used a ternary diagram (Fig. 1) to show the relationships between crackle breccias, mosaic breccias, chaotic breccias, and cave sediments. Crackle breccias are highly fractured rock, with thin fractures separating the clasts and only minor displacement existing between the clasts. Mosaic breccias show more displacement than crackle breccias, but the clasts can still be ftted back together. Chaotic breccias are com- ton §3 Crackle breccia I gravel | *ä •• % »!_•• Cave-sediment fill Matrix-rich: clast-supported: chaotic breccia Matrix-supported chaotic breccia cd Cave sediment with chips, slabs, and blocks Fig. 1: Cave-sediment flls and breccias can be separated into three end members: crackle breccia, chaotic breccia, and cave-sediment fll. modifed from Loucks (1999) and reprinted by permission of the AAPG whose permission is required for further use.” posed of mixtures of clasts that have been transported vertically by collapse or laterally by fuvial or density-fow mechanisms. Clasts show no inherent association with their neighbors. Chaotic breccias grade from matrix-free, clast-supported breccias; through matrix-rich, clast-supported breccias; to matrix-rich, matrix-supported breccias. Cave-sediment fll can consist of any material, texture, or fabric. Loucks and Mescher (2001) proposed a classifcation of six common paleocave facies (Fig. 2): (1) Undisturbed strata, which are interpreted as undisturbed host rock. In this facies bedding continuity is excellent for tens of hundreds of meters. (2) Disturbed strata that are disturbed host rock around the collapsed passage. Bedding continuity is high, but it is folded and ofset by small faults. It is commonly overprinted by crackle and mosaic brecciation. (3) Highly disturbed strata, which is collapsed host rock adjacent to or immediately above passages. (4) Coarse-clast chaotic breccia that is interpreted as collapsed-breccia cavern fll produced by ceiling and wall collapse. It is characterized by a mass of very poorly sorted, granule- to boulder-sized chaotic-breccia clasts approximately 0.3 to 3 m long that form a ribbon-to tabular-shaped body as much as 15 m across and hundreds of meters long. It is commonly clast 122 TIME in KARST – 2007 A REVIEw OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS AND ASSOCIATED SUPRASTRATAL DEFORMATION supported, but can contain matrix material. (5) Fine-clast chaotic breccia interpreted as laterally (hydro-dynamically) sorted, transported-breccia cavern fll. Characterized by a mass of clast-supported, moderately sorted, granule- to cobble-sized clasts with varying amounts of matrix. Clasts can be imbricated or graded. Resulting bodies are ribbon-to tabular-shaped and are as much as 15 m across and hundreds of meters long. (6) Cave-sediment cavern fll that can be carbonate and/or silici-clastic debris of any texture or fabric and commonly displaying sedimentary structures. Fig. 2: Six basic cave facies are recognized in a paleocave system and are classifed by rock fabrics and structures. modifed from Loucks and mescher (2001) and reprinted by permission of the AAPG whose permission is required for further use.” EVOLUTION OF CAVE PASSAGES Knowledge of the processes by which a modern cave lapsed paleocave passage in the subsurface is necessary passage forms at the surface and evolves into a col- to understand the features of paleocave systems. Loucks (1999) described this evolutionary process (Fig. 3), and the review presented here is mainly from that investigation. A cave passage is a product of near-surface karst processes that include dissolutional excavation of the passage, partial to total breakdown of the passage, and sedimentation in the passage (Fig. 4). During later-burial cave collapse, mechanical compaction takes place. Cave-ceiling crackle breccia Breakout dome Burial cave-ceiling crackle breccia Crackle/mosaic breccia Burial cave-wall crackle breccia breakdown breccia Cave-ceiling collapse and further dissolution Transported breccia and sediment Mechanical compaction Sag, faults, and fractures Fig. 3: Schematic diagram showing evolution of a single cave passage from its formation in the phreatic zone of a near-surface karst environment to burial in the deeper subsurface. modifed from Loucks (1999) and reprinted by permission of the AAPG whose permission is required for further use.” TIME in KARST – 2007 123 ROBERT G. LOUCKS Initial passages form in phreatic and/or vadose zones (Fig. 3). Passages are excavated where surface recharge is concentrated by preexisting pore systems, such as bedding planes or fractures (Palmer, 1991), that form a continuous link between groundwater input, such as sinkholes, and groundwater output, such as springs (Ford, 1988). Cave passages are under stress from the weight of Karst towers Vadose canyon (passage) Cave-sediment fill Solution-enlarged fractures Phreatic zone Phreatic tube (passage) Stream sediment Chamber (room) Cave-ceiling Cave-floor crackle breccia breakdown breccia Fig. 4: block diagram of a near-surface modern karst system. Te diagram depicts four levels of cave development (upper-right corner of block model), with some older passages (shallowest) having sediment fll and chaotic breakdown breccias. modifed from Loucks (1999) and reprinted by permission of the AAPG whose permission is required for further overlying strata. A tension dome, or zone of maximum shear stress, is induced by the presence of the passage or cavity (white, 1988). Stress is relieved by collapse of the rock mass within the stress zone. Tis collapse produces chaotic breakdown breccia on the foor of the cave passage (Figs. 3 and 4). Te associated stress release around the cavity produces crackle and mosaic breccias in the adjacent host rock. As cave-bearing strata are buried, extensive mechanical compaction begins, resulting in collapse of the remaining void (Fig. 3). Multiple stages of collapse occur over a broad depth range. Meter-scale bit drops in wells (indication of cavernous pores) are not uncommon down to depths of 2,000 m and are observed to occur to depths of 3,000 m (Loucks, 1999). Te collapsed passages become pods of chaotic breccia (Fig. 3). Te areal cross-sectional extent of brec-ciation and fracturing afer burial and collapse is greater than that of the original passage because the adjacent fractured and brecciated host rock has become part of the brec-ciated pod. Sag features, faults, and fractures (Fig. 3) occur over the collapsed passages. Sediment-filled passages Breakout dome Breakdown pile EVOLUTION OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS A coalesced, collapsed-paleocave system can be divided into two parts: (1) a lower section of strata that contains collapsed paleocaves and (2) an upper section of strata that is deformed to varying degrees by the collapse and compaction of the section of paleocave-bearing strata (Fig. 5). Te deformed upper section of strata is termed suprastratal deformation (Loucks, 2003) and is discussed in a later section. Cave systems are composed of numerous passages. If the areal density of passages is low, the collapsed cave system will feature isolated, collapsed passages (nonco-alescing paleocave system; Fig. 6). If the cave system has a high density of passages, as is common at composite third-order unconformities (Esteban, 1991; Lucia, 1995; Fig. 6: Schematic diagram showing burial and collapse of low-density cave system (noncoalescing, collapsed-cave system) and reprinted by permission of the AAPG whose permission is required for further use.” 124 TIME in KARST – 2007 A REVIEw OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS AND ASSOCIATED SUPRASTRATAL DEFORMATION Active cave systems Unconformity 4 Phase 1: Modern cave system Phase 2: Multiple near-surface cave systems developed below composite Sg2k Composite unconformity Phase 3: Coalesced, collapsed-paleocave system 100 to >1000 m Loucks, 1999), then upon burial and collapse the system can form large-scale, coalesced, brecciated and fractured breccia bodies upon burial and collapse that are the amalgamation of many passages and intervening disturbed host rock (coalescing paleocave system; Fig. 5). Te bodies are hundreds to several thousands of meters across, thousands of meters long, and tens of meters to more than 100 m thick. Internal spatial complexity is high, resulting from the collapse and coalescence of numerous passages and cave-wall and cave-ceiling strata. Fig. 5: Schematic diagram showing the stages of development of a coalesced, collapsed-paleocave system. modifed from Loucks et al., (2004) and reprinted by permission of the AAPG whose permission is required for further use.” SUPRASTRATAL DEFORMATION Collapse and compaction of cave systems provide potential for development of large-scale fracture/fault systems that can extend from the collapsed interval upward to more than 700 m (Kerans, 1990; Hardage et al., 1996a; Loucks, 1999, 2003; McDonnell et al., in press). Tese fracture/fault systems are not related to regional tectonic stresses. Large-scale suprastratal deformation occurs above the collapsed-cave system. As the cave system collapses during burial, overlying strata will sag or subside over the collapsed area. Tis phenomenon is well documented in mining literature (Kratzsch, 1983; wittaker and Reddish, 1989). Kratzsch (1983, p. 147) presented a diagram (Fig. 7) that shows the stress feld above a collapsed mine passage and associated subsidence. Te overlying stress feld widens from the edges of the excavation, and the overlying strata are under compression directly over the excavation. Near the edges of the excavation, between a vertical line extending from the edge of the cavity and the limit line, strata are under extension (tension). within this zone of stress the overlying strata have the potential to sag, creating faults and fractures for some distance upward, depending on the mechanical properties of the strata and the thickness of the beds within the strata. Fig. 8 is a scatterplot showing a number of examples of the magnitude of subsidence over coal mines. Te graph indicates that subsidence is recorded at horizons more than 800 m above the cavity. Tese data indicate the magnitude of the efect that the collapse of a cavity can have on overlying strata. et Extension »I Compression ,- . . Extension Collapsed mine Fig. 7: diagram of a collapsed mine showing collapsed breccia zone and suprastratal deformation. Te center of the subsidence trough is under compression, whereas the wings are under extension. modifed from Kratzch (1983). Applying the above concept of stress felds over cavities to the collapse of a cave passage during burial sug- TIME in KARST – 2007 125 ROBERT G. LOUCKS gests that similar stress felds will develop. As the cave passage collapses, it has the potential to afect a consider- 1000 500 400 300 200 100 50 40 30 20 10 Mm Wittaker and Reddish (1989) U^ + •^ • • • •- • « « ^• • si • 2 3 Subsidence (m) able number of overlying strata. within a cave system, numerous passages will collapse with burial. Each passage will develop a stress feld above it, and these stress felds will interact to create a larger, combined stress feld. Tis concept was presented by wittaker and Reddish (1989; p. 47), who detailed instances in which multiple mining excavations are collapsing. Te stress feld above a collapsing cave system will be complex because the different cave passages do not collapse and compact uniformly over time. As local areas collapse, diferent stress felds will develop, producing fractures and faults related to that individual stress feld. Resulting suprastratal deformation will show variable fracture and fault patterns within an overall subsidence sag. A unique circular fault pattern above collapsed cave systems is recognized by cylindrical faults (Hardage et al., 1996a; Loucks, 1999; McDonnell et al., in press). Fig. 8: Scatterplot showing thickness of overburden that can be afected by mine collapse. Graph shows a trend of greater subsidence with less overburden. MEGASCALE ARCHITECTURE PATTERNS OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS Coalesced, collapsed-paleocave systems are megascale diagenetic/structural features that can afect more than 700 m of section and be regional in scale. As discussed earlier, the karsted section refects the coalescing of collapsed breccias that formed by collapse of passages and associated disturbed host rock. Te vertical extent of the breccias commonly afects the upper 100 m of section (Loucks and Handford, 1992; Loucks 1999) and as much as 300 m of the total section (Lucia, 1996). Te intensity of brecciation can vary throughout the afected interval. Kerans (1990), Loucks (1999), Loucks et al., 2004), and many others have published descriptions of collapsed, brecciated paleocave zones. Fig. 9 shows examples of cave facies from the Lower Ordovician Ellenburger Group in central Texas (Loucks et al., 2004). Te regional pattern of the collapsed paleocave system is commonly rectilinear (Loucks, 1999). Tis rectilinear pattern is probably an artifact of the original cave system developing along an early-formed fracture system. In a detailed study of a paleocave system in the Fig. 10: Slice map through a collapsed-paleocave system in the Lower Ordovician Ellenburger Group in central texas. modifed from Loucks (2004) and reprinted by permission of the AAPG whose permission is required for further use.” \ Quarry wall 1000 ft 300 m Undisturbed host rock Fractured disturbed host rock Fractured and brecciated rock (coalesced, collapsed cavern) 126 TIME in KARST – 2007 0 1 4 5 A REVIEw OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS AND ASSOCIATED SUPRASTRATAL DEFORMATION Lower Ordovician in central Texas, Loucks et al. (2004) presented maps (Fig. 10) and cross sections of the three-dimensional, fne-scale architecture of a coalesced, col-lapsed-paleocave system. Te coalesced, collapsed-pas-sage breccias range in size to as much as 350 m and are separated by disturbed and undisturbed host rock ranging in size up to 200 m. Lucia (1995) also presented a map of brecciated collapsed passages (Fig. 11) from outcrops in the Franklin Mountains of far west Texas, which displays a crude rectilinear pattern. Tis rectilinear pattern can be seen on seismic data as well. Loucks (1999) presented seismic-based maps from Benedum feld in west Texas that display a rectilinear pattern of sags and circular faults induced by collapse of the Ellenburger paleocave system below (Fig. 12). A similar rectilinear pattern is evidenced on seismic data in Boonsville feld (Fig. 13) in the northern Fort worth Basin in Texas (Hardage et al., 1996a; McDonnell et al., in press). In both the Benedum and Boonesville datasets, suprastratal deformation afects up to 700 m of section above the karsted interval (Figs. 12 and 13). (a)" TJ J '¦ ^Jl 1 ¦ä _^2 ¦ — 1 ' -* 1 ? mm 1 _t * . i 1 * ¦ -- U^ — 5 err i ¦ - ____. (b) (c) Fig. 9: Representative cores from paleocave facies. (a) Crackle-fractured disturbed host rock. (b) Collapsed chaotic breccia with large slabs and cave-sediment fll. (c) transported chaotic breccias in carbonate cave-fll matrix. Sample on right is under Uv light. Samples from Lower Ordovician Ellenburger Group in central texas. modifed from Loucks (2004) and reprinted by permission of the AAPG whose permission is required for further use.” TIME in KARST – 2007 127 ROBERT G. LOUCKS (a) Sag (suprastratal deformation) Upper Ordovidan Montoya Silurian Ftisseiman Upper Ortoviasn Montoya Paleocave trends Approximate area of Great McKelligon Sag (above photograph) ¦ Brecciated McKelligon Canyon, Cindy, and Ranger Peak Formations ] Brecciated Ranger Peak Formation ___ Unbrecciated El Paso Group Fig. 11: (a) Photograph of the Great mcKelligon Sag in the Franklin mountains of far West texas. Photograph and general interpretation are from Lucia (1995) but have been modifed by current author. Tis outcrop is an outstanding example of a collapsed-paleocave system with associated overlying suprastratal deformation. (b) map produced by Lucia (1995) of several paleocave systems within the Franklin mountains. Paleocave trend lines are by current author. CONCLUSIONS Coalesced, collapsed-paleocave systems are megascale diagenetic/structural features that can afect more than 700 m of section and be regional in scale. Te architecture of the complete system can be divided into the lower collapsed zone, where the dense system of caves formed and collapsed with later burial, producing a complex zone of brecciation. Te upper, suprastratal deformation section formed during the collapse of the karsted section. Te overlying strata were generally lithifed, but the sag also afected concurrent sedimentation patterns (Hard-age et al., 1996b). Te deformation in the deformed su-prastratal zone consists of normal, reverse, and cylindrical faults and fractures (Loucks, 1999; McDonnell et al., in press). It is important to emphasize that large-scale structural features can develop above karsted zones and not be related to regional tectonic stresses. 128 TIME in KARST – 2007 j 3000 ft j 900 m A REVIEw OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS AND ASSOCIATED SUPRASTRATAL DEFORMATION (a) (b) fusse -EUenburger Marker Collapsed-paleocave zones -<----------» (shown by missing reflections) ~300 m Fig. 12: 3-d seismic example over an Ellenburger paleocave system from benedum feld in West texas. (a) Second-order derivative map in the Fusselman interval displaying sag zones produced by Ellenburger paleocave collapse. (b) Seismic line showing missing sections (collapse in Ellenburger section), cylindrical faults, and sag structures. Suprastratal deformation is >1,000 f thick in this section. modifed from Loucks (1999) and reprinted by permission of the AAPG whose permission is required for further use.” Coalesced, collapsed-paleocave systems and associated suprastratal deformation are complex systems, and large-scale outcrops or datasets are necessary to defne them. However, with the model presented in this paper, individual data points can lead to recognition that the system is a coalesced, collapsed-paleocave feature. TIME in KARST – 2007 129 ROBERT G. LOUCKS (a) N \ 1500 m (b) |Sag| Top Üiddo-i- Forestburg^i lop Ellenburger ^~ 200 m ^—^ - * 7. '-ml Fig. 13: Suprastratal deformation sag features in post-Lower Ordovician Ellenburger strata in Fort Worth basin in north texas. (a) Curvature map at mississippian Forestburg Limestone horizon displaying sag features and faults produced by collapse in the Ellenburger interval. From mcdonnell et al. (in press). (b) 3d seismic line at 1:1 scale showing sag features produced by paleocave collapse in the Ellenburger section. Line-of-section location is shown by dashed line in Fig. 13a. ACKNOwLEDGEMENTS I would like to express my appreciation to Angela McDonnell for reviewing this manuscript. Lana Deiterich edited the text. Published with the permission of the Di- rector, Bureau of Economic Geology, John A. and Kath-erine G. Jackson School of Geosciences, Te University of Texas at Austin. 130 TIME in KARST – 2007 A REVIEw OF COALESCED, COLLAPSED-PALEOCAVE SySTEMS AND ASSOCIATED SUPRASTRATAL DEFORMATION REFERENCES Candelaria, M. P. & C. L. Reed, eds., 1992: Paleokarst, karst related diagenesis and reservoir development: examples from Ordovician-Devonian age strata of west Texas and the Mid-Continent.- Permian Basin Section SEPM Publication No. 92-33, p. 202. Combs, D. M., R. G. Loucks, & S. C. 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Anderson: 1985, Depositional fa-cies, diagenetic terrains, and porosity development in Lower Ordovician Ellenburger Dolomite, Puckett Field, west Texas.- in P. O. Roehl & P. w. Choquette, eds., Carbonate petroleum reservoirs: Springer-Verlag, 19-38. Loucks, R. G. & R. H. Handford, 1992: Origin and recognition of fractures, breccias, and sediment flls in paleocave-reservoir networks.- in M. P. Candelaria & C. L. Reed, eds., Paleokarst, karst related diagenesis and reservoir development: examples from Or-dovician-Devonian age strata of west Texas and the Mid-Continent: Permian Basin Section SEPM Publication No. 92-33, 31-44. TIME in KARST – 2007 131 ROBERT G. LOUCKS Loucks, R. G. & P. Mescher, 2001: Paleocave facies clas-sifcation and associated pore types.- American Association of Petroleum Geologists, Southwest Section, Annual Meeting, Dallas, Texas, March 11-13, CD-ROM, p.18. Loucks, R. G., P. Mescher, & G. A. McMechan, 2000: Architecture of a coalesced, collapsed-paleocave system in the Lower Ordovician Ellenburger Group, Dean word quarry, Marble Falls, Texas.- Final report prepared for the Gas Research Institute, GRI-00/0122, CD-ROM. Loucks, R. G., P. Mescher, & G. A. McMechan, 2004: Tree-dimensional architecture of a coalesced, col-lapsed-paleocave system in the Lower Ordovician Ellenburger Group, Central Texas.- American Association of Petroleum Geologists Bulletin, 88, 545-564. Lucia, F. J., 1968: Sedimentation and paleogeography of the El Paso Group.- in w. J. Stewart, ed., Delaware basin exploration: west Texas Geological Society Guidebook No. 68-55, 61-75. Lucia, F. J., 1995: Lower Paleozoic cavern development, collapse, and dolomitization, Franklin Mountains, El Paso, Texas.- in D. A. Budd, A. H. Saller, and P. M. Harris, eds., Unconformities and porosity in carbonate strata: American Association of Petroleum Geologists Memoir 63, 279-300. Lucia, F. J., 1996: Structural and fracture implications of Franklin Mountains collapse brecciation.- in E. L. Stoudt, ed., Precambrian-Devonian geology of the Franklin Mountains, west Texas-Analogs for exploration and production in Ordovician and Silurian karsted reservoirs in the Permian basin: west Texas Geological Society 1996 Annual Field Trip Guidebook, wTGS Publication No. 96-100, 117-123. Lucia, F. J., Charles Kerans, & G. w. Vander Stoep, 1992: Characterization of a karsted, high-energy, ramp-margin carbonate reservoir: Taylor-Link west San Andres Unit, Pecos County, Texas.- Te University of Texas at Austin, Bureau of Economic Geology Report of Investigations No. 208, p. 46. Mazzullo, S. J. & G. V. Chilingarian, 1996: Hydrocarbon reservoirs in karsted carbonate rocks.- in G. V. Chil-ingarian, S. J. Mazzullo, & H. H. Rieke, eds., Carbonate reservoir characterization: a geologic-engineering analysis, Part II: Elsevier, 797-685. McDonnell, A., R.G. Loucks, & T. Dooley, (in press): quantifying the origin and geometry of circular sag structures in northern Fort worth Basin, Texas: paleocave collapse, pull-apart fault systems or hydrothermal alteration?.- American Association of Petroleum Geologists Bulletin. McMechan, G. A., R. G. Loucks, P. A. Mescher, & xia-oxian Zeng, 2002: Characterization of a coalesced, collapsed paleocave reservoir analog using GPR and well-core data.- Geophysics, 67, 1148-1158. McMechan, G. A., R. G. Loucks, x. Zeng, & P. A., Me-scher, 1998: Ground penetrating radar imaging of a collapsed paleocave system in the Ellenburger dolomite, Central Texas.- Journal of Applied Geophysics, 39, 1-10. Orchard, R. J., 1975: Prediction of the magnitude of surface movements.- in Proceedings, European Congress on Ground Movement, 39-46. Palmer, A. N., 1991: Origin and morphology of limestone caves.- Geological Society of America Bulletin, 103, 1-21. white, w. B., 1988: Geomorphology and hydrology of karst terrains.- Oxford University Press, New york, p. 464. wilson, J. L., R. L. Medlock, R. D. Fritz, K. L. Canter, & R. G. Geesaman, 1992: A review of Cambro-Orodovi-cian breccias in North America.- in M. P. Candelaria & C. L. Reed, eds., Paleokarst, karst related diagenesis and reservoir development: examples from Or-dovician-Devonian age strata of west Texas and the Mid-Continent: Permian Basin Section SEPM Publication No. 92-33, 19-29. wittaker, B. N. & D. J. Reddish, 1989: Subsidence; Occurrence, Prediction and Control: Elsevier, Development in Geotechnical Engineering, No. 56, p.528. wright, V. P. , M. Esteban, & P. L. Smart, eds., 1991: Pal-aeokarst and palaeokarstic reservoirs: Postgraduate Research for Sedimentology, University, PRIS Contribution No. 152, p.158. 132 TIME in KARST – 2007 COBISS: 1.01 THE wORLD’S OLDEST CAVES: - HOw DID THEy SURVIVE AND wHAT CAN THEy TELL US? NAJSTAREJŠE JAME NA SVETU: KAKO SO SE OHRANILE IN KAJ NAM LAHKO POVEDO? R. Armstrong L. OSBORNE1 Abstract UDC 551.44(091) R. Armstrong L. Osborne: Te world’s oldest caves: - how did they survive and what can they tell us? Parts of an open cave system we can walk around in today are more than three hundred million years old. Common sense tells even enthusiasts like me that open caves this old should not still exist, but they do! Teir survival can be partly explained by extremely slow rates of surface lowering, but this is not sufcient by itself. Isolation by burial and relative vertical displacement by faults are probably also required. Now one very old set of caves have been found, are there more of them? what can they tell us? Key words: speleology, oldest cave, survival of old caves. Izvleček UDK 551.44(091) R.A.L. Osborne: Najstarejše jame na svetu: kako so se ohranile in kaj nam lahko povedo? Deli odprtega jamskega sistema, po katerem se lahko danes sprehajamo, so stari več kot 300 milijonov let. Zdrav razum celo takemu navdušencu, kot sem jaz, pove, da tako stare odprte jame ne morejo obstajati, a vendar so! Da so se ohranile, je lahko deloma vzrok v izredno počasnem zniževanju površja, toda to samo po sebi ni dovolj. Jama je morala biti najbrž tudi zasuta in s tem odrezana od sveta, potreben pa je bil tudi relativen navpičen premik ob prelomih. Zaenkrat je bil najden en sam niz zelo starih jam, ali jih je morda še več? Kaj nam lahko povedo? Ključne besede: speleologija, najstarejša jama, ohranitev starih jam. INTRODUCTION In June 2004, when I last spoke here at Postojna about dating ancient caves and karst I found it difcult to not to reveal the exciting discovery which this paper follows (see Osborne, 2005). My collaborators and I had been convinced since mid 2001 that sections of Jenolan Caves in eastern Australia had formed 340 million years ago. we had to ensure that our story was published and that we could convince others. Te issue was not whether the dates themselves were correct, but did the evidence really mean that the caves containing the clays were of such a great age. Tis took four years of intensive work on the clays and additional dating. Now afer the publication of the results (Osborne et al, 2006), and the following media interest; it seems appropriate to refect on the signifcance and implications of the survival of Early Carboniferous open caves. 1 R.A.L. Osborne, Faculty of Education and Social work, A35, University of Sydney, NSw 2006, Australia; e-mail: a.osborne@edfac.usyd.edu.au Received/Prejeto: 27.11.2006 TIME in KARST, POSTOJNA 2007, 133–142 R. ARMSTRONG L. OSBORNE THE POTENTIAL FOR CAVES/SECTIONS OF CAVE TO HAVE A GREAT AGE Despite many years of working on palaeokarst, I initially found the Early Carboniferous (340 Ma) K-Ar dates for unlithifed clays in Jenolan Caves incredible (Figure 1). Fig. 1: Plastic illite–bearing clay, mustard yellow, in the River Cave, jenolan Caves, NSW Australia. Te < 2µm fraction of this clay was K-Ar dated by Osborne et al., (2006) at 357.30 ± 7.06 ma. As I pointed out in 2004 (Osborne, 2005), some Permian landforms do survive relatively intact in Australia. Even a Late Carboniferous age would not have been too surprising, as a Late Carboniferous landsurface has been exhumed at Jenolan from below the overlying Permo-Trias-sic Sydney Basin. An Early Carboniferous age seemed challenging for two main reasons: 1 Te 340 Ma age sits in the middle of the accepted timing for the last folding event in the area (350-330 Ma). Not only the caves, but also the relatively undeformed and well-lithifed caymanite deposits they intersected had to be younger than this event. Te clay dates upset the accepted chronology for the area and suggested that the last folding was older than previously thought. 2 Te 340 Ma age is older than the accepted emplacement age for the adjacent Carboniferous granites (320 Ma). Te plateau surface adjacent to the caves intersects granite plutons. why didn’t the process that exposed the plutons wipe out the ancient caves? My opponents believed that while other landforms in Australia were old, the caves were not. Tey argued that there was no demonstrably old sediment in the caves. I have already discussed this argument elsewhere (Osborne, 1993a, 2002, 2005). Te Early Carboniferous clays from Jenolan are the frst evidence for ancient sediments in Australian caves accessible to humans, but they make the problem of the survival of ancient caves even more difcult, because they are so very old. If we think about the geological history of karstif-cation at Jenolan then the formation of caves in the Carboniferous should not be surprising. Te best dates for the Jenolan Caves Limestone put it in the Latest Silurian (Pridoli, 410-414 Ma)(Pickett, 1982). As well as telling us about the 340 Ma event, the KAr clay dating indicated that the limestone underwent a pre-tectonic period of cave development in the Early Devonian before 390 Ma when the caves were flled with the unconformably overlying volcaniclastics. Tere was also a post-tectonic period of ancient speleogenesis before a marine transgression flled the second generation of caves with lime-mud and crinoidal debris. I suspect if we had announced a third-phase of lithifed palaeokarst some 340 million years old at Jenolan, there would have been little reaction, although the problem of its survival and the problem with the timing of folding would have been the same as the problems with our relict sediments. It would not be surprising for limestone anywhere in the world to have undergone speleogenesis some 70 Ma afer its deposition. Te development of a modern cave in Late Cretaceous limestone is hardly unusual. So, what is the problem? I suspect that while geo-morphologists think surface lowering will destroy old caves, many geologists expect that: 1 open caves fail relatively quickly by breakdown (by analogy with mines and quarries) 2 palaeokarst caves only survive because they are flled with rock; the rock supports the roof preventing destruction due to breakdown. 3 cave sediments become lithifed quickly, so old unlithi-fed relic sediments cannot exist Tese ideas are refuted by the fndings of palaeo-karst workers, surface cavers and the oil industry so I will not expand on them here, rather I will concentrate on geomorphological challenges to the survival of 340 million year old caves. 134 TIME in KARST – 2007 THE wORLD’S OLDEST CAVES: - HOw DID THEy SURVIVE AND wHAT CAN THEy TELL US? HOw COULD THEy SURVIVE? wHy CAVES MAy SURVIVE LONGER THAN SURFACE LANDFORMS Landforms are always under threat from the processes of weathering, incision and surface lowering. weathering in the normal sense of the word is irrelevant in karst since, except in the case of Nadja’s incomplete solution (Zupan-Hajna, 2003), carbonate weathering results in almost total removal of the rock mass. Incision may re-activate or expose ancient caves, but will rarely afect enough of the rock mass to lead to the destruction of ancient caves. It is surface lowering that is the greatest threat to ancient caves and the main process that leads to their late stage modifcation into unroofed caves. what processes may protect caves from surface lowering? Protection by the rock mass Since caves form below the surface, there is a thickness of rock between them and the zone where surface lowering is progressively removing the surface of the Earth. Tis means that caves have a head start in survival compared with surface landforms of the same age. Caves unroofed at the surface are always substantially older than the surface in which they are exposed. Isolation and “karst resistance” Not a lot happens once a cave space enters the vadose zone, there may be breakdown or speleothem deposition, but many cave openings just sit there, inactive while the water is directed through active conduits at a lower level. Te “god” that protects cave walls Apart from speleothem and lithifed sediments that may outlive all of the cave they formed in (Figure 2), it is the walls of a cave that survive the longest, right up to the very last stage of an unroofed cave (Figure 3). why don’t the cave walls fail and simply fall into the void beside them and why don’t they allow the whole cave to fll with speleothem during its siesta in the vadose zone? Some process must protect cave walls from failure and penetration by potentially lethal vadose fow. I am indebted to Andrej Mihevc for the concept of a ‘”god” that protects cave walls’. I am sure this god is a useful addition to the karst panoply. Tree factors are probably important for the survival of cave walls, particularly in teleogenic karsts: - • rock strength • Slow and gentle cave excavation, leading to gradual stress release (caves are not mines or tunnels) • Degassing and precipitation from seeping water makes cave walls self-sealing Fig. 2: Speleothem, exposed on surface above dip Cave, Wee jasper, NSW, Australia. Cave entrance can be seen top of photo. Tis speleothem has outlived all of the cave it formed in. Some cave walls do fail for a variety of reasons. we can observe this in many breakdown chambers and it is possible to recognise the sources of the weakness in the walls that resulted in their failure. Lack of substantial entrances Some caves, e.g. cryptokarst caves of thermal /hydrother-mal origin, may have no entrances or very poor connection to the surface. If there is no entrance or surface connection then surface processes cannot get in and modify the cave. Entrance Blockages It is very easy for cave entrances to become blocked. Pro-grading entrance facies talus cones reaching the ceiling, talus from the surface or breakdown, growth of fow-stone masses, logs, vegetation and biogenic deposits such as guano piles can all easily block cave entrances. with a small amount of vadose cementation, these blockages can become efectively permanent and the cave can become isolated. TIME in KARST – 2007 135 R. ARMSTRONG L. OSBORNE Fig. 3: Looking towards the surviving cave wall from the foor of an Carso, Italy. Protection by flling If a cave is flled with easily removed material, it is possible for the cave to remain “fossilized” for a geologically signifcant time and then become exhumed. If the fll is impermeable to vadose seepage, it will not become cemented. Even if it is cemented, if the fll contains minerals that are unstable when exposed to oxygen-rich vadose water it can be removed from the cave with little efect on the enclosing walls. Protection by cover/burial Cover by sediments, volcaniclastics or lava fows can protect not only the caves, but also surface karst land-forms. For the process to be efective, the cover must be removed without a great efect on the underlying older karst. It helps if the cover consists of relatively weak rock or of rock that is easily weathered. An outstanding example of this process is the burial by Permian basalt and later exhumation of the Shinlin karst in southern China. DENUDATION RATES Both biblical prophets and geomorphological pioneers predicted a fat future, the “rough places a plain” of Isaiah 40:4 and the peneplanation of w. M. Davis. while peneplanation may be out of favour, surface lowering is a real phenomenon. Te problem for survival of old caves is that even with the slowest rates of surface lowering most Mesozoic and all Palaeozoic caves should have been destroyed, except those that have been deeply buried and later exhumed following tectonic movements. In some parts of Australia, extremely low denudation rates apply. wilford (1991) reported rates as low as 0.5 metres per million years in the Ofcer Basin of western Australia over the last hundred million years. unroofed cave, trieste Surface lowering rates in the eastern Australian highlands, where Jenolan Caves are located, are said to range between 1-10 metres per million years (Bishop 1998). If this is so, then the limestone exposed at the surface today in these areas was between 65 and 650 metres below the surface at the end of the Meso-zoic. while these rates are slow by world standards, they are not slow enough to account for the survival of extremely old features. Surface lowering and early incision may be slower than we think Studies of past erosion rates in the Shoalhaven Catchment in eastern Australia by Nott et al., (1996) show that we must approach incision and denudation with some care. Teir relevant fndings are that: • summit lowering and scarp retreat were insignifcant when compared to the process of gorge extension • the rate of summit lowering was 250 times less and the rate of scarp retreat was 15 times less than the rate of headward advancement of gorges • stream incision in the plateau upstream of the erosion head is very slow compared to the rate of gorge extension • there was “insignifcant lowering of the interfuves throughout the Cainozoic” (Nott et al., 1996, p 230) • “Over the long term, the highlands…will become considerably more dissected well before they decrease substantially in height or are narrowed” (Nott et al., 1996, p 224) Te stream incision rate is important when we consider the age of relict caves. If incision rates early in the history of the landscape are much slower than at later stages, present incision rates will lead us to seriously underestimate the age of relict caves located high in the sides of valleys. If lowering of interfuves, i.e. surface lowering, is much slower than incision, scarp retreat and nick-point recession then plateau karst, high level caves and surface caves exposed on hilltops could be very much older than we have previously thought. In dissected terrains the caves will not just be as old as the hills, but considerably older. TECTONIC PROCESSES ARE NECESSARy FOR ExTREME SURVIVAL Low denudation rates, low relief and low rainfall, the Australian trifecta, can only go so far to preserve old 136 TIME in KARST – 2007 THE wORLD’S OLDEST CAVES: - HOw DID THEy SURVIVE AND wHAT CAN THEy TELL US? landforms. Stephen Gale recognised this point: “Although low rates of denudation are an important factor in ensuring the survival of ancient landscapes, this alone is inadequate as an explanation of the maintenance of landforms over ten and even hundreds of millions of years” (Gale, 1992, p 337). Gale went on to discuss how denudation needed to be localized if old landsurfaces were to survive. One way the landsurface can be isolated from surface lowering is through the relative adjustment of adjacent blocks by faulting. Te Fault-Block Shufe Te problem at Jenolan is the elevation of the old caves relative to the adjacent plateau surface. Te plateau surface to the south of Jenolan Caves exposes and intersects post-tectonic Carboniferous granites, thought to be 320 million years old. Figure 4 is a cartoon drawn to explain in simple terms how the caves may have survived. Te caves must have been relatively close to the surface when the cupolas formed and the volcanic ash that formed our old clays entered them (Step 1 in Figure 4). Fig. 4: Cartoon of postulated events at jenolan Caves to explain the survival of caves with Carboniferous clays 1 Cave excavated by thermal processes following folding of limestone 2 volcano erupts; tephra falls to ground and enters caves. 3 Fine tephra begins to fll caves and reacts with water in caves to produce clay minerals. Tese clays have been dated at 340 million years. 4 volcano stops and begins to be eroded. Te caves are full of clay. Granite intrudes the rock near the caves (? 320 ma). 5 Te rock mass containing the granite moves up along the fault, while the rock mass containing the caves moves down. 6 Late Carboniferous: At least 8 kilometres thickness of rock is eroded away, probably partly by glaciation. Tis cuts of the top of the granite and brings the cave back close to the surface. 7 Late mesozoic: valleys erode into the surface and a new stream cave forms below the level of the flled cave. Te clays, still sof, are undermined. Tey fall down and are carried way by the stream. 8 today: Almost all of the 340 million year old clay has now been removed from the caves, small remnants are found and dated. TIME in KARST – 2007 137 R. ARMSTRONG L. OSBORNE Even if the granites did form close to the surface, something between hundreds of metres and a few kilometres of rock must have been removed from the plateau surface to expose the granite. Tis amount of surface lowering should have removed any older caves, particularly those shallow enough to fll with surface-derived sediment. For the caves to survive there must have been a relative change in elevation between the mass of rock intruded by the granite and the mass of rock hosting the caves (Step 5 in Figure 4) before signifcant regional denudation took place. For the sake of simplicity and because the history is not well understood, several steps have been lef out when speaking here in 2004 (Osborne, 2005) I suggested a number of characteristics of localities where one might expect to fnd very old caves, interestingly Jenolan has only some of these. So how might we recognize “funny old caves” and ancient cave sediments? “ABNORMAL CAVES” AND “ABNORMAL” SECTIONS OF “NORMAL” CAVES My work on palaeokarst in caves and on non-fuvial cave morphology frequently takes me to caves that others regard as unusual. Te Carboniferous clays from Jenolan are found in cupolas and other non-fuvial sections of the caves. Interestingly, these same sections of cave also intersect caymanite palaeokarst. Fieldwork on non-fuvial morphology in Europe during 2005 took me to Belianska Cave in Slovakia and Račiška pečina in Slovenia. Co-incidentally, (or not) these are the same localities where Pavel Bosak and co-workers have found the oldest relict cave sediments in Europe (see Bella et al., 2005 & Bosák et al., 2005). Non-fuvial caves, the per ascensum caves of Ford (1995), are characterised by being isolated from or poorly integrated with the modern hydrological system. Some have no natural entrances, while others have poor connection or secondary breakdown entrances. Tis gives them a head start in the survival stakes when compared with fuvial caves. Generally odd caves may survive longer than normal ones. THE OLDEST CAVES ARE NOT ALwAyS AT THE TOP when I frst discovered the caymanite deposits in Jenolan Caves in the 1980s, I could not understand why they were intersected by cave passages at low levels in the limestone mass, not by (older) high-level passages. I did not realize in Figure 4 between Step 6 and Step 7. In the Late Carboniferous, the upper sections of the present valleys were incised and fuvial caves formed. Tese flled with gla-ciofuvial sediment and the whole landscape was buried under the Sydney Basin. In the late Mesozoic, the Sydney Basin was stripped back and the valleys re-juvenated. New fuvial caves formed below the level of the old flled ones (Step 7 in Figure 4). Underhand stoping has now removed most of the old clay and only tiny remnants of clay remain in the caves. then that while level in the landscape is a good indicator of the age of fuvial caves, it has little to do with the age Fig. 5: Palaeokarst sandstone flling spar-lined tube intersected by more recent cave in the entrance area of Lucas Cave, jenolan Caves, NSW, Australia. Te strongly cemented sandstone is younger than the plastic clay shown in Figure 1. wHERE ARE THE OTHER OLD CAVES? 138 TIME in KARST – 2007 THE wORLD’S OLDEST CAVES: - HOw DID THEy SURVIVE AND wHAT CAN THEy TELL US? of non-fuvial caves. In fuvial caves you look to the top for the old sections of cave, but in non-fuvial caves, you must look high and low. RECOGNISING OLD SEDIMENTS How can we recognise very old relict sediments in caves? Te old clays at Jenolan were not found by looking for old material, we were originally looking for unusual minerals. Te clays that looked diferent contained larger than normal amounts of illite and so we were able to date them. Afer the frst old date, samples were chosen strategically, to get the maximum amount of chronological information from the minimum number of samples. Tis was only possible because there were existing pal-aeokarst and cave morphology stratigraphies to test (Os-borne, 1999). GEOLOGICAL HISTORy OF THE CAVES During the 1980s and 1990s, the aim of my research on palaeokarst was to show that speleogenesis and karstif-cation in eastern Australia had a geological history (Os-borne 1984, 1986, 1991b, 1993 a & b, 1995, 1999). Tat is, palaeokarst deposits intersected by “modern” accessible (open) caves indicate repeated periods of cave development at the same locality over periods of hundreds of millions of years. Cavities flled with strongly lithifed palaeokarst deposits represented the older periods of cave development. Te discovery of 340 million year old clays in open accessible caves at Jenolan (Osborne et al., 2006) demonstrated something signifcantly diferent. Te open caves themselves, not just cavernous karsts, can have developmental histories extending over geologically signifcant periods of time (i.e. hundreds of million years). Not much happens during the life of an old cave; they just snooze like an old pet cat. Sometimes dramatic events above, below or beside the cave may wake it from its slumber and leave their mark for us to fnd in the future. GEOLOGICAL HISTORy FROM THE CAVES Much has been said about the potential of the strati-graphic, geomorphic and climatic record in caves. Even the most generous previous estimates for the age of caves (not palaeokarst) suggested that such evidence would be limited largely to the younger end of the Cainozoic, and might perhaps in places like eastern Australia with old landscapes extend to the late Mesozoic. Te survival of Palaeozoic open caves presents a new vista of using caves Unconsolidated Relict Sediments May Be Older than Lithifed Palaeokarst Deposits In my last presentation here, I raised the idea of the lithi-fcation trap: the idea that strongly lithifed cave deposits and palaeokarsts may be younger than some unconsoli-dated or uncemented cave sediments (Osborne, 1995). Tis makes sense if we think about fowstone growing over mud and recognise that cementation, rather than compaction is the main agent of lithifcation in caves. Above ground geologists ofen fnd this idea conceptually challenging. At Jenolan Caves, a crystal-lined cave passage is flled with strongly cemented sandstone (Figure 5). we have no problem with the sandstone being younger than the crystal, but stratigraphy suggests that this sandstone is younger than the unconsolidated clay shown in Figure 1. as a source of geological information. Both ancient caves and palaeokarst deposits could contain records of “missing sequences” for which there is no other record. while there has been signifcant progress in reading the ancient record of palaeokarst, lack of suitable dating techniques and a lack of expectation make geological history from the caves an open and uncultivated feld. Evidence for Global Events Cave sediment research, particularly in the UK and Australia, began with a focus on a geological problem of global signifcance. Today we call it the Pleistocene extinction. Te protagonists at the time saw it in terms of the “deluge” and the extinction or not of “antediluvian” faunas (see Osborne 1991a). Caves were an obvious focus for this research as Pleistocene vertebrate fossils occur in great abundance in the red earths of caves throughout the globe. If the surface of some interfuves dates back to the Mesozoic, then ancient caves have the potential to contain evidence of the K-T boundary. what signal should we expect to fnd in the caves from the K-T event and how would we recognise it? Commentators have suggested that the K-T event involved dramatic changes in the pH of meteoric water, with strongly acidic rain falling from the sky. If this were sustained it should have lef an imprint of extreme surface karstifcation and enhanced vadose and fuvial speleogenesis. Given how efectively caves have trapped Pleistocene loess, we might also expect to fnd iridium-rich silt in caves that were open at the K-T boundary. I don’t know if anyone has looked, but perhaps they should. wHAT CAN THEy TELL US? TIME in KARST – 2007 139 R. ARMSTRONG L. OSBORNE Caymanites & unknown transgressions Lazlo Korpas has been able to make great progress in understanding the evolution of the karst of Hungary by dating caymanites, because these contain fossils and they correlate with magnetostratigraphy (Korpas, 1998, Korpas et al., 1999). Caymanites provide very useful evidence for marine transgressions (Korpas, 2002). Caves intersect caymanites in at least six karst areas in eastern Australia. None of the caymanites have been directly dated. Te 340 Ma old caves at Jenolan intersect caymanites, indicating a minimum age. Te eastern Australian caymanites indicate one or more marine transgressions, probably in the Early Carboniferous for which there is no other geological evidence. Volcaniclastic cave sediments/palaeokarst Given the close physical relationship between stratovol-canoes and carbonate terrains in island arcs and active margins, volcaniclastic cave sediments and palaeokarst deposits should be common in both modern and ancient island arcs and active margins. Tere seems, however, to we still know very little about extremely ancient caves. Tere are good prospects for making new geological discoveries in very old caves. All we have to do is identify funny old sediments in funny old caves, ascertain their meaning and fnd ways to date them. Tis sounds easy, but it is not. Te Jenolan team consisted of a karst geologist, a dating guru (essential so there is no argument about the technical aspects of the dates) and two mineralogists. It be scant reference to such deposits in the literature. Perhaps this is due to the concentration of karstological effort on Tethyan karsts. Volcaniclastic cave sediments and palaeokarst deposits should be expected to occur around the Pacifc rim, particularly in volcanically active island arcs e.g. Indonesia, Philippines, Malaysia, Japan, New Zealand and in southern Europe (Mts Etna and Vesuvius). Tey should also be expected where I work in the early Palaeozoic island arc environments of the Tasman Fold Belt of eastern Australia. while andesitic and silicic stratovolca-noes are likely to be the most common sources of tephra for volcaniclastic deposits in caves and karst depressions, basaltic tephra can also fll caves. Five volcaniclastic palaeokarsts and volcaniclastic relict sediment deposits, including the 340 million year old clays, have now been recognised in eastern Australia (Table 1). It seems likely that more will be recognised, given that many of the cavernous Palaeozoic limestones are overlain by volcaniclastics. took six frustrating years and a sponsor with deep pockets to get the work completed and published. A new world of geology of and from ancient caves awaits those with a stout heart, a thick skin, a good sponsor and eyes for caves and sediments that don’t seem quite right; something like the qualifcations for Antarctic explorers. tab. 1: volcaniclastic Palaeokarst and Relict Cave Sediments in eastern Australia Type Likely Age Karst Area Chemistry Reference Pk ? Tertiary Crawney Pass Basaltic observed by author Pk Mid Devonian Jenolan Silicic Osborne et al. 2006 R Early Carboniferous Jenolan Silicic Osborne et al. 2006 Pk Mid Devonian Wombeyan Silicic Osborne, 1993 Pk ? Wellington Silicic Osborne in prep Pk = palaeokarst R= relict cave sediment SPECULATION 140 TIME in KARST – 2007 THE wORLD’S OLDEST CAVES: - HOw DID THEy SURVIVE AND wHAT CAN THEy TELL US? ACKNOwLEDGEMENTS Tis paper was presented at the Time in Karst symposium at the Karst Research Institute, Postojna, Slovenia in March 2007. Te University of Sydney Overseas Travel Grant Scheme and Top-Up funding for the Faculty of Education and Social work supported attendance at the symposium. Tis paper arises from the dating of clays at Jenolan Caves (Osborne et al., 2006). Many thanks are due to my co-workers, Horst Zwingmann, Ross Pogson and David Colchester. Final corrections to the Jenolan Clay paper were made in Europe in the second half of 2005. I wish to thank colleagues in the Czech Republic, Hungary, Slovakia and Slovenia for their assistance and support. work on Belianska Cave with Pavel Bella, Peter Gazik, Jozef Psotka and Stanislav Pavlarčik assisted in developing the ideas presented here. Other inspiration came when Karel Žác showed me the caves of the Bohemian Karst and Lazlo Korpas showed me his caymanite sequences and well-dated unconsolidated old sediment. Andrej Mihevc engaged in lively discussions about surface lowering, the “god that protects cave walls” and the origins and survival of Račiška pečina. Penney Osborne read the drafs. REFERENCES Bella, P., P. Bosák, P. , J. Glazek, D. Hercman, T. Kiciniska, & S. Pavlarcik., 2005: Te antiquity of the famous Belianska Cave (Slovakia). Abstracts, 40th International Speleological Congress, Athens-Kalamos 21-28 August 2005: 144-145. Bishop, P. , 1998: Te eastern highlands of Australia: the evolution of an intraplate highland belt. Progress in Physical Geography 12, 159-182. Bosák, P. , P. Pruner, A. Mihevc, N. Zupan-Hajna, I. Hora-cek, J. Kadlec, O. Man, & P. Schnabl., 2005: Palaeo-magnetic and palaeontological research in Račiška pečina Cave, Sw Slovenia. Abstracts, 40th International Speleological Congress, Athens-Kalamos 21-28 August 2005: 204. Ford, D.C., 1995: Paleokarst as a target for modern karst-ifcation. Carbonates and Evaporites 10, 2, 138-147. Gale, S.J., 1992: Long-term landscape evolution on Australia. Earth Surface Processes and Landforms,17, 323-343. Korpas, L., 1998: Palaeokarst Studies in hungary. Geological Institute of Hungary, Budapest. Korpas, L., 2002: Are the palaeokarst systems marine in origin? Caymanites in geological past, pp.415-424, in F. Gabrovšek [ed.] Evolution of Karst: From Prek-arst to Cessation, Založba ZRC, Ljubljana. Korpas, L., M. Lantos, & A. Nagymarosy., 1999: Timing and genesis of early marine caymanites in the hydrothermal palaeokarst system of Buda Hills, Hungary. Sedimentary Geology 123, 9-29. Nott, J., R. young, & I. McDougall., 1996: wearing down, wearing back and gorge extension in the long-term denudation of a highland mass: quantitative evidence from the Shoalhaven catchment, southeast Australia. journal of Geology 104, 224-232. Osborne, R.A.L., 1984: Multiple karstifcation in the Lachlan Fold Belt in New South wales: Reconnaissance evidence. journal and Proceedings of the Royal Society of New South Wales 107, 15-34. Osborne, R.A.L., 1986: Cave and landscape chronology at Timor Caves, New South wales. journal and Proceedings of the Royal Society of New South Wales 119, 1/2, 55-76. Osborne, R.A.L., 1991a: Red earth and bones: Te history of cave sediment studies in New South wales, Australia. journal of Earth Sciences history 10, 1, 13-28. Osborne, R.A.L., 1991b: Palaeokarst deposits at Jenolan Caves, N.S.w. journal and Proceedings of the Royal Society of New South Wales 123, 3/4, 59-73. Osborne, R.A.L., 1993a: A new history of cave development at Bungonia, N.S.w. Australian Geographer 24,1, 62-74. Osborne, R.A.L., 1993b: Te history of karstifcation at wombeyan Caves, New South wales, Australia, as revealed by palaeokarst deposits. Cave Science 20, 1, 1-8. TIME in KARST – 2007 141 R. ARMSTRONG L. OSBORNE Osborne, R.A.L., 1995: Evidence for two phases of Late Palaeozoic karstifcation, cave development and sediment flling in southeastern Australia. Cave and Karst Science 22, 1, 39-44. Osborne, R.A.L., 1999: Te origin of Jenolan Caves: Elements of a new synthesis and framework chronology. Proceedings of the Linnean Society of New South Wales 121, 1-26. Osborne, R.A.L., 2002: Paleokarst: Cessation and Rebirth?, pp. 97-114. In F. Gabrovšek [ed.], Evolution of karst: from prekarst to cessation, Založba ZRC, Ljubljana. p. 97-114. Osborne, R.A.L., 2005: Dating ancient caves and related palaeokarst. Acta carsologica 34, 1, 51-72. Osborne, R.A.L., H. Zwingmann, R. E. Pogson, & D.M. Colchester., 2006: Carboniferous Cave Deposits from Jenolan Caves, New South wales, Australia. Australian journal of Earth Sciences 53, 3, 377-405. Pickett, J., 1982: Te Silurian System in New South wales. bulletin of the Geological Survey of New South Wales 29, 1-264 wilford, G.E. 1991: Exposure of land surfaces, drainage age and erosion rates, pp. 93-107. In C.D. Ollier [ed.], Ancient Landforms. Belhaven, London. Zupan-Hajna, N., 2003: Incomplete Solution: Weathering Of Cave Walls And Te Production, transport And deposition Of Carbonate Fines, Založba ZRC, Ljubljana. p. 167. 142 TIME in KARST – 2007 COBISS: 1.01 CLASTIC SEDIMENTS IN CAVES – IMPERFECT RECORDERS OF PROCESSES IN KARST KLASTIČNI SEDIMENTI V JAMAH – NEPOPOLNI ZAPIS KRAŠKIH PROCESOV Ira D. SASOwSKy1 Abstract UDC 552.517:551.7 Ira D. Sasowsky: Clastic sediments in caves – imperfect recorders of processes in karst Clastic sediments have played an important role in deciphering geologic history and processes since the inception of the discipline. Early studies of caves applied stratigraphic principles to karst deposits. Te majority of cave deposits are breakdown and alluvium. Te alluvial materials have been successfully investigated to determine ages of caves, landscape evolution, paleoen-vironmental conditions, and paleobiota. Rapid stage changes and the possibility of pipe-full fow make cave deposits diferent than surface deposits. Tis and other factors present difculties with interpreting the cave record, but extended preservation is aforded by the “roofng” of deposits. Dating by magnetism or isotopes has been successful in many locations. Caves can be expected to persist for 10 Ma in a single erosive cycle; most cave sediments should be no older than this. Key words: clastic sediments, paleoclimate, sedimentology, stratigraphy, dating. Izvleček UDK 552.517:551.7 Ira D. Sasowsky Klastični sedimenti v jamah – nepopolni zapis kraških procesov Že od nekdaj so klastični sedimenti pomembno orodje pri razbiranju geološke zgodovine. V zgodnjih študijah so uporabili načela stratigrafje tudi pri raziskovanju jamskih sedimentov . Glavnino jamskih sedimentov sestavljajo podori in aluvij. Raziskave aluvija so se uspešno izkazale pri dataciji jam, določanju razvoja površja, paleookolja in paleontologije. Zaradi možnega tlačnega toka in hitrih sprememb stanj, so jamski sedimenti drugačni od površinskih. To, poleg ostalih dejavnikov, predstavlja težave pri interpretaciji zapisov, ki jih hranijo jame. Po drugi strani pa je obstojnost jamskih sedimentov daljša zaradi zavetja, ki jim ga nudi jama. Po vsem svetu poznamo številne uspešne datacije jamskih sedimentov z magnetizmom ali izotopi. Jame znotraj erozijskega cikla vzdržijo do10 milijonov let, zato naj jamski sedimenti ne bi bili znatno starejši. Ključne besede: klastični sedimenti, paleoklima, sedimen-tologija, stratigrafja, datiranje. INTRODUCTION Geology is undeniably a science of history, and since the earliest practice of the discipline, that history has been revealed in clastic sedimentary deposits. william Smith, for example, created maps of the sedimentary rocks in England in the late 1700’s, and established a relative chronology of their deposition using stratigraphic position and fossils. It has been natural, therefore, that karst scientists examine clastic deposits in caves, in order to explore geologic time. In doing so, they are in large part applying the same principles and techniques developed by classical stratigraphers. An early example of this was a study by Kukla and Ložek (1958) examining the processes of cave sediment deposition and preservation. In the present day, work such as that by Granger et al. (2001) and Polyak et. al. (1998) builds upon those classical techniques and applies laboratory methods to develop absolute chronolo- 1 Ofce for Terrestrial Records of Environmental Change, Department of Geology and Environmental Science, University of Akron, Akron, OH 44325-4101, USA. Received/Prejeto: 24.01.2007 TIME in KARST, POSTOJNA 2007, 143–149 IRA D. SASOwSKy gies. Tese chronologies in turn have allowed insight to such processes as river incision, water-table lowering, and landscape/climate linkages. Tis paper is a brief evaluation of clastic sediments as they apply to deciphering historical processes and events MATERIALS A Te processes that result in clastic sedimentation in caves are quite varied. Reviews and details including classif-cation of deposits are presented in several texts (white, 1988; Ford and williams, 1989; Sasowsky and Mylroie, 2004). A perspective is given here. A useful broad-level classifcation is genetic, and based upon whether the clastic material originated within the cave (autogenic) or was carried in from the surface (allogenic). Te former class is mainly bedrock breakdown (incasion), but encompasses fne grained sediments sourced from insoluble residue during phre-atic enlargement, collapse of secondary mineralization (speleothems), and so forth. Allogenic sediments include alluvium, windblown material, animal feces, fossil matter, till, etc. In practice, the most commonly occurring materials by far are bedrock breakdown and alluvium. Consequently, autogenic cave sediments are mainly limestone. Allogenic sediments are usually resistant siliciclastics, because carbonates do not typically persist in the fuvial environment. Tere is no satisfying overall term for the clastic deposits found in caves. Te word “soil” has been applied to the fne grained deposits, but this is a misnomer by most defnitions, and is not recommended. Cave fll and cave earth have also been used. Regolith seems applicable in spirit, but, because this material does not strictly “….form(s) the surface of the land ....” (Jackson, 1997) some may object to such usage. BREAKDOwN Te collapse of cave bedrock walls and ceilings results in material that is angular, and ranges in size from sand to boulders. It is possible many times to visually ft larger blocks to their point of origin on the adjacent cave walls and ceilings. Te process of breakdown is not a common occurrence on human timescales. Only a few cases of present-day natural failure have been documented. For example, in Mammoth Cave, Kentucky only one large collapse was noted in 189 years of mining and tourism (May et al., 2005). However, on geologic timescales, the proc- in karst terrane. Advantages and problems of working with these unique deposits are presented. For purposes of this paper, the “age” of a given cave sediment refers to the time of deposition of the material in the cave. D PROCESSES ess is pervasive and evident in most caves. Failures occur along existing planes of weakness (joints, faults, bedding planes). Causes of collapse can include removal of underlying support (particularly loss of buoyancy caused by the transition from phreatic to vadose conditions), removal of overlying arch support, cryoclastism (wedging by ice), and secondary mineral wedging (white and white, 2003). Triggering by earthquakes has also been observed, for example in Sistem Zeleške Jame-Karlovica (personal communication, F. Drole). Davies (1951) published an early analysis of expected collapse parameters in the cave environment. Tis was expanded on by white (1988, p. 232) to evaluate stability of ceilings relative to limestone bed thickness. Greater spans can be maintained by thicker beds. Jameson (1991) provides a comprehensive overview and classifcation of breakdown. Breakdown is frequently most prolifc at 1) the intersections of cave passages, presumably due to the greater span lengths present at such points, and 2) where the cave is close to the surface, due to lack of thinning of the span and resulting decreased competency. In evaluations of causes for passage terminations (white, 1960) it was noted that many cave passages ended in breakdown blockage (referred to by explorers as “terminal breakdown”). Although pervasive, breakdown has not found sig-nifcant utility for deciphering earth history in karst ter-ranes. ALLUVIUM Alluvium enters caves by sinking stream, and occasionally by colluvial mechanisms. Te transport processes are for the most part similar to those in surface channels. Te full range of sediment sizes are seen, structures such as cross-bedding and pebble imbrications develop, and cut-and-fll stratigraphy is possible. However, there are two important diferences exhibited for stream fow in caves when compared to most surface channels. First, channel width is severely constrained by bedrock walls. Tis promotes rapid stage increase during fooding, akin to that of slot canyons in surface streams (Fig. 1). Second, 144 TIME in KARST – 2007 CLASTIC SEDIMENTS IN CAVES – IMPERFECT RECORDERS OF PROCESSES IN KARST Fig. 1: Subterranean stream channels are typically narrow, and have no foodplain (a). Tis leads to rapid stage changes. Similar conditions in the surface environment are only seen in slot canyons such as the virgin River, Utah, USA (b). because the channel is roofed over, it is possible to have confned (pipe-full) rather than open channel fow. Taken in conjunction, the results of these two conditions are the likelihood of high fow velocities, and the possibility of upwards phreatic fow. A striking example of rapid stage change is seen in Hölloch, Switzerland, where rises of 250 m in a single food have been recorded (wildberger and Preiswerk, 1997; Jeannin, 2001). Cases of phreatic lifs are seen in many cave systems. In Castleguard Cave (Rocky Mountains, Canada) a seasonally active lif of 9 m is observed (Schroeder and Ford, 1983). In that situation well-rounded cobbles are accumulated at the base, where they reside until communition reduces them suf-ciently to allow transport up the lif tube. Te composition of the alluvium refects the source of the material, as well as some other factors. It is interesting to note that a high proportion of clay sized material found in cave alluvium is actually fne-grained silica, not a clay mineral (white, 1988). Te residuum found on the surface of many karst terranes frequently contains high amounts of clay and chert. Te clay results from insoluble residues of the weathered limestone. Te chert behaves in a very persistent way, being found throughout cave passages. INFORMATION REVEALED In the investigation of clastic sedimentary deposits, either cave related or not, answers are sought to such questions as: How old? what was the paleoenvironment? what was the fow direction? what organisms were present? Tese in turn allow an understanding of geologic history, environments of deposition, past climates, and potential for sedimentary deposits to act as mineral and fuel reservoirs. In the case of cave studies, it is primarily the frst question which has been addressed. Caves can only be numerically dated by the deposits that they hold, and this age is usually reported as a minimum value. Alluvial materials are considered superior to speleothems in this undertaking, because they are emplaced much earlier in the existence of the cave. Once a date has been obtained, subsequent inferences such as rates of river incision, denudation, and so forth, can then be made based upon the relation of the cave to the landscape. Dating has been accomplished by radiocarbon, magnetism, and cosmo-genic isotopes. Paleoenvironmental information is revealed through studies of sedimentary structures and sequences, as well as via analyses of clay mineralogy and environmental magnetism. Paleohydrology can be deduced using traditional stratigraphic indicators such as cross-bedding, pebble imbrication, etc. Fossil deposits of organisms are actually rather rare within caves – most cave depsits are barren of these materials. Signifcant deposits are known, though, and many excavations made in caves (particularly in the entrance facies) serve as irreplaceable records of terrestrial fauna. TIME in KARST – 2007 145 IRA D. SASOwSKy LIMITS ON TIMESCALE Caves are erosional landforms, which have a limited period of existence. Excluding those caves which have been subjected to burial, this places a practical limit on their duration as potential recorders of nearby processes. In any case, the cave sediments can be no older than the cave they are emplaced in (Sasowsky, 1998). Terefore, the ultimate limit on preservation of sediments within a cave is the persistence (lifetime) of the cave in the environment. In most limestone terranes epigenetic processes occur, with dissolution taking place both at the surface (forming pavements, dolines, etc.) and in the subsurface (forming caves). As base level lowers, denudation of the upland surfaces is also occurring and uppermost caves are eventually breached and destroyed. In certain settings examples of various states of decay can be seen in the landscape, and the sedimentary flls of breached (unroofed) caves may even be observed (e.g. Šušteršič, 2004). In settings such as the Appalachian Valley and Ridge, hundreds of meters of carbonate have been denuded from anticlinal valleys (white, 1988), and one may imagine extensive systems of caves which have been obliterated with no remaining trace. Bounds on the expected lifetime of an epigene cave may be evaluated by considering the two main control- Fig. 2: Teoretical persistence of caves in an erosional environment. Te length of time that a given cave will exist depends upon the initial depth of formation (position on y-axis) and the denudation rate (slope of line). Gray regions envelope a range of reasonable denudation pathways for two examples. In case A, a cave formed at 200 m depth, the expected lifetime is 2.5 to 10 ma. For a cave formed at 100 m depth (case b), the lifetime is reduced to 1.25 to 5 ma. Solid sloping lines are the average denudation rate, 69 m/ma, for 33 major drainage basins (calculated from data in Summerfeld and hulton, 1994). ling factors: initial depth of formation and rate of land surface lowering (denudation, Fig. 2). Although caves may form at any depth, a practical limit of 300 m is reasonable, and the majority of caves are much shallower (Milanovic, 1981). Note that this “depth” is not correlative to the frequently reported mapped depth of caves, which refers to the maximum vertical extent of survey. In the context of the present evaluation, depth is the position below surface (thickness of overlying rock) at a given point in the cave. Denudation rates can be quite variable, and tend to correlate with rainfall (white, 1988, p. 218). Envelopes of expected cave persistence can be constructed (Fig. 2) using these 2 parameters. Based upon this calculation, epigene caves would usually exist in the erosive environment for up to 10 Ma. In practice, dating has not yet resulted in identifca-tion of caves this old within the present erosional cycle. Paleomagnetic dating has been used back to 4.4 Ma (Cave of the winds, Colorado, USA; Luiszer, 1994). Cosmogen-ic isotope dating has documented cave sediments as old as 5.7 (±1.1) Ma (Bone Cave, Tennessee, USA; Anthony and Granger, 2004). Te absence of older values may be a consequence of limitations of dating methods, or refect the relative dearth of older caves in the environment, or both. Te challenges of paleomagnetic dating include absence of fne-grained sediments, lack of uninterrupted sedimentation, and uncertainties of correlation with the global magnetic polarity scale. Cos-mogenic dating is constrained by the absence of quartzose sediments, uncertainties in parent isotope values, and the cost/efort of analyses. If consideration is extended beyond the present erosional cycle, flled and buried caves (paleokarst) are found in the rock record. Such materials have been recognized in many places, and the flls described in some detail (e.g. Loucks, 1999). Interest has been strong in the context of exploration for minerals or petroleum. Tese deposits also represent a potential trove of information on far past hydrologic and environmental conditions because of their capacity to preserve. 146 TIME in KARST – 2007 CLASTIC SEDIMENTS IN CAVES – IMPERFECT RECORDERS OF PROCESSES IN KARST RESOLUTION, CONTINUITy, AND VERACITy Stratigraphers have traditionally examined marine or paralic sediments because of their resolution, continuity, and veracity. Compared to terrestrial deposits, marine/ paralic strata are much more laterally and vertically extensive, they are of economic interest, and they potentially function as continuous recorders for long periods of time. Terrestrial deposits are of interest though, particularly because they contain information about the on-continent setting. within the terrestrial environment lacustrine deposits and fuvial terraces have seen the greatest attention as recorders of Cenozoic paleo-conditions. Lakes probably represent the highest quality records in the terrestrial environment – their environment many times is one of high preservation potential. Lacustrine deposits can be sampled by coring; duplication of cores can serve as a quality control; accumulation rates can be rapid; sediment properties are well tied to local environmental conditions; and spatial variability is usually well understood. Terraces tend to preserve a partial record of the fuvial environment, depending upon regional uplif or down-cutting of the stream. In comparison, most caves contain spatially irregular deposits that can be afected by factors such as plugging of swallets, extreme fow events, and back-fooding. Hydrologic complexity is common (Bosák et al., 2003), even more so than surface fuvial environments. Analysis of the paleohydrology of the depositional setting through cave passage morphometry is usually necessary, and may be quite time consuming if detailed maps are not available. Stratigraphic sections may be discontinuous, and require compilation. Caves are difcult sampling locations, due to logistics, remoteness, lack of light, and constraints on sampling equipment transport. Nevertheless, the cave environment is one that provides some advantages in recording the history of a region. Te greatest advantage is that of potential preservation. Because caves are “roofed over” deposits are likely to be protected (at least on intermediate time scales), from Fig. 3: Comparison of sedimentary records from Lake baikal, Russia (3 columns on lef), and Cave of the Winds, USA (3 columns on right). baikal data used with permission from King and Peck, 2001. Cave of the Winds data used with permission from Luiszer, 1994. TIME in KARST – 2007 147 IRA D. SASOwSKy Fig. 4: Episodic inflling and removal of sediments is commonly observed in caves. In this section of Windy mouth Cave (West virginia, USA) a diamict was almost completely removed afer being covered with fowstone. Te conduit is presently dry. surfcial erosion. Tis is particularly germane for the fu-vial deposits. weathering and erosion of surface fuvial terraces is commonplace. In the cave, such materials may sit undisturbed for years. For example, in xanadu Cave, Tennessee, USA, a pristine, non-indurated fuvial deposit that is greater than 780 ka was sampled (Sasowsky, et al., 1995). Although rare, in exceptional settings the quality of the cave record may approach that of lakes (Fig. 3). Conditions amenable to this are stable recharge confguration, diffuse recharge, minimal variation of discharge, and deep circulation. In Figure 3 two exceptional records are compared. Te Lake Baikal record was constructed from cores taken on watercraf. In that setting, about 40 m of sediment accumulate in 1 Ma. In contrast, at Cave of the winds the accumulation rate is slower by more than an order or magnitude. In many settings caves appear to undergo episodic flling and excavation (Fig. 4). In certain cases this may be locally controlled by catastrophic storms (e.g. Doehring and Vierbuchen 1971). However, the presence of broadly similar deposits/incisions within many caves in a region supports the idea that cave clastic materials refect regional paleoclimatic conditions. Tese deposits hold much information that will be revealed with continued advances in conceptual frameworks and improved laboratory methods. REFERENCES Anthony, D.M. & D.E. 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TIME in KARST – 2007 149 COBISS: 1.01 ANALySIS OF LONG-TERM (1878-2004) MEAN ANNUAL DISCHARGES OF THE KARST SPRING FONTAINE DE VAUCLUSE (FRANCE) ANALIZA DOLGOČASOVNEGA (1878-2004) POVPREČNEGA LETNEGA PRETOKA KRAŠKEGA IZVIRA FONTAINE DE VAUCLUSE (FRANCIJA) Ognjen BONACCI1 Abstract UDC 556.36(44) Ognjen Bonacci: Analysis of long-term (1878-2004) mean annual discharges of the karst spring Fontaine de Vaucluse (France) Statistical analyses have been carried out on a long-term (1878-2004) series of mean annual discharges of the famous karst spring Fontaine de Vaucluse (France) and the mean annual rainfall in its catchment. Te Fontaine de Vaucluse is a typical ascending karst spring situated in the south-eastern region of France. Te spring has an average discharge of 23.3 m3/s. Te average annual rainfall is 1096 mm. Its catchment area covers 1130 km2. Using the rescaled adjusted partial sums (RAPS) method the existence of next fve statistically signifcant difer-ent sub-series was established: 1) 1878-1910: 2) 1911-1941; 3) 1942-1959: 4) 1960-1964; 5) 1965-2004. Te diferent spring discharge characteristics during this long period (1878-2004) can be caused by natural climatic variations, by anthropogenic infuences, and possibly by climate changes. At this moment it should be stressed that objective and scientifcally based reasons for diferent hydrological behaviour in fve time sub-periods could not be found. Keywords: karst hydrology, mean annual discharges, annual catchment rainfall, karst spring, Fontaine de Vaucluse, France. Izvleček UDK 556.36(44) Ognjen Bonacci: Analiza dolgočasovnega (1878-2004) povprečnega letnega pretoka kraškega izvira Fontaine de Vaucluse (Francija) V prispevku predstavim statistično analizo časovne vrste povprečnega letnega pretoka in letnih padavin v zaledju slavnega izvira Fontaine de Vaucluse v Franciji. Fontaine de Vaucluse je tipični kraški izvir pri katerem voda priteka iz velike globine. Nahaja se v jugovzhodni Franciji. Povprečni pretok izvira je 23,4 m3/s. Povprečna količina letnih padavin v zaledju, ki meri 1130 km2, je 1096 mm. Z uporabo metode umerjenih delnih vsot (RAPS) smo določili pet statistično pomembnih različnih podobdobij: 1) 1878-1910: 2) 1911-1941; 3) 1942-1959: 4) 1960-1964; 5) 1965-2004. Vzrokov za različne pretoke preko celotnega obdobja (1878-2004) je lahko več; npr. klimatske spremembe in antropogeni vplivi. V tem trenutku moramo poudariti, da objektivne znanstvene razlage za različne hidrološke značilnosti v petih podobdobjih še ne poznamo. Ključne besede: hidrologija krasa, povprečni letni pretok, količina letnih padavin, kraških izvir, Fontaine de Vaucluse, Francija. INTRODUCTION Te Fontaine de Vaucluse represents one of the most et al., 1992b). Te karst system of the Fontaine de Vauc-famous and most important karst springs on the Earth. luse is characterised by an approximately 800 m unsatu-It is located in the south-eastern karst region of France rated zone. Emblanch et al., (1998) and Emblanch et al., (Figure 1), about 30 km eastward of the town of Avignon. (2003) stressed important role of this zone for the transIt represents the only fow exit from the 1500 m thick formation of rainfall into runof. Te Fontaine de Vauc-karst aquifer of Lower Cretaceous limestone (Blavoux luse karst spring catchment area is estimated to be 1130 1 Faculty of Civil Engineering and Architecture, University of Split, 21000 Split, Matice hrvatske 15, Croatia, E-mail: obonacci@ gradst.hr Received/Prejeto: 27.11.2006 TIME in KARST, POSTOJNA 2007, 151–156 OGNJEN BONACCI km2 (Cognard-Plancq et al., 2006a; 2006b). Te average catchment altitude is 870 m a. s. l. Te average annual air temperature of the catchment is 9,6 °C. Fig. 1: Location map of karst spring Fontaine de vaucluse. Te Fontaine de Vaucluse is typical ascending karst spring (Michelot & Mudry 1985; Blavoux et al., 1991/1992; 1992a). Its limestone channel ranges in diameter from 8 to 30 m (Mudry & Puig, 1991). Te lowest Te climate in the catchment is Mediterranean. Rainfall distribution over the year as well as over the large spring catchment is irregular. Intensive and signifcant rainfall events occurred during autumn and spring, while summer and winter are generally dry. Interannual fuctua-tions of rainfall on the catchment are very high. In order to defne an historical homogeneous catchment rainfall database Cognard-Plancq et al., (2006b) used six rainfall gauging stations. Te mean elevation of these stations is 445 m a. s. l., while the mean elevation of the spring catchment is 870 m a. s. l. Transformation of the measured monthly rainfall to the altitude of 870 m a. s. l. was made. Te average annual catchment rainfall in the 1878-2004 period is 1096 mm, while the minimum and maximum observed values were 641 mm (1953) and 1740 mm (1977) respectively. Data series with linear trend line of the annual rainfall on the Fontaine de Vaucluse catchment for the period 1878-2004 are presented in Figure 2. Te increasing depth reached by diver was -308 m below the gauging station datum of 84.45 m a. s. l. Tis depth is still not at the bottom of the ascending karst channel. Te maximum water level measured at the gauging station was 24.10 m above the datum, the minimum was a few centimetres below the datum. Te rate of the maximum discharge of the spring has never been precisely measured, but it is estimated that it cannot exceed 100 m3/s (Blavoux et al., 1991/1992; 1992a). Cognard-Plancq et al. (2006b) state that maximum spring discharge varies between 100 and 120 m3/s. Tis surmise identifes a karst spring with limited discharge capacity (Bonacci 2001). Te historical minimum discharge is 3.7 m3/s (Blavoux et al., 1991/1992). Every karst aquifer has complex hydrodynamic behaviour. Te Fontaine de Vaucluse karst system responses to rainfall quite rapid in comparison with the large recharge area. Te peak of hydrograph occurred 24 to 72 hours afer the rainfall events. Te spring water level and discharge recessions are slow, which can be explained by the existence of a large storage capacity of the aquifer (Cognard-Plancq 2006b). Te primary objective of the investigation was to defne sub-periods with diferent hydrological behaviour of the Fontaine de Vaucluse karst spring during 127 years period (1878-2004), analysing time series of mean annual spring discharges. It should be the frst step in explanation of this extremely important and interesting phenomenon. trend of the catchment rainfall of 1.045 mm per year is not statistically signifcant but should not be neglected in further analyses. Fig. 2: time data series of annual rainfall P at the Fontaine de vaucluse catchment with trend line for the period 1878-2004. ANALySIS OF CATCHMENT ANNUAL RAINFALL TIME SERIES 152 TIME in KARST – 2007 ANALySIS OF LONG-TERM (1878-2004) MEAN ANNUAL DISCHARGES OF THE KARST SPRING FONTAINE DE VAUCLUSE ANALISIS OF MEAN ANNUAL DISCHARGES TIME SERIES Data series with linear trend line of the mean annual spring discharges q for the period 1878-2004 are presented in Figure 3. Te decreasing trend of the mean annual discharges of 0.0468 m3/s per year is not statistically signifcant. Te average annual catchment discharge in the 1878-2004 period was 23.3 m3/s, while the minimum and maximum observed values were 7.61 m3/s (1990) and 53.4 m3/s (1915) respectively. Fig. 3: time data series of mean annual discharges Q at the Fontaine de vaucluse karst spring with trend line for the period 1878-2004. It should be stressed that annual catchment rainfall during the same period has an increasing trend. In Figure 4 linear regression between the mean annual the Fontaine de Vaucluse discharges q and the Fontaine de Vaucluse catchment annual rainfall P is shown. Te linear correlation coefcient is only 0.713, which is relatively low. A special problem is that the regression line cut abscissa line at 222 mm of annual rainfall P, which is relatively low value. Explanation of so unusual rainfall-runof relationship can be found in fact that accuracy of discharges and rainfalls are not very high, and maybe the value of catchment area of 1130 km2 is not precisely defned. It should be stressed that determination of exact catchment area in karst is one of the greatest and very ofen unsolved problems. Tis may be the case with the catchment of the Fontaine de Vaucluse spring. Te weak relationship between runof and rainfall means that some others factors (probably: air temperature, groundwater level, interannual rainfall distribution, changes of catchment area during the time, preceding soil wetness, anthropological infuences, climate change etc) have infuence on it. A time series analysis can detect and quantify trends and fuctuations in records. In this paper the Rescaled Adjusted Partial Sums (RAPS) method (Garbrecht & Fernandez 1994) was used for this purpose. A visualisation approach based on the RAPS overcomes small systematic changes in records and variability of the data values themselves. Te RAPS visualisation highlights trends, shifs, data clustering, irregular fuctuations, and periodicities in the record (Garbrecht & Fernandez 1994). It should be stressed that the RAPS method is not without shortcomings. Te values of RAPS are defned by equation: t-l ^Y where Y is sample mean; is standard deviation; n is number of values in the time series; (k=1, 2…,n) is counter limit of the current summation. Te plot of the RAPS versus time is the visualisation of the trends and fuctuations of yt. Time data series of Rescaled Adjusted Partial Sums (RAPS) for mean annual spring discharges in the period 1878-2004 are given in Figure 5. Terefore, the total data Fig. 4: Linear regression between the mean annual the Fontaine de vaucluse discharges Q and annual the Fontaine de vaucluse catchment rainfall P. Fig. 5: time data series of the Rescaled Adjusted Partial Sums (RAPS) for mean annual discharges Q for the period 1878-2004 with designated next fve sub-periods: 1) 1878-1910; 2) 1911-1941; 3) 1942-1959; 4) 1960-1964; 5) 1965-2004. TIME in KARST – 2007 153 OGNJEN BONACCI series was divided into next fve subsets: 1) 1878-1910; 2) 1911-1941; 3) 1942-1959; 4) 1960-1964; 5) 1965-2004. Cognard-Plancq et al., (2006a; 2006b) defned the same fve stationary sub-periods using diferent methodology. Five time data sub-series of the Fontaine de Vau-cluse karst spring mean annual discharges q with trend lines for fve defned sub-periods are shown in Figure 6. In order to investigate statistically signifcant diferences between the averages of fve time sub-series for q and P the t-test was used. Te neighbouring averages of discharges for all fve sub-series are statistically signifcant at Fig. 6: Five time data sub-series of the Fontaine de vaucluse karst spring mean annual discharges Q with trend line for fve defned sub-periods. Te rescaled adjusted partial sums (RAPS) method established existence of next fve statistical, and hydrologi-cal signifcant diferent time sub-series: 1) 1878-1910: 2) 1911-1941; 3) 1942-1959: 4) 1960-1964; 5) 1965-2004. Variations in the Fontaine de Vaucluse karst spring hy-drological regime during relatively short period of 127 years are very strong and cannot be neglected. Anthropogenic impacts are probably the main cause of such behaviour of the mean annual spring discharges time series analysed, but the natural pattern of drought and wet years is also possible. Land-use changes and overexploitation of surface water and groundwater at the spring catchment on hydrological regime of the Fontaine de Vaucluse spring certainly exists. Teir exact quantifcation during analysed period is extremely questionable due to missing of many parameters. Strict division of natural and anthropogenic infuences on the hydrological regime is hardly possible. The significant changes of spring discharge characteristics during 127 years long period (1878-2004) can be caused by natural climatic variations, Fig. 7: Linear regressions between mean annual discharges Q and annual catchment rainfall P defned for fve diferent sub-periods. the 5 % and even more 1 %. At the same time the neighbouring sub-series averages of the catchment rainfall are not statistically signifcant. Figure 7 shows fve linear regressions between mean annual discharges q and annual catchment rainfall P de-fned for fve diferent sub-periods. It can be seen that linear correlation coefcients for all sub-series, except for third (1942-1959) and fourth (1960-1964) ones are higher than the linear correlation coefcient for whole time series. by anthropogenic influences, and possibly by climate changes. It is extremely hard, but at the same time extremely practically and theoretically important, to find correct and scientifically based explanation of this phenomenon. Cognard-Plancq et al., (2006a) consider that rain-fall-runof data have shown the large impact of clima-tologic variations on the hydrogeological system. Tey conclude that the underground storage zone is an important infuence on karst spring outfow, which depends on rainfall amount over 2 or 3 previous years. Investigations made in this paper do not confrm this statement. Correct answers on many questions dealing with changes in hydrological-hydrogeological regime of the Fontaine de Vaucluse karst spring cannot be done using only annual data. Some processes can be explained measuring and analysing climatologic, hydrologic, hydrogeo-logical and geochemical interactions in shorter as well as larger time increments. Te problem is that most of parameters required for these analyses were not monitored in the past. 154 TIME in KARST – 2007 ANALySIS OF LONG-TERM (1878-2004) MEAN ANNUAL DISCHARGES OF THE KARST SPRING FONTAINE DE VAUCLUSE More accurate and precise delineation and defni-tion of the Fontaine de Vaucluse spring catchment should be done. It is possible that its catchment area changes as a function of groundwater level. Tis means that ground-water level measurements in deep piezometers should be organized across the catchment. Te second task which should be considered in further analyses is detailed analysis of infuence of rainfall distribution during the year on the spring runof. Tis can have very strong infuence on the relationship between rainfall and runof, especially in karst areas. Te author thanks to Anne-Laure Cognard-Plancq and Christophe Emblanch from Laboratoire d’Hydrogéologie, Faculté des Sciences, Université d’Avignon et Pays de Blavoux, B., Mudry, J. & Puig, J.-M., 1991/1992: Bilan, fonctionnement et protection du systeme karstique de la Fontaine de Vaucluse (sud-est de la France). Geodinamica Acta, 5 (3), 153-172., Paris. Blavoux, B., Mudry, J. & Puig, J.-M., 1992a: Te karst system of the Fontaine de Vaucluse (Southeastern France). Environ. Geol. water. Sci., 19 (3), 215-225. Blavoux, B., Mudry, J., & Puig, J.-M., 1992b: Role du con-texte geologique et climatique dans la genese et le fonctionnement du karst de Vaucluse (Sud-Est de la France). In: H Paine, w Back (eds.) Hydrogeology of Selected Karst Regions. IAH International Contributions to Hydrogeology, Vol. 13, 115-131. Bonacci, O., 2001: Analysis of the maximum discharge of karst springs. Hydrogeol. J., 9, 328-338. Cognard-Plancq, A.-L., Gévaudan, C. & Emblanch, C., 2006a: Apports conjoints de suivis climatologique et hydrochimique sur le rôle de fltre des aquiferes karstiques dans l’étude de la problématique de changement climatique; Application au systeme de la Fontaine de Vaucluse. Proceedings of the 8th Conference on Limestone Hydrogeology. Neucha-tel, Sep. 21-23, 2006, 67-70. It can be stated that main dilemmas about variations of mean annual discharges of the Fontaine de Vaucluse karst spring during 127 years long period have not been solved. Tey should be explained using number of different procedures and climatic as well as other indicators, and performing further detailed measurements and analyses. Te paper presents the need for interdisciplinary analyses incorporating several approaches and techniques. For the sustainable development and the protection of such valuable water resource it is very important to establish prerequisites for the defnition of a causes and consequences of its hydrological changes. Vaucluse 84000 Avignon, 33 Rue Louis Pasteur, France, which kindly provide me with data analysed in this paper. Cognard-Plancq, A.-L., Gévaudan, C., &, Emblanch, C., 2006b: Historical monthly rainfall-runof database on Fontaine de Vaucluse karst system: review and lessons. IIIéme Symposium International Sur le Karst „Groundwater in the Mediterranean Coun-tries“, Malaga, Spain. In: J J Duran, B Andreo, F y Carrasco (eds.) Karst, Cambio Climatico y Aguas Subterraneas. Publicaciones des Instituto Geological y Minero de Espana. Serie: Hidrogeologia y Aguas Subterrraneas, N°18: 465-475. Emblanch, C., Puig, J. M., Zuppi, G. M., Mudry, J., & Bla-voux, B., 1998: Comportement particulier lors des montées de crues dans les aquiferes karstiques, mise en évidence d’une double fracturation et/ou de circulation profonde: Example de la Fontaine de Vau-cluse. Ecologae Geol. Helv., 92: 251-257. Emblanch, C., Zuppi, G. M., Mudry, J., Blavoux, B. & Batitot, C., 2003: Carbon 13 of TDIC to quantify the role of the unsaturated zone: Te example of the Vaucluse karst systems (Southeastern France). J. of Hydrol., 279 (1-4): 262-274. ACKNOwLEDGEMENT REFERENCE TIME in KARST – 2007 155 OGNJEN BONACCI Garbrecht, J. & Fernandez, G. P. , 1994. Visualization of Mudry, J., Puig, J.-M., 1991: Le karst de la Fontaine de trends and fuctuations in climatic records. water Vaucluse (Vaucluse, Alpes de Haute-Provence, Resources Bulletin, 30 (2): 297-306. Drôme). Karstologia, 18 (2): 29-38. Michelot, C. & Mudry, J., 1985: Remarques sur les exu-toires de l’aquifere karstique de la Fontaine de Vau-cluse. Karstologia, 6(2): 11-14. 156 TIME in KARST – 2007 COBISS: 1.01 TIMING OF PASSAGE DEVELOPMENT AND SEDIMENTATION AT CAVE OF THE wINDS, MANITOU SPRINGS, COLORADO, USA ČASOVNO USKLAJEVANJE RAZVOJA JAMSKIH PROSTOROV IN SEDIMENTACIJA V JAMI CAVE OF THE wINDS, MANITOU SPRINGS, COLORADO, ZDA Fred G. LUISZER1 Abstract UDC 551.3:551.44:550.38 550.38:551.44 Fred G. Luiszer: Timing of Passage Development and Sedimentation at Cave of the Winds, Manitou Springs, Colorado, USA. In this study the age of the onset of passage development and the timing of sedimentation in the cave passages at the Cave of the winds, Manitou Springs, Colorado are determined. Te amino acid rations of land snails located in nearby radiometri-cally dated alluvial terraces and an alluvial terrace geomorphi-cally associated with Cave of the winds were used to construct an aminostratigraphic record. Tis indicated that the terrace was ~ 2 Ma. Te age of the terrace and its geomorphic relation to the Cave of the winds was use to calibrate the magne-tostratigraphy of a 10 meter thick cave sediment sequence. Te results indicated that cave dissolution started ~4.5 Ma and cave clastic sedimentation stopped ~1.5 Ma. Key words: Cave of the winds, Manitou Springs, magneto-stratigraphy, aminostratigraphy, land snails. .Izvleček UDK 551.3:551.44:550.38 550.38:551.44 Fred G. Luiszer: Časovno usklajevanje razvoja jamskih prostorov in sedimentacija v jami Cave of the Winds, Manitou Springs, Colorado, ZDA Članek se osredotoča na začetek razvoja jamskih prostorov in časovno sosledje sedimentacije v jami Cave of the winds, Manitou Springs, Kolorado. V bližini jame se nahajajo aluvialne terase, ki so bile datirane z radiometrično metodo. Z geomorfološko metodo so bile povezane z jamo Cave of the winds. V teh aluvialnih terasah so bili najdeni fosilni ostanki kopenskih polžev, na katerih so bile opravljene datacije z aminokislinami, ki so pokazale starost ~ 2 Ma let. Starost aluvialnih teras in njihova geomorfološka povezava z jamo Cave of the winds, sta služila kot izhodišče za natančnejšo časovno umestitev 10 metrov debele sekvence jamskih sedimentov, ki so bili magnetostratigrafsko opredeljeni. Raziskava je pokazala, da se je raztapljanje v jami pričelo pred ~4.5 Ma leti, medtem ko se je odlaganje klastičnih sedimentov prenehalo pred ~1.5 Ma let. Ključne besede: Cave of the winds, Manitou Springs, ZDA, magnetostratigrafja, aminostratigrafja, kopenski polži. INTRODUCTION Cave of the winds, which is 1.5 km north of Manitou Springs (Figure 1), is a solutional cave developed in the Ordovician Manitou Formation and Mississippian williams Canyon Formation. Commercialized soon afer its discovery in the1880s it has been visited by millions of visitors in the last 125 years. As part of an extensive study (Luiszer, 1997) of the speleogenesis of the cave the timing of passage development and sedimentation needed to be determined. Te task of dating the age of caves has always been an enigma because dating something that has been removed is not possible. Sediments deposited in the cave passages, however, can be dated, which then can be used to estimate the timing of the onset of cave dissolution and when the local streams abandoned the cave. 1 University of Colorado, Boulder, Department of Geological Sciences, Campus Box 399, Boulder, CO 80302, USA. Received/Prejeto: 13.12.2006 TIME in KARST, POSTOJNA 2007, 157–171 FRED G. LUISZER A specially constructed coring device was utilized to core several locations in the cave. Te natural remnant magnetization (NRM) of samples taken from the cores were use to construct a magnetostratigraphic record. Tis record by itself could not be used to date the age of the sediments because sedimentation in the cave stopped sometime in the past and part of the record was missing. An alluvial terrace, which overlies the Cave of the winds, is geomorphically related to the cave. Te age of the alluvial terrace, which had not been previously dated, can be used to determine the age of the youngest stream deposited sediments in the cave. An abundant number and variety of land snails were found when this alluvium was closely searched. Biostratigraphy could not be used to determine the age of the terrace because all of the snail species found were extant, however, the amino acid rations of the snails collected from this terrace and nearby radiometrically dated terraces were used to construct an aminostratigraphy that was used to date the alluvium. Once the age of the terrace was determined the age of the youngest magnetic chron of the magnetostratigraphic record could be assigned thus enabling the dating of cave dissolution and sedimentation. FIELD AND LABOR Amino Acid Dating Snails were collected from outcrops of the Nussbaum Alluvium, and from younger radiometrically dated alluvia (Fig. 2) for the purpose of dating the Nussbaum Alluvium by means of amino-acid racemizatio. Approximately 50 kg of sandy silt was collected at each site. To minimize sample contamination, washed plastic buckets and fresh plastic bags were used. Te samples were loaded into containers with a clean metal shovel and with minimal hand contact. In the lab, the samples were disaggregated by putting them in buckets flled with tap water and letting them soak overnight. Te samples were then washed with tap water through 0.5-mm mesh scree. Following air drying, the mollusks were hand picked from the remaining matrix by means of a small paint brush dipped in tap water. Te mollusks were then identifed. Only shells that were free of sediment and discoloration were selected for further processing. Tese shells were washed at least fve times in distilled water while being sonically agitated. Te amino-acid ratios were determined on a high-performance liquid chromatograph (HPLC) at the Institute of Arctic and Alpine Research (University of Colorado, Boulder). Colorado oDenver / ^/El Paso /^ County \^ (25) Cave'.of the Winds j^24s-ö^—Colorado Springs Manitou Springs v25/ Fig. 1: Location of study area. ORy PROCEDURES Paleomagnetism A coring device was used to sample the cave sediments at six cored holes in the Grand Concert Hall (Fig. 3). Te core samples were obtained by means of a coring device in which a hand-powered hydraulic cylinder drives a stainless-steel, knife-edged barrel down into the sediments. Up to 40 cm of sediment could be cored each trip into the hole without sediment distortion. Samples were also collected from hand-dug pits at Mummys Alcove and Sniders Hall (Fig. 3). Additionally, samples were collected from a vertical outcrop in Heavenly Hall (Fig. 3). Te pits and outcrops were sampled for paleomagnetic study by carving fat vertical surfaces and pushing plastic sampling cubes into the sediment at stratigraphic intervals ranging from 3.0 to 10.0 cm. Te samples were oriented by means of a Brunton compass. Te core barrel and all pieces of drill rod that attached to the barrel were engraved with a vertical line so that the orientation of the core barrel could be measured with a Brunton compass within ±2°. A hand-operated hydraulic device was used to extract the sediment core from the barrels. As the core was extruded, a fxed thin wire sliced it in half, lengthwise. Plastic sampling cubes were then pushed into the sof sediment along the center line of the fat surface of the core half at regular intervals 158 TIME in KARST – 2007 TIMING OF PASSAGE DEVELOPMENT AND SEDIMENTATION AT CAVE OF THE wINDS, MANITOU SPRINGS, COLORADO, USA Figure 2. GEOLOGY MAP OF COLORADO SPRINGS AND MANITOU SPRINGS AREA with locations of snail collection sites. Geology adapted from Trimble and Machette, (1979). (EARLY PLEISTOCENE) (CRETACEOUS) TRIASSIC, (precambian) or amino acid 4 KILOMETERS (generally ~5.0 cm). Te samples at Sniders Hall, Mum-mys Alcove, and Hole 1 were taken with 3.2 cm3 sampling cubes; all other samples were taken with 13.5 cm3 cubes. In the lab, the NRM (Natural Remanent Magnetization) of all samples was initially measured. Subsequently, the samples were subjected to alternating-feld (A. F.) demagnetization and remeasured. All samples were frst demagnetized at 10, and then at 15 millitesla (mT). Some samples at the bottom of Hole 5 that displayed aberrant inclinations and declinations were additionally demagnetized at felds up to 30 mT. All remanence measurements were made on a Schonstedt SSM 1A spinner magnetometer with a sensitivity of 1x10-4 A/m. Repeat measurements indicate an angular reproducibility of ~2° at an intensity of 1x10-6 A/m2. Age Of Cave Passages Because Cave of the winds is an erosional feature, its exact age cannot be determined. However, geologic and geomorphic features related to the cave can be used to bracket the age of incipient and major cave development. Solution breccia in the Manitou Formation indicate that there may have been some Middle Ordovician to Devonian cave development (Forster, 1977). Sediment-flled paleo-caves and paleo-sinkholes at Cave of the winds in- dicate Devonian to Late Mississippian karst development (Hose & Esch, 1992). Subsequent Cenozoic dissolution along some of these paleokarst features has resulted in the formation of cave passage (Fish, 1988). Between the Pennsylvanian and Late Cretaceous, about 3000 m of sediments, which contain abundant shale beds, were deposited over the initial cave. Very little, if any, cave development could take place during this period of deep burial under the thick blanket of the nearly impervious rock. Te Laramide Orogeny, beginning in the Late Cretaceous (~75 Ma, Mutschler et al., 1987), was associated with the uplif of the Rocky Mountains. Te up-lif, which included the Rampart Range and Pikes Peak, caused the activation of the Ute Pass and Rampart Range Faults (Morgan, 1950; Bianchi, 1967). In the Manitou Springs area, movement on the Ute Pass Fault resulted in the folding, jointing and minor faulting of the rocks (Hamil, 1965; Blanton, 1973). Te subsequent fow of corrosive water along the fractures related to the folding and faulting would produce most of the passages in Cave of the winds and nearby caves. Uplif during the early Laramide Orogeny increased the topographic relief in the Manitou Springs area, resulting in the initiation of erosion of the overlying sediments and also increased TIME in KARST – 2007 159 jp '. es (LATE jb jlo 3s 2\ jrf i N

168 TIME in KARST – 2007 2. 4. Limestone TIMING OF PASSAGE DEVELOPMENT AND SEDIMENTATION AT CAVE OF THE wINDS, MANITOU SPRINGS, COLORADO, USA South A. ~2 Ma Fountain Creek North Williams Canyon Creek Text in green are sediments that are being deposited in the cave. * = Grand Concert Hall, Cave Of The Winds ----------------- = outline of cavern I = Nussbaum Alluvium South B. ~1.8 Ma North Williams Canyon Creek Fountain Creek South C. Present Fountain Creek Figure 7. Schematic cross sections showing the sequence of changes in the water table, topographic setting, and depositional phases over the last ~2 Ma. Nussbaum Alluvium was being deposited at the same time that clay was being deposited in the Grand Concert Hall the (Fig. 7A). As Fountain Creek downcut and moved to the south, the water table dropped (Fig. 7B). Te drop in the water table coincided with drop in the water depth in rooms like the Grand Concert Hall. As the water depth dropped, the velocity of the water passing through the room increased. Te increased stream en- ergy changed the sedimentation regime from clay deposition to silt, sand, and gravel deposition (Fig. 7B). Fluvial sedimentation at Cave of the winds stopped as Fountain Creek moved further to the south and downcut further (Fig. 7C). Te relationship between the Nussbaum Alluvium and the sediments in the cave indicate that the silt-clay interface in the Grand Concert Hall took place afer the Nussbaum was deposited. More specifcally, the silt-clay interface should be the same age as the Nussbaum Alluvium minus the time it took for Fountain Creek to downcut and drop the water table to the level of the Grand Concert Hall (Fig. 7B). Te sediment foor of the Grand Concert Hall, where the paleomagnetic data was obtained, is about 20 m below the Nussbaum Alluvium. Te age of the Nussbaum Alluvium (~1.9 Ma) and its height above modern streams (200 m) provides an estimate of the average down-cutting rate of 10.5 cm/1000 years. Accordingly, accumulation of coarse sediments in the cave 20 m below the Nussbaum Alluvium probably would have begun ~1.7 Ma. Te estimated 1.7 Ma age of the clay-coarse sediment interface correlates well with the onset of the Olduvai Subchron at 1.9 Ma ( ~2.2 m depth, Fig. 6). Tis is the most probable correlation. Alternatively, one could match the normal-polarity sequence (1.0 to 2.2 m depth, Fig. 6) with the Jaramillo Subchron (Harland et al., 1982) or the Gauss Chron (Fig. 6). Tese correlations, however, would result in an age of ~1.0 Ma or ~ 2.6 Ma, respectively, for the clay-coarse-sediment interface, which is estimated to be 1.7 Ma, thereby making these alternate correlations unlikely. North TIME in KARST – 2007 169 FRED G. LUISZER Te complete paleomagnetic correlation shown on Fig. 6 follows from correlation of the normal-polarity interval between 1.0 and 2.2 m in depth with the Olduvai Subchron. According to the correlation suggested here, the oldest cave sediment was deposited about 4.3 Ma, a date that agrees quite well with the previously discussed probable age of the major onset of cave formation (7 Ma to 4 Ma). CONCLUSIONS Cave of the winds is a phreatic cave dissolved from the calcite-rich Manitou, williams Canyon, and Leadville Formations. Dissolution occurred along joints associated with Laramide faulting and folding. Paleokarst features, such as sediment-flled fssures and caves, indicate that some of the passages at Cave of the winds are related to cave-forming episodes that started soon afer the deposition of the Ordovician Manitou Formation and continued to the beginning of the Cretaceous Laramide Orogeny. Most speleogenesis, however, occurred in the last ~5.0 Ma. Te Nussbaum Alluvium was assigned an age of ~1.9 Ma by means of aminostratigraphy. Te age of the Nussbaum Alluvium and its relation to coarse grained sediments at Cave of the winds were used to fx an age of ~1.7 Ma for the onset of coarse grained sedimentation in the cave. Tis enabled the identifcation of the Olduvai Polarity Subchron in the coarse grained sediments. Correlation of the magnetostratigraphy of cave sediments with the accepted polarity time scale indicates that the dissolution of cave passage started ~4.2 Ma and stopped ~1.5 Ma. REFERENCES Blanton, T. L., 1973: Te Cavern Gulch Faults and the Fountain Creek Flexure, Manitou Spur, Colorado [M.S. thesis]: Syracuse University, New york, 90 p. Bianchi, L., 1967: Geology of the Manitou-Cascade Area, El Paso County, Colorado with a study of the permeability of Its crystalline rocks [M.S. Tesis]: Golden, Colorado School of Mines. Cande, S. C., and D. Kent., 1992: A new geomagnetic polarity time scale for the Late Cretaceous and Ceno-zoic: Journal of Geophysical Research, 97, 10, 13- 17. Epis, R. C., and C.E. Chapi, 1975: Geomorphic and tectonic implications of the Post-Laramide, Late Eocene Erosion surface in the Southern Rocky Mountains, in Curtis, B.F., ed., Cenozoic History of the Southern Rocky Mountains: Geological Society of America Memoir 144, 45-74. Fish, L., 1988: Te real story of how Cave of the winds Formed: Rocky Mountain Caving, 5, 2, 16-19. Forster, J. R., 1977: Middle Ordovician subaerial exposure and deep weathering of the Lower Ordovician Manitou Formation along the Ute Pass Fault zone: Geological Society of America Abstracts with Programs, 9, 722. Goodfriend, G. A., 1987: Evaluation of amino-acid race-mization/epimerization dating using radiocarbon-dated fossil land snails: Geology 15, 698-700. Hailwood, E. A., 1989: Te role of magnetostratigraphy in the development of geological time scales; Pale-oceanography, 4, 1, 1-18. Hamil, M. M., 1965: Breccias of the Manitou Springs area, Colorado [M.S. thesis]: Louisiana State University, 43 p. Harland, w. B., et al., 1982: A geologic time scale: Cambridge, Great Britain, Cambridge University Press, 66 p. Hose, L. D., & Esch, C. J., 1992: Paleo-cavity flls formed by upward injection of clastic sediments to lithostat-ic load: exposures in Cave of the winds, Colorado [abs.]: National Speleological Society Convention Program, Salem, Indiana, p.50 Izett, G. A., Obradovich, J. D., & H.H. Mehnert., 1989: Te Bishop Ash Bed (Middle Pleistocene) and some older (Pliocene and Pleistocene) chemically and mineralogically similar ash beds in California, Nevada, and Utah: U. S. Geological Survey Bulletin, 1675, 37 p. Luiszer, F. G., 1997: Genesis of Cave of the winds, Mani-tou Springs, Colorado, [Ph. D. thesis]: Boulder, University of Colorado, 112 p. Machette, M. M., 1975: Te quaternary geology of the Lafayette quadrangle, Colorado, [M. S. thesis]: Boulder, University of Colorado, 83 p. 170 TIME in KARST – 2007 TIMING OF PASSAGE DEVELOPMENT AND SEDIMENTATION AT CAVE OF THE wINDS, MANITOU SPRINGS, COLORADO, USA Mankinen, E. A., & Dalrymple, G. B., 1979: Revised geomagnetic polarity time scale for the interval 0-5 m. y. B. P. ; Journal of Geophysical Research, 84, B2, 615-626. Miller, G. H., & Brigham-Grette, J., 1989: Amino acid geochronology: Resolution and precision in carbonate fossils in INqUA quat. Dating Methods, Rutter and Brigham-Grette Eds. Pergamon Press. Mitterer, R. M., & Kriausakul, 1989: Calculation of amino acid racemization ages based on apparent parabolic kinetics: quaternary Science Reviews, 8, 353-357. Morgan, G. B., 1950: Geology of williams Canyon area, north of Manitou Springs, El Paso County, Colorado (Masters thesis): Golden, Colorado School of Mines, 80 p. Mutschler, F. E., Larson, E. E., & R.M. Bruce: 1987: Laramide and younger magmatism in Colorado-New petrologic and tectonic variations on old themes: Colorado School of Mines quarterly 82, 4, 1-47. Sawyer, D. A. et al., 1995: New chemical criteria for quaternary yellowstone tephra layers in central and western North America: Geological Society of America Abstracts with Programs, 27, 6, 109. Scott, G. R., 1963, Nussbaum Alluvium of Pleistocene(?) age at Pueblo, Colorado. U. S. Geological Survey Professional Paper, 475-C, C49-C52 Scott, G. R., 1975, Cenozoic surfaces and deposits in Curtis, B. F., ed., Cenozoic History of the Southern Rocky Mountains: Geological Society of America Memoir 144, 227-248. Soister, E., 1967, Relation of Nussbaum Alluvium (Pleistocene) to the Ogallala Formation (Pliocene) and to the Platte-Arkansas divide, Southern Denver Basin, Colorado. U. S. Geological Survey Professional Paper 575-D, p.D39-D46. Szabo, B. J., 1980, Results and assessment of uranium-series dating of vertebrate fossils from quaternary alluviums in Colorado: Arctic and Alpine Research, 12, 95-100. Tarling, D. H., 1983, Palaeomagnetism; principles and applications in geology, geophysics and archaeology: Chapman and Hall Ltd., London, 379 p. Trimble, D. E., & Machette, M. M., 1979, Geologic map of the Colorado Springs-Castle Rock Area, Front Range Urban Corridor, Colorado; U. S. Geological Survey, 1:100,000, Map I-857-F Tweto, O., 1975, Laramide (Late Cretaceous-Early Tertiary) Orogeny in the Southern Rocky Mountains in Curtis, B.F., ed., Cenozoic History of the Southern Rocky Mountains: Geological Society of America Memoir 144, 1-44. Verosub, K. L., 1977, Depositional and post-depositional processes in the magnetization of sediments: Reviews of Geophysics and Space Physics, 15, 129-143. TIME in KARST – 2007 171 COBISS: 1.01 HOw LONG DOES EVOLUTION OF THE TROGLOMORPHIC FORM TAKE? ESTIMATING DIVERGENCE TIMES IN ASTyANAx MExICANUS KAKO DOLGO TRAJA EVOLUCIJA TROGLOMORFNIH OBLIK? OCENJEVANJE DIVERGENČNIH ČASOV PRI ASTyANAx MExICANUS Megan L. PORTER1, Katharina DITTMAR2 & Marcos PéREZ-LOSADA3 Abstract UDC 551.44:597 591.542 Megan L. Porter, Katharina Dittmar & Marcos Pérez-Losada: How long does evolution of the troglomorphic form take? Estimating divergence times in Astyanax mexicanus Features including colonization routes (stream capture) and the existence of both epigean and cave-adapted hypogean populations make Astyanax mexicanus an attractive system for investigating the subterranean evolutionary time necessary for acquisition of the troglomorphic form. Using published sequences, we have estimated divergence times for A. mexicanus using: 1) two diferent population-level mitochondrial datasets (cyto-chrome b and NADH dehydrogenase 2) with both strict and relaxed molecular clock methods, and 2) broad phylogenetic approaches combining fossil calibrations and with four nuclear (recombination activating gene, seven in absentia, forkhead, and ?-tropomyosin) and two mitochondrial (16S rDNA and cytochrome b) genes. Using these datasets, we have estimated divergence times for three events in the evolutionary history of troglomorphic A. mexicanus populations. First, divergence among cave haplotypes occurred in the Pleistocene, possibly correlating with fuctuating water levels allowing the colonization and subsequent isolation of new subterranean habitats. Second, in one lineage, A. mexicanus cave populations experienced introgressive hybridization events with recent surface populations (0.26-2.0 Ma), possibly also correlated with Pleistocene events. Finally, using divergence times from surface populations in the lineage without evidence of introgression as an estimate, the acquisition of the troglomorphic form in A. mexicanus is younger than 2.2 (fossil calibration estimates) – 5.2 (cytb estimate) Ma (Pliocene). Key words: Astyanax mexicanus, divergence time, troglomor-phy, subterranean, evolution. Izvleček UDK 551.44:597 591.542 Megan L. Porter, Katharina Dittmar & Marcos Pérez-Losada: Kako dolgo traja evolucija troglomorfnih oblik? Ocenjevanje divergenčnih časov pri Astyanax mexicanus Značilnosti, ki vključujejo tudi kolonizacijske poti in obstoj tako epigejičnih kot hipogejičnih populacij vrste Astyanax mexica-nus, ji omogočajo, da predstavlja zanimiv sistem za proučevanje evolucije in časa, potrebnega za razvoj podzemeljskih troglo-morfnih oblik. Za A. mexicanus smo na podlagi že objavljenih sekvenc ocenili divergenčni čas ob uporabi: 1) dveh različnih populacijskih mitohondrialnih podatkovnih baz (citokrom b in NADH dehidrogenaze 2), obe z natančno in sproščeno metodo molekularne ure, in 2) razširjenega flogenetskega pristopa v kombinaciji s fosilno kalibracijo ter štirimi jedrnimi geni (rekombinacijski aktivacijski gen, »forkhead kontrolni gen« in ?-tropomiozin) in dvema mitohondrialnima genoma (16S rDNA in citokrom b). Ob uporabi navedenih podatkovnih baz smo ocenili divergenčni čas za tri dogodke v zgodovini razvoja troglomorfnih populacij A. mexicanus. Prvič, razhajanje med podzemeljskimi haplotipi se je zgodilo v Pleistocenu, verjetno v odvisnosti od nihanja vode, ki je omogočilo kolonizacijo in posledično izolacijo v novih podzemeljskih habitatih. Drugič, verjetno je v povezavi s pleistocenskimi dogodki pri eni liniji podzemeljskih populacij A. mexicanus prišlo do introgresivne hibridizacije s takratnimi površinskimi populacijami (0.26-2.0 Ma). Z uporabo divergenčnega časa površinskih populacij tistih linij, ki ne kažejo introgresije ocenjujemo, da je troglomorfna oblika A. mexicanus mlajša od 2,2 (ocene fosilne kalibracije) do 5,2 milijona let (cytb ocena) (Pliocen). Ključne besede: Astyanax mexicanus, divergenčni čas, troglo-morfzem, podzemlje, speleobiologija, evolucija. 1 Dept. of Biological Sciences, University of Maryland Baltimore County, Baltimore, MD, USA; e-mail: porter@umbc.edu 2 Dept. of Molecular Biology, University of wyoming, Laramie, wy, USA 3 GENOMA LLC, 50E woodland Hills, Provo, UT 84653-2052, USA Received/Prejeto: 06.12.2006 TIME in KARST, POSTOJNA 2007, 173–182 MEGAN L. PORTER, KATHARINA DITTMAR & MARCOS PéREZ-LOSADA INTRODUCTION Understanding the evolution of the cave form has fascinated biologists interested in subterranean faunas since Darwin. Termed ‘troglomorphy’, the suite of progressive and regressive characters associated with cavernicolous animals can be observed in the worldwide convergence of form found in the cave environment, exhibited in similar structural, functional, and behavioral changes across diverse taxonomic groups. Much of the debate over troglo-morphy has centered on the evolutionary mechanisms responsible for character regression, generally argued to be either neutral mutation or natural selection. Several studies, (Gammarus minus - Culver et al., 1995; Astyanax mexicanus – Jefery, 2005) have shown eye degeneration is the result of selection, and, in the case of A. mexica-nus, is caused by the pleiotropic efects of natural selection for constructive traits. Another, less studied, aspect of understanding troglomorphy is the evolutionary time required to gain the cave form. Because it is generally dif-fcult to pinpoint the time of subterranean colonization and isolation from surface ancestors, few troglomorphic species ofer the opportunity for quantitative estimates of the evolutionary time spent in the subterranean realm. Terefore, the time of cave adaptation is thought of in relative terms, where the degree of eye and pigment reduction indicates the period of cavernicolous evolution and therefore the relative phylogenetic age of each species (Aden, 2005). In evolutionary studies of cave adaptation, Asty-anax mexicanus has become a model system (Jefery, 2001). Te advantageous features of A. mexicanus as a model system include the existence of both surface and troglomorphic cavefsh populations, with several cave fsh populations having evolved constructive and regressive changes independently (Jefery, 2001). Furthermore, since the discovery of the species in 1936 (Hubbs & Innes, 1936), there has been an extensive amount of research devoted to characterizing developmental, phylogenetic, taxonomic, and biogeographic aspects of the species (Jef-fery, 2001; Mitchell et al., 1977; wiley & Mitchell, 1971;). In terms of being a model system for understanding the evolution of the troglomorphic form, A. mexicanus has at least one additional favorable attribute. Te primary mode of A. mexicanus subterranean colonization is via stream capture, with most of the captured surface drainages no longer supporting epigean populations (Mitchell et al., 1977). Tese captures provide discrete colonization events correlated with divergence time from surface populations and therefore with the time of subterranean evolution. Molecular studies that have looked at A. mexicanus phylogeography indicate that at least two independent invasions of surface Astyanax have occurred (Dowling et al., 2002a; Strecker et al., 2003, 2004). Tese two distinct A. mexicanus genetic lineages consist of cave fsh from La Cueva Chica, La Cueva de El Pachón, El Sótano de yerbaniz, El Sótano de Molino, El Sótano de Pichijumo, and La Cueva del Río Subterráneo (lineage A) and from La Cueva de los Sabinos, El Sótano de la Tinaja, La Cueva de la Curva, and El Sótano de Las Piedras (Lineage B) with diferent evolutionary histories - Lineage A clusters with closely related epigean populations while lineage B has no closely related epigean counterparts. Te close association of Lineage A to epigean populations (as estimated by mitochondrial markers) is thought to be the result of either recent subterranean colonization or refect recent introgressive hybridization with surface populations, while lineage B is considered to be a more ancient colonization event from surface populations that are extinct in the region (Dowling et al., 2002a; Strecker et al., 2004). Although the evolutionary histories of different hypogean A. mexicanus populations are complex, the two lineages ofer the unique opportunity to estimate the divergence time required for the evolution of the tro-glomorphic form based on discrete times of colonization and the previous molecular studies of their phylogeogra-phy. At least one other study has estimated lineage ages in A. mexicanus populations; however, this study was based on a single gene molecular clock estimate and did not specifcally estimate the divergence times of the cave populations (Strecker et al., 2003). Here we use three different sets of publicly available sequence data and known fossil calibrations and apply multiple phylogenetic approaches to estimate the age of cave colonization and stream capture events, and to provide an estimate of the time necessary to acquire the troglomorphic form in A. mexicanus. 174 TIME in KARST – 2007 HOw LONG DOES EVOLUTION OF THE TROGLOMORPHIC FORM TAKE? ESTIMATING DIVERGENCE TIMES ... METHODS Sequence Data Data were acquired from Genbank (http://www.ncbi. nlm.nih.gov/) from previously published studies of A. mexicanus and characiform fshes (Tab. 1). Tese studies provided three diferent datasets, consisting of: 1) population-level haplotype datasets for the mitochondrial cy-tochrome b (cytb; Strecker et al., 2004) and NADH dehy-drogenase 2 (ND2; Dowling et al., 2002a) genes, and 2) a species-level dataset of four nuclear (recombination activating gene – RAG2; seven in absentia – sina; forkhead – fh; and ?-tropomyosin - trop) and two mitochondrial genes (16S rDNA and cytb) from representatives within the Otophysi (Calcagnotto et al., 2005). Divergence times from all three data sets were estimated and compared. Species-level Phylogenetic Analyses Te species-level dataset included selected Otophysi, Characiformes, and Characidae sequences (see Tab. 1), and was analyzed using Anotophysi species as outgroups. Representative A. mexicanus cytb haplotype sequences from the Strecker et al., (2004) study were included in the dataset of characiform species to estimate divergence times based on fossil calibrations for comparison with population-based estimates utilizing substitution rates. Alignments of protein-coding regions were trivial and were accomplished using amino acid translations. Sequences of the trop gene spanned an intron, which was removed due to signifcant length variation (70-836 bp) leading to ambiguous alignments. Te alignment of the 16s rDNA gene was generated using the E-INS-i accuracy-oriented strategy of MAFFT v.5 (Katoh et al., 2005). All of the individually aligned genes were then concatenated to form a single dataset consisting 3770bp in length. Te concatenated dataset was analyzed with PAUP* 4.0b10 (Swoford, 2000) using maximum parsimony and implementing the parsimony ratchet method (Nixon, 1999) using a batch fle generated by PAUPRat with the default parameters for 5000 replicates (Sikes & Lewis, 2001). Divergence time estimation Population analysis. Dates of divergence were inferred for A. mexicanus lineage A and B cave fsh populations using the cytb and ND2 datasets with BEASTv1.4 (Drummond & Rambaut, 2003). Because the cytb and ND2 haplotype datasets were generated from diferent studies, they cannot be combined. Terefore, each dataset was used to independently estimate the divergence times of the A. mexicanus cave-adapted haplotype sequences. Each dataset was analyzed using both strict and relaxed clock models (Drummond et al., 2006) tested under constant and skyline models of population growth. As part of BEAST divergence time estimation, either a calibration point (fossil or geologic) or a gene-specifc substitution rate is required. Because there are no geologic dates corresponding to A. mexicanus populations invading subterranean systems, substitution rates were used. For each gene, the range of substitution rates calculated for other freshwater fsh were used. For cytb, mean substitution rates ranged from 0.005 to 0.017 substitutions/site/million year (my) (Bermingham et al., 1997; Burridge et al., 2006; Dowling et al., 2002b; Perdices & Doadrio, 2001; Sivasundar et al., 2001; Zardoya & Doadrio, 1999) and for ND2 mean substitution rates ranged from 0.011 to 0.026 substitutions/site/my (Near et al., 2003; Mateos, 2005). Tese independent rates were used to calibrate the rate of evolution of our datasets by either fxing the rate to the lowest and highest value estimated for each gene or using strong prior distributions on the substitution rates. Two independent MCMC analyses 2x107 steps long were performed sampling every 2,000th generation, with a burn-in of 2x106 generations. All the Bayesian MCMC output generated by BEAST was analyzed in Tracer v1.3 (Drummond & Rambaut, 2003). Likelihood-based AhRS method. we used the likelihood heuristic rate-smoothing algorithm of (yang, 2004) as implemented in PAML3.14 (yang, 2001). Sequence data were analyzed using the F84+? model. Branches at each locus were classifed into four rate groups according to their estimated rates. Te oldest known fossil representatives of major lineages within the Ostariophysi are well established in recent literature (see Briggs, 2005 and references therein), and have been used in recent studies estimating molecular-based divergence times of Otocephalan clades (Peng et al., 2006). Tese fossil representatives were used as calibration points for the AHRS divergence time analysis (Fig. 1, Tab. 2,). Fossil calibrations were accommodated as fxed ages and mapped to the basal node of the clade of interest. Given that most fossils are dated to an age range, the minimum and maximum ages of each fossil were used for divergence time estimations under separate analyses. Fossil dates were determined using the 1999 GSA Geologic Time Scale. TIME in KARST – 2007 175 MEGAN L. PORTER, KATHARINA DITTMAR & MARCOS PéREZ-LOSADA tab. 1: taxonomy, gene data, and Genbank accession numbers for sequences used in Characiformes phylogeny reconstruction. Abbreviations of mitochondrial gene sequences: 16S = 16S rdNA, cytb = cytochrome b; abbreviations for nuclear gene sequences: fh = forkhead, RAG2 = recombination activating gene, sina = seven in absentia, trop = ?-tropomyosin. Anotophysi (outgroup) Chanidae Chanos chanos Gonorynchidae Gonorynchus greyi Kneriidae Cromeria nilotica Parakneria cameronensis Otophysi (ingroup) CHARACIFORMES 16S cytb fkh RAG2 sina trop NC004693 NC004693 NC004702 NC004702 NC007881 NC007881 NC007891 NC007891 Anostomidae Leporinus sp. Chilodontidae Chilodus punctatus Prochilodontidae Prochilodus nigricans Hemiodontidae Hemiodus gracilis Parodontidae Parodon sp. Serrasalmidae Colossoma macropomum Cynodontidae Hydrolycus pectoralis Characidae Acestrorhynchus sp. Aphyocheirodon sp. Astyanacinus sp.1 Astyanacinus sp.2 Astyanax bimaculatus Astyanax mexicanus (Brazil) Astyanax mexicanus (haplotype AB) Astyanax mexicanus (haplotype AL) Astyanax mexicanus (haplotype EA) Astyanax mexicanus (haplotype FA) Astyanax mexicanus (haplotype GA) Astyanax mexicanus (haplotype GB) Astyanax scabripinis Brycon hilarii Bryconamericus diaphanus Bryconops sp. Chalceus erythrurus Chalceus macrolepidotus Cheirodon sp. Cheirodontops sp. Creagrutus sp. Exodon paradoxus Gephyrocharax sp. Hemibrycon beni Hemigrammus bleheri AY788044 AY791416 AY817370 AY804095 AY790102 AY817252 AY787997 AY788075 AY788027 AY788065 AY788000 AY788033 AY787956 AY787966 AY787969 AY787987 AY787955 AY817325 AY790056 AY817215 AY791437 AY791405 AY791427 AY791386 AY817400 AY804120 AY790133 AY817278 AY817353 AY804084 AY790086 AY817240 AY817390 AY804110 AY790123 AY817269 AY817328 AY804061 AY790059 AY817218 AY817359 AY804088 AY790091 AY817244 AY791353 AY791363 AY791365 AY817288 AY817298 AY817301 AY817317 AY804026 AY804031 AY804033 AY804051 AY790014 AY790025 AY790028 AY790046 AY817181 AY817190 AY817209 AY817287 AY804025 AY790013 AY817180 AY177206 AY639041 AY639051 AY639075 AY639084 AY639089 AY639090 AY787967 AY787976 AY787984 AY787985 AY787990 AY787999 AY787995 AY787996 AY788001 AY788013 AY788014 AY788020 AY788017 AY791370 AY791375 AY791376 AY791379 AY791385 AY791382 AY791383 AY817299 AY817307 AY817314 AY817315 AY817320 AY817327 AY817324 AY791397 AY791398 AY791402 AY817340 AY817341 AY817346 AY817343 AY804040 AY804048 AY804049 AY804053 AY804060 AY804057 AY804058 AY804062 AY804072 AY804073 AY804079 AY804076 AY790026 AY790035 AY790043 AY790044 AY790049 AY790058 AY790054 AY790055 AY790060 AY790072 AY790073 AY790079 AY790076 AY817188 AY817198 AY817206 AY817207 AY817211 AY817217 AY817219 AY817227 AY817228 AY817234 AY817231 176 TIME in KARST – 2007 HOw LONG DOES EVOLUTION OF THE TROGLOMORPHIC FORM TAKE? ESTIMATING DIVERGENCE TIMES ... 16S cytb fkh RAG2 sina trop Hemigrammus erythrozonus Hemigrammus rodwayi Hyphessobrycon eques Inpaichthys kerri Knodus sp. Moenkhausia sanctaphilomenae Mimagoniates lateralis Prodontocharax sp. Roeboides sp. Salminus maxillosus Triportheus angulatus Ctenolucidae Ctenolucius hujeta Lebiasinidae Nannostomus beckfordi Crenuchidae Characidium fasciatum Erythrinidae Hoplias sp. Alestidae Arnoldichthys spilopterus Brycinus nurse Phenacogrammus aurantiacus Hepsetidae Hepsetus odoe Citharinidae Citharinus citharus Distichodontidae Distichodus sexfasciatus Neolebias trilineatus AY788023 AY788034 AY788022 AY788039 AY788041 AY788054 AY788051 AY788064 AY787994 AY788080 AY788082 AY787998 AY788059 AY787992 AY788031 AY787968 AY787970 AY788066 AY788030 AY787989 AY788012 AY788063 AY817349 AY817360 AY817348 AY817365 AY791414 AY817367 AY791420 AY791426 AY791381 AY791438 AY791384 AY791380 AY791409 AY791364 AY791366 AY791428 AY791408 AY791378 AY791396 AY791425 AY817377 AY817389 AY817323 AY817405 AY817407 AY817326 AY817384 AY817322 AY817357 AY817300 AY817302 AY817391 AY817356 AY817319 AY817339 AY817388 AY804081 AY804089 AY804080 AY804093 AY804094 AY804104 AY804101 AY804109 AY804056 AY804124 AY804125 AY804059 AY804055 AY804087 AY804032 AY804034 AY804111 AY804086 AY790082 AY790092 AY790081 AY790097 AY790099 AY790112 AY790109 AY790122 AY790053 AY790137 AY790139 AY790057 AY790117 AY790051 AY817236 AY817245 AY817235 AY817248 AY817249 AY817261 AY817259 AY817214 AY817282 AY817283 AY817216 AY817265 AY817213 AY790090 AY817242 AY804071 AY804108 AY790027 AY790029 AY790124 AY790089 AY790048 AY790071 AY790121 AY817189 AY817191 AY817270 AY817241 AY817226 AY817268 CYPRINIFORMES Cobitidae Misgurnus sp. Cyprinidae Danio rerio Labeo sorex Gyrinocheilidae Gyrinocheilus sp. SILURIFORMES Callichthyidae Corydoras rabauti Loricariidae Ancistrus sp. Bagridae Chrysichthys sp. Heptapteridae Pimelodella sp. Ictaluridae Ictalurus punctatus AY788053 AY817379 AY804103 AY790111 AY788011 — AY817338 AY804070 AY790070 AY817225 AY788043 AY791415 AY817369 — AY790101 AY817251 AY788015 AY791399 AY804074 AY790074 AY817229 NC004698 NC004698 AY787958 AY791354 AY817290 AY787957 AY791355 AY787953 AY791351 AY817285 AY788040 AY791413 AY817366 AY790016 AY817183 AY790017 AY817193 AY790011 AY817178 AY790098 TIME in KARST – 2007 177 MEGAN L. PORTER, KATHARINA DITTMAR & MARCOS PéREZ-LOSADA Fig. 1: Characiform divergence time chronogram estimated using a representative topology chosen from the set of 867 most parsimonious trees. White branches indicate branches where less than 75% of the most parsimonious trees were topologically congruent. Te grey box indicates the clade of Astyanax mexicanus sequences. Fossil calibration nodes are numbered and correspond to tab. 2. Te major geologic periods are mapped onto the phylogeny. 178 TIME in KARST – 2007 HOw LONG DOES EVOLUTION OF THE TROGLOMORPHIC FORM TAKE? ESTIMATING DIVERGENCE TIMES ... tab. 2: taxonomy and ages of fossils used as calibrations for divergence time estimation. Node # refers to Fig. 1. Taxonomy Reference Geologic age (MYA) Node # Otophysi Characiformes Gayet, 1982 Late Cretaceous (65-99) 1 Cypriniformes Catostomidae Cavender, 1986 Paleocene (54.8-65) 4 Siluriformes Gayet & Meunier, 2003 late Campanian-early Maastrichtian (68.2-77.4) 3 Corydoras Cockerell, 1925 Late Palaeocene (61-65) 2 RESULTS Population-level divergence time estimations. Estimates of the mean divergence times were not signifcantly different between strict and relaxed clock and population growth models and calibration methods of the substitution rate, but confdence intervals under the fxed substitution rate approach were narrower, as expected. Hence only the time estimates under the strict clock model, constant population size and minimum and maximum mean substitution rates for both genes are provided. Comparing the cytb and ND2 estimates of divergence times for the A. mexicanus A and B lineages show several features. First, the estimated ranges of divergence for cave hap-lotypes within each lineage were similar between genes (cytb and ND2) and lineages (A and B), placing the divergence among hypogean populations between 0.141-0.885 Ma for lineage A, and 0.084-0.575 Ma for lineage B (Tab. 3). when comparing the estimates among genes within a lineage, however, the divergence times of hypo-gean and epigean haplotypes are diferent, with cytb estimates providing generally older estimates. Species-level divergence time estimation. Using the maximum parsimony ratchet, the selected Characidae, Characiform, and Otophysi sequences generated 867 trees of score 11758. Te 50% majority rule consensus of these trees was similar to the published research that generated the data (Calcagnotto et al., 2005). Because a fully resolved tree with branch lengths is required for AHRS divergence time estimation and because very few branches in the consensus tree collapsed (e.g. were in confict), a random tree from the set of 867 was used (Fig. 1). Te A. mexicanus sequences included in the analysis clustered with other Characidae species, although were not monophyletic with other Astyanax species (A. bi-maculatus and A. scabripinnis). Te divergence time estimates for the representative A. mexicanus cave fsh populations generated using this phylogeny with Oto-physi fossil calibrations agreed well with the estimates of hypogean haplotype divergence from cytb and ND2 using substitution rates (Tab. 3). However, the estimates of cave versus surface population divergence times based on fossil calibrations were in better agreement with ND2 than with cytb estimates. Tis is particularly interesting, as the only gene included in this dataset for A. mexica-nus was cytb. tab. 3: Comparison of divergence time estimates using substitution rates and molecular clock methods for cytochrome b (cytb) and NAdh dehydrogenase 2 (Nd2) mitochondrial genes, and for molecular methods incorporating fossil dates as calibrations. Substitution Rates Fossil Calibration Cytb ND2 Min – Max (Ma) Min – Max (Ma) Min – Max (Ma) Lineage A cave 0.261 - 0.885 0.141 - 0.331 0.2-0.3 cave vs. surface 0.588 - 2.00 0.256 - 0.599 0.4-0.5 Lineage B cave 0.169 - 0.575 0.084 - 0.196 0.1-0.1 cave vs. surface 1.524 - 5.181 0.877 - 2.055 1.7-2.2 Lineage A vs. Lineage B 1.741 - 5.922 1.053 - 2.472 1.7-2.2 TIME in KARST – 2007 179 MEGAN L. PORTER, KATHARINA DITTMAR & MARCOS PéREZ-LOSADA DISCUSSION Previous molecular studies of A. mexicanus phylogeog-raphy indicate that at least two independent invasions of surface Astyanax have occurred (Dowling et al., 2002a; Strecker et al., 2003, 2004). Our estimates of divergence time from two diferent methods and three diferent datasets are in general agreement about the divergence times among the cave haplotypes in each lineage (Tab. 3). Tese estimates place cave haplotype divergence times in the Pleistocene, when it is suggested that climatic cooling of surface waters led to the extinction of Astyanax in North America (Strecker et al., 2004). In particular, our data show an interesting pattern for lineage B haplotypes, which are proposed to be the older of the two lineages. Te recent divergence times estimated for lineage B hap-lotypes (0.084-0.575 Ma) supports the hypothesis that afer the initial colonization event, subterranean routes of colonization were associated with fuctuating ground-water levels in the Pleistocene (Strecker et al., 2004). Te fact that estimated times of within lineage divergence were similar also suggests that the divergence of subterranean haplotypes in both lineages were infuenced by the same processes. In order to determine the evolutionary age of the subterranean lineage, and therefore estimate the time required for evolution of the troglomorphic form, the divergence of the hypogean haplotypes from epigean populations is needed. However, the estimates from our three datasets did not agree, with cytb molecular clock methods estimating older divergence times than either ND2 or fossil calibrated estimates. Some of the discrepancy is due to the fact that diferent sets of surface populations were sampled in each study (Dowling et al., 2002a; Strecker et al., 2004). For example, the most closely related surface population in the cytb study were from Belize (Strecker et al., 2004) while there were no closely related surface populations to lineage B haplotypes in the ND2 study (Dowling et al., 2002a). However, this makes the older cytb estimates even more notable because lineage B haplotypes have no evidence of introgressive hybridization with surface populations. If we consider just lineage B hypogean divergence from surface ancestors as an estimate of subterranean evolution, the estimated time for acquisition of the troglomorphic form is 0.877-2.055 Ma (quaternary – Tertiary boundary) based on ND2 and fossil calibrations, while it is 1.524-5.181 Ma (Pliocene) based on cytb. Although the estimates of divergence times among the three diferent datasets did not agree, comparison of estimates between the lineages show that lineage A diverged from surface ancestors more recently than lineage B (Tab. 3). Tis more recent divergence from epigean populations is congruent with previous hypotheses, that either lineage A populations represent a more recent subterranean invasion, or that they are an older invasion masked by more recent mitochondrial intro-gressive hybridization with surface forms (Dowling et al., 2002a). In the few studies that have looked at other markers (allozymes, microsatellites, and RAPDs), it has been suggested that at least Chica and Pachón populations are the result of surface introgression (Avise & Se-lander, 1972; Espinasa & Borowsky, 2001; Strecker et al., 2003). Furthermore, based on the degree of variability in troglomorphic features of each lineage A population, it has been suggested that diferent populations represent diferent degrees and patterns of surface introgression. In order to more accurately determine both the patterns of introgression in the lineage A populations, as well as the underlying relationships of the cave populations to each other in order to estimate subterranean evolutionary times, studies investigating more types of markers are needed. Previous research of A. mexicanus populations throughout Mexico (including cavefsh lineages A and B) estimated haplotype divergences to range from 1.8 – 4.5 Ma (Strecker et al., 2004). Our estimates suggest that divergence times among cave haplotypes and between lineage A cave and epigean haplotypes are much younger than this; however, hypogean divergences from surface ancestors in lineage B are concordant with these older dates. Te evolutionary history of cave adaptation in A. mexicanus is complex. Based on mitochondrial molecular clock estimates, our estimates of divergence times are congruent with previous hypotheses by showing lineage B to be a phylogenetically older subterranean lineage, with more recent divergence among subterranean systems. However, this study also provides quantitative dates for these events. Lineage A populations are estimated to be younger; however, these dates only represent mito-chondrial lineages. Several of the populations in lineage A have been shown to be introgressed with surface forms (Chica, Pachón, and Subterraneo). To our knowledge, the hypothesis of surface introgression has not been investigated in the remaining lineage A populations (Molino, Pichijumo, and yerbaniz). Understanding the patterns of introgression in all of the lineage A populations, and estimating the actual subterranean evolutionary time, requires investigating additional nuclear markers. 180 TIME in KARST – 2007 HOw LONG DOES EVOLUTION OF THE TROGLOMORPHIC FORM TAKE? ESTIMATING DIVERGENCE TIMES ... CONCLUSIONS Features including colonization routes (stream capture) and the existence of both epigean and cave-adapted hy-pogean populations make A. mexicanus an attractive system for investigating the subterranean evolutionary time necessary for acquisition of the troglomorphic form. If it is possible to estimate the divergence time of closely related cave versus surface populations, we can estimate the age of subterranean occupancy. Tis same divergence time also has relevancy to geologic processes in the karst system by providing a rough estimate of the age of subterranean stream capture in particular regions. Based on published sequence data, we have estimated divergence times for three events in the evolutionary history of troglomorphic A. mexicanus populations. First, divergence times among cave haplotypes in both lineages occurred in the Pleistocene, possibly correlating with fuctuating water levels allowing the colonization, and subsequent isolation of, new subterranean habitats. Second, in lineage A, A. mexicanus cave populations experienced introgressive hybridization events with surface populations recently. Finally, using divergence times of lineage B from surface populations as an estimate, the acquisition of the troglomorphic form in A. mexicanus is younger than 2.2 (fossil calibration) – 5.2 (cytb) Ma (Pliocene). Given that there are at least 30 caves known to contain populations of A. mexicanus (Espinasa et al., 2001; Mitchell et al., 1977), the number of independent invasions and instances of introgressive hybridization may be even higher than currently understood. In order to fully understand the number of independent invasions, the history of introgression with surface populations, and the divergence times of cave and surface populations, a broader survey of cave fsh populations and of both nuclear and mitochondrial markers is needed. LITERATURE CITED Aden, E., 2005: Adaptation to darkness. In Culver, D.C., & white, w.B., (eds.), Encyclopedia of Caves, Elsevier Academic Press, pp.1-3. Avise, J.C., & R.K. Selander., 1972: Genetics of cave-dwelling fshes of the genus Astyanax. –Evolution, 26, 1-19. Bermingham, E., McCaferty, S.S., & A.P. Martin., 1999: Fish biogeography and molecular clocks: perspectives from the Panamanian isthmus. 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Doadrio., 1999: Molecular evidence on the evolutionary and biogeographical patterns of European cyprinids. -Journal of Molecular Evolution, 49, 227-237. 182 TIME in KARST – 2007 COBISS: 1.01 AGE ESTIMATES FOR SOME SUBTERRANEAN TAxA AND LINEAGES IN THE DINARIC KARST OCENE STAROSTI ZA NEKATERE PODZEMELJSKE TAKSONE IN ŽIVALSKE LINIJE NA DINARSKEM KRASU Peter TRONTELJ1, Špela GORIČKI1, Slavko POLAK2, Rudi VEROVNIK1, Valerija ZAKŠEK1 & Boris SKET1 Abstract UDC 575.8:551.442(234.422.1) 591.542(234.422.1) Peter Trontelj, Špela Gorički, Slavko Polak, Rudi Verovnik, Valerija Zakšek & Boris Sket: Age estimates for some subterranean taxa and lineages in the Dinaric Karst Using a comparative phylogeographic approach and diferent independent molecular clocks we propose a timescale for the evolution of troglobionts in the Dinaric Karst that is relatively consistent over a wide taxonomic range. Keystone events seem to belong to two age classes. (1) Major splits within holodinaric taxa are from the mid-Miocene. Tey present the potential upper limit for the age of cave invasions. (2) Regional diferentia-tion, including speciation, which can at least in part be associated with a subterranean phase, took place from early Pliocene to mid-Pleistocene. we suggest two to fve million years as the time when most of the analyzed lineages started invading the Dinaric Karst underground. Key words: subterranean, molecular clock, molecular phylog-eny, phylogeography, Dinaric Karst. Izvleček UDK 575.8:551.442(234.422.1) 591.542(234.422.1) Peter Trontelj, Špela Gorički, Slavko Polak, Rudi Verovnik, Valerija Zakšek & Boris Sket: Ocene starosti za nekatere podzemeljske taksone in živalske linije na Dinarskem krasu Z uporabo primerjalnega flogeografskega pristopa in neodvisnih molekularnih ur smo predlagali časovni potek evolucije troglobiontov Dinarskega krasa, ki velja za sorazmerno veliko število taksonov. Zdi se, da ključni dogodki pripadajo dvema obdobjema. (1) Glavne razdelitve znotraj holodinarskih tak-sonov so iz obdobje srednjega miocena. Predstavljajo zgornji potencialni časovni limit za naselitev jam. (2) Regionalna diferenciacija, vključno s speciacijo, ki je lahko vsaj deloma povezana s podzemeljsko fazo, naj bi se zgodila med zgodnjim in srednjim pleistocenom. Ocenjujemo, da se je začela invazija večine proučevanih živalskih linij v podzemlje Dinarskega krasa v obdobju med dvema in petimi milijoni let. Ključne besede: podzemlje, molekularna ura, molekularna flogenija, flogeografja, Dinarski kras. INTRODUCTION Te use of new molecular and systematic techniques using allozymes and DNA sequences has enabled us to see a new picture of the evolution and diversity of subterranean fauna (e.g. Avise & Selander 1972; Sbordoni et al., 2000; Caccone & Sbordoni 2001; Leys et al., 2003; Verovnik et al., 2004; Gorički & Trontelj 2006; Lefébure et al., 2006; Zakšek et al., 2007). Molecular clock ap- proaches should, at least in theory, enable us to date, to verify or to falsify previous hypotheses about the age of subterranean species. To be exact, it is usually not the age of a lineage or a taxon itself that is of special interest or under dispute, but the time since it has attained its subterranean nature, making it even more challenging. Hypotheses and models explaining cave invasions 1 Oddelek za biologijo, Biotehniška fakulteta, Univerza v Ljubljani, Večna pot 111, 1000 Ljubljana, Slovenia, fax: +386 1 2573390, e-mail: peter.trontelj@bf.uni-lj.si 2 Notranjski muzej Postojna, Ljubljanska 10, 6230 Postojna, Slovenia. Received/Prejeto: 30.01.2007 TIME in KARST, POSTOJNA 2007, 183–189 PETER TRONTELJ, ŠPELA GORIČKI, SLAVKO POLAK, RUDI VEROVNIK, VALERIJA ZAKŠEK & BORIS SKET and speciation in caves are well-elaborated (e.g. Rouch & Danielopol 1987; Holsinger 2000; Trajano 2005) and should thus ofer good grounds for the timing of such events and for testing their correlation with geographical, geological and hydrographical counterparts. For example, Leys et al., (2003) have shown that all evolutionary transitions to subterranean life in Australian dytiscids took place during the Late Miocene and Early Pliocene as a result of aridifcation. However, reliable data on the age of these events is surprisingly scarce. when such data are available, the accuracy is ofen below that of molecular clock rates. In fact, the use of molecular dating methods itself has introduced considerable uncertainty about how old subterranean species might be. while the youngest estimation, based on “classical” biological reasoning, is no more than 10,000 years (Sket 1997), the upper limit for the divergence of two subterranean sister species has been pushed to an incredible 110,000,000 years (Buhay & Crandall 2005). Boutin and Coineau (2000) have argued that dating of cladogenetic events by a molecular clock is particular- Te presented data were taken from several phylogeo-graphic studies of subterranean animals in the Dinaric Karst, including the ubiquitous aquatic isopod Asellus aquaticus Linne (Verovnik et al., 2004, 2005), the cave salamander Proteus anguinus Laurenti (Gorički 2006, Gorički & Trontelj 2006), and the cave shrimp troglo-caris s. lato (Zakšek et al., 2007). Further, we included unpublished sequences from studies that are in progress, including leptodirine cave beetles and aquatic sphaero-matid isopods from the genus monolistra. Te age estimations for the last two groups should be regarded as preliminary because in-depth analyses of phylogenetic relationships and corroboration by further loci are still under way. we were only interested in a small number of well-supported splits and therefore used straightfor- Te split between major geographically defned lineages Te geographical distribution of troglobiotic (including stygobiotic) sister taxa can be used to infer independent cave invasions. For example, if the present-day ranges of two troglobionts are separated by large areas of non-karstic terrain without hypogean habitat, we can ly useful in the case when the dates are corroborated by other methods. Since the obvious problem of the Dinaric Karst area is that reliable dating for clearly defned vicari-ant events or the age of available subterranean habitat is lacking, it has been impossible to corroborate molecular clock divergence by independent data. In this case a comparative phylogeographic approach might provide the means for an independent validation of age estimates. Comparative phylogeography seeks, as does historical biogeography, concordant geographical patterns of codistributed lineages (e.g. Arbogast & Kenagy 2001). Te evolution of codistributed phylogeographic groups of diferent taxa is likely to have been driven by the same historical factors, like vicariant events or climatic shifs. In this contribution we (1) identify common phylo-geographic patterns among those troglobiotic taxa from the Dinaric Karst for which such data are available, and (2) estimate the timeframe of the corresponding cladoge-netic events using a global molecular clock approach. ward minimum evolution searches with bootstrapping as implemented in MEGA (Kumar et al., 2004). Divergence time estimates are based on available clock-rate data for groups that are as closely related as possible (Caccone & Sbordoni [2001] for leptodirines, Ketmaier et al., [2003] for Asellus aquaticus, and Sturmbauer et al., [1996] and Schubart et al., [1998] for monolistra). To assure compatibility between molecular divergences we used the same models as were used in the original works describing the rates (Tamura-Nei distances with a gamma distributed rate variation among sites). where more than one hap-lotype per population or lineage was analyzed we used net between group distances to correct for ancestral in-traspecifc diversity. postulate an epigean last common ancestor. Examples of that kind can be found in the shrimp genus troglo-caris, with the hercegovinensis lineage inhabiting Trans-caucasian and SE parts of the Dinaric Karst where it is sympatric with the SE populations of the Anophthalmus lineage (Zakšek et al., 2007). Teir split estimated at 6–11 MATERIAL AND METHODS RESULTS 184 TIME in KARST – 2007 AGE ESTIMATES FOR SOME SUBTERRANEAN TAxA AND LINEAGES IN THE DINARIC KARST Myr ago is the oldest, although unlikely, possible time of cave invasion. Te youngest split that could be reliably inferred from the phylogenetic tree and probably still occurred in surface waters, was the one between the Bosnian lineage and other “Anophthalmus” lineages. Because the karst area in Bosanska Krajina, to which the Bosnian clade is restricted, is so remote and isolated from the rest of the Dinaric populations, it is reasonable to assume that an underground connection between them could never have existed. Te estimated time of this split, 3.7–5.3Myr ago, is hence the oldest possible age at which troglocaris anophthalmus might have invaded the Dinaric Karst underground (Tab. 1). For the cave salamander Proteus anguinus, exhibiting a distribution pattern similar to that of troglocaris, the corresponding age of the Bosanska Krajina lineage was estimated at 4.4–5.4 Myr (Gorički 2006). However, older lineages exist that, theorethically, might have invaded caves even as early as 8.8–16 Myr ago (see also Fig. 1). Another troglobiotic group restricted to the Dinaric Karst area and having a non-troglomorphic sister group is the Dinaric clade of Asellus aquaticus (see Verovnik et al., 2005). Te time of this split, and hence the maximum possible age of cave invasion is 3.8–4.8 Myr. TIMING OF MORPHOLOGICAL CHANGES where possible, we tried to combine the biology (e.g. degree of troglomorphism, lack of gene fow) of taxa with corresponding data on paleogeography and paleo-hydrography to infer speculative scenarios on how and when lineages might have switched to subterranean life and evolved troglomorphic traits. For example, we have some indication about how long at most it takes a salamander population to become troglomorphic. Since the subspecies P. a. parkelj Sket et Arntzen has retained its ancestral, non-troglomorphic characteristics, it is reasonable to conclude that its sister lineage must have evolved troglomorphoses independently from other, less related troglomorphic lineages (Sket & Arntzen 1994; Gorički & Trontelj 2006; see Fig. 1). Te split between the non-troglomorphic lineage and its last troglomorphic sister lineage was estimated at 0.5–0.6 Myr based on mitochon-drial rDNA sequences, 1.1–2.4 Myr based on the mtDNA control region (Gorički 2006), and at 1.1–4.5 Myr by an allozyme clock (Sket & Arntzen 1994). Asellus aquaticus has evolved several separate subterranean and troglomorphic populations. One of them, from the subterranean Reka River below the Kras/Carso Plateau, is genetically completely isolated from epigean populations at the Reka resurgence while there are no epi-gean populations in the Reka before the sink (Verovnik et al., 2003, 2004, 2005; Fig. 2). Further, it has no mtDNA tab. 1. Estimated time (in million years) of some keystone events in the evolution of troglobionts in the dinaric Karst. Taxon Age of holodinaric group Age of merodinaric group Mid-Dinaric split Northwest split Troglocaris (Dinaric and Caucasian lineages)1 7.9-15.1 n.a. n.a. n.a. Troglocaris anophthalmus agg.1 n.a. 3.7-5.3 1.3-2.3 1.5-2.1 Troglocaris hercegovinensis agg.1 n.a. 3.8-4.8 n.a. n.a. Proteus anguinus2 8.8-16.0 n.a. 8.8-16.0 4.2-5.2 Asellus aquaticus (Dinaric clade)3 n.a. 3.8-4.8 n.a. 0.8-1.2 Microlistra4 n.a. 1.1-2.3 n.a. n.a. Pseudomonolistra hercegoviniensis4 n.a. 0.3-1.0 n.a. n.a. Monolistra caeca4 n.a. 1.8-3.7 n.a. n.a. Leptodirus hochenwartii hochenwartii et L. h. reticulatus5 n.a. 1.9-2.0 n.a. n.a. 1Using COI clock for shrimps (see Knowlton & Weigt 1998; zakšek et al., 2007) 2Using 12S and 16S rdNA clock for Newts (see Cacconesee et al., 1997; Gorički 2006) 3Using COI clock for subterranean Asellota (see Ketmaier et al., 2003; verovnik et al 2005) 4Using 16S r-RNA clock for fddler crabs (Sturmbauer et al 1996) and land crabs (Schubart et all 1998) 5Using COI clock for subterranean leptodirine beetles (Caccone & Sbordoni 2001) TIME in KARST – 2007 185 PETER TRONTELJ, ŠPELA GORIČKI, SLAVKO POLAK, RUDI Fig. 1: A simplifed view of the phylogenetic relationships obetween troglomorphic and non-troglomorphic Proteus anguinus populations (from Gorički and trontelj 2006). Postulating a non-troglomorphic ancestor and unidirectional evolution toward troglomorphism, we can take the split between the black subspecies (non-troglomorphic) and its unpigmented sister lineage to estimate the maximal time (t1) needed for a salamander lineage to evolve the entire array of cave-related traits known in this taxon. If one accepts the notion of multiple independent cave invasions for Proteus, than t2 is the potentially oldest time since it has become subterranean. and nuclear rDNA haplotypes in common with hypoge-an populations from the Ljubljanica River drainage with which the Reka drainage has been connected many times during the Pleistocene and occasionally even nowadays (Habič 1989). It is thus reasonable to assume that the ancestor of the subterranean Reka River population invaded hypogean waters and became cave-adapted before any secondary contact could occur. Te estimated age of the Reka River lineage is 3.1–4.1 Myr (Verovnik et al., 2004), making it a pre-Pleistocene troglobiotic relict (Verovnik et al., 2004). monolistra, a troglobiotic group of freshawater sphaeromatid isopods, shows a high taxonomic and morphological diversity restricted to the Dinaric Karst and parts of the Southern Calcareous Alps. According to our preliminary results of a molecular phylogenetic analysis based on nuclear and mitochondrial DNA sequences, there are at least three well-supported monophyla. Tese are the subgenus m. (microlistra), m. (monolistra) caeca Gerstaecker, and the polytypic m. (Pseudomonolistra) hercegoviniensis Absolon. Several lines of evidence suggest that the common ancestors of each of these groups invaded cave waters polytopically (Sket 1986, 1994). while we remain ignorant about when and how ofen ancestral monolistra lineages invaded subterranean waters, we can expect that the radiation of at least some of the three groups took place in the underground. Teir ages VALERIJA ZAKŠEK & BORIS SKET Fig. 2: Te case of troglomorphic and non-troglomorphic lineages of Asellus aquaticus in the dinaric Karst, highly simplifed (from verovnik et al 2004, 2005). Te Reka and the Ljubljanica (Ljub) basin lineages have independently invaded subterranean waters and thus constitute separate taxa, although traditionally assigned to the same subspecies, A. a. cavernicolus. Te subterranean Reka River population presents the oldest stygobiotic lineage of Asellus aquaticus. because it is genetically completely distinct, it must have escaped interbreeding during various times of hydrological contact with surface populations. We therefore believe that it became a specialized stygobiont soon afer the split at time t1. Te Ljub lineage from the subterranean Ljubljanica River, although morphologically distinct, is still sharing mtdNA haplotypes with surface populations and thus represents a younger invasion. Eur and din denote various epigean European and dinaric lineages, respectively. (maximally 0.4–3.7 Myr) give us an idea for how long some monolistra lineages have been dwelling in the Di-naric Karst underground. Leptodirus hochenwartii Schmidt, a highly troglo-morphic leptodirine cave beetle, is the only terrestrial Dinaric troglobiont with available molecular dating. Using a leptodirine COI clock calibration by Caccone and Sbordoni (2001) we estimated the age of the Leptodirus lineage by dating the split with Astagobius angustatus Schmidt, its slightly less troglomorphic sister lineage. Te estimated time of this split (8.7–9.8 Myr ago) is the oldest possible age at which the extremely specialized morphology of Leptodirus could have started evolving. Moreover, taking into account recent unpublished phy-logenetic fndings based on nuclear and mitochondrial gene sequences, the traditional subspecies of Leptodirus in fact represent distinct lineages with divergences well in the range of between species comparisons. Tese lineages all share the same constructive apomorphic troglomor-phic characters, and it seems probable these troglomor-phies have already existed at least at the time of their last common ancestor. Te time of divergence between basal Leptodirus lineages hence represents the youngest possible age at which Leptodirus has evolved its full array 186 TIME in KARST – 2007 AGE ESTIMATES FOR SOME SUBTERRANEAN TAxA AND LINEAGES IN THE DINARIC KARST of troglomorphic characters. Based on a yet incomplete taxonomic sample (L. h. hochenwartii Schmidt and L. h. reticulatus J. Müller) we tentatively dated it at 1.9–2.0 Mya. TIMING OF PALEOHyDROGRAPHIC CHANGES For some stygobiotic taxa with a broader Dinaric range, we identifed two concordant geographic patterns possibly pointing to common underlying historical events, like changes in hydrographic connections. Tese vicari-ant patterns include (1) a split between a northwestern and southeastern Dinaric clade (mid-Dinaric split), and (2) a younger subdivision of the northwestern clade (or of a part thereof) into a western and eastern Slovenian lineage (Tab. 1). It has been stated that some stygobiotic species inhabit areas that are hydrographically fragmented. Te Before we reach any conclusions we would like to note that dating of keystone events in the evolution of subterranean life, as well as anywhere else in evolution (e.g. Graur & Martin 2004), remains a highly speculative enterprise. Of central concern should be the fact that we are relying on a more or less global clock within certain taxonomic boundaries. Tese clocks usually rely on single calibration points (e.g. the separation of the Sardinia-Corsica microplate from the Iberian Peninsula; Ketmaier et al., 2003) and have mostly not been tested against independent geological events. Further, all our timings assume linear accumulation of substitutions over time, i.e. the existence of a valid molecular clock. Although we can be quite sure that this assumption is violated to a certain extent, we can mitigate the problem by excluding those taxa from the analysis that violate the linearity assumption most. More sophisticated and realistically modeled approaches use a relaxed clock allowing for diferent local rates on diferent branches of the tree (e.g. Sanderson 2002). However, with single calibration points only, such approaches yield quite hopeless and certainly unrealistic intervals. For example, the age of the deepest split in the Niphargus virei (subterranean amphipod from France) complex was estimated at 14–19 Myr using a global Stenasellus clock, whereas the relaxed clock estimate was 22–71 Myr (Lefébure et al., 2006). Tird, it should be noted that even with the aid of molecular phylogenetic tools the timing is still susceptible to incorrect estimations of relationships and incomplete taxonomic coverage. For example, the timing of the most parsimonious explanation of such distributions is that their ranges were hydrographically interconnected in the past. Tis may as well include the surface paleohy-drography that was heavily fragmented by karstifcation. Te (polytopic) immigration underground could thus have proceeded simultaneously with the separation of ancestral populations. we can illustrate this scenario by the case of some monolistra lineages, namely of the subgenus microlistra and of the species m. (m.) caeca. Some ten microlistra spp. are perfectly allopatric in distribution, mainly bound to actual watersheds. Another group, m. caeca, inhabits at least three watersheds, in which four named subspecies have evolved. According to a 16S rDNA molecular clock (Sturmbauer et al., 1996, Schubart et al., 1998), the system began to fragment about three million years ago. origin of the highly troglomorphic morphologies in Lep-todirus depends on the most basal split in the taxon. By not having included all known subspecies, we are facing the risk that some other subspecies might have branched of earlier than the studied ones. One potentially useful way to improve our informal confdence in the timing of evolutionary events in subterranean animals is to look for phylogeographic correspondence of timings derived from independent taxa with independent molecular clocks. At the present stage of most of our analyses such comparisons can only be preliminary. we can nevertheless notice that specifc groups of events belong to diferent age classes, most markedly the gap between the age of holodinaric troglobionts and those with narrower distributions within the Dinaric Karst (Tab. 1). Te recent lineages of Proteus and troglocaris probably both originate from the Miocene Dinaride Lake System (Krstić et al., 2003), and the age of both taxa refects their diferentiation long before they invaded the hypogean environment (Sket 1997; Gorički 2006; Zakšek et al., 2007). Regional diferentiation, including speciation, which can at least in part be associated with a subterranean phase, appears to be much younger, ranging from Pliocene to mid-Pleistocene. Based on these estimates plus the estimated age of the Reka River lineage of Asellus aquaticus (see above) we, tentatively, suggest two to fve million years as the time when most of the analyzed lineages started invading the Dinaric Karst underground. Te mid-Dinaric split of Proteus and troglocaris anopthalmus does not seem to originate from the same TIME in KARST – 2007 187 PETER TRONTELJ, ŠPELA GORIČKI, SLAVKO POLAK, RUDI VEROVNIK, VALERIJA ZAKŠEK & BORIS SKET vicariant event as the latter was estimated to be younger by an order of magnitude. Another commonality of the phylogeographic pattern, the division between a western and an eastern clade in the Slovenian Dinaric Karst, might have a common hydrogeological cause in two sty-gobiotic crustaceans (A. aquaticus and t. anophthalmus) somewhere in the middle of the Pleistocene. In Proteus, however, the same split appears to be substantially older. In the Dinaric Karst we were, so far, unable to fnd reliable time estimates for paleogeographic events to calibrate local molecular clocks in diferent lineages. Con- versely, the timing of phylogenetic events can serve, inasmuch as we rely on global molecular clocks, to estimate the date of geographical, hydrographical, and geological changes (Sket 2002). Te comparative phylogeographic approach and the use of diferent independent molecular clocks have enabled us for the frst time to propose a tim-escale for the evolution of troglobionts that is relatively consistent over a wide taxonomic range. 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Verovnik, R., Sket, B., Prevorčnik, S. & Trontelj, P., 2003: Random amplifed polymorphic DNA diversity among surface and subterranean populations of Asellus aquaticus (Crustacea: Isopoda).– Genetica, 119, 155–165. Zakšek, V. , Sket, B. & Trontelj, P. , 2007: Phylogeny of the cave shrimp troglocaris: evidence of a young connection between Balkans and Caucasus.– Molecular Phylogenetics and Evolution, 42, 223–235. TIME in KARST – 2007 189 COBISS: 1.01 THE CHALLENGE OF ESTIMATING THE AGE OF SUBTERRANEAN LINEAGES: ExAMPLES FROM BRAZIL IZZIV OCENJEVANJA STAROSTI PODZEMELJSKIH ŽIVALSKIH LINIJ: PRIMERI IZ BRAZILIJE Eleonora TRAJANO1 Abstract UDC 551.44:597(81) 591.542(81) Eleonora Trajano: Te challenge of estimating the age of subterranean lineages: examples from Brazil Te applicability and efectiveness of diferent kinds of evidence used to estimate the age of lineages – morphological, molecular, phylogenetic, biogeographical, geological – are discussed. Examples from the Brazilian subterranean fauna are presented, using mainly fshes, one of the best studied groups, as a model. Only three taxa including troglobites are object of molecular studies, all in progress. Terefore, molecular clocks cannot be applied yet, and indirect evidence is used. Few phylogenies are available, e.g. for the catfsh families Heptapteridae and Tricho-mycteridae. Teoretically, basal troglobitic clades are older than apical ones, but the possible existence of extinct epigean taxa belonging to such clades hampers the comparison. As well, the limitations of the use degrees of troglomorphism to estimate phylogenetic ages are analyzed with focus on the complexity of the mechanisms underlying morphological diferentiation. Paleoclimatic reconstructions based on dating of speleothems from caves in northeastern and southeastern Brazil are available, but limited up to the last 200,000 years, thus useful for relatively recent lineages. Topographic isolation, probable for some fsh groups from Central Brazil, is also within the time range of 105 years. Older dated events (in the order of 106 years or more) that may represent vicariant events afecting aquatic lineages with subterranean derivatives are related to the establishment of the modern South American main river basins. In view of the paucity of data useful for estimating the age of Brazilian troglobitic lineages, combined evidence, including morphology, systematics and biogeography, seems to be the best approach at the moment. Key words: evolution of troglobites, degree of troglomorphism, Brazil, subterranean fshes, diferentiation rates. Izvleček UDK 551.44:597(81) 591.542(81) Eleonora Trajano: Izziv ocenjevanja starosti podzemeljskih živalskih linij: primeri iz Brazilije V prispevku je opisana uporabnost in učinkovitost različnih pristopov za ocenjevanje starosti živalskih linij s pomočjo morfologije, molekularne flogenije, biogeografje in geologije. Predstavljeni so primeri podzemeljske favne iz Brazilije, predvsem rib kot najbolj raziskane skupine. Molekularno-biološke raziskave, ki vključujejo tudi troglobionte, opravljamo na zgolj treh taksonih. Molekularne ure zaenkrat še ne moreme uporabiti, vendar zgolj posredne dokaze. Na voljo imamo le nekaj flogenetskih podatkov, npr. za morske zmaje iz družin Hep-tapteridae in Trichomycteridae. Teoretično so bazalni troglo-bitski kladi starejši od apikalnih, čeprav verjeten obstoj, sicer izumrlega epigejičnega taksona, ki pripada takim kladom, ovira primerjavo. Omejitve uporabe troglomorfzma za ocenjevanje flogenetske starosti smo analizirali s poudarkom na kompleksnosti mehanizmov, ki so osnova morfološkemu razločevanju. Razpoložljiva paleoklimatska rekonstrukcija, ki temelji na dat-iranju kapnikov iz jam severovzhodne in jugovzhodne Brazilije, je omejena na zadnjih 200.000 let in je kot taka uporabna le za relativno recentne linije. Topografska izolacija, ki verjetno velja za nekaj skupin rib iz osrednje Brazilije, spada v časovno obdobje 105 let. Starejši datirani dogodki (obdobje 106 let ali več), ki naj bi predstavljali vikariantske dogodke in ki so pomembni za vodne linije podzemeljskih sorodnikov, so povezane z razvojem današnjih glavnih južnoameriških porečij. Trenutno je, zaradi maloštevilnih podatkov, najboljša metoda za ocenjevanje starosti brazilskih troglobitskih linij kombinacija pristopov, ki vključujejo morfologijo, sistematiko in biogeografjo. Ključne besede: evolucija troglobiontov, speleobiologija, stopnja troglomorfzma, Brazilija, podzemeljske ribe, razločevalno razmerje. 1 Departamento de Zoologia, Instituto de Biociencias da Universidade de Sao Paulo, Sao Paulo, BRASIL; e-mail: etrajano@usp.br Received/Prejeto: 06.12.2006 TIME in KARST, POSTOJNA 2007, 191–198 ELEONORA TRAJANO INTRODUCTION Te problem of estimating ages for subterranean or any other lineages starts with the very defnition of age, whether the time since the isolation from the immediate sister-group (age of the cladogenetic event) or the beginning of diferentiation, either the genetic or the morphological one (see Boutin & Coineau, 2000, for a discussion about the concept of phylogenetic ages). Diferent kinds of evidence have been used to establish ages of lineages, but their applicability depends on the aspect of age considered. Molecular studies may provide ages of genetic diferentiation, independently of morphological change. Dating of potential geological isolation events, such as periods of climatic stress and large scale geological changes, may be used to infer the time in isolation. Inferences about relative times of isolation or diferentiation also come out from comparative morphological studies within a phylogenetic and biogeographic framework. Ideally, all evidence should be combined to produce coherent hypotheses about the evolution of subterranean lineages in the temporal scale. In Brazil, robust molecular studies encompassing exclusively subterranean (troglobitic) taxa started very recently and focus on a few fsh groups with very specialized troglomorphic derivatives. Basically three groups are under study with focus on populations or species: the phreatobitic characiform Stygicthys typhlops, from a karst area in eastern Brazil (studied by F. P. L. Marques & C. R. Moreira); the Amazonian catfsh genus Phreatobius, with phreatobic species collected in wells situated in alluvial plains (studied by J. Muriel Cunha); and the hep-tapterid subterranean catfsh from Chapada Diamantina, northeastern Brazil, belonging to the genus Rhamdiopsis (F.A.Bockmann, pers. comm.), previously cited as a “new genus” (studied by R. Borowsky & M. E. Bichuette). Few phylogenetic studies with biogeographic analyses of larger groups including Brazilian troglobites are available. Studies aiming to establish the ages of paleoclimatic fuctuations based on speleothem dating are also recent in Brazil, but are progressing quickly. Important climatic changes have been recorded in diferent karst areas, from the presently semiarid northeast to wet areas in the subtropical southeast. However, these studies are restricted to the late quaternary, imposing limits to its application to the problem of establishing ages for subterranean lineages because many of these lineages probably have a more ancient origin. Older geological events, such as the Miocene – Plio-Pleistocene important changes that produced the modern Amazon River system, are useful to estimate the age of some Brazilian lineages. Classically, the degree of troglomorphism, basically the reduction of eyes and pigmentation, has been used as a measure of the phylogenetic age for troglobitic animals (Poulson, 1963; wilkens, 1973, 1982; Langecker, 2000). In spite of the many restrictions to its generalized application (see below), the degree of morphological specialization may, in certain cases, provide relative ages of isolation in the subterranean environment, being a supplement to molecular and geological evidence. In the phylogenetic context, a lineage is a branch which departs from one node to another (hypothetical “ancestor”), from a node to a terminal, or an “ancestral” branch plus all the derived terminals, including extinct taxa (which remain unnoticed unless a fossil is known). Te present discussion deals lineages including terminals. It must be noted that the ever present possibility of extinction of epigean terminals in a lineage leading to a troglobitic taxon is a source of bias that may produce overestimations of its time of isolation in the subterranean environment. Among Brazilian subterranean taxa, fshes are by far the best studied group with focus on the currently discussed aspects. Tus, I took basically examples from these animals. For the sake of simplicity, I use herein the term “subterranean” as synonym of “troglobitic” (exclusively subterranean) species, to the exclusion of the equally subterranean, although not exclusively, troglophilic and trogloxenic populations. DEGREE OF TROGLOMORPHISM AND PHyLOGENETIC AGE: Te use of the degree of troglomorphism to infer relative phylogenetic ages is based on the assumption that the rates of morphological diferentiation are fairly constant among subterranean taxa, at least those regarding eyes and pigmentation, which tend to be lost along the isolation in subterranean habitats. To accept this notion, it is necessary to assume that the mechanisms of reduction are the same for each of these characters and that their reduction progress in parallel. However, there is strong evidence in contrary. Te occurrence of diferent mosaics of character states in closely related taxa suggests diferent mecha- 192 TIME in KARST – 2007 THE CHALLENGE OF ESTIMATING THE AGE OF SUBTERRANEAN LINEAGES: ExAMPLES FROM BRAZIL nisms acting at diferent rates in each population. An illustrative example is provided by the armored catfshes, Ancistrus cryptophthalmus, from Central Brazil: in the large population found in Angélica Cave, pigmentation is more reduced but eyes are less reduced than in the much smaller population from Passa Tres Caves (Reis et al., 2006). A study based on geometric morphomet-rics showed that the four known populations also difer in general body shape, with a mosaic in the deformation axes, indicating divergence probably due (at least partially) to topographic isolation (Reis et al., op. cit.). Other mosaics are also observed among related heptapterids – among the Rhamdiini, Pimelodella kronei presents eyes more reduced than Rhamdia enfurnada, the opposite being observed for melanic pigmentation. Such mosaics may encompass a larger number of characters, including behavioral and physiological ones. Tis is the case with the troglobitic amblyopsids, traditionally ranked in order of increasing degree of reduction of eyes and pigmentation as: typhlichthys subterraneus < Amblyopsis spelaea < A. rosae (Poulson, 1963). Nevertheless, A. spelaea presents more specialized life history traits and feeding behavior, while A. rosae is more derived as regards to agonistic behavior and metabolic rates (both subject to reduction); the otherwise less derived typhlichthys subterraneus is intermediate in relation to agonistic behavior and metabolic rates (Poulson, 1963; Bechler, 1983). Distinct selective pressures are likely to explain such mosaics. For this reason, attempts to rank species like these according to their degree of “adaptation” or specialization to the cave life are unconvincing. In fact, the reduction of melanic pigmentation in subterranean fshes results from diferent, independent mechanisms, which may superpose. Morphological mechanisms afect the size and number of melanocytes, whereas physiological ones afect the ability to synthesize melanin. Apparently, this ability may be lost due to diferent mutations afecting at least distinct two steps in the synthesis of eumelanin, one upstream and the other downstream the synthesis of DOPA: the frst corresponds to completely depigmented fsh which respond to the administration of L-DOPA by synthesizing melanin, referred as DOPA(+) by Trajano & Pinna (1996) and tyrosinase-positive by Jefery (2006); the second correspond to depigmented fsh which to not respond to L-DOPA (DOPA(-) albinos; Trajano & Pinna, op. cit.). Among Brazilian completely depigmented subterranean fshes, Stygichthys typhlops, the new Rhamdiopsis from Chapada Diamantina and the armored catfsh, Ancistrus formoso are DOPA(+), the heptapterid “taunayia” sp. (actually a Rhamdiopsis – F.A. Bockmann, pers. comm.) is DOPA(-) (M.A. Visconti and V. Felice, pers. comm.), and one third of the population of the trichomycterid trichomycterus itacarambiensis is DOPA(-), whereas the remaining two thirds have functional melanophores reduced in density. Te morphological mechanism is based on an additive polygenic system (wilkens, 1988), resulting in a continuous variation in the frst evolutionary steps and progressing towards complete depigmentation throughout the population at slower rates than that caused by the loss of the ability to synthesize melanin, which is based on monogenic systems (wilkens, 1988). For instance, it has been demonstrated that albinism in diferent populations of Mexican Astyanax is caused by independent mutations in the same gene, Oca2 (Protas et al., 2005). Terefore, very pale but still pigmented fsh species, with scattered micromelanophores (such as the trichomycter-us undescribed species respectively from Bodoquena and from Serra do Ramalho karst areas, and the Ituglanis spp. from Sao Domingos karst areas) may be younger than any of those DOPA(+) “albinos”. Tus, the use of troglo-morphic pigmentation as a measure of relative age should be restricted to related taxa retaining melanin (i.e., to the exclusion of DOPA albinos), where the degree of paleness is due to mutations in the additive polygenic system underlying the morphological, gradual mechanism. Regression of eyes is also due to complex genetic systems. In the blind Mexican tetra characins, genus As-tyanax, it has been shown that regression is caused by the inactivation of several genes that take part in the developmental control, and that growth factors acting at a lower level of this control appear to be involved in the degeneration of the eyes (Langecker, 2000). Clearly, studies on a much large sample of troglobitic species are needed before any inference about diferentiation rates can be made. Two other factors infuence the rates of divergence: population sizes and life cycle strategies. Small populations tend to diferentiate faster due to phenomena as genetic drif. Population sizes are highly infuenced by ecological factors such as nutrient availability and the extent of habitats suitable for colonization. It is noteworthy that energy is higher in streams (higher carrying capacity), but phreatic habitats occupy larger areas and volumes. Because there is no taxonomic correlation with these factors, related species may difer in population sizes (for instance, populations respectively with 20,000 and 1,000 individuals were estimated for A. cryptophthalmus in An-gélica and in Passa Tres caves – Trajano, 2001a), thus in divergence rates. As well, nutrient availability may also be “perceived” diferently even by taxonomically related species, depending on the efciency of energy use. Such efciency may be improved along the adaptation to the subterranean life, allowing for increase in population sizes, then in lowered diferentiation rates. TIME in KARST – 2007 193 ELEONORA TRAJANO K-selected life strategies imply lower diferentia-tion rates due to delayed ages for frst maturation and low reproductive rates (few individuals reproducing at given times), which work on opposite directions: delayed frst maturation implies slow divergence rates (longer reproductive generations), whereas low reproductive rates result in lowered efective populations, which would accelerate divergence rates.. Dating of paleoclimatic events based on growth phases of speleothems and similar deposits may be applied to subterranean lineages within the framework of the paleocli-matic model (Barr, 1968; wilkens et al., 2000). However, its cyclical nature imposes serious limitations because, without biological data (molecular, morphological, phy-logenetic), it is not possible to establish in which phase the isolation frst took place. As a matter of fact, isolation with diferentiation may occur along several subsequent unfavorable phases intercalated with coalition phases, thus what really counts to produce genetic and/or morphological divergence is the sum of isolation periods (Trajano, 1995), and not simply the time since the frst isolation event. For instance, in northeastern Brazil there were nine dry phases (no speleothem growth) in the last 210,000 years, intercalated with short wet phases lasting from several hundreds to a few thousand years each. Overall, these periods of speleothem growth occupied only 8% of the studied period, i.e., around 20,000 years in contrast with 190,000 years with dry conditions, like the one prevalent nowadays in the region (wang et al., 2004). Hence, at least in the late Pleistocene, there was a much extended period of isolation for the hypogean fauna in northeastern Brazil – for lineages already established in subterranean habitats, from 190,000 to 210,000 years, depending on the occurrence or not of introgression with epigean relatives during the wet periods. As a matter of fact, several of the most highly specialized Brazilian tro-globites have been found in this region (e.g., Rhamdiopsis catfshes, Spelaeogammarus amphipods, Pongicarcinia xi-phidophorum isopods, Coarazuphium beetles), as well as the only Brazilian troglobitic scorpions, cockroaches and Ctenid spiders. On the other hand, climatic changes were not as dominant in the subtropical southeast Brazil and dry phases were shorter, at least for the last 116,200 years (Cruz-Jr. et al., 2005). Terefore, total time of isolation in subterranean habitats during the late Pleistocene was shorter in SE than in NE Brazil. Hypothetically, a pop- In conclusion, there is a complex balance between diferent genetic, ecological and biological factors, which may act in diferent directions to produce the actual divergence rates. Such rates may difer among related taxa, and even among diferent characters. Terefore, the degree of troglomorphism as a measure of age of subterranean lineages should be used with extreme caution. ulation that became frst isolated at a given time in the northeast would be much more diferentiated, both genetically and morphologically, than another population frst isolated at the same time in the southeast. If one considers “age” as the time of the frst isolation, these two lineages have the same age; if “age” is the total time in isolation, then the frst one is older. It is clear that, in a cyclical model, the degree of genetic diferentiation do not provide a good evidence of age without a precise determination of the duration of each phase. Geological and geographical events over larger temporal scales may provide more robust evidence. Te genus Phreatobius is distributed around the Amazon basin, in tributary basins from both margins of the Amazon River. Te frst described species, P. cisternarum, lives underground in the alluvial fan around the Amazon delta, being collected in shallow hand-dug wells. Much latter, in the 1990´s, other species were found deeply buried in submerged litter banks in shallow “igarapés” (small tributaries) along the lef margin of the Negro and Amazon rivers. More recently, a second phreatobic species was discovered in wells in the State of Rondônia, Rio Madeira basin, in the right margin of the Amazon drainage (J. Muriel-Cunha & J. Zuanon, pers. comm..; description in progress by J. Muriel-Cunha & M. de Pinna). Tis wide, peripheral distribution of the Phreatobius genus around the Amazon basin may be explained by an origin between the late Miocene and the late Pliocene (~2.5 Ma), when a gigantic lake, or a series of interconnected mega-lakes occasionally united to cover most or all of lowland Amazonia to a shallow depth (Campbell et al., 2006). In fact, Phreatobius cat-fshes are adapted to shallow, hypoxic conditions, with dark pink to red skin indicating cutaneous breathing; since all known species exhibit this conspicuous trait, this is probably an ancestral condition for the genus. I suggest that the fragmentation of the lacustrine habitat during the late Pliocene, leading to the establishment of the modern Amazon River drainage system, may have been an isolation event for the ancestors of the extant GEOLOGICAL, PALEOCLIMATIC AND BIOGEOGRAPHICAL EVIDENCE: 194 TIME in KARST – 2007 THE CHALLENGE OF ESTIMATING THE AGE OF SUBTERRANEAN LINEAGES: ExAMPLES FROM BRAZIL species. Nevertheless, an older origin for cannot be ruled out. On the other hand, P. cisternarum has been found not only north and south of the Amazon River mouth but also in the large Marajó Island in between, with no unequivocal morphological diferentiation so far detected between these localities (Muriel-Cunha & Pinna, 2005). Tese populations were isolated during the formation of the Amazon delta, ~2.5 Ma ago, suggesting a high evolutionary stability, at least at the morphological level, possibly due to the environmental stability of the subterranean habitat. Te disjunct distribution also points to a very ancient origin for the Calabozoidea isopods. So far, this taxon is composed exclusively by three extant phreatobic species, one from the Orinoco basin, in Venezuela (Calabozoa pellucida), e two from Brazil, respectively from the Sao Francisco (Pongycarcinia xiphidiourus) and the Paraguay (undescribed species) river basins. Te only connection between these regions is through the Amazon basin, and I speculate that the ancestors may have dispersed during In order to be minimally reliable and useful, molecular clocks must be based on well corroborated phylogenies with at least one node correlated to geographic or geological isolation events of known age. In cyclical models, such correlation is hampered when cycles are relatively short and repetitive, as is the case with the paleoclimatic fuctuations in the late Pleistocene in Brazil, adding a great deal of uncertainty to the molecular clock. Marine transgressions, which have been used to establish dates for vicariant events in epigean Brazilian taxa such as freshwater fshes, are of no use for subterranean lineages because almost all karst areas in Brazil are above the maximum sea levels. In any case, the conclusion of the molecular studies on Phreatobius spp., S. typhlops and Rhamdiopsis sp. from Chapada Diamantina will certainly open new interesting avenues in this feld. As already mentioned, few phylogenetic studies of groups including Brazilian troglobites are available, most at the genus level and incomplete in terms of taxa encompassed. Among fshes, the heptapterid catfshes were object of a phylogenetic study, but the cave species were not included (Bockmann, 1998). Phylogenetic and molecular studies on heptapterids are in progress, but the position of the Phreatobius genus and of the troglobitic Rhamdi-opsis species within this genus are still unclear. Recently analyzed morphological data indicate that, within the genus Rhamdiopsis, “taunayia” sp. is basal whereas the species from Chapada Diamantina have a more apical posi- or prior to the formation of the huge Lago Amazonas. Actually, the Sao Francisco lineage would be older, at least 5 Ma, which is the estimated age of separation of this basin based on studies of the biogeographical patterns in Brazilian freshwater fshes (Hubert & Renno, 2006). Messana et al., (2002) argue for a close relationship between the Calabozoidea and the Oniscoidea isopods, thus both lineages have the same phylogenetic age, which goes back to the Jurassic-Cretaceous (gondwanic origin – L. A. Souza, pers. comm.). A phylogenetic study, that could add more light to this interesting problem, is waiting for the collection of additional specimens, what is proving to be very difcult in spite of the eforts of biologists and cave divers. Apparently these animals are very rare and/ or live mainly in inaccessible, deep phreatic habitats. Geomorphological events as alluvial erosion producing waterfalls that split populations (topographical isolation), once dated, also provide data useful to estimate the age of lineages such as the diferent populations of the armored catfsh, A. cryptophthalmus. tion in the phylogeny (F. A. Bockmann, pers. comm.). Tese two species independently adapted to the same kind of habitat, the upper phreatic zone connected to the surface through caves (Trajano, 2001b), having developed advanced characters states related to the hypogean life, including miniaturization. “taunayia” sp., however, is even more specialized, presenting a hypertrophied lateral line system in the head, with behavioral evidence of enhanced mechano-sensory sensitivity. Tis, associated with its putative basal position in the Rhamdiopsis phy-logeny, points to an older age for the lineage to which the troglobitic “taunyaia” sp. belongs, much anterior to the late Pleistocene. Te phylogeny of the catfsh family Trichomycteri-dae was also studied (wosiack, 2002), but only one among 10+ troglobitic species presently known, trichomycterus itacarambiensis, was included. It is an apical taxa in the phylogeny, indicating a relatively recent origin. A recent derivation of t. itacarambiensis from an epigean ancestor from the Upper Sao Francisco River basin is consistent with the morphological variation observed in eyes and pigmentation and also with the notion of a quick fxation of genes for albinism, since one third of the population is made of albinos. However, in the absence of a correlation between some node and dated geographic or geological isolation events, it is not possible to estimate an absolute age, even approximate, for this cave lineage. PHyLOGENETIC AND MOLECULAR EVIDENCE: TIME in KARST – 2007 195 ELEONORA TRAJANO COMBINED EVIDENCE: For extremes in the inter-taxa variation, the troglomor-phism degree may provide good evidence of relative ages. For instance, it is reasonable to suppose that fshes with slightly reduced eyes and pigmentation such as the heptapterids Rhamdiopsis sp. from Cordisburgo (eastern Brazil) and Pimelodella spelaea, from Sao Domingos (Central Brazil), are younger than the highly troglomor-phic Rhamdiopsis sp. from Chapada Diamantina and “taunaya” sp., from Campo Formoso. Te two former species probably have been isolated topographically because they inhabit streams several meters above the base level, and an isolation period in the order of 105 years (estimated time for the erosional processes lead to the current landscape – A. Auler, pers. comm..) may be estimated. Te two latter species inhabit presently semiarid karst areas in northeastern Brazil subject to extended periods of isolation at least during the last 210,000 years, but they probably became isolated well before. Tus, an estimate in the order of 105-106 years seems reasonable. A molecular study focusing on the hypervariable Region I of MtDNA did not fnd any evidence of divergence between the cave populations of Ancistrus cryptophthalmus (Moller & Parzefall, 2001). However, geometric morphometric analyses showed a clear, statistically signifcant diference between these populations, but with some superposition with the epigean closest relatives (Reis et al., 2006). Taken together, these data indicate a recent isolation of the cave populations from the epigean ones and also from each other, in the order of 104 -105 years. Preliminary molecular studies on Ituglanis species from Sao Domingos karst area are consistent with the observed morphological diferences (Bichuette et al., 2001) justifying the recognition of four species, each one in a separate microbasin that runs parallel westwards (Bichuette & Trajano, 2004). Tese catfshes are sym-patric with the morphologically less specialized A. cryp- tophthalmus, P. spelaea and Eigenmannia vicentespelaea (Gymnotiformes), making Sao Domingos karst area a world hotspot of biodiversity for subterranean fshes. All the Ituglanis catfshes have eyes more reduced and are paler than the other species, presenting scattered mela-nophores, i.e., they are not DOPA albinos. Tree among these Ituglanis species occupy a very specialized habitat, with adaptations to the phreatic environment that include miniaturization. Moreover, I. epikarsticus, and probably also I. bambui and I. ramiroi (Trajano & Bichuette, unpubl. data), live and disperse through the epikarst, whereas the other species are typical stream-dwellers, like their epigean relatives. In spite of intensive collecting eforts, no epigean Ituglanis catfsh was found in Sao Domingos (the same is true for Pimelodella; Bichuette & Trajano, 2003). Taken together, these evidences indicate a longer time in isolation for the Ituglanis catfshes. In conclusion, the rich troglobitic ichthyofauna from Sao Domingos seems to be the result of anachronous isolation events, including both the extinction of epigean relatives due to unknown factors (for Ituglanis and Pimelodella) and topographic isolation (for Ituglanis spp. and also A. cryptophthalmus). Anachronous isolation, possibly in association with diferent divergence rates, may also explain the disparity in troglomorphic degree observed for the subterranean fauna from the Upper Ribeira Valley karst area, SE Brazil. Tis fauna includes very specialized species, such as the pseudoscorpion Spelaeobochica muchmorei and the decapod Aegla microphthalma, to moderately troglomorphic species, such as the opilionid Pachylospeleus strinatii, the carabid beetle Schizogenius ocellatus and the catfsh Pimelodella kronei. within the framework of the paleo-climatic model, in view of the short isolation periods (= dry phases) during the late Pleistocene (see above) it is probable that all these species became frst isolated in caves before this period. ACKNOwLEDGEMENTS I am grateful to Augusto Auler, Fernando P. L. Marques and Janice Muriel Cunha for the discussion of ideas and criticisms, and to Richard Borowsky for the critical reading and revision of the English style of an early version of the manuscript. Many data were gathered during stud- ies sponsored by the Fundaçao de Amparo a Pesquisa do Estado de Sao Paulo – FAPESP (grant n. 03/00794-5, among others). 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Tabin., 2005: Genetic analysis of cavefsh reveals molecular convergence in the evolution of albinism. Nature Genetics, Advance Online Publication, Letters, p. 1-5 [published online 11 December 2005] Reis, R. E., Trajano, E. & E. Hingst-Zaher., 2006: Shape variation in surface and cave populations of the armoured catfsh Ancistrus (Siluriformes: Lori-cariidae) from the Sao Domingos karst area, Upper Tocantins River, Brazil. Journal of Fish Biology, 68, 414-429, London. Trajano, E., 1995: Evolution of tropical troglobites: Applicability of the model of quaternary climatic fuc-tuations. Mémoires de Biospéologie, 22, 203-209, Moulis. Trajano, E., 2001: Habitat and population data of tro-globitic armoured cave catfshes, Ancistrus cryp-tophthalmus Reis 1987, from Central Brazil (Silu-riformes: Loricariidae). Environmental Biology of Fishes, 62, 1-3, 195-200, Dordrecht. Trajano, E., 2001: Ecology of subterranean fshes: an overview. Environmental Biology of Fishes, 62, 1-3, 133-160, Dordrecht. TIME in KARST – 2007 197 ELEONORA TRAJANO Trajano, E. & M.C.C. Pinna., 1996: A new cave species of trichomycterus from eastern Brazil (Siluriformes, Trichomycteridae). Revue française d’Aquariologie, 23, 3-4, 85-90, Nancy. wang, x., Auler, A. S., Edwards, R. L., Cheng, H., Cris-talli, P. S., Smart, P. L., Richards, D. A. & C.-C. Shen., 2004: wet periods in northeastern Brazil over the past 210 kyr linked to distant climate anomalies. Nature, 432, 740-743, London. wilkens, H., 1973: Ancienneté phylogénetique et degrees de reduction chez les animaux cavernicoles. Annales de Spéléologie, 28, 2, 327-330, Paris. wilkens, H., 1982: Regressive evolution and phylogenetic age: the history of colonization of freshwaters of yucatan by fsh and Crustacea. Texas Memorial Museum Bulletin, 28, 237-243. wilkens, H., 1988: Evolution and genetics of epigean and cave Astyanax fasciatus (Characidae, Pisces). Evolutionary Biology, 23, 271-367, New york. 198 TIME in KARST – 2007 COBISS: 1.01 PATTERN AND PROCESS: EVOLUTION OF TROGLOMORPHy IN THE CAVE-PLANTHOPPERS OF AUSTRALIA AND HAwAI’I – PRELIMINARy OBSERVATIONS (INSECTA: HEMIPTERA: FULGOROMORPHA: CIxIIDAE) VZOREC IN PROCES: EVOLUCIJA TROGLOMORFNOSTI PRI JAMSKIH MREŽEKRILNIH ŠKRŽATKIH IZ AVSTRALIJE IN HAVAJEV – PRELIMINARNE UGOTOVITVE (INSECTA: HEMIPTERA: FULGOROMORPHA: CIxIIDAE) Andreas wESSEL1, Petra ERBE1,2 & Hannelore HOCH1 Abstract UDC 591.542(94+739,9) Andreas Wessel, Petra Erbe & Hannelore Hoch: Pattern and process: Evolution of troglomorphy in the cave-planthoppers of Australia and Hawai’i ? Preliminary observations (Insecta: Hemiptera: Fulgoromorpha: Cixiidae) Te evolution of troglobites comprises three distinct problems: cave colonization by an epigean ancestor, the evolution of tro-glomorphies, and intra-cave speciation. Te study of cave-dwelling planthoppers has contributed much to our understanding of troglobite evolution and provides useful model systems to test various aspects of the theoretic framework developed in recent years. Most promising in this respect are taxa with several closely related but independently evolved troglobiontic lineages, such as on the Canary Islands, in queensland/Australia and on the Hawaiian Archipelago. Closely related species ofen occur in caves with comparable ecological parameters yet difer in their age. Here we use comparative age estimates for Australian and Hawaiian cave cixiids to assess the dynamics of reductive evolutionary trends (evolution of troglomorphy) in these taxa and cave planthoppers in general. we show that the degree of troglomorphy is not correlated with the age of cave lineages. Morphological alteration may not be used to draw conclusions about the phylogenetic age of cave organisms, and hypotheses based on such assumptions should be tested in light of these fndings. Key words: adaptive shif, cave adaptation, climatic relict, founder efect, reductive evolutionary trends, troglobites, tro-glomorphies. Izvleček UDK 591.542(94+739,9) Andreas Wessel, Petra Erbe & Hannelore Hoch: Vzorec in proces: Evolucija troglomorfnosti pri jamskih mrežekrilnih škržatkih iz Avstralije in Havajev Preliminarne ugotovitve (Insecta: Hemiptera: Fulgoromorpha: Cixiidae) Evolucija troglobiontov zajema tri značilne korake: kolonizacija jame s površinskim prednikom, razvoj troglomorfnosti ter podzemeljska speciacija. Študija podzemeljskih mrežekrilnih škržatkov je prispevala veliko k našemu razumevanju evolucije troglobiontov in hkrati predstavlja uporaben modelni sistem za testiranje različnih teoretičnih pristopov, ki so bili razviti v zadnjih letih. V tem pogledu so najobetavnejši tisti taksoni, ki so si sicer sorodni, toda pripadajo evolucijsko neodvisnim troglobiontskimi linijami, kot so npr. tisti na Kanarskih otokih, v državi queensland (Avstralija) in na havajskem arhipelagu. Bližje sorodne vrste se v jamah pogosto pojavijo v primerljivih ekoloških pogojih, vendar se razlikujejo v starosti. Za ugotavljanje dinamike trendov redukcijske evolucije (evolucija tro-glomorfzmov) teh taksonov in jamskih škržatkov na splošno, smo v prispevku uporabili ocene primerjalnih starosti za avstralske in havajske jamske škržatke. Ugotavljamo, da stopnja troglomorfnosti ni v korelaciji s starostjo jamskih linij. Zgolj morfološke spremembe pri organizmih se ne bi smele uporabljati za prikazovanje flogenetske starosti jamskih organizmov. Hipoteze, ki temeljijo na takšnih predpostavkah, bi morale biti preverejene v luči pričujočih ugotovitev. Ključne besede: prilagoditveni premik, prilagoditve na podzemlje, klimatski relikt, učinek osnovatelja, redukcijski evolucijski trendi, troglobiti, troglomorfzmi. 1 Museum für Naturkunde der Humboldt-Universität zu Berlin, Biosystematics Research Group, Invalidenstrasse 43, D-10115 Berlin, Germany; e-mail: andreas.wessel@museum.hu-berlin.de 2 Chiang Mai University, Te Uplands Program, Faculty of Agriculture, Chiang Mai, Tailand Received/Prejeto: 30.01.2007 TIME in KARST, POSTOJNA 2007, 199–206 ANDREAS wESSEL, PETRA ERBE & HANNELORE HOCH INTRODUCTION Te origin of troglobites has fascinated evolutionary biologists since Darwin remarked on their curious and strong modifcation (1859: 177-178). He did not provide a ‘Darwinistic’ explanation for their evolution1, however, this was subsequently supplied by August weismann (1886)2. Troglobite evolution, i.e. the process leading to diferent, sometimes closely related species, which are highly adapted to life in subterranean spaces, comprises three somewhat independent phenomena and problems: (i) initial cladogenesis of a cave species, or the origin of a cave-dwelling bio-species from an epigean ancestor, which is basically the problem of isolation or rather (spatial) separation of a cave population from its epigean relatives; (ii) subsequent anagenetic transformation, which comprises the dynamics and driving forces of cave adaptation, the ofen so-called regressive evolution or reductive evolutionary trend, and, in some cases, (iii) subterranean (intra-cave) radiation. RELICTS OR ExPLORERS? A widely accepted concept aiming to explain the specia-tion event giving rise to a cave-dwelling and reproduc-tively isolated bio-species, was developed by Tomas Barr in the 1960s, commonly known as the Climatic Relict Hypothesis (CRH): “A s it is difcult to imagine that eyes, though useless, could be in any way injurious to animals living in darkness, I attribute their loss wholly to disuse.” (Darwin 1859: 177) „As soon as such a cave immigrant has developed the ability to obtain food without the help of eyes a reduction of the eyes must commence, since as soon as the same are no longer neces-sary for the animals’ existence, they are not infuenced anymore by natural selection, because now it does not matter whether the eyes are a little worse or a little better. Now, no more selec-tion will take place between individuals with better and those with worse eyes, but both will have an equal chance to be preserved and reproduce. Individuals with better and those with worse eyes will cross from now on, and the result can only be a general degradation of the eyes. Pos-sibly this is helped by the circumstance that smaller and stunted eyes can even present an ad-vantage, since this allows other organs such as sensory and olfactory organs, which are more important for the animal now, to develop more strongly. Even without such efect, though, the lack of natural selection maintaining the eye’s high level of organization will necessarily lead to its degradation, slowly or even very slowly, especially at the beginning of this process, but in-exorably.” (Translated from the German; weismann 1886: 16-17) “Troglobites have evolved from colonies of troglo-philes which became isolated in caves through extinction of surface populations of the troglophiles” (Barr 1968: 96). According to Barr, the evolution of troglobites is a two-step process: at frst, it involves a preliminary, tro-glophilic stage without apparent troglomorphies or a disruption of genefow between cave-dwelling and epigean populations. Following this initial cave colonization, the cave-dwellers become geographically separated, and thus genetically isolated, due to the extinction of parental epi-gean populations (supposedly caused by climatic change), at least in the region of the cave. Over time, reproductive isolation will inevitably follow as a side efect of genetic change by drif and natural selection (Barr 1968). Support and evidence for this concept was gained from the observed relict distribution of most troglobites known at that time, which were almost exclusively confned to temperate regions. Glaciation during the ice ages was suggested as the most important factor for the change of surface conditions (Barr 1968, Sbordoni 1982, Barr & Holsinger 1985). Tis hypothesis remained without alternatives until the early 1970s, when Francis G. Howarth discovered the Hawaiian cave ecosystems (Howarth 1972). Te lava tubes host, among other taxa, highly troglomorphic plan-thoppers that are parapatrically distributed with respect to their close epigean relatives, which are still extant, i.e. they are non-relictual troglobites. Consequently, How-arth (1981, 1986, 1987) formulated the Adaptive Shif Hypothesis (ASH): “[...] potential food resource provides the driving force for the [...] evolution of cave species. Troglomor-phic populations [...] evolve from pre-adapted habitual accidentals which [...] establish temporary populations in marginal underground habitats. Once an adaptive shif occurs, allowing a reproducing population to establish itself underground, then it is both the efects of strong new selection pressures and the release from previously strong selection pressures that bring about [...] troglo-morphy” (Howarth 1986: 155). while the exploitation of a large new habitat with new food resources may be the driving force in the evolution of troglobites according to the ASH, a major challenge for survival underground is probably the ability to locate mates and reproduce in the dark. A change in mating behaviour might thus have been the most important adaptive shif necessary for a successful colonization of caves, and would almost inevitably lead to reproductive isolation of the incipient cavernicolous species. Te Hawaiian cave planthoppers provide a striking example for 200 TIME in KARST – 2007 PATTERN AND PROCESS: EVOLUTION OF TROGLOMORPHy IN THE CAVE-PLANTHOPPERS OF AUSTRALIA AND HAwAI’I this process, and consequently played a pivotal role in the formation of the Adaptive Shif Hypothesis (Howarth 1986, Howarth & Hoch 2005). A principal acceptance of the ASH does not necessarily invalidate the CRH, especially not in cases where the preconditions for the CRH are met, i.e. cave taxa displaying a relict distribution. However, a relict distribution observed today is not sufcient evidence to unconditionally accept the CRH, given the alternative present in the ASH 3. Te predictions arising from both hypotheses must be tested for every single system. For the CRH, we expect the closest epigean relatives at least to be allopatrically distributed compared to the cave species, while the ASH predicts a parapatric distribution of cave and epigean species, which are necessarily sistergroups (adelphotaxa). Conclusive evidence for a decision between both hypotheses may be gained from a well-founded phylogeny in conjunction with a sound knowledge of the geographic distribution of both cave and epigean taxa. Te last requirement is ofen problematic, though, as the sampling of epigean relatives for some cave species is frequently insufcient. For some groups no epigean relatives are known at all, and it is only through intensive, directed search eforts that this obstacle may be overcome (see e.g. Stone 2004). REDUCTIVE EVOLUTIONARy TRENDS Once a population has shifed towards a permanently cav-ernicolous mode of living, the second problem of troglo-bite evolution – subsequent anagenetic transformation – arises. A basic assumption since weismann (1886) has been a correlation between the degree of troglomorphy of a taxon and its residence time in caves. Cave adaptation is accordingly described as an orthogenetic, time-dependent process, which is an overall slow, gradual adaptation towards a stage of ‘absolute troglomorphy’; see e.g. wilkens (1986), for review see Barr (1968) and Howarth “Te evidence suggests that troglobites evolve from pre-adapted habitual visitors or accidentals in the cave rather than from well-adapted troglophiles. Te former group requires an adaptive shif in order to fully exploit the cave resources. Tis adaptive shif may lead to the evolution of a troglobitic lifestyle. well-adapted troglophiles on the other hand tend to remain opportunistic exploiters of the cave environment. Some temperate troglobites may ft the scenario of isolation by changing climates (Barr, 1968). However, many species including those in the tropics probably do not. I postulate that adaptive shifs led to the colonization of caves and evolution of troglobites, including most of those in temperate caves, but that the complex geological history of the continents including glaciations has obscured the early history and obfuscated the earlier distribution and the evolution of troglobites there.” (Howarth 1981: 540) (1987). Traditional explanations for the mechanisms of this process includes (i) the accumulation of neutral mutations, (ii) pleiotropic efects, and (iii) natural selection for energy economy (Sket 1986, Culver 1982). Both the CRH (Barr 1968) and ASH (Howarth 1986), however, contain some notion of a founder efect: Barr with an explicit quotation of Mayr’s genetic revolution (Mayr 1954) and Howarth with reference to the Carson model of founder efects (Carson 1968, 1975). Te process of cave adaptation is infuenced by several parameters – such as availability of food, population density, microclimate of the caves and other biotic and abiotic factors of the cave ecosystem – , which make comparisons even between closely-related species exceedingly difcult, and generalisations even more so. An excellent opportunity to test the assumption of gradual and increased troglomorphy over time may nevertheless be found in radiations of cave-dwelling planthoppers inhabiting caves of diferent age. CAVE-ADAPTATION IN PLANTHOPPERS Studies during the last three decades have revealed numerous cases of evolution of cave-adapted planthoppers in tropical and subtropical caves. Among the Fulgoro-morpha, 53 cave-dwelling species have been described from many parts of the world, four-ffh of them cixiids including the Australian taxa Solonaima and Undarana and the Hawai’ian Oliarus species (Hoch 1994, Hoch & wessel 2006). Te adaptation to similar environments in cave planthoppers has led to the evolution of a very similar external morphology in diferent parts of the world and represents a striking example of parallel evolution. Te morphological modifcations of cave planthoppers are characterized by reductive evolutionary trends, as in most obligately cavernicolous animals. Te degree of adaptation to a subterranean life varies greatly, primarily depending on their habitat in the cave or soil (Fig. 1). Most conspicuous are the reduction and loss of compound eyes and ocelli, tegmina, wings and bodily pigment. It has also been suggested that apparently non-troglomor-phic characters have an increased adaptive value in the underground environment, such as e.g. the specialized spine confgurations of hind tibiae and tarsi, which may possibly enhance walking on wet or rocky surfaces (Hoch & Howarth 1989a, 1989b, Hoch 2002). Te closest epigean relatives of cavernicolous Fulgo-romorpha species all have immature stages living close to the soil, e.g. under the dead bark of rotting logs, in leaf litter or moss, or even within the soil, feeding on roots or perhaps on fungi (Remane & Hoch 1988). Tis mode of life has been considered an ecological pre-adaptation TIME in KARST – 2007 201 ANDREAS wESSEL, PETRA ERBE & HANNELORE HOCH Habitat Abiotic factors Life cycle Morphology epigeic (surface) habitats unstable physical parameters entirely epigeic / epigean no troglomorphies (including leaf litter) (ambient climatic influence high) troglophilic (facultative soil & cave-dwellers, may live & reproduce underground as well as in surface domain) troglomorphies hypogeic (subterranean) habitats T stable / constant physical parameters (ambient climatic influence low) of varying degrees e.g. .''*¦ + +¦+¦ , ;'' soil- *\ . [ interstitial [ • chambers ) ; meso-cavernous rock stratum \ endogeic / endogean (obligatory soil dwellers) - reduction of eyes, body pigmentation & wings - decreasing cuticle sclerotization I caves '• (deep cave zone) troglobitic (obligatory cavernicoles, restricted to cave environment) - specialized sensory organs - elongate appendages Fig. 1: terminology of interdependence between physical parameters of the habitat and organismic adaptations (From hoch et al. 2006). to a later switch to a permanent (adult) life underground (Hoch 2002, Howarth & Hoch 2005). CAVE-PLANTHOPPERS OF AUSTRALIA AND HAwAI’I In Australia, closely related Solonaima and Undarana species have colonized old karst caves as well as younger lava tube systems. Te four epigean Undarana species occur in the (rain)forest at the south of queensland’s east coast, while the two cave-dwelling species (U. rosella, Bayliss & Pinwills cave, Undara lava tube; U. collina, Collins cave) inhabit the lava caves of the McBride Formation in the dry grasslands westward of the Great Dividing Range (Hoch & Howarth 1989a). Te epigean Solonaima species can be found all along the east coast (rain)forest, while the cave species inhabit lava tubes within the Mc-Bride Formation, too (S. baylissa, sympatric with U. rosella), as well as karst caves of the Chillagoe Karst Towers (S. pholetor, S. stonei, S. halos, S. irvini) and Mount Mul-grave (S. sullivani) (Hoch & Howarth 1989b). Tus, epi-gean and cavernicolous species of both Australian genera show an allopatric distribution. On the Hawaiian islands the cave-dwelling species of the endemic, monophyletic Oliarus clade represent independent cave colonizations on islands of diferent age. with about 80 described epigean taxa (species and subspecies), Oliarus is the most speciose planthopper genus on the Hawaiian islands (Zimmerman 1948, Asche 1997). Based on morphological data, this diversity has been hypothesized to stem from a single colonization event (Asche 1997, Hoch & Howarth 1993). Te frst cave-dwelling species of the genus, Oliarus polyphemus Fennah, 1973 and Oliarus priola Fennah, 1973 (Fennah 1973) were discovered by Howarth (1972) on Hawai’i Island and Maui where they are endemic. Later, fve more troglobitic taxa were discovered on the archipelago (Hoch & Howarth 1999). Te seven cave-dwelling species owe their origin to several independent colonization events on three islands; on Molokai, one adaptive shif (O. kalaupapae); on Maui, three adaptive shifs (O. priola, O. gagnei, O. waikau); on Hawai’i Island, at least three adaptive shifs (O. polyphemus, O. lorettae, O. makaiki) (Hoch & Howarth 1999). Te closely related epigean species of all cavernicolous Oliarus taxa occur parapatrically at the surface. Both the Australian and the Hawaiian cave species complexes exhibit diferent degrees of troglomorphy. Figure 2 shows the heads of six Australian Solonaima, one epigean (1), three facultative cavernicolous (2-4), and two obligate cavernicolous species (5,6). Figure 3 depicts the habitus of six Hawai’ian Oliarus species, one epigean relative on the lef (note the diferent scale), and fve troglobitic species. Te varying degree of eye reduction is clearly visible; two of the Hawaiian species even show a complete loss of eyes. Te same pattern is seen in wing reduction. Te time factor is crucial for assessing the dynamics of troglobite evolution. Unfortunately, though, it is 202 TIME in KARST – 2007 PATTERN AND PROCESS: EVOLUTION OF TROGLOMORPHy IN THE CAVE-PLANTHOPPERS OF AUSTRALIA AND HAwAI’I Fig. 2: Australian Solonaima species, heads, dorsal view. 1, S. solonaima (epigean); 2. S. sullivani; 3, S. pholetor; 4, S. stonei; 5, S. irvini; 6, S. baylissa. (From hoch & howarth 1989b). rarely possible to obtain direct estimates of the age of the cave lineages. Rather, the maximum age of the habitat is usually employed - at least if an active colonization of caves sensu Howarth is assumed - , or even just the maximum age of the underlying geological structure. By these measures, the maximum age for the troglobitic Oliarus lineages on Hawai’i is the age of the islands: 1.8 myr for Molokai, 1.3 myr for Maui, and less than 400,000 y for Hawai’i Island. In the case of the Australian troglobitic cave plan-thoppers, the situation is even more complex. At frst sight their distribution fts the Climatic Relict Model sensu Barr very well assuming a late Miocene desertifca-tion, i.e. replacement of the rain forest by dry savannah or grassland east of the Great Dividing Range (see Kemp 1978, Truswell 1990). while not per se refuting the relict hypothesis, we do not exclude the possibility of adaptive shifs for the Australian cave planthoppers as well. In that case, Australian cave taxa may be much older than hitherto assumed. Also, the late Miocene climatic change is not necessarily be regarded as the sole reference point for the calculation of the maximum age of the Australian cave planthoppers. what could matter instead is the availability of the caves as a suitable novel habitat. Te limestone of the Chillagoe Tower Karst and Mitchell-Palmer Karst are presumably of Silurian origin, and the current main caves were formed by phreatic solution during the last 5-10 million years (Ford 1978, Jennings 1982, Pearson 1982). Remnant older passages and solution breccias near the tops of many towers indicate the existence of caves since the area was uplifed and the limestone was exposed in the mid-Tertiary about 20-25 million years ago (Howarth 1988). Te much younger Undara lava fow (190,000 years old) covers portions of older fows within the McBride Formation (Atkinson et al., 1976), some of which may date back from the Pliocene, i.e. more than 2.5 million years ago (Best 1983). Te cave animals could have migrated through the mesocavernous systeme into young basalt and colonized new caves in each fow in succession. “Te troglobitic species could be, and probably are, older than the age of their caves” (Howarth 1988). PHyLOGENETIC AGE AND TROGLOMORPHy Against this background we here attempt to assess the problem of the dynamics of reductive evolutionary Fig. 3: hawai’ian Oliarus species, habitus, dorsal view. 1, epigean Oliarus species (O. tamehameha); 2, O. kalaupapae; 3, O. lorettae; 4, O. gagnei; 5, O. waikau; 6. O. polyphemus. (1, from zimmermann 1948; 2-5, from hoch & howarth 1999; 6, hoch, Original). TIME in KARST – 2007 203 ANDREAS wESSEL, PETRA ERBE & HANNELORE HOCH Fig. 4: variability of relative eye width and relative tegmina length in epigean and cave-dwelling species (O. polyphemus, O. lorettae, O. makaiki (no wing measurements), O. gagnei, O. waikau, O. kalaupapae; S. pholetor (stonei), S. irvini (halos), S. baylissa; U. collina, U. rosella). Fig. 5: Correlation of indices for troglomorphic characters with the maximum age of inhabited cave formations (O. polyphemus, O. lorettae, O. gagnei, O. waikau, O. kalaupapae; S. pholetor (stonei), S. irvini (halos), S. baylissa; U. collina, U. trends (troglomorphies) or regressive evolution. A ma- approach should aim at analyzing character evolution jor obstacle in this context is the poor comparability of in monophyletic groups where similar (morphological) characters across diferent taxonomic groups. A strict pre-conditions or pre-adaptations for parallel evolution 204 TIME in KARST – 2007 PATTERN AND PROCESS: EVOLUTION OF TROGLOMORPHy IN THE CAVE-PLANTHOPPERS OF AUSTRALIA AND HAwAI’I may safely be assumed. while we are aware of these problems, we nevertheless found it useful to employ a quanti-fcation of troglomorphic characters in order to achieve at least a preliminary idea of the possible correlation between troglomorphy and lineage age. we computed two ‘troglomorphy indices’ for all Hawaiian and Australian taxa from which data were available by using two characteristic troglomorphic characters in cavernicolous plan-thoppers: the reduction of eyes and the reduction of the tegmina. Eye reduction is apparently coupled with an obvious broadening of the vertex (see Fig. 2), so the index eye diameter: vertex width gives a clear statistical signal, ranging from 2-5 in epigean species with fully developed eyes to 0 in eyeless species. For the second index relative tegmina length we computed the absolute tegmina length: mesonotum width. Values ranges from 4.6-5.5 in epigean and from 1.1 to 5.3 in facultative and obligatory cavernicolous species. Te conspicuous diferences between epigean and cave-dwelling species are clearly refected in both indices (Fig. 4). If the data are plotted against the maximum age Asche, M., 1997: A review of the systematics of Hawaiian planthoppers (Hemiptera: Fulgoroidea). - Pac. Sci., 51(4), 366-376, Honolulu. Atkinson, A., Grifn, T. J. & Stephenson, P. J., 1976: A major lava tube system from Undara volcano, North queensland. - Bull. Volcanol., 39(2), 1-28, Napoli. Barr, T. C., 1968: Cave ecology and the evolution of tro-globites. - Evol. Biol., 2, 35-102, Amsterdam. Barr, T. C. & Holsinger, J. R., 1985: Speciation in cave faunas. - Ann. Rev. Ecol. Syst., 16, 313-337, Palo Alto. Best, J. G., 1983: 1:250,000 geological series, explanation notes. 2nd printing. - Geological Survey of queensland, p. 36, Atherton, qld. Carson, H. L., 1968: Te population fush and its genetic consequences. In: Lewontin, R. C., ed.: Population biology and evolution. - Syracuse University Press, 123-137, Syracuse, Ny. Carson, H. L., 1975: Te genetics of speciation at the dip-loid level. - Am. Nat., 109, 83-92, Chicago. of the cave species, the a priori expectation is a clear negative correlation, at least for the Hawaiian taxa: the oldest cave lineages should exhibit the highest degree of troglo-morphy. In contrast, we surprisingly found a weak (not signifcant) positive trend (Fig. 5). Te same unexpected trends are seen in the Australian planthopper species. Our results presented here, although preliminary, do not provide any evidence for cave-adaptation as a gradual orthogenetic process. Instead, we rather postulate that founder efects indeed play an important role in the origin of cave species. A correlation of the observed trends with particular ecological parameters of the cave environment cannot be excluded based on our data, but clearly this hypothesis needs further testing, especially in respect to selection pressures exerted by the conditions in high stress environments (such as caves) (Hoch & How-arth 1989b, Howarth 1993). we can conclude with some certainty, however, that even in closely related species the degree of troglomorphy cannot be employed to infer the phylogenetic age of the cave lineages. Culver, D. C., 1982: Cave life. Evolution and ecology. -Harvard University Press, p. 190, Cambridge, MA. Darwin, C. R., 1859: On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. - John Murray, p. 502, London. Fennah, R. G., 1973: Te cavernicolous fauna of Hawaiian lava tubes, 4. Two new blind Oliarus (Fulgoroi-dea: Cixiidae). – Pac. Ins., 15, 181-184, Honolulu. Ford, T. D., 1978: Chillagoe – a tower karst in decay. -Trans. Brit. Cave Res. Assoc., 5(2), 61-84, Bridge-water. Hoch, H., 1994: Homoptera (Auchenorrhyncha, Fulgoroi-dea). In: Juberthie, C. & Decu, V. , eds.: Encyclopaedia biospeologica. tome I. - Société de Biospéologie, 313-325, Moulis-Bucarest. ACKNOwLEDGEMENTS we thank Dr. Tomas von Rintelen, Museum für Naturkunde der Humboldt-Universität zu Berlin, for helpful discussions and many useful suggestions. REFERENCES TIME in KARST – 2007 205 ANDREAS wESSEL, PETRA ERBE & HANNELORE HOCH Hoch, H., 2002: Hidden from the light of day: planthop-pers in subterranean habitats (Hemiptera: Auche-norrhyncha: Fulgoromorpha). In: Holzinger, w. & Gusenleitner, F., eds.: zikaden. Leafoppers, plan-thoppers and cicadas (Insecta: hemiptera: Auchenor-rhyncha). (Denisia 4) - Oberösterreichisches Landesmuseum, 139-146, Linz. Hoch, H. & Howarth, F. G., 1989a: Reductive evolutionary trends in two new cavernicolous species of a new Australian cixiid genus (Homoptera Fulgoroidea). -Syst. Entomol., 14, 179-196, Oxford. Hoch, H. & Howarth, F. G., 1989b: Six new cavernicolous cixiid planthoppers in the genus Solonaima from Australia (Homoptera Fulgoroidea). - Syst. Ento-mol., 14, 377-402, Oxford. Hoch, H. & Howarth, F. G., 1993: Evolutionary dynamics of behavioral divergence among populations of the Hawaiian cave-dwelling planthopper Oliarus polyphemus (Homoptera: Fulgoroidea: Cixiidae). - Pac. Sci., 47, 303-318, Honolulu. Hoch, H. & Howarth, F. G., 1999: Multiple cave invasions by species of the planthopper genus Oliarus in Hawaii (Homoptera: Fulgoroidea: Cixiidae). - Zool. J. Linn. Soc., 127, 453-475, Oxford. Hoch, H., Asche, M., Burwell, C., Monteith, G. M. & wessel, A., 2006: Morphological alteration in response to endogean habitat and ant association in two new planthopper species from New Caledonia (Hemiptera: Auchenorrhyncha: Fulgoromorpha: Delphacidae). - J. Nat. Hist., 40(32-34), 1867-1886, London. Hoch, H. & wessel, A., 2006: Communication by substrate-borne vibrations in cave planthoppers. In: Drosopoulos, S. & Claridge, M. F., eds.: Insect sounds and communication. Physiology, behaviour, ecology and evolution. - CRC-Taylor & Francis, 187-197, Boca Raton, London, New york. Howarth, F. G., 1972: Cavernicoles in lava tubes on the Island of Hawaii. – Science, 175, 325-326, washington, DC. Howarth, F. G., 1981: Non-relictual troglobites in the tropical Hawaiian caves. - Proc. 8th Int. Cong. Spe-leol., 539-541, Bowling Green. Howarth, F. G., 1986: Te tropical cave environment and the evolution of troglobites. - Proc. 9th Cong. Int. Speleol., 2, 153-155, Barcelona. Howarth, F. G., 1987: Te evolution of non-relictual tropical troglobites. - Int. J. Speleol., 16, 1-16, Bologna. Howarth, F. G., 1988: Environmental ecology of North queensland Caves: why there are so many troglo-bites in Australia. - Te 17th Australian Speleological Federation Biennial Conference, TROPICON, 77-84, Lake Tinaroo, Far North queensland, Australia. Howarth, F. G., 1993: High-stress subterranean habitats and evolutionary change in cave-inhabiting arthropods. - Amer. Nat., 142(Suppl.), 65-77, Chicago. Howarth, F. G. & Hoch, H., 2005: Adaptive shifs. In: Culver, D. C. & white, w. B., eds.: Encyclopedia of caves. - Elsevier Academic Press, 17-24, Amsterdam. Jennings, J. N., 1982: Karst of northeastern queensland reconsidered. - Tower Karst, 4, 13-52, Chillagoe. Kemp, E. M., 1978: Tertiary climatic and vegetation history of the Southeast Indian Ocean region. - Palaeo-geography, Palaeoclimatology, Palaeoecology, 24, 169-208, Amsterdam. Mayr, E., 1954: Change of genetic environment and evolution. In: Huxley, J., Hardy, A. C. & Ford, E. B., eds.: Evolution as a process. - Allen & Unwin, 157-180, London. Pearson, L. M., 1982: Chillagoe Karst solution and weathering. - Tower Karst, 4, 58-70, Chillagoe. Remane, R. & Hoch, H., 1988: Cave-dwelling Fulgoroi-dea (Homoptera Auchenorrhyncha) from the Canary Islands. - J. Nat. Hist., 22, 403-412, London. Sbordoni, V. , 1982: Advances in speciation of cave animals. In: Barigozzi, C., ed.: mechanisms of specia-tion. - Liss, 219-240, New york. Sket, B., 1986: why all cave animals do not look alike – A discussion on adaptive value of reduction processes. - NSS Bulletin, 47, 78-85, Huntsville. Stone, F. D., 2004: Blattodea in the genus Nocticola from Australian cave & surface habitats. In: LaSalle, J., Patten, M. & Zalucki, M., eds.: Entomology – Strength in diversity. (xxII International Congress of Entomology, Brisbane 2004) - Austr. Entomol. Soc., S14w66 (abstract on CD-ROM), Brisbane. Truswell, E. M., 1990: Australian rainforests: the 100 million year record. In: webb, L. J. & Kikkawa, J., eds.: Australian tropical rainforests: science, values, meanings. - CSIRO, 7-22, Melbourne. weismann, A., 1886: Ueber den Rückschritt in der Natur. – Ber. Naturforsch. Ges. Freiburg i. Br., 2, 1-30, Freiburg/Br. wilkens, H., 1986: Te tempo of regressive evolution: Studies of the eye reduction in stygobiont fshes and decapod crustaceans of the Gulf Coast and west Atlantic region. - Stygol., 2, 130-143, Leiden. Zimmerman, E. C., 1948: Insects of hawaii. A manual of the insects of the hawaiian Islands, including an enumeration of the species and notes on their origin, distribution, hosts, parasites, etc. vol. 4, homoptera: Auchenorhyncha. - University of Hawaii Press, p. 268, Honolulu. 206 TIME in KARST – 2007 ABSTRACTS RULES OF CLIMATE, SOIL AND VEGETATION ON DEVELOPMENT OF THE KARSTSySTEM Ilona BÁRÁNy –KEVEI1 Department of Climatology and Landscape Ecology, University of Szeged, 6722. Szeged, Egyetem u. 2. POBox 653, Hungary; e-mail: keveibar@earth.geo.u-szeged.hu In the evaluation of environmentally sensitive karst regions for nature conservation value the most useful information is supplied by the changes in the ecological conditions of the climate-soil-vegetation system. Te changes in the system determine matter and energy cycles. A change in any of the three factors involves changes in the other two and eventually in the future functioning of the whole karst system. Climate infuences the physical, chemical and biological processes of the karst system. Air temperature, humidity, precipitation and evaporation infuence the water and matter cycles. Temperature regulates life processes of the biota. Matter transport is a function of soil, vegetation, relief and climatic parameters. Te karst regions of various nature are characterised by diferent processes. In landscape planning and management this mechanism of interactions has to be taken into consideration in every case in the future. TIME AND KARST PROCESSES: SOME CONSIDERATIONS Pavel BOSÁK1, 2 Institute of Geology, Academy of Science of the Czech Republic, Rozvojová 269, 165 02 Praha 6-Lysolaje; email: bosak@gli.cas.cz Karst Research Institute, ZRC SAZU, Titov trg 2, 6230 Postojna, Slovenia Karst evolution is particularly dependent upon the time available for process evolution and on the geographical and geological conditions of the exposure of the rock. Te time scale for the development of karst features cannot be longer than that of the rocks on which they form. Te longer the time, the higher the hydraulic gradient and the larger the amount of solvent water entering the karst system, the more evolved is the karst (Tab. 1). In general, stratigraphic discontinuities, i.e. intervals of nondepo-sition (disconformities and unconformities), directly infuence the intensity and extent of karstifcation. Te higher the order of discontinuity under study, the greater will be the problems of dating processes and events. Te order of unconformities infuences the stratigraphy of the karst through the amount of time available for suba-erial processes to operate. Results of paleokarst evolution are best preserved directly beneath a cover of marine or continental sediments, i.e. under sediments, which terminated karstifcation periods or phases. Te longer the stratigraphic gap the more problematic is precise dating of the age of the paleokarst, if it cannot be chronostrati-graphically proven. Terefore, ages of paleokarsts has been associated chiefy with periods just or shortly before the termination of the stratigraphic gap. Te characteristic time scale for the development of a karst surface landform or a conduit is 10 to 100 ka. Determining the beginning and the end of the life of a karst system is a substantial problem. In contrast to most of living systems development of a karst system can be „frozen“ and then rejuvenated several times (polycy-clic and polygenetic nature). Te principal problems may include precise defnition of the beginning of karstifca-tion (e.g. inception in speleogenesis) and the manner of preservation of the products of karstifcation. Terefore, karst and cave flls are relatively special kinds of geologic materials. Te end of karstifcation can also be viewed from various perspectives. Te fnal end occurs at the moment when the host rock together with its karst phenomena is completely eroded/denuded (tze end of the karst cycle) or sunken into the subduction zone. In such cases, nothing remains to be dated. Karst forms of individual evolution stages (cycles) can also be destroyed by erosion, denudation and abrasion, complete flling of epikarst and covering of karst surface by impermeable sediments, without the necessity of the destruction of the whole sequence of karst rocks. Temporary and/or fnal interruption of the karstifcation process can be caused by the fossilisation of karst due to loss of its hydrological function. Such fossili-sation can be caused by metamorphism, mineralisation, marine transgressions, burial by continental deposits or volcanic products, tectonic movements, climatic change TIME in KARST – 2007 207 ABSTRACTS tab. 1: Evolution of selected karst features in time on the background of transgression-regression set within one hypothetical karst period related to unconformity order FealufeJOtdef 1 2 3 4 5 UtiCOtifttffaity' MegaurtCönfüimily SupönurtCönfünTiily Regional Lin content hty Parastqui&noe öüundary ¦eeddmg plana' Cam'frean mcufef* Interregional kars-l Local karst Deposit kj rial karsi General model" Karst penod Karst phase Type 1 Karst pnase TvwZ Gät>k>QiCal SGHirtQ Cralan/Platfomn - CralDri.flaHurnfi + c&nire margins DBpoalional basin Ttms {Mb} XM-XÜ XÜ-X x-o.x O.X-fl.OX 0.OX-O.0DX Fnsshw-Bler lEni Protoso) Caliche Soil W&Blhefing BroJile Karren 1 Sinkhole Cava Cave s-yslem HvpogenitKarsI HYdml hernial k. Early tarsi" Mjilure lonjf* BuneC ksnst" Reju>analBd k " Relict karsl" Unnco^gd ia^gr <----------------------- Transgression Regression -------------------------------------------------------------------------------------------------------------¦> SJKßV! Ja a>" SffiH^S + sensu Esteban (1991); * sensu Choquette & James (1988); ** sensu Bosák et al, (1989); § sensu Mihevc (1996); weathering profile = more evolved weathering cover (like laterite, bauxite, kaoline, etc.); Hypogenic karst = deep-seated karst, interstratal karst, intras-tratal karst, subjacent karst, subrosion. a etc. Nevertheless, in contrast to living organisms, the development of the karst system can be „frozen“ and rejuvenated even for a multiplicity of times (polycyclic and polygenetic nature of karst). Further, the dynamic nature of karst can cause redeposition and reworking of classical stratigraphic order, making the karst record unreadable and problematic for interpretation. Known karst records for the 1st and 2nd orders of stratigraphic discontinuity cover only from 5 to 60 % of geological time (time not recorded in any correlated sediments in old platforms usually represents 40 to 90 % of time). Te shorter the time available for karstifcation, the greater is the likelihood that karst phenomena will be preserved in the stratigraphic record. while products of short-lived karstifcation on shallow carbonate platforms can be preserved by deposition during the immediately succeeding sea-level rise, products of more pronounced karstifcation can be destroyed by a number of diferent geomorphic processes. Te longer the duration of sub-aerial exposure, the more complex are those geomorphic agents. Te dating of cave initiation and evolution, i.e. the origin of the void within the bedrock is more problematic. Te age of the erosional cave falls between the age 208 TIME in KARST – 2007 ABSTRACTS of the host rock and that of the oldest dated fll. with the inception theory, the true start of speleogenesis can hardly be estimated. Many caves contain only very young flls, older ones having been excavated during repeating cave exhumations/rejuvenations caused by changes in hydraulic conditions, spring position, climate, etc. Te minimum age for the cave initiation phase is estimated to be a minimum of 10 ka and cave enlargement up to accessible diameters usually takes about 10-100 ka under favourable conditions. URANIUM-THORIUM AGES OF STALAGMITES FROM KATERLOCH CAVE (STyRIA, AUSTRIA) Ronny BOCH1, Christoph SPÖTL1 & Jan KRAMERS2 1 Institut für Geologie und Paläontologie, Leopold-Franzens-Universität Innsbruck, Innrain 52, 6020 Innsbruck, Austria; e-mail: Ronny.Boch@uibk.ac.at 2 Institut für Geologie, Universität Bern, Baltzerstraße 1, 3012 Bern, Switzerland Speleothems recently experienced growing importance in the light of paleoenvironmental research. Absolute age determination using Uranium-series methods allow for precise age constraints of environmental information preserved in speleothems, particularly in stalagmites and fowstones. Katerloch Cave, situated within the Styrian Karst Province near Graz was chosen as an object of extensive paleoenvironmental studies. Abundantly decorated cave chambers show a clear dominance of stalagmites over stalactites, many of them being several meters in length. Te majority of stalagmites are of the candle-stick type, a morphology suggesting fast growth. Five inactive stalagmites were recovered and dated using the U-T method. In addition, drill cores were obtained from the base of in-situ growing stalagmites. Te dating of calcite sub-samples of these cores give an age of growth inception of the respective stalagmites. Age measurements were carried out using Multi Collector-ICP-Mass Spectrometry at the Institute of Geology at Bern, Switzerland. Our dating campaign allowed detecting several speleothem generations: stalagmites from the current Interglacial, the Marine Isotope Stage 3 and the Last Interglacial (MIS 5.5), indicating major speleothem growth during relatively warm (and wet?) climate conditions. Te ages of two stalagmites are beyond the range of the U-T method, i.e. they are older than ~ 450 kyr. Two Holocene stalagmites, 73 and 139 cm in length, yielded ages between 10.32 ± 0.13 and 7.02 ± 0.14 kyr and between 9.80 ± 0.24 and 7.88 ± 0.09 kyr, respectively. Age models derived from dating of multiple sub-samples along the stalagmite growth-axis suggest growth rates of 0.2 to 0.7 mm/yr. Tese rates are very high compared to caves in the alpine region. Two stalagmites from the preceding Interglacial (MIS 5.5) suggest similar growth rates than those of the Holocene stalagmites. Furthermore, fast growth of speleothems in Katerloch Cave is corroborated by stalagmite petrography, displaying a lamination of alternating white-porous and compact-translucent layers. COPEPOD CRUSTACEAN DIVERSITy IN SOUTH FLORIDA KARST, U.S.A. Maria Christina BRUNO1 & Janet w. REID2 1 Museum of Natural Sciences of Trento, Section of Hydrobiology and Invertebrate Zoology, Via Calepina 14, 38100, Trento, Italy. 2 Virginia Museum of Natural History, 21 Starling Avenue, Martinsville Virginia 24112, USA. Southern Florida is mostly occupied by the Everglades, an extensive subtropical wetland ecosystem that formed during the past 5,000 years when peat and marl were deposited within a pre-existing limestone depression in the southern peninsula. Te high porosity of the limestone of the aquifer allows for considerable fux between surface water and ground water. Hydrologically, karst systems in South Florida are very open, and numerous epigean invertebrates ofen penetrate the aquifer by means of sinkholes, some of them establishing permanent populations in the aquifer. Te inventory of free-living freshwater copepods recorded from peninsular Florida includes a total of 65 taxa: 9 calanoids, 41 cyclopoids, and 15 harpacticoids. TIME in KARST – 2007 209 ABSTRACTS Two-thirds (44) of these are known from Everglades National Park and adjacent areas, partly as a result of more intensive sampling in this area; 10 have so far been found only in the Everglades. Of the species collected in central and northern Florida, 2 calanoids and 1 cyclopoid have been found only in the state, whereas all the others are widespread in North America and beyond. South of the Everglades, in the Florida Keys, recent collections from small permanent or ephemeral surface waterbod-ies, some of them brackish, yielded 2 species of calanoids, 27 cyclopoids, and 11 harpacticoids, adding 1 calanoid, 2 cyclopoids and 9 harpacticoids to the list for Florida. Ten species of cyclopoids and 1 harpacticoid collected in the Florida Keys were already known from the Everglades. In peninsular Florida, the Nearctic fauna is predominant, but a small Neotropical component is present (1 calanoid, 6 cyclopoids, and 5 harpacticoids); 1 cyclopoid species is considered to be introduced. In the Florida Keys, the assemblage consisted mainly of cosmopolitan or Neotropical continental cyclopoids (1 introduced), and Neotropical, coastal harpacticoids, with only 2, Neotropical continental calanoids. Because the Florida peninsula is relatively young geologically, we did not expect that a diverse endemic groundwater fauna would be present. However, several taxa that are usually known elsewhere from surface water, were collected in subterranean water in the Florida Everglades during the dry season, likely an adaptation to survive the drought. Tis happened for the 2 species of calanoids, 16 of the 27 cyclopoids, and 4 of the 13 harpac-ticoids; among them, stygophile taxa were represented by 6 cyclopoids and 3 harpacticoids. Te species occurring in groundwater in the Everglades are either widely distributed elsewhere in North America or in the neotropics, members of speciose genera with both epigean and hy- 1 ”Emil Racovitza” Institute of Speleology, Bucharest, Romania. 2 University of Bergen, Department of Geology, Norway. Several speleothem samples from Piatra Cave (Dobrogea, South-East Romania) were dated by means of alpha and mass spectrometry, in order to obtain absolute datings of the episodes during which speleotheme formation oc-cured here. Piatra Cave is a submerged cave located a few kilometers far from the Black Sea shore at an absolute altitude of 1-2 m, thus being suitable for recording the sea pogean species, or members of predominantly marine groups with some brackish- or freshwater representatives. Te low diversity of harpacticoids in groundwater habitats in the Everglades is surprising, because harpacticoids include the largest number of stygobitic forms within the copepods. Te closest area in which true stygobitic cope-pods have been found is northern Florida, which has an older geological history. Te relatively young age of the Everglades may not have allowed some taxa enough time to disperse here, or may not have allowed sufcient time for groundwater colonizers to evolve adaptations to life in subterranean habitats. Besides the young geological age, historical and environmental conditions may also account for the Everglades’ relatively depauperate groundwater copepod fauna. Both the higher-elevation marshes and the deeper central sloughs undergo periodic droughts of varying severity, which afect the composition of the copepod fauna. Te hydrological regime is irregular because of variations in rainfall and water-management activities, and ground-water communities are dominated by surface copepods that colonize groundwater mainly during the dry season. Te abundance and species richness decrease with depth, and sharply below the 3-m depth, due to high permeability of the limestone above 3 m and to the presence of a semipermeable layer at lower depths. Groundwater copepod communities are more similar on a local scale: when local surface-water populations enter the shallow aquifer by following the receding water table, they do not generally disperse widely through the groundwater system. Te dissimilarity in communities over larger distances may refect diferences in surface habitats, as well as limitations on dispersal by diferent local porosities of the limestone. level oscillations that took place in the past. Nine dates were performed by alpha spectrometry and one by mass spectrometry. In spite of a low Uranium content (0.04 – 0.1 ppm), all subsamples showed good chemical extraction yields, thus providing analytical reliable dates. One of the stalagmites appears to have been grown very fast during the Eemian (OI 5e), betwen 112 ± 10 ka and 122 BLACK SEA LEVEL FLUCTUATIONS BASED ON THE STUDy OF SUBMERGED SPELEOTHEMES FROM DOBROGEA (ROMANIA) Silviu CONSTANTIN1, Virgil DRAGUSIN1, Cristian LASCU1 & Stein Erik LAURITZEN2 210 TIME in KARST – 2007 ABSTRACTS ± 13.5 ka ( 1? ). Another sample, dated by TIMS method yielded a date of 597 +108/-53 ka. Although these dates are not sufcient for a detailed analysis of the Black Sea oscillations, they reveal two periods of time during which environmental conditions here were favourable for spele-otheme deposition as the cave was emerged. Most interesting are the samples dated from the warm OI substage 5e during which, the Black Sea level is supposed to have been 10 m higher than the present one. Te area, during the quaternary, being virtually stable from the tectonic point of view (except for a narrow coastal area), leads us to three possible situations that allowed speleotheme formation: either there was a slight subsidence in the area (around 15 m), or the sea-level was more than 15 meters lower than previously believed. Te third possibility could be that during the main transgression that rose the level 10 meters above the present one, there has been a short period of low-stand. HIGH-RESOLUTION SPELEOTHEM RECORDS FROM SOqOTRA ISLAND (yEMEN), AS RECORDERS OF INDIAN OCEAN MONSOON VARIABILITy Pierre De GEEST1 Department of Geology, Vrije Universiteit Brussel, Plenlaan 2, 1050 Brussels, Belgium; e-mail: Pierre.De.Geest@vub.ac.be Te arid tropical island Soqotra is situated in the Indian Ocean between the Horn of Africa and the Arabian Peninsula. Here a bi-annual rainy season is active, due to the passing of the inter-tropical convergence zone (ITCZ) twice each year, known as the Indian Ocean Monsoon system. Only recently more than 35 km of underground cave passages in around 50 caves were discovered and explored, revealing an important karst potential for paleo-climate and paleoenvironmental research. By evaluating the frst available meteorological data (rainfall and temperature) from the last 5 years, we calculated that approximately 85% of the rainfall is related to the NE Monsoon period, while only 15% is related to the Sw Monsoon period with an important irregular geographical distribution over the island. Diferences in the oxygen isotope composition of meteoric water versus groundwater are used to estimate amount and timing of karst aquifer recharge, because seasonal fuctuations of rainwater oxygen isotopic composition are related to the amount of rainfall. Karst aquifer recharge at the NE lime- stone plateau takes only place during the NE Monsoon rainy period when a rainfall threshold of 80-90mm is exceeded, explaining the more negative cave drip waters and groundwater in general. Because the controls on the isotopic composition of nowadays forming speleothem calcite could be monitored in this region, two caves in this area were chosen as research location. A sampled speleothem from Hoq Cave (S STM1) and one from Casecas Cave (S-STM5) have formed over the last 6 ka BP and the last 1 ka BP (U/T dating) respectively. Carbon and oxygen isotopic measurements were performed at a resolution up to 50 m, corresponding to a time resolution of up to one month. Te two sampling locations, distant of 6km, display similar isotopic changes in both speleothems over the last 1 ka. Te speleothems also clearly registered seasonal variations, coinciding with the alternation of dark compact and white porous layers, representing annual banding in both stalagmites. Tese observations suggest that the speleothems reliably registered Monsoon climate variability. EVOLUTIONARy CHANGE IN CAVE ADAPTED ORGANISMS Katharina DITTMAR1 1 Department of Molecular Biology, University of wyoming, 1000 E. University Avenue, 82072 Laramie, wyoming, USA; e-mail: katharinad@gmail.com Change of selective constraint on a gene may be expect- coding genes across cave- and non cave related taxa to ed following changes in the environment or life history. assess general patterns of evolutionary changes in dif-Tis is especially true for switches to the cave environ- ferent lineages, and whether (and to what extend) they ment. Te purpose of this work is to explore selective depend on environmental conditions. Protein coding se-pressures and functional change on a variety of protein quences that appear to be undergoing adaptive evolution TIME in KARST – 2007 211 ABSTRACTS or changes of function along specifc branches of the tree will be analyzed in detail as to determine the specifcity and functionality of those changes. Results of this work will be incorporated into the publicly accessible TAED database (http://www.sbc.su.se/~liberles/TAED2002/) and will help pinpoint target genes for future studies on cave organisms and their environment (e.g., karst). TEMPO OF CHANGES IN KARST BIOLOGICAL VERSUS PHySICAL PROCESSES Louis DEHARVENG1 1 UMR5202 CNRS, Origine, Structure et Evolution de la Biodiversité, USM601, CP50 Museum National d’Histoire Naturelle, 45 rue Bufon, 75005 Paris (France); e-mail: deharven@mnhn.fr Time is an essential parameter for understanding any ecosystem. Lansdcape, habitats, community composition, genetic pools and species distribution change with time, in a more or less connected way. yet, integrative approaches linking physical and biological karst processes in a time perspective are scarce in the literature. One of the intriguing property of subterranean karst systems is that time required for biological or physical processes may difer considerably from that required for similar processes in surface karst or non-karst systems. Essential aspects of this peculiar tempo of changes in karst are presented here. Teir interest for understanding paleogeog-raphy and organism evolution are discussed. KARST AS A MODEL SySTEM TO ExAMINE TERRESTRIAL MICROBIAL BIOGEOGRAPHy PATTERNS THROUGH TIME: AN ExAMPLE FROM THE EPSILONPROTEOBACTERIA Annette SUMMERS ENGEL1, Megan L. PORTER2 & Barbara J. CAMPBELL3 Department of Geology & Geophysics, Louisiana State University, Baton Rouge, LA 70803 USA; e-mail: aengel@geol.lsu.edu Department of Sciences, University of Maryland Baltimore County, Baltimore, Maryland 21250, USA; e-mail: porter@umbc.edu College of Marine and Earth Sciences, University of Delaware, Lewes, DE 19958 USA; e-mail: bjc@udel.edu Te evolutionary mechanisms that govern the distribution of microbes on Earth are poorly understood. Te concept that geographic isolation among microorganisms can lead to endemism is controversial. Because microbes are small in size, are metabolically tenacious, and are overwhelmingly abundant on Earth, microbes may be distributed everywhere and therefore may not be afected by geography. However, some studies reveal that various taxa have restricted distribution patterns. Many micro-bial biogeography studies have been conducted to test for microbial species endemism, but these studies have been conducted in habitats with high dispersion (e.g., aquatic, and specifcally oceanic, habitats). Consequently, there have been few studies of microbial biogeography in terrestrial habitats. Karst, in particular, is a discontinuous habitat typifed by diferent hydrostratigraphic units in distinct geologic provinces formed over time. while the karst habitat may be similar in its physicochemical and geological conditions from one locale to another, microbial groups colonizing karst may refect biogeography because of genetic exchange limitations. In the separated karst habitats, speciation events could be high. Considering the ubiquity of karst worldwide, understanding the types of microbial communities in cave and karst systems, as well as determining what geologic and geochemical processes may control microbial species distribution and diversity, are critical aspects to preserving the integrity of the karst ecosystem and to predicting ecosystem changes that may occur following disturbances. Te Epsilonproteobacteria taxonomic class is an ideal microbial group to investigate endemism, biogeographic diversity, and possible mechanisms controlling bacterial evolution in karst. Members of this class are found in many diferent sulfur-rich environments, including marine and terrestrial aquatic settings. Although investigated less, e-Proteobacteria from terrestrial settings are potentially 212 TIME in KARST – 2007 ABSTRACTS widely distributed. Our previous investigations diagnosed novel evolutionary lineages within the ?-Proteobacteria from one terrestrial subsurface environment, Lower Kane Cave (LKC) in wyoming (USA). Two lineages dominated the subterranean communities and were related to organisms identifed from other sulfur-rich terrestrial habitats (contaminated groundwater and caves), but not to microbes from marine habitats, including deep-sea hydrothermal vent sites and marine sediments. Microbial mat samples from six cave and nine karst spring locations were collected, including the Frasassi Caves (Italy) and Movile Cave (Romania). 16S rRNA gene sequences were amplifed, cloned, and sequenced from total community DNA extractions using general and ?-pro-teobacterial lineage-specifc primers. From a total of 336 sequences used for phylogenetic analyses, 71 ?-proteobac-terial species-level lineages (operational taxonomic units, OTUs) were identifed from cave and spring habitats. we recognize the fact that biogeographic interpretations based on the presence or absence of 16S rRNA genes should be made cautiously. However, fve OTUs, consisting of sequences from up to nine locations, came from diferent continents and from karst systems with varying speleoge-netic histories. Analyses of single copy genes (rpoB and aclB) from selected karst locations are currently underway to test distribution and occurrence pattern diferences between the multiple copy genes (e.g., 16S rRNA gene) and single-copy genes. we expect the number of OTUs will be more than that observed with the 16S rRNA gene. Cave-adaptation, the shif to a life underground, can provide critical information to understand evolutionary change in general, but the process of cave-adaptation is still not well understood and controversially discussed. Planthoppers, especially of the family Cixiidae (In-secta: Homoptera: Fulgoromorpha) inhabited caves in many parts of the Tropics and Subtropics independently. Tese evolutionary lineages are ideally suited models to study the concepts of genetic diferentiation and spe-ciation dynamics. One of those planthopper taxa is the Australian genus Solonaima. Tis genus is endemic to queensland and New South wales and contains epigean Our initial 16S rRNA gene sequence investigations do not reveal a correlation between the relative speleogenetic age of the cave, the age of the carbonate host rocks, and the microbial group (or OTU). Although most of the systems have formed within the past 500 ka (thousand years) in carbonate rocks ranging up to 400 Ma (millions of years old), all of the cave and spring locations are currently inundated by modern sulfdic waters that are the consequence of distinct regional hydrostratigraphic conditions. Tese data suggest that extremely ancient, large-scale geological processes and events (e.g., depositional basin sedimentol-ogy and geochemistry, plate tectonics, regional karstifca-tion events) must have infuenced the ancestral distribution of these groups, which were likely marine in origin. Once ?-Proteobacteria colonized these terrestrial habitats, limited dispersal capabilities (i.e. no cyst or spore formation) and hydrostratigraphic barriers to genomic exchange led to site-specifc lineage evolution. In our analyses, this is indicated by more than 89% of the OTUs consisting of sequence groups from single sampling locations. Furthermore, three OTUs (comprised of sequences from more than one location) were retrieved from the same geographic region, illustrating the potential importance of biogeography in terrestrial ?-Proteobacteria distribution patterns. Te large percentage of site- and region-specifc OTUs detected in our study indicates potentially high site-specifc endemism efects for some terrestrial ?-proteobac-terial lineages and possibly some recent (on an evolutionary/geologic time scale) exchange on a regional scale. as well as cavernicolous species. Te epigean species occur in the rainforest habitats along the east coast, while the cave-dwelling species are restricted to more arid areas western to the Great Dividing Range (Chillagoe & Mitchell Palmer Karst and Undara Lava Tubes). Tis relictous pattern seems to be due to the extinction of epi-gean species in the Outback according to the retreat of habitat starting with the desertifcation of inner Australia in Miocene, Pliocene. Te observed species display diferent degrees of troglomorphy, which are incongruent with the age of the caves, as the highest derived species occur in young lava tubes, while less cave-adapted species occur in older RADIATION SCENARIOS FOR AUSTRALIAN CAVE-ADAPTED PLANTHOPPERS Petra ERBE1, Andreas wESSEL2 & Hannelore HOCH2 Uplands Program, Faculty of Agriculture, Chiang Mai University, 50200 Chiang Mai, Tailand; petra.erbe@gmx.net Museum für Naturkunde der Humboldt-Universität zu Berlin, Department of Research, Biosystematics Group, Invalidenstrasse 43, D-10115 Berlin, Germany; e-mail: andreas.wessel@museum.hu-berlin.de TIME in KARST – 2007 213 ABSTRACTS Tower Karsts, which also leads to the assumption of several cave invasion events. According to the cladogram only two cave-invasions are requisited, one for S. sullivani and one for the other cavernicolous species, although the problem of homopla-sis in cave-adaptation has to be considered. If more than one ancestral epigean Solonaima species inhabited once the Outback, from which all cavernicolous species (except S. sullivani) descended, all hints for this are vanished with these hypothetic ancestral species and/or veiled by cave inhabiting conditioned homoplasis. Hence, morphometrical statistics and variation studies concerning aedeagal characters were conducted additionally, allowing to reconstruct scenarios for the invasion history of the extant populations: 1. Multiple cave-invasion scenario 2. Single initial cave-invasion scenario To accommodate these results a hypothesis merging the relict hypothesis (Barr 1968) and the adaptive-shif hypothesis (Howarth 1986) had to be formulated. CAVE-DwELLING PLANTHOPPERS OF THE GENUS SOLONAIMA (INSECTA: HEMIPTERA: CIxIIDAE) IN AUSTRALIA: RELICTS OR ExPLORERS? Petra ERBE1,2, Andreas wESSEL2 & Hannelore HOCH2 1 Te Uplands Program, Hohenheim Ofce, Faculty of Agriculture, Chiang Mai University, 50200 Chiang Mai, Tailand; e-mail: petra.erbe@gmx.net 2 Museum für Naturkunde der Humboldt-Universität zu Berlin, Department of Research, Biosystematics Group, Invalidenstrasse 43, D-10115 Berlin, Germany. Te evolution of obligately cavernicolous terrestrial organisms is still controversially discussed. Traditionally, cave colonization has been interpreted as a response to adverse (climatic/ecological) conditions on the surface leading to the extinction of epigean populations, “trapping” troglo-philic populations inside underground environments. Tese populations subsequently acquired morphological (troglomorphies) and behavioral alterations. Te patterns commonly observed in temperate regions seem to support this assumption. Te discovery of terrestrial obligate cav-ernicolous invertebrates (mainly arthropods) in the tropics, however, with close epigean relatives still extant, challenged this view. Here, cave colonization and -adaptation could also be the result of an active process, driven by the exploitation of novel food resources, such as roots. Here we present a model system which allows us to test these hypotheses. Te Australian planthopper genus Solonaima, endemic with ca. 15 species in queensland and New South wales, contains epigean as well as facultative and obligatory cavernicolous species. Te cave-dwelling species display varying degrees of troglomorphy pertaining to eye- and wing reduction. Epigean Solonaima species occur in rainforest habitats along the East Coast while the cave-dwelling species are restricted to more arid areas west of the Great Dividing Range. To test the existing hypotheses on our model system, the following criteria should be applied, while these hypotheses are not necessarily considered to be opposi-tional: Relict hypothesi s - deterioration of climatic conditions on the surface - close epigean relatives not extant - cavernicolous taxa are of the same phylogenetic age Adaptive shif hypothesis - stable climatic conditions on the surface - close epigean relatives extant - cavernicolous taxa are of diferent phylogenetic age Te current distribution pattern seems to support the relict hypothesis in the case of the cavernicolous Solonaima species: Epigean and cavernicolous species occur allopatrically. It is conceivable that in the course of the gradual desertifcation of central Australia during Miocene/Pliocene, when rainforests retreated eastwards, being replaced by a more xerophilic fora, Solonaima species survived in (the still moist) cave refugia and subsequently acquired troglomorphies. Te observation, however, of varying degrees of troglomorphies in cavernicolous Solonaima species may account for diferent phylogenetic age of these taxa and thus, be an indication for an adaptive shif. Hence it cannot be excluded that cave colonization and -adaptation in Solonaima occurred before the period of desertifcation, making an initial adaptive shif prior to the development of the extant relictual pattern assumable. 214 TIME in KARST – 2007 ABSTRACTS STyGOFAUNA FROM A KARSTIC ECOSySTEM IN THE PONOR MOUNTAINS, wESTERN BULGARIA: PRESENT KNOwLEDGE AND RESEARCH CHALLENGES Vesela VASILEVA EVTIMOVA1 1 Department of Hydrobiology, Institute of Zoology, Bulgarian Academy of Sciences, 1 Tsar Osvoboditel blvd., 1000 Sofa, Bulgaria; e-mail: evtimova@zoology.bas.bg Te Ponor Mountains (western Bulgaria) are part of the Balkan Ridge, with altitude 400 - 1473 m a.s.l. and more than 120 caves and chasms. Te massif is formed mainly by Mesozoic rocks. Two basic rock complexes can be distinguished regarding to the karstifcation: Triassic karstic complex, formed by lime and dolomites with maximum depth 600m; and upper Jurassic complex with depth 100-150m. Te two aquifers are hydrologically isolated. Basic collector of underground waters is the Triassic complex with annual outfow 2900 dm3/s from which 80% is leaving the system through the Iskretski izvori spring. Tis is the biggest spring in Bulgaria (maximum discharge 35000 dm3/s). Te Jurassic complex (average outfow 120 dm3/s) is lacking superfcial outfow which determines its precipitation alimentation. Te stygofauna is very heterogeneously distributed and its composition varies greatly from one station to another. From the aquifer with rocks with Triassic age are found 21 species while from the Jurassic one are found only 7. Te common species for the two aquifers are 5 amongst which with higher frequency of occurrence are 1 Earth & Planetary Sciences, washington University in St. Luis, e-mail: cfadem@levee.wustl.edu Te formation of and sedimentation processes in poljes (large, elongate, fat-bottomed karstic valleys) are uncertain and may be unique to each polje – dependent on climate, bedrock chemistry, and groundwater. One aim of our work on the Early Farming in Dalmatia Project at the Danilo Bit-inj and Pokrovnik archaeological sites is to elucidate polje sedimentary history. Analysis of his sediment record will inform the conditions of Neolithic settlement and earliest agriculture. To this end we are examining granulometric and ground-penetrating radar data from these valleys, and reconciling this GPR data with ground-truthing from soil profles to create a three-dimensional subsurface map. we Niphargus bureschi, Speocyclops lindbergi and Stygoela-phoidella elegans. At present 25 stygobionts are known from the Ponor Mountains: from Mollusca (2), Hirudin-ea (1), Acari (1), Copepoda (16), Syncarida (1), Isopoda (1), Amphipoda (3) groups. Crustacean assemblages are natural indicators of the typological characteristics of the system (hydrogeological division of karst, potential subsurface water connections, hydrological regime, contaminants transformation and bioaccumulation). Some of the main challenges to be faced in future concern understanding regional and fundamental functioning and structure of subterranean aquatic ecosystems. Te issues to be solved with the help of invertebrate communities are: origin and circulation pattern of groundwater fow in a fractured karstic aquifer; relationships between ground-water hydrodynamics and contamination; connections among the underground areas and understanding differences that may make individual areas unique in terms of fauna; assessment of the contemporary condition and extent of human perturbation on the Ponor Mountains ecosystem; measures for protection and management. Brookings Dr, Campus Box 1169, St. Louis MO 63130 USA; will use both the texture and 3-D form of the deposits to reconstruct the geomorphic setting prior to and during Neolithic occupation. we hope that ground-truthing the GPR data, along with textural and soils analyses will enable further interpretations concerning the presence and extent of debris fans, the presence of erosive surfaces in the bedrock and sediments, and evidence for high-energy sedimentation events. Large limestone clasts (5-50 cm) present in valley soils (on- and of-site) may be the product of in situ bedrock weathering or the result of high-energy or high-viscosity events. x-ray fuorescence analysis of bedrock and of valley clasts may also help resolve this question. N M F 1 A CONTExT FOR EARLy FARMING IN THE CENTRAL DALMATIAN POLJE: EVIDENCE FROM SEDIMENTS AND SOILS Cynthia FADEM1 TIME in KARST – 2007 215 ABSTRACTS MORPHOLOGICAL EVIDENCE OF MULTIPLE CAVE LEVEL DEVELOPMENT IN THE UPPER BASIN OF SOMESUL CALD RIVER, BIHOR MTS., ROMANIA Ioana FEIER1 1 Faculty of Geography – Geology, Al. I. Cuza University, B-dul Carol I 22, 700506 Iasi, Romania; e-mail: ioanafeier@yahoo.com Te karst on the upper basin of Somesul Cald River is situated in the Apuseni Mts., Romania, on the NE border of Padis karst plateau. It covers a relative small surface, superimposed on the drainage basins of Ponor, Alunu Mic, Alunu Mare and Firii valleys, all right hand afuents of Somesul Cald River. Two main types of karst are found in the area: slope karst and plateau karst. Te slope karst includes numerous caves situated at diferent elevations on the slopes of the above-mentioned valleys, vertical clifs and partly denudated caves. Te plateau karst is less extended, being present mainly on the northern sector of the area, at the springs of the Firii and Ponor valleys. Numerous dolines punctuate the surface, and some shafs gave access to deep and long cave systems, among them being the Avenul din Poienita-Humpleu karst system, the second longest in Romania. Down cutting of Somesul Cald river and subsequent lowering of the base level triggered the genesis of a multi- stage, well-defned karst system, developed along the rivers that cross this area. Here we report the results of a long-term cave survey project, aiming to correlate cave levels from the upper basin of Somesul Cald river, and construct a chronology of the karstifcation processes in the area. By combining morphologic observations (both in caves and at the surface) with detailed mapping of the caves we were able to link the quaternary evolution of the Somesul Cald River and its tributaries with the cave systems presents on the slopes of the rivers, and also establish a relative chronology of the karstifcation processes that afected the area. Moreover, morphological observations in the caves show that the early stages of cave development took place mainly under phreatic conditions, while later ones were in vadose conditions, as the entire area was uplifed. HyDROSTRATIGRAPHy OF THE KARST AqUIFERS OF FLORIDA Lee FLOREA1 & Kevin CUNNINGHAM1 United States Geological Survey, Florida Integrated Science Center, 3110 Sw 9th Ave, Ft. Lauderdale, FL 33315 (USA); e-mail: lforea@usgs.gov In the United States, two highly-productive carbonate aquifers occur within peninsular Florida: 1) the Floridan aquifer, primarily composed of Eocene to Oligocene carbonates generally unconfned in the northwestern part of the peninsula and confned in the remainder; and 2) the Biscayne aquifer of southeastern Florida, predominately unconfned and mostly Pleistocene in age. Combined, these two aquifers provide potable water for about 17 million Florida residents. Much of the unconfned Floridan and Biscayne aquifers are in low-lying, coastal, and subtropical environments. Both are composed of karstifed eogenetic carbonates that have not been deeply buried and therefore retain substantial primary porosity. Permeability within eogenetic aquifers is highly heterogeneous, with the ma- trix as much as 107 times as permeable as the matrix of telogenetic carbonates. Te karst features manifest within the unconfned Floridan and Biscayne aquifers ofer contrasting examples of cavernous-scale (>2-cm diameter) porosity found within eogenetic karst. Both aquifers, however, difer considerably from epigenic karst in telogenetic limestones, where water fows from sinkholes to springs through fractures and discrete conduits. Groundwater in eogenetic karst similarly can travel through large conduits and fractures, but in contrast, also through mazes of stratibound touching-vugs and rock matrix. Detailed petrophysical and geophysical studies demonstrate an organization to the hydrostratigraphy of both aquifers. For example, cavernous porosity is com- 216 TIME in KARST – 2007 ABSTRACTS monly layered. Te origins of the cavernous porosity are primarily associated with fuctuations in sea level. In the unconfned Floridan aquifer, stratiform cavities at 5 m, 12-15 m, 21 m, and 30 m above modern sea level occur at elevations similar to geomorphic terraces in Florida, suggesting cavity formation during higher paleo-altitudes of sea level and water table. Likewise, cavities at depths of 15 m, 40 m, 70 m, and 90-120 m below the modern water Located in the Leiria district (Portugal centre), the municipality of Alvaiázere it has an area of 160 km2 and it dominates the karst landscape with many characteristic karstic features. It is an area that presently is facing many problems concerning local development, a decreasing and ageing population are causing a dramatic scenario for its future as a municipality, which implies a big challenge for it’s own future, concerning economy, social and environmental policies. Other threats like infrastructures are a problem for the cultural, historic and geomorphologic heritage in this area. Despite this scenario, this area has many characteristics and features that can be a good opportunity for the development of a sustainable development strategy, based in its own values. Its an area in which culture, geo-morphology and landscape are closely linked and also an excellent opportunity to develop research at the interface between geomorphology and human sciences. Many values can also if not identifed be lost forever, not only geomorphologic values but also cultural, historic, among many others. table generally agree with depths of marine terraces submerged in the Gulf of Mexico, which formed at previously lower altitudes of sea level and water table. Distinctly diferent from the unconfned Floridan aquifer, cavernous porosity in the Biscayne aquifer commonly occurs as horizons of cm-scale, touching-vug porosity within the upper and lower boundaries of depositional cycles. A recent research has showed that this territory has a good potential for the development of a local strategy, similar to a geopark, based in this known and unknown values, but only afer being identifed this values. Te mountains of Alvaiázere and Ariques are probably the most important area in this municipality concerning to the existence of some of this important values. Karst features like lapiés, caves, doline, a karst valley are present here among many others. Also a rich biodiversity exists here, that´s why this area is included in the place Natura 2000 – Sicó/Alvaiázere. Other value with great importance is the late Bronze Age walled settlement of the mountain of Alvaiázere, one of the largest known habitats from this age in the west of the Iberian Península. Dinosaur footprints are also present in this area, not only in the mountains but also in the surrounding area. Te municipality of Alvaiázere from these values can create and stimulate a global strategy for sustainable territorial development along with geoconservation in this beautiful karst area and a door for the future. A LOCAL DEVELOPMENT OPPORTUNITy FOR A KARST AREA – THE MUNICIPALITy OF ALVAIÁZERE Joao FORTE1 Environment department, Municipality of Alvaiázere, Rua Conselheiro Furtado Santos, 3250-100 Alvaiázere, Portugal; e-mail: joao.forte@cm-alvaiazere.pt TIME in KARST – 2007 217 ABSTRACTS GEOCHEMICAL AND PHySICAL PROPERTIES OF STALAGMITES AS A MARKER OF PALEOENVIRONMENTAL CHANGES Lisa FULLER1 Hofman Environmental Research Institute, western Kentucky University, 1906 College Heights Blvd., Bowling Green, Ky 42101-3576, USA; e-mail: lisa.fuller@wku.edu Stalagmites are archives of palaeoclimatic information and many geochemical and physical properties can be used to interpret former environmental conditions that may have occurred during their growth. when calibrated against the present conditions in the cave system, the composition and growth rate of stalagmites can be interpreted in terms of former surface climate. An extensive 2-year monitoring of the modern cave system shows that isotopic ?18O of the cave drips is equal to the mean annual ?18O of the local precipitation (- 7.2‰); the ?13C a product of soil organic matter, whereas Mg and Sr concentration is determined and multiple entrances, although mean temperature is equal to that above the cave (7.2°C). Cave air pCO2 is similarly variable. A newly sampled stalagmite is analysed together with SU967 a sample previously documented within the literature. Te lamina thickness chronology duplicates that of SU967. High-resolution records of ?13C and ?18O are shown together with high and low-resolution Mg and Sr acquired using a variety of new and well-established techniques. Inter and intra-stalagmite variation in ?18O, Sr and Mg is good with excellent replication demonstrated. Te ?13C is more variable between samples. Statisti- primarily by the dolomitic bedrock. Te complex karst cally signifcant co-variations are found between these hydrology modifes the fnal drip composition through stalagmite records, instrumental records and climate re-diferential transport, storage and mixing, whilst cave air constructions. Finally T/P and summer temperature are temperature is seasonally variable due to links with the reconstructed producing a multi-proxy record of climate external atmosphere provided through the cave stream for Nw Scotland. HISTORICAL BIOGEOGRAPHy OF THE GENUS MESONISCUS CARL, 1906 Andrei GIURGINCA1, Cristian-Mihai MUNTEANU2 1“Emil Racovita” Institute of Speleology, 13 Septembrie Road, No. 13, 050711, Bucharest, Romania; e-mail: sankao2@yahoo.com 2“Emil Racovita” Institute of Speleology, 13 Septembrie Road, No. 13, 050711, Bucharest, Romania; e-mail: criscarst@yahoo.com Te genus Mesoniscus is a morphologically well-defned group, clearly apart from the other groups of the Onisci-dea, with only two species - mesoniscus alpicola (Heller, 1858) and mesoniscus graniger (Frivaldsky, 1865) - and a distribution strictly limited to the Alpino-Carpathic Chain and, as such, particularly interesting from a bio-geographical point of view. Te spreading of the genus Mesoniscus allows us to notice that the two species are clearly spatially separated: mesoniscus alpicola is found exclusively in the Alps; mesoniscus graniger is spread in the whole Carpathic Chain, from the Northern and the Romanian Carpathians, South-Danube Carpathians, the Dinarids and the Julian Alps. Tere are two hypotheses regarding the origin of the species mesoniscus graniger: a Northern Carpathian origin / a Bohemian one (implying a southward spread- ing along the Carpathic Chain to the Dinarids and the Julian Alps) or, of contrary, an Illyric origin, followed by a northward migration from the Dinarids and the South-Danube Carpathians, through the Romanian Carpathians, up to the Northern Carpathians. Te Northern Carpathian origin hypothesis of the species mesoniscus graniger is argued by an earlier and longer connection between the Alps and the Carpathians and also by a later connection between the Dinarids and the Carpathians. But recurring land bridges between the Alps and the Dinarian-Pelagonian-Anatolian landmass, anterior to the connection Alps-Bohemian Massif-Carpathians, suggest and argue for an Illyric origin, also sustained by the subsequent isolation of the Dinarids from the Alps and the Carpathians. 218 TIME in KARST – 2007 ABSTRACTS IMPACTS OF ACIDIC PEAT BOG DRAINAGE ON HOLOCENE KARST DEVELOPMENT IN SOUTHEAST ALASKA, USA Melissa HENDRICKSON1 Department of Geology and Geography, western Kentucky University, 1906 College Heights Blvd, Bowling Green 42101, Kentucky, USA; e-mail: Melissa.hendrickson@wku.edu Te Tongass National Forest of Southeast Alaska, USA covers nearly 6.9 million hectares of mountainous of-shore archipelago with extensive mature temperate rain forests. Because of the accretionary terrane geologic setting, the geology there is extremely complex and heterogeneous, and includes numerous blocks of limestone that have been intensively karstifed. Tese extensive areas of carbonate bedrock are focused mainly on the northern portion of Prince of wales Island. Te last glacial activity in the area occurred with the wisconsinan (Marine Isotope Stage 2) glacial advance during the Pleistocene epoch. Tis approach strongly infuenced the karst landscape. Te development of muskeg peatlands has occurred in poor drainage areas where compacted glacial sediments and silts have been deposited over the bedrock below. Te decomposition of the Sphagnum mosses leads to highly acidic waters with pH as low as 2.4. Te measurement of continuous water chemistry at a muskeg input location and the down gradient karst resurgence found that the carbonate karst system acts as a bufer for the highly acidic muskeg waters. Over the gradient of the system, the pH increases from 3.89 to 7.22 and the predicted and measured dissolution rates drop from the insurgence to the resurgence of the system. Tese organic acids from the muskeg waters at the insurgence contribute to the highest recorded dissolution rates for natural karst systems. Depending upon the model used to calculate dissolution, rates ranged from 0.09 cm/yr to 2.5 cm/yr of wall retreat. Te karst resurgence does not differ signifcantly from other karst springs that do not have highly acidic inputs. As such, the acidic muskeg waters are rapidly depleted upon entering the karst system and do not propagate very far down gradient. Tis is supported by the presence of pits located where the muskeg waters run onto the carbonate areas. Tese pits have formed since the last glaciation which is backed up by both the dissolution rates and by the geomorphology in the area. TIME IN KARST: A BIOSTRATIGRAPHIC PERSPECTIVE Ivan HORÁČEK1 Department of Zoology, Charles University, Viničná 7, CZ 128 44 Praha, Czech Republic; e-mail: horacek@natur.cuni.cz Almost 150 years ago Louis Aggassiz demonstrated that there are only three essential methods by which the natural sciences can operate: comparative, ontogenetic and palaeontologic. Major diferences between them are in the way in which the time component of phenomena is considered. Te aspect of time is excluded from the scope of the comparative method: an act of comparison as well as its results are the mathematic structures open to further comparisons and testing. Tey are instant sources of formally objective nomological-deductive hypotheses largerly invariant to contextual changes. Te ontogenetic method operates with the phenomena whose temporality can be eywitnessed by our personal presence and immediately expressed in terms of a smooth continuous time scale, corresponding to exposure of time in common life. Te outputs of the ontogenetic method are the historical-narrative hypotheses whose reliability is entirely de- termined by the reliability of particular observer and the observation techniques. Te paleontological method is the way to treat the phenomena for which a temporality is an essential trait of them but stays far beyond the scope of the ontogenetic method. Almost all the phenomena within the scope of this symposium fall in the domain of the palaeontological method and our comprehension of them is thus potentially biased by essential inconsistency of palaeontological method. Tese phenomena can be well mutually compared but their temporality can only be derived of the assumption that the diference revealed by an act of comparison is an instant function of time. Of course, the fossil record, including that relevant for study of karst phenomena, is almost never continuous. On contrary, it is fragmentary in more respects and its density exponentially decreases with time distance. Moreover, as pointed out by Eldredge TIME in KARST – 2007 219 ABSTRACTS and Gould (1972), in large time scales, the fossil record exhibits the punctuated equilibria pattern: alternation of sudden large-scale rearrangements with long periods of stasis. In other words, the time scale generated by fossil record is in no way smoth and continuous but discrete and non-linear. Exactly the same is apparently valid for dynamics of karst development and the record that is available for its study. Te major problem of the paleontologic method and dating karst phenomena lies in absence of a direct feedback control over reliability of the empirical ordinance relations between outputs of comparative analysis and time. Of course, these relations can be biased in many ways and the proposed time datum is never exact but a mere estimate which confdence limits should be specif-cally discussed in every single case. Te methodological syllogism ofen applied in order to prevent possible dating bias is „count from top stratigraphy“: choosing the upper confdence limit , least distant from now, as a real datum. Traditional application of that technique in „safe“ dating is seemingly the more pertinent the less complete the record is and less robust its dating power is. Tis may lead to extensive underestimation of the real time span of the phenomena being studied. In dating of karst phenomena the respective bias is even more serious because what is available for study is not the true karstifcation events and/or the processes producing them but in most instances merely the epiphenomena of their past incidence (infllings of underground cavities, speleothemes etc.). Any data obtained from karst infllings and/or any other karst phenomenon is thus necessarily „a possible minimum age“ , i.e. the upper most confdence limit of karstifcation stage. Regardless of various instrumental techniques, bio-stratigraphy remains to be the most signifcant source of dating particularly for time slices of distant past. Of course, just from them, the fragmentarity of fossil record is greatly pronounced, in general, and any efort of biostratigraphic dating must hence be performed in full respect to this fact. Te procedure of biostratigraphic dating includes the following steps: (i) a careful comparative analysis undertaken both with the morphometric characteristics of all items composing the respective record and with taxonomic and structural composition of the sample. Te reliability of the result clearly depends not only on extent of the data obtained from the record under study but even more importantly on the quality and extent of the data taken in comparison (both recent and fossil), (ii) transcription of the specifcities of the record revealed by step (i) into terms of its time distance from well dated records. As the rules of the respective transcriptions are essentially quite specifc for each particular taxon and even each trait under study, the procedure (ii) actually results in a large set of diferent data with greatly variegated con-fdence limits. Te next step should hence include (iii) a comparison of them and application of the techniques discriminating their actal meaning and producing the consensus date. An instant summary of previous comparisons is provided in form of a biostratigraphic system: a set of ordinance rules regarding the phenomena repeatedly revealed in multiple previous analyses. with a formal biostratigraphic system, a practical performance of the step (iii) is essentially simplifed namely in that it reduces amount of the comparative efort to be performed onto answering few questions put by defnitions of particular units of the respective stratigraphical system. Such an approach works quite well and efectivelly if the fossil record in study is rich and reliable in respect to representation of particular index fossils and, at the same time, of course, if the biostratigraphic system applied is actually responding to the purpose. Tis means that its units must be sufciently well defned and balanced with respect to local and temporal variations in the criteria discriminating them. Te fner the time scale on which the biostratigraphic system operates the lesser is its reliability beyond the geographic limits of its type area. Simply said, the absolute reliability is not granted for any biostratigraphic system and its dating power is the smaller the less representative is the actual fossil record. Unfortunately, this is the typical case in the study paleokarst phenomena. Ofen we obtain only few poor fragments belonging to the taxa not representing real index fossils. Such cases requires application careful case-specifc analyses by a well trained palaeontologist, including ad hoc reconsiderations of confdence limits for any possible date that would come in account. It rests upon the karst scientists, of course, to claim such a challenging approach from the palaeontologists cooperating in the study and to expect from them a detailed accounts of the possible confdence limits for the fnal date they propose. with respecting the above rules, the biostratigraphic approach will undoubtedly continue to play its essential role among the tools by which the information on time in karst is gained. REFERENCES Eldredge, N., Gould, S.J., 1972: Punctuated equilibria: An alternative to phyletic gradualism. pp. 82-115 In: Schopf, T.J.M. (ed.): Models in paleobiology. Freeman, Cooper and Co., San Francisco. 220 TIME in KARST – 2007 ABSTRACTS BIOGEOGRAPHy OF STyGOBIOTIC CyCLOPOIDS (CRUSTACEA: COPEPODA) FROM BALKAN PENINSULA FOCUSED ON ACANTHOCyCLOPS KIEFERI LINEAGE Sanda IEPURE1 & Tanja PIPAN2 Institutul de Speleologie »Emil Racoviţă«, Clinicilor 400006, Cluj Napoca, Romania; e-mail: siepure@hasdeu.ubbcluj.ro Karst Research Institute ZRC SAZU, Titov trg 2, 6230 Postojna, Slovenia; e-mail: pipan@zrc-sazu.si Evolution of subterranean fauna in Balkan Peninsula has mainly been governed by the geographical patterns of region and its geological history. Te Balkan region boasts high level of endemism, particularly in caves, which are well known as Pleistocene glacial refugia. we studied the biogeography of subterranean Cyclopida (Crustacea: Copepoda) of the Balkans on a regional scale. By far the richest sub-region is the Dinaric Mountains, which form the western part of the peninsula (defned by the political borders of the states from former yugoslavia), followed by the eastern (Bulgaria and S-SE Romania) and the southern Balkans (Greece and Aegean Islands). For the stygobiotic cyclopoids, the average observed number of species per country is 12.7 and the total diversity is 69. Tus, ?-diversity accounts for 18.4% of the regional diversity and ß-diversity contributed for 81.6% to regional diversity. Te Cyclopinae fauna is comprised of Nearctic-derived epigean forms (i.e. Acanthocyclops, diacyclops and Speocyclops) of which the most diversifed genus is diacyclops with 27 taxa known. Te faunistic connection between western and central Europe is supported by the species widespread in Mediterranean region (e.g. Spain and southern France) as well as cosmopolitan stygobi-onts found throughout Europe. Bray-Curtis similarity coefcients indicate that the Transylvanian Plateau (Nw Romania) is closely related to western Balkans (similarity level 41.38%) than to that of northern part of Bulgaria (30.3%). Te Acanthocyclops kieferi lineage was analyzed in detail. Tirteen of 17 species in the lineage are endemic to the Balkans. we hypothesize that these 13 species have a common ancestor and arose as a result of vicariant events in the Miocene. Tese resulted in their isolation in subterranean waters. AGE CONSTRAINTS FOR KARST FORMS AND PROCESSES IN APULIA (SOUTHERN ITALy) Vincenzo IURILLI1,2, Giuseppe MASTRONUZZI2, Giovanni PALMENTOLA2, Paolo SANSO3 & Gianluca SELLERI3 Dottorato in Geomorfologia e Dinamica Ambientale, Universita degli Studi di Bari, via Orabona 4, 70125 Bari - Italia; e-mail: e.iurilli@geo.uniba.it Dipartimento di Geologia e Geofsica, Universita degli Studi di Bari, via Orabona 4, 70125 Bari, Italia. Dipartimento di Scienza dei Materiali, Universita degli Studi di Lecce, via per Arnesano, 73100 Lecce, Italia. Apulia region, in the south-eastern part of Italy, has three main karst reliefs, named Gargano, Murge and Salento; they show in outcrop a wide part of the apulian foreland (Ricchetti et al., 1992). Teir morphological history started in the upper Cretaceous (Luperto Sinni et al., 1991), followed on in the Tertiary age, and developed its major karst forms during the quaternary (Neboit, 1975; Grassi et al., 1982). Recently, diferent kind of researches have been collecting data in Gargano (Sauro, 2000; Caldara & Palmen-tola, 1993) as in Murge plateau (Sauro, 1991; Bruno et al., 1995; Castiglioni & Sauro, 2000; Palmentola & Iurilli, 2002) and in Salento peninsula (Mastronuzzi & Sanso, 1991; 2001). Tese recent works, with the help of studies still in progress, are giving further contributions for reconstructing the evolution of karst. For instance, in TIME in KARST – 2007 221 ABSTRACTS hypogean geomorphology and speleological research, karst morphosequences (Dramis & Bisci, 1998) are used, relating cave forms with deposits, some of which could be dated (Iurilli et al., 2005). On the other hand, surface geomorphology can relate karst forms with the local geological setting giving further constraints to apulian karst history (Marsico & Selleri, 2005). REFERENCES Bruno, G., Del Gaudio, V. , Mascia, U. & Ruina, G., 1995: Numerical Analysis of morphology in relation to coastline variations and karstic phenomena in the southeastern murge (Apulia, Italy). Geomorphology, 12, 313-322. Caldara, M. & Palmentola, G., 1993: Lineamenti geomor-fologici del Gargano con particolare riferimento al carsismo. Bonifca VIII (3), 43-52. Castiglioni, B. & Sauro,U., 2000: Large collapse dolines in Puglia (Southern Italy): the cases of “dolina Poz-zatina” in the Gargano plateau and of “Puli” in the murge. Acta Carsologica 29/2, 16, p. 83-93. Dramis, F. & Bisci, C., 1998: Cartografa geomorfologica, p. 105. Pitagora Editore, Bologna. Grassi, D., Romanazzi, L., Salvemini, A. & Spilotro, G., 1982: Grado di evoluzione e ciclicita del fenomeno carsico in Puglia in rapporto all’evoluzione tettonica. Geol. Appl. e Idrogeol., vol. xVII (2), Bari, 55-73. Iurilli, V. , Mastronuzzi, G., Palmentola, G. & Selleri, G., 2005: Indizi di tettonica recente in cavita carsiche della murgia meridionale (Puglia). Conv. “Mon-tagne e pianure – recenti sviluppi della ricerca in geografa fsica e geomorfologia” AIGEO, Padova 15-17/2/2005, Preprints, 120-121. Luperto Sinni, E., Reina, A. & Santarcangelo, R., 1991: Il ruolo della tettonica nel processo di carsifcazione: l’esempio dei giacimenti di bauxite di Spinazzola (murge baresi, Puglia). Proc. Int. Conference on Environmental Changes in Karst Areas. I.G.U. – U.I.S. – Italy 15-27 sept. 1991. quaderni del Dipartimento di Geografa n. 13, 1991 – Universita di Padova, pp. 399-404. Marsico, A. & Selleri, G., 2005: Il paesaggio carsico del-la Puglia meridionale. Conv. “Montagne e pianure – recenti sviluppi della ricerca in geografa fsica e geomorfologia” AIGEO, Padova 15-17/2/2005, Preprints, 135-136. Mastronuzzi, G. & Sanso P. , 1991: Cenni sul paesaggio carsico della penisola salentina. Itinerari Speleologici s.II n.5, Castellana Grotte (Ba)., ???? Mastronuzzi, G. & Sanso P. , 2001: Pleistocene sea-level changes, sapping processes and development of valley networks in the Apulia region (southern Italy). Geo-morphology, 46 (2002), p. 19-34. Neboit, R., 1975: Plateaux et collines de Lucanie orientale et des Pouilles. Etude morphologique. Libr. Honore Champion, Paris, 715 p. Palmentola, G. & Iurilli, V. (coll.), 2002: Il carsismo pug-liese, problemi e prospettive di ricerca. Grotte e Din-torni, riv. d. Museo delle Grotte di Castellana (Ba), n. 4, p. 203-220. Ricchetti, G., Ciaranf, N., Luperto Sinni, E., Mongelli, F. & Pieri, P.,1992: Geodinamica ed evoluzione sedi-mentaria e tettonica dell’avampaese apulo. Mem. Soc. Geol. It., 41 (1988), pp. 57-82. Sauro, U. ,1991: A polygonal karst in Alte murge (Puglia, Southern Italy). Zeitschrif für Geomorf. N. 35 (juni 1991) pp. 207-223. Sauro, U., 2000: Coastal speleogenesis and collapsing by emptying of karst breccia-pipes on the marine clifs of the Gargano peninsula (Apulia, Italy). Acta Carso-logica 29/2, 16, p. 185-193. 222 TIME in KARST – 2007 ABSTRACTS “AGE OF KARST IN GLACIATED TERRAIN”, wITH ExAMPLES FROM NORwAy AND SVALBARD Stein-Erik LAURITZEN1 Department of Geosciences, University of Bergen, Allegt. 41, Bergen 5007, Norway; e-mail: stein.lauritzen@geo.uib.no Karstifcation of the Caledonian metacarbonates in Scandinavia and Svalbard developed in pace with a landscape that was heavily eroded during the quaternary glacia-tions. On the Norwegian mainland, nappe tectonics led to the formation of stripe karst, and regional metamorphosis efectively destroyed most primary stratigraphic and fabric structures which normally guide cave inception processes. Instead, we are lef with coarsely crystalline marbles that are almost as impermeable as granites. Karstifcation and speleogenesis is therefore dictated by late tectonic fracturing in the brittle regime and by chemical contrasts at lithological contacts. Karstifcation occurred both during ice cover (subglacial speleogenesis) and in ice-free periods. During stadials, water supply was dictated by the thermal conditions within the icesheets. Due to the chemistry and immense water supply in the ice-contact environment, enlargement of pre-existing caves (speleogenesis sensu lato) was very efcient, whilst the formation of a proto-cave from a fracture (speleogenesis sensu stricto) is slower than in a non-glacial situation. Te present-day conditions on Svalbard may serve as a model for how caves developed on the Norwegian mainland during the quaternary. Most relict caves in the present landscape may be explained by both interglacial and subglacial evolution phases, but a small number of very large passages may have survived since the Tertiary. THREE-DIMENSIONAL ARCHITECTURE AND ASSOCIATED STRUCTURES wITHIN A LOwER ORDOVICIAN ELLENBURGER COALESCED, COLLAPSED PALEOCAVE SySTEM Robert. G. LOUCKS1 Bureau of Economic Geology, John A. and Katherine G. Jackson School of Geosciences, Te University of Texas at Austin, University Station Box x, Austin, Texas 78713-8924, USA; e-mail: bob.loucks@beg.utexas.edu Te three-dimensional, interwell-scale architecture of a Lower Ordovician Ellenburger coalesced, col-lapsed-paleocave system was constructed by integrating ground-penetrating radar (GPR), shallow core, and outcrop data. Te data were collected near Marble Falls in central Texas over an ~800- × ~1,000-m area. Integration of rock facies from core descriptions with GPR-refec-tion response identifed several paleocave facies that can be deciphered and mapped using GPR data alone: (1) continuous refections image undisturbed strata; (2) relatively continuous refections (tens of meters or more), characterized by faults and folds, image disturbed strata; and (3) chaotic refections having little to no perceptible continuity image heterogeneous, collapsed, cave-related facies that cannot be individually resolved using GPR data. Tese latter facies include highly disturbed strata, coarse-clast chaotic breccia, fne-clast chaotic breccia, and sediment fll. Te three-dimensional architecture of the coalesced, collapsed-paleocave system, according to core and GPR data, indicates that trends of brecciated bodies are as much as 350 m wide, 1,000+ m long, and tens of meters high. Tese brecciated bodies are coalesced, collapsed paleocaverns. Between the brecciated bodies are areas of disturbed and undisturbed host rock that are jointly as much as 200 m wide. As a cave system is buried, many structural features form by mechanical compaction, such as folds, sags, and faults. Te folds and sags measure from a few meters to several hundreds of meters in width. Collapse-related faults are numerous and can have several meters of throw. Most are normal faults, but some reverse faults also occur. TIME in KARST – 2007 223 ABSTRACTS RESERVOIR-MODEL ANALOGS AND PORE-NETwORK SUMMARy FOR ELLENBURGER COALESCED, COLLAPSED-PALEOCAVE SySTEMS Robert. G. LOUCKS1 Bureau of Economic Geology, John A. and Katherine G. Jackson School of Geosciences, Te University of Texas at Austin, University Station Box x, Austin, Texas 78713-8924, USA; e-mail: bob.loucks@beg.utexas.edu Paleocave systems are not easy to describe, and no simple description can be applied to these reservoir types. Edges of the productive karsted reservoirs are generally structurally controlled, extent and magnitude of porosity are difcult to defne, and pore networks evolve with depth. Collapsed-paleocave reservoirs commonly display an internal rectilinear pattern in which trends of porous breccias (chaotic and crackle breccias) are separated by tighter rock. Te pattern probably refects penecontem-poraneous-karst regional fracture patterns. Te breccias may be several thousands of meters across, kilometers long, and 100+ meters thick. Tese scales are larger than individual caves, indicating the collapse and coalescing of cave systems that formed at composite unconformities. A detailed description of a paleocave system in central Texas can be used as an analog for understanding In spite of its supposedly protection from major climatic changes, the subterranean domain in northern Europe seems to have been badly afected by the quaternary glacial activity during the Pleistocene, resulting in a impoverished fauna. In Belgium, two complementary processes (“dispersal” and “refugial”), of unknown relative explanatory value, have probably played a role in shaping the composition of post-glacial groundwater fauna. Faunal and ecological characterizations of the Belgian groundwater biodiversity were carried out in order to assess in what extent the present-day stygobiotic fauna can be attributed to one of these processes. A total of 202 sampling sites were selected in four hydrogeographic basins within the catchment basin of breccia (reservoir) distribution and reservoir heterogeneity. Te three main facies are undisturbed host rock, disturbed host rock (crackle brecciated), and collapsed cave passages (chaotic breccias). Te brecciated reservoir zones are separated by tighter, nonbrecciated zones. A complication in understanding paleocave reservoirs is that the pore network evolves from a megapore system near the surface to a crackle-breccia-dominated pore system with deep burial. Delineation of reservoir burial history, therefore, helps us comprehend the pore network present. Te reservoir may be fairly well connected because of the large amount of fracturing, and strong heterogeneity of reservoir quality should be expected. Fortunately, many Ellenburger reservoirs are dolomitized, and the dolomite promotes preservation of pores into the very deep subsurface (>7,000 m). the Meuse River. Sites were equally divided among the saturated and unsaturated zones of fractured aquifers (karst) and within the hyporheic and the phreatic zones of porous aquifers. Selected environmental parameters were gathered in parallel (17 variables). More than 140 species were recorded inhabiting Belgian groundwater environment, with representatives of the Amphipoda, Cladocera, Copepoda, Hydrachnidia, Isopoda, Oligochaeta, Ostracoda, Mollusca, Syncarida and Nematoda. Tirty stygobiont species were identifed. To date, the total number of stygobiotic species recorded in Belgium is 41, of which 10 species were new to the Belgian fauna. Te number of occurrences for stygobiotic species was always low; and 40% of sampled sites had no stygobi- BIODIVERSITy OF BELGIAN GROUNDwATERS AND CHARACTERIZATION OF THEIR STyGOBIOTIC FAUNA FROM A HISTORICAL AND ECOLOGICAL PERSPECTIVE Patrick MARTIN1, Claude De BROyER1, Frank FIERS1, Georges MICHEL2, Rose SABLON1 & Karel wOUTERS1 Royal Belgian Institute of Natural Sciences, 29 rue Vautier, B-1000 Bruxelles, Belgium; e-mail: Patrick.Martin@naturalsciences.be Commission wallonne d’Etude et de Protection des Sites souterrains (CwEPSS), 20 avenue Gilbert, B-1050 Bruxelles, Belgium. 224 TIME in KARST – 2007 ABSTRACTS onts. A few species were exclusive to one zone although no statistically signifcant diferences were observable in species richness between the diferent stratifcation levels considered (basin, aquifer-type, and hydrological zone). It appeared justifed to consider the stygobiotic fauna of Belgium as species-poor and mostly constituted of widely distributed species, with broad ecological tol- 1 Royal Belgian Institute of Natural Sciences, Brussels, Belgium Niphargus is a widespread stygobiont amphipod genus, of which six species are known to occur in Belgium. Tese species are: N. schellenbergi, N. fontanus, N. kochi-anus dimorphopus, N. virei, N. aquilex and N. pachypus. (Te animals can be caught in wells, caves, sources and resurgences.) Belgian Niphargus populations live close to the north or north-western border of the species’ distributions. Peripheral populations ofen show a decrease in genetic diversity relative to central populations, from which they may also be more or less isolated. In addition, they may be subject to diferent selection pressures, possibly reinforcing genetic diferentiation and in some cases even speciation. Terefore, we want to investigate to what extent the Belgian Niphargidae are genetically comparable and/or diferentiated from populations elsewhere in Europe. Tis information is required to assess the conservation value of the Belgian populations and to uncover eventual cryptic diversity. we started this research with a mtDNA analysis of Belgian N. schellenbergi, N. fontanus and N. kochianus dimorphopus. Individual DNA was extracted with the qIAamp DNA Mini kit (qiagen), using the legs from one side of a specimen, so that the remainder of the body could still be used for morphological study. Several sets of primers were tested for the mitochondrial gene fragments cytochrome oxidase c subunit I (COI), cyto-chrome B (cytB), 12S and 16S. Te COI primers (LCO 1490: 5’-GGTCAACAAATCATAAAGATATT GG-3’, HCO 2198: 5’-TAAACTTCAGGGTGACCAAAAAAT-CA-3’) yielded successful amplifcations in the three species. Te standard primers for cytB (UCytB 151F: 5’-TGT GGRGCNACyGTwATyACTAA -3’ and UCytB 270R: 5-AGGAARTAyCAyTCNGG yTG-3’) only worked for N. schellenbergi and N. kochianus dimorphopus, and the 12S primer pair (kindly provided by P. Trontelj) only erance. Tis corresponds to the view of a post-glacial colonisation of the walloon karst by eurytopic species dispersed from refugia south of the ice and permafrost (Dispersal hypothesis). Te nearly absence of endemic species suggests that the scenario of an ancient fauna having survived in local refugia (Refugial hypothesis) remains marginal. yielded good amplifcation products in N. schellenbergi. Hence more primers will have to be tested. Amplifed DNA fragments were purifed using a GFx DNA and Gel Band Purifcation Kit (Amersham Biosiences) or the wizard SV Gel and PCR Clean-up System (Promega), according to the manufacturers’ protocols. Sequencing was performed using an automated DNA sequencer (Applied Biosystems 3130 Genetic Analyzer). In this way we screened (up till now) 150 individuals for COI (approximately 650 bp long), 60 individuals for cytB (app. 350 bp long) and 140 individuals for 12S (app. 510 bp long). For COI, we found 12 haplotypes within N. schel-lenbergi (18 populations, sequence divergence (p-dis-tance) within the species 0,14%), 2 haplotypes within N. kochianus dimorphopus (one population, sequence divergence 6,3 %) and one haplotype within N. fontanus (one population). Te divergence found in the N. kochianus dimorphopus population (sequence divergence 6,3 %) is high compared to the other species studied and to some other subterranean amphipods, especially since we only analysed one population. Finston et al., (Mol. Ecol. 16 (2007): 355-365) found sequence divergences for COI of less than 1,8 % between subterranean amphipod populations (genus Pilbarus) caught within a tributary. Te three taxa that we studied were well-diferentiated by sequence divergences of 31,1 % between N. schellenbergi and N. fontanus, 37,5 % between N. schellenbergi and N. kochianus dimorphopus, and 22,9 % between N. fontanus and N. kochianus dimorphopus. For 12S, we found 9 haplotypes within N. schellen-bergi (18 populations) with an overall mean sequence divergence of 1,2 %. So the sequence divergence for 12S is tenfold higher than the sequence divergence for COI, within the same populations. MTDNA ANALySIS IN THE GROUNDwATER AMPHIPOD NIPHARGUS FROM THE MEUSE VALLEy (BELGIUM) S. MATTHIJS1, T. BACKELJAU1 & Frank FIERS1 TIME in KARST – 2007 225 ABSTRACTS while amplifying cytB, we ofen obtained a pattern that looked like a ‘double’ sequence within a single individual. Sometimes one of both was readable and so it turned out that there were indeed two kinds of sequences, with a sequence divergence between the two groups of 15,1 % (sequence divergence within the groups are 1,1 % and 0,6 %). whether these two kinds of sequences represent a case of heteroplasmy or involve nuclear cytB copies (numts) remains to be investigated. qUANTIFyING PALEOCAVE COLLAPSE FROM 3D SEISMIC DATA: ExAMPLES FROM THE PALEOZOIC SECTION IN THE NORTHERN FORT wORTH BASIN, TExAS Angela McDONNELL1, Robert G. LOUCKS1 & Tim DOOLEy2 Bureau of Economic Geology, John A. and Katherine G. Jackson School of Geosciences, Te University of Texas at Austin, University Station Box x, Austin, Texas 78713-8924, USA; e-mail: angela.mcdonnell@beg.utexas.edu Bureau of Economic Geology, Jackson School of Geosciences, Austin, Tx, USA; e-mail: tim.dooley@beg.utexas.edu Te Lower Ordovician Ellenburger Group of western and central Texas displays excellent paleokarst, both in outcrop and subcrop (e.g., cores), refecting multiple exposure events. In the northern Fort worth Basin, circular collapse structures representing suprastratal deformation above these collapsed-paleocave systems are well-known drilling hazards for tight-shale gas wells (Lower Carboniferous Barnett Formation). It is, consequently, important to understand their geometries in detail; the collapse features are therefore imaged using 3D seismic data. Tis study provides quantitative analysis of the timing of collapse and geometrical characteristics of these largely concentric structures. Concentric faults extend vertically 760 to 1,060 m from the Lower Ordovician Ellenburger Group. Te largest structures remained active longer, in-fuencing deposition of a younger, Upper Carboniferous fuvial system. A defning parameter of these features is the upward-narrowing trough of the sag zone. A rectilinear paleokarst system is identifed, with coalesced passage and cavern systems aligning along Nw-SE and NE-Sw trends. Tis orthogonal trend refects the rectilinear fracture and joint system within the Ellenburger Group that was preferentially exploited by karst weathering processes. Collapse appears to have been incremental, with signifcant collapse occurring afer about 300 m of overburden had been deposited. A set of criteria are outlined for quantifying the geometries of collapse-generated sag structures identifed on seismic data (as opposed to sags generated by other processes—e.g. pull-apart basins). Tis template provides valuable information for defning seismically resolvable karst-collapse features worldwide. STyGOBIOTIC ISOPODA: A TOOL TO ASSESS wORLD CHANGES Giuseppe MESSANA1, Mariella BARATTI1 Istituto per lo Studio degli Ecosistemi del CNR Via Madonna del Piano, 10, 50019 Sesto F.no, Firenze, Italy. Te fauna of the subterranean aquatic realm is rich, widely distributed and certainly underestimated, in number and, moreover, in the potential to provide a good insights into the evolution of each taxon, of the environment they colonize and of the paleogeographic events that shaped them. within this fauna the Isopoda constitute an important portion. Tey are present in all the subterranean environments so far explored. Several diferent taxa of various families do colonize diferent areas with several species whose number is mostly related to the research efort. Te populations are composed mostly by endemic species, a common aspect of the subterranean habitats, due in general to the long time since each of them have colonised these ecosystems. Some of these taxa are ancient relicts, while others are more recent colonisers. we will concentrate on few of these. Relict taxa: 1. Calabozoidea, South American, restricted to few localities in Venezuela, and Brazil. 226 TIME in KARST – 2007 ABSTRACTS 2. Phreatoicoidea, colonizing Australia, in a great variety of freshwater habitats, wells and caves in India, and caves and riverbeds in South Africa. 3. Stenasellidae widely distributed, they colonize diferent subterranean ecosystems from the Far East to America, accounting for an old conquest of the subterranean aquatic habitats. Ancient and recent colonisers: the Mediterranean genera of the Cirolanidae with strict relationships to genera of other regions. Te family has diferent levels of endemic areas, is widely distributed with several stenoendemic genera. THE RECONSTRUCTION OF THE PLIOCENE PALEOENVIRONMENT OF THE UNROOFED CAVES wITH FOSSIL REMAINS OF MARIFUGIA CAVATICA FROM THE ČRNOTIČE qUARRy, PODGORSKI KRAS, w SLOVENIA Andrej MIHEVC1 Karst Research Institute, ZRC SAZU, Titov trg 2, Sl – 6230 Postojna; e-mail:Andrej.Mihevc@guest.arnes.si From the morphology of the caves, cave sediments and remnants of fossil cave animals and other fossils reconstructions of the paleoenvironments in which they were formed can be made. However it is difcult not to mix the evidences from diferent environmental settings which followed during the whole speleogenesis of a cave. Here we present the reconstruction of the early Pliocene paleogeography of the Podgora karst and caves with fossil remains of cave animal marifugia cavatica. Te unroofed caves and other caves discovered in Črnotiče quarry on the edge of the Podgora karst show a very complex speleogenesis and proved to be very important for the understanding of the evolution of SE part of the Kras plateau. Fill in the largest of fossil horizontal unroofed caves was roughly dated back to 4.2–5.2 Ma (Bosak et.al., 1999). Later work in the quarry reveal a new part, a side passage of the same cave in which the cave animal serpulid marifugia cavatica tubes attached to the walls were preserved (Mihevc 2000, Mihevc et. al., 2001, 2002). Sediments were dated by paleomagnetic methods to 2.5–3.6 Ma (Bosak et. al., 2004). Most important part of the cave is phreatic or epi-phreatic, more than 17 m high and 4-8 m wide water passage. Tere are large scallops on the walls of the passage with attached marifugia cavatica tubes indicating rather slow fow and stable environment. It was located in low position with small gradient and was already a result of a long speleogenesis. water with discharge several hundred litres per second fowing trough the cave system was coming from a sinking river, ponors of the river were close to the preserved part of the cave. At least part of catchment area of the sinking river was on Eocene fysch marls. Tere were fsh living in a river and foods were washing in drowned small mammals. Filling of the passage which preserved the sessile tubes of marifugia followed was probably a result of a fast change, maybe collapse in other part of the cave. Later gradient both outside and in karst enlarge, the fne sediments in the upper part of the profle were eroded and replaced by coarser sediments. In that part of the cave wall no serpulid tubes were found. water lef the cave and in a dry passage more than 7 m thick fowstone layer was deposited. Today the surface is at 420 m a.s.l. and is cutting the fowstone fll of the passage exposing it on the surface as a typical unroofed cave. Nearest water table caves where Marifugia cavatica still lives are karst springs about 2 km away in elevation about 50 m a.s.l. REFERENCES: Bosák, P. , Mihevc, A., Pruner, P. , Melka, K., Venhodová, D. & Langrová, A., 1999: Cave fll in the Črnotiče quarry, Sw Slovenia: Palaeomagnetic, mineralogi-cal and geochemical study. – Acta Carsologica, 28/2, 2, 15-39, Ljubljana. Bosák, P. Mihevc, A. & Pruner, P. , 2004: Geomorphologi-cal evolution of the Podgorski Karst, Sw Slovenia: contribution of magnetostratigraphic research of the Črnotiče II site with Marifugia sp. Acta carsol., 2004, letn. 33, št. 1, str. 175-204, ilustr. Mihevc, A., 2000: Fosilne cevke iz brezstrope jame – verjetno najstarejši ostanki jamskega cevkarja Mari-fugia (Annelida: Polychaeta). – Acta Carsologica, 29/2, 261-270, Ljubljana. TIME in KARST – 2007 227 ABSTRACTS Mihevc, A. Sket, B., Pruner, P. & Bosák, P., 2001: Fossil remains of a cave tube worm (Polychaeta: Serpuli-dae) in an ancient cave in Slovenia. Proc., 13th International Speleological Congress, Brasilia, 2001, str. 20-24, Brasilia. Mihevc, A., Bosak, P. , Pruner, P. & Vokal, B., 2002: Fossil remains of the cave animal Marifugia cavatica in the unroofed cave in the Černotiče quarry, w Slovenia. Geologija, 45, 2, str. 471-474. FORMATION AND EVOLUTION OF THE PERI-MEDITERRANEAN KARST DURING THE MESSINIAN SALINITy CRISIS AND THE PLIOCENE: EVIDENCE FROM THE ARDECHE VALLEy, SOUTHERN FRANCE Ludovic MOCOCHAIN1 CEREGE, Europole Méditerranéen de l Arbois, 13 545 Aix-en-Provence cedex 4, France. During the Messinian–Pliocene eustatic cycle, the Mediterranean Sea was characterized by a short lived (5.95– 5.32 Ma) sealevel fall, which attained -1500 m in some areas. Te study of benchmark levels permits the chronology and dynamics of this event to be established. In the Rhône’s middle valley, our investigations allow a new interpretation for the genesis of the Ardeche endokarst. A fall in base-level was responsible for both the incision of the so-called Messinian canyons as well as a deep karst development. Karst systems were formed in association with the Messinian canyons of the Ardeche and Rhône Rivers. During the fooding of the Mediterranean Basin (5.32 Ma), these karst systems were flled by water and plugged by sedimentary inflling of the rias. Tis mechanism pushed groundwater backward through the karst system, which in turn formed diagnostic “chimney-shafs”. Tese pathways were geometrically connected to the position of the Pliocene benchmark levels. Consequently, the Messinian Salinity Crisis was responsible for two karst responses. Te frst was concomitant with the crisis itself and corresponds to the formation of a karst system. Te second followed the Messinian Salinity Crisis and corresponds to the per ascensum adaptation of this karst system in Vauclusian karsts by the formation of “chimneyshafs”. THE CARBONATE HyDROGEOCHEMISTRy OF THE KRKA RIVER, CROATIA Natalie NAHILL1 Department of Earth and Environmental Sciences, University of Pennsylvania, 240 S. 33rd Street, Philadelphia, USA; e-mail: nahilln@sas.unpenn.edu Te Krka River is incised into a broad plateau underlain by folded carbonate rocks along the Dalmatian coast of Croatia. Te Krka River drains a surface watershed of ~2,610 km2, and receives fow contributions from a network of interconnected subsurface watersheds of unknown confguration. At eight separate locations, tufa structures have developed across the river perpendicular to fow, and have grown to a size that interferes with that fow. Te purpose of this investigation is to characterize the hydrogeochemical condition of the Krka River to defne the geometry of subsurface components of the watershed, the chemistry of the water delivered by those individual components, and the efect of subsurface contributions on the process and location of tufa precipitation. To determine the mechanisms responsible for tufa precipitation, we investigated the carbonate chemistry, temperature, pH, and topography of the Krka. we determined the carbonate content by in situ titration every ~2.0 km along the 72-km length of the Krka. Values of pH ranged from 7.4 to 8.4; the concentration of total carbonate ranged from 300 to 500 ppm. we collected 32 samples of modern tufa from sites close to the titration stations, 228 TIME in KARST – 2007 ABSTRACTS and analyzed those samples by x-Ray Difraction. within limits of analytical precision, all samples were identical in mineralogy, with calcite the predominant mineral. Aragonite, magnesium calcite, magnesite, stenonite, and quartz were identifed at much smaller percentages. In tufa-precipitating river systems with well characterized hydrologic inputs, such as Havasu Creek, AZ, pH and carbonate content follow a systematic trend of rising pH and decreasing carbonate content with distance from the source. In the Krka system, that simple pattern is not Djara Cave is one of the few caves in the western Desert of Egypt containing speleothems. It is located close to the Farafra–oasis in western Egypt and was frst visited by the German GERHARD ROHLFS on 24th December 1873, who found an extensive speleothem (stalactites, stalagmites and fowstones) development in this cave, part of them are covered by sand Te cave was then visited and studied by many research groups (geological, geomor-phological and archeological) due to its importance as a paleoclimatic indicator for the desert. Te cave is located on an Eocene plateau to the west of the Nile valley, which consists of highly fractured carbonates of the Naqb Formation. Today, the western Desert of Egypt, where the cave is located, is part of the hyperarid Eastern Sahara and belongs to the subtropical desert climate zone. High temperatures, low humidity and strong winds cause high potential evaporation rates in excess of 5000mm per year. In contrast, the interpolated annual precipitation sum is less than 5mm with sparse rain on only 1–5 days per year on average. Te cave temperature is 23°C year- round and appears to stay very close to the external mean temperature. realized; sequential subsurface inputs apparently alter the hydrochemistry and, ultimately, the process of tufa precipitation. within the Krka system, we identifed three short segments of the river through which systematic rise of pH and decline in total carbonate content document distance downstream from a discrete subsurface source. Tose segments are separated by reaches in which variation of those chemical parameters follows no clear pattern, suggesting that subsurface inputs to the Krka vary both in space and time. U/T datings of speleothems by ?–spectrometry yielded ages ranging from 140 ± 15.9 kyr to 283 ± 56 kyr (Brook et al., 2002). One TIMS date yielded 201.05 ±2.1 kyr. A number of samples was beyond the U/T dating limit (~ 500 kyr for TIMS dating, ~ 350 kyr for ?–spec-trometry) (Holzkämper, 2004). Isotopically depleted values, which were also measured within earlier investigations, suggest that enhanced African summer monsoon and westerly circulation during the winter months advected precipitation to the western Desert enabling speleothems to form in the recently hyper–arid region. A number of covered hearths in the close vicinity of the cave have been found by previous expeditions, some of which were dated with the radiocarbon method. Tese and additional datings of ostrich egg shells yielded ages between 9.7 kyr to 5.5 kyr, indicating that wetter conditions prevailed in the area during this time interval. So Djara Cave is a great source of information for the paleo-climate of the Sahara, showing the change from wet, dry to hyperarid conditions through time. DJARA CAVE (wESTERN DESERT OF EGyPT) AS A PALEOENVIRONMENTAL, AND PALEOCLIMATIC INDICATOR Tamer NASSAR1 Geology Department, Cairo University, Faculty of Science, 12613 Giza-Cairo, Egypt; e-mail: nassar@ndara.com TIME in KARST – 2007 229 ABSTRACTS MORPHOMETRIC ANALySIS OF THE LEPTODIRINAE BEETLES wITH RESPECT TO THEIR HyPOTHETIC SUCCESSIVE HISTORICAL COLONISATION OF SUBTERRANEAN HABITATS IN SLOVENIA Tone NOVAK1, Slavko POLAK2 & Franc JANŽEKOVIČ1 1 Department of Biology, University of Maribor, Koroška 160, SI-2000 MARIBOR, Slovenia; e-mails: tone.novak@uni-mb.si; franc.janzekovic@uni-mb.si 2 Notranjska Museum Postojna, Ljubljanska 10, 6230 Postojna, Slovenia; e-mail: slavko.polak@guest.arnes.si Te animal morphology is, besides recent molecular investigations, still an unavoidable classical approach in the analysis of taxonomic groups in space and time. Te understanding of the adaptatiogenesis for living in hypo-gean habitats can contribute to the research of the karst evolution. In Slovenia, within the subfamily Leptodirinae (Cholevidae = Leiodidae), three morphological groups have been established representing unique lineages with respect to their adaptations to preferred habitats. Tree species of the epigean lineage live in moos and litter, while two troglobitic lineages have been recognized among 34 species and 38 subspecies. Most of them belong to the bathyscioid type sensu Jeannel with an ovoid body and relatively robust short appendages resembling adaptations in edaphic beetles. Te second troglobitic leptodirine lin- eage represents the ultra evolved leptodiroid type sensu Jeannel with specially transformed, mostly enlarged body and gracile, long thin appendages. Te species of the frst troglobitic group live more or less in all karstic regions of Slovenia till Karavanke/Karawanken Mts., known to be the border of the continuous distribution of troglobites in the Central Europe. Te species of the second group are limited to the Dinaric Karst. In this study, the mor-phometric comparisons between selected species of the three groups as well as the outgroup of Cholevinae were carried out to fnd statistical diferences between them. Te authors suggest that the detailed analysis of the two evolutionary successive troglobitic lineages can support the discussion about the karst evolution. THE TIME wHEN NEANDERTHALS VISITED ROMANIAN CAVES Bogdan P. ONAC1,2, Iosif VIEHMANN1, Joyce LUNDBERG3, Stein-Erik LAURITZEN4 & Chris STRINGER5 “Emil Racoviţă” Institute of Speleology, Clinicilor 5, 400006 Cluj / quaternary Research Group, “Babeş-Bolyai” University, Kogal-niceanu 1, 400084 Cluj, Romania; e-mail: bonac@bioge.ubbcluj.ro Department of Geology, University of South Florida, 4202 E. Fowler Ave. SCA 528 Tampa, FL 33620 USA. Department of Geography and Environmental Studies, Carleton University, Ontario K1S 5B6, Canada. Department of Geosciences, University of Bergen, Allegt. 41, Bergen 5007, Norway Department of Palaeontology, Te Natural History Museum, London, Sw7 5BD, Great Britain. ABSTRACT Humanoid footprints in the fossil record are rare. A survey of the literature reveals only two well documented, dated cases. Te one, from ~325 ka in Italy (Mietto et al., 2003), represents a very early, pre-Neanderthal human. Te other, from ~117 ka in Africa (Gore, 1997), is likely a homo sapiens sapiens print. Here we report the frst clearly homo sapiens neanderthalensis footprint. It was found in Vârtop Cave, Romania. Te person stepped into calcareous mud that later hardened. Te 22 cm long print suggests a body height of ~1.46 m; a gap of 1.6 cm marks the separation of big toe and second toe. Te date of the footprint is constrained by the date of the deposition of the mud (~97 ka, dated by U-T isochron method) and the date on the base of a nearby stalagmite on top of the mud (~64 ka). Tus the Vârtop Cave person lived in Romania sometime between 97 and 64 ka, long before the earliest known homo sapiens sapiens remains in Europe (~35-30 ka) (Carciumaru & Anghelinu, 2000; Trinkaus et al., 2003). To our knowledge, this is the frst discovered and dated homo sapiens neanderthalensis footprint. 230 TIME in KARST – 2007 ABSTRACTS REFERENCES Ford, D.C. & williams, P.w., 1989: Karst Geomorphol-ogy and Hydrology. London. Unwin Hyman. 601 pp. (Times New Roman, 11, single spaced, fully jus-tifed) Cârciumaru, M. & Anghelinu, M., 2000: Te Carpathian Mousterian and the transition from middle to upper Palaeolithic in southern Romania. in Neanderthals and modern humans -- discussing the transition: Central and Eastern Europe from 50,000 - 30,000 B. P. (J. Orschiedt, G.-C. weniger, Eds.) Mettmann, Neanderthal Museum, pp. 190-195. Gore, R., 1997. Te dawn of humans: tracking the frst of our kind. National Geographic magazine. 192: 92-99 Mietto, P. , Avanzini, M. & Rolandi, G., 2003. Human fottprints in Pleistocene volcanic ash. Nature. 422: 133. Trinkaus, E., Moldovan, O., Milota, S., Bilgar, A., Sarcina, L., Athreya, S., Bayley, S.E., Rodrigo, R., Gherase, M., Higham, T., Ramsey, C.B. & van der Plicht, J., 2003. An early modern human from the Pestera cu Oase, Romania. Proc. Natl. Acad. Sci. U.S.A. 100: 11231-11236. ISOTOPIC INVESTIGATIONS OF CAVE DRIP wATERS AND PRECIPITATION: APPLICATIONS TO HyDROGEOLOGICAL AND PALEOCLIMATE STUDIES IN FLORIDA, USA Kali PACE-GRACZyK1 Department of Geology, University of South Florida, 4202 E. Fowler Avenue, Tampa, Florida, USA; e-mail: kpacegra@mail.usf.edu Precipitation and in-cave drip waters from speleothems at three caves that transect the Gulf coast of the Florida peninsula from North to South have been collected to satisfy three research goals. 1) quantify the variability of the isotopic signal of precipitation and drip waters at each cave and isolate the primary factors that control that isotopic variability both at the site and regional scale. 2) Determine the fraction of meteoric waters that originate in the Atlantic Ocean and Gulf of Mexico at each site. 3) Calculate the recharge rates through the vadose zone of the aquifer matrix at each site. In addition to weekly water samples, instrumentation inside and outside each cave monitors hourly data on temperature and relative humidity; acoustic loggers acquire data on drip rates beneath active speleothems attached to unfractured blocks of the aquifer matrix. Te relationship between the iso-topic composition (?18O and ?2H) of cave waters and meteoric sources is critical for reconstructing paleoclimates using speleothem data. Tis study will serve as a frst of its kind in the southeastern US where the impacts of con- founding variables such as temperature, vapor source, storm frequency and intensity, distance from shore, and soil/water/rock interactions are poorly understood. Determining precipitation sources will shed light on how the climate of the Florida Peninsula is afected by regional weather patterns such as the Atlantic Multi-decadal Oscillation, El Nino, and the Intertropical Convergence Zone. For example, rainfall from tropical systems in the summer show a depleted ?18O signal when compared to frontal precipitation events during the winter. Calculating recharge rates will help us better understand the transport of meteoric waters through the epikarst and into the underlying Floridan Aquifer. All three caves formed in the eogenetic Ocala limestone where matrix permeabilities are high, 10-11 m2 to 10-13.8 m2. Te lag times between precipitation events and changes in drip rate from the aquifer matrix are short at all three caves, on the order of days to weeks, compared to caves in the telogenetic, low-permeability limestones of the mid-continent. TIME in KARST – 2007 231 ABSTRACTS SPELEO-FAUNA OF THE MONTI SIMBRUINI REGIONAL NATURAL PARK Federica PAPI1 1 Via delle Cave 42, 00181 Rome, Italy. In the region of Lazio there live 50% of total Italian species due to its environmental variety. A sampling and a study about speleo-fauna has been done for two years (starting in 2002) in 30 diferent caves (24 of then biospaeleologically unknown) in the Central Appennino in the Monti Simbruini Natural Park. During this research probably new species belong of Orders of Araneae and Coleoptera are to be found. Te Coleoptera is a Curculionidae Otiorhynchus (Lixorhyn-chus) similar in morphological characters to O. gianquin-toi (F. Solari, 1936) (det. G. Osella); the Spider is a Cen-tromerus considered, from Dr. P. Pantini, similar to C. puddui (known only in a Sardegna’s cave). Tis sample needs a comparison with samples of C. cinctus (known in Corsica and Algeria). Specialization level of cave fauna of Monti Sim-bruini and Volsci, in particular Monti Lepini (the better biospeleologically known mountains complex in central Italy). Te Specialization Index (Sbordoni et al. 1977) has been calculeted. Tis Index uses the report between the number of troglobiontes and the total of eutrglobious species (the total of troglobiontes and trglophilus). Tis index appears independent both of cave morphology and development. From preliminary observations it is seen that tro-globiontes of Monti Simbruini are fewer than the tro-globiontes of Lepini. Moreover in Monti Lepini endemic species are more and with areas of endemicity larger than the Simbruini’s species. Te specialization Index is higher for Monti Lepini fauna than for Monti Simbruini. Te diferent kinds of cave fauna in the two mountains complex could be due to two diferent reasons, probably together: 1) Te geological kinematics of the zone: Te mountain chain of Volsci emerged 1.5 million years before than Monti Simbruini, and so cave fauna of Monti Simbruini could be in a more in arrears evolutive level than Volsci one. Te diferent climatic conditions: In Volsci is usual to fnd uncovered karst with mediterranean clima. Tis condition simplify the separation between fauna of humid environment, typical of caves, and external fauna; so allopatric division is favoured and speciation too. GENESIS AND AGE OF ICE ACCUMULATIONS IN CAVES Aurel PERSOIU1 Department of Geology, Univeristy of South Florida, 4202 E. Fowler Avenue, SCA 528, 33620 Tampa, Florida, USA; e-mail: persoiu@hasdeu.ubbcluj.ro & apersoiu@mail.usf.edu Perennial ice exists in a signifcant number of caves in Europe and elsewhere in the world. Ice in caves accumulates through complex freezing and melting processes, which eventually lead to the development of laminated deposits, with clear ice layers alternating with impurity-rich ones. Tis deposition style ofers good perspectives for the use of ice accumulations in caves as paleoclimatic archives, as they contain a large number of climatic proxies: O, H and C stable isotopes, pollen, beetle remains etc. However, in order to be able to make best use of these proxies, a good age model is needed for the hosting ice. Using present-day ice formation as a model, we propose a genetic and evolutionary model for the formation of ice deposits in selected caves from Apuseni Mountains, Romania. In order to achieve our goal, we frstly developed a model for the present-day accumulation of ice, using glaciological, meteorological and geochemical (tritium dating) analyses. we found that the ice has a well structured response to external climatic forcing, its melting or building-up being controlled mainly by variations in precipitation (i.e., availability of dripping water), while temperature is acting as a second order controller (e.g., dripping water can determine the accumulation of ice, if air temperature is bellow O°C, or can lead to ice melting, if air temperature is above 0°C). Air temperature itself has a limited infuence on the dynamics of ice, due to peculiar morphology of caves, witch do not allow warm air penetration inside the caves during summer (cold air traps). winter air is entering the cave at a high rate, but in 232 TIME in KARST – 2007 ABSTRACTS the absence of dripping water has a reduced infuence on the dynamics of ice. Tis model, combined with detailed stratigraphic measurements of the existing ice bodies enabled us to reconstruct the past dynamics of the ice. By plotting the data obtained against 14C values, we were able to build a good age-depth scale for the last 1000 years, as basis for further paleoclimatic reconstructions, using ice in caves as sources of information PALEOCLIMATIC SIGNIFICANCE OF THE MAMMAL CAVE FOSSILS Alexandru PETCULESCU1 Department of Geospeleology and Paleontology, Institut of Speleology, 13. Septembrie 1-15, Bucharest, Romania; e-mail: alexpet@gmail.com Te importance of the fossils from caves sediments is obviously from the paleoclimatic interpretations point of view. Te paleontological studies above the mammals from the Upper Pleistocene from Romania reveal a distinct particularity in comparison with other mammals associations from the rest of the Europe. we will reconstruct the paleoenvironment from the last glacial cycle (wurm) on the basis of the mammal cave fossils discovered in few cave from Romania. Te research made in carstic deposits from the last glacial cycle, is based on systematic survey in few of the most important paleontological sites. Te remains accumulated here have diferent origins: bones abandoned here by the primitive hunters, scavengers and from natural causes (death) when the cave was used like shelter (Ursus spelaeus and Crocuta spelaea). For the small mammals the main remains sources was the existence, next to the entrance of the caves, of the carnivorous bird’s nests. Te remains was accumulated in chronological positions and represent a mirror for the mammalian associations from the outside, especially for the small mammals. OPTICALLy STIMULATED LUMINESCENCE ANALySIS OF THE SPELEOTHEMS IN CROATIA Srđan PICHLER1 & Dalibor PAAR2 Geofzika d.d., Savska cesta 64, Zagreb, Croatia; e-mail: spichler@irb.hr Department of Physics, Faculty of Science and Mathematics, Bijenička 31, p.p. 331, 1002 Zagreb, Croatia; e-mail: dpaar@phy.hr Te luminescence dating covers a range of analytical methods that can be applied over diferent time periods in diferent minerals and in diferent environmental conditions. Te optical stimulation allows very small luminescence signals to be detected. In most cases the minerals analyzed with this method were quartz and feldspars, because they are able to store energy in a crystal structure. Tis energy is deposited mainly by ionising radiation from environment and by cosmic radiation. Te analysis of this energy is the base of the luminescence dating method. In our work we have analyzed the speleothems mainly consist of calcite or aragonite. Te sources for the optical stimulated luminescence were the laser coherent light and the non coherent light emitting diodes (LED) sources. TIME in KARST – 2007 233 ABSTRACTS KARST AS A MODEL SySTEM TO ExAMINE TERRESTRIAL MICROBIAL BIOGEOGRAPHy PATTERNS THROUGH TIME Megan PORTER1 Department of Sciences, University of Maryland Baltimore County, Baltimore, Maryland 21250, USA; e-mail: porter@umbc.edu Te evolutionary mechanisms that govern the distribution of microbes on Earth are poorly understood. Continental plate motion and geologic processes have changed the distribution of terrestrial and marine life throughout Earth’s history. Confnement to certain locations has resulted in speciation events and even endemism. For microscopic life, however, the concept that geographic isolation plays a role in microbial species diversity is controversial. Because microbes are small in size, are meta-bolically tenacious, and are overwhelmingly abundant on Earth, microbes may be distributed everywhere and therefore may not be afected by geography. Many micro-bial biogeography studies have been conducted to test for microbial species endemism, but these studies have been conducted in habitats with high dispersion (e.g., aquatic, and specifcally oceanic, habitats). Tere have been rela- tively few studies to characterize microbes living in terrestrial habitats where it is assumed that there has been limited dispersal due to the geographic and hydrostrati-graphic barriers. Hypothetically, similar to what has been observed for organisms living on islands, microbial communities in the terrestrial subsurface, and specifcally in caves and karst settings, would have had less opportunity to exchange genetic information because of barriers to gene fow and exchange; therefore, speciation events would be higher. Considering the ubiquity of karst worldwide, understanding the types of microbial communities in cave and karst systems, as well as determining what geologic processes may control microbial species distribution and diversity, are critical aspects to preserving the integrity of karst ecosystem and to predicting changes in ecosystem that may occur following disturbances. SPELEOGENESIS IN 15 DAyS Mitja PRELOVŠEK1 Karst Research Institute, SI-6230 Postojna, Slovenia; e-mail: mitja.prelovsek@zrc-sazu.si Although corrosion is very important from the speleoge-netical point of view feld measurements of corrosion are still very scarce. while the number of long-lasting speleo-genetical interpretations and evolution modeling of karst aquifers still grows, actual values of corrosion are not yet well established. One should ask, if these interpretations of cave development ft into the feld acquired data. On the basis of this question we started to measure the intensity of corrosion at several diferent underground streams, which fow in the epiphreatic zone. Measurements were done with limestone tablets as their preparation is easy and accuracy of the method (up to 0.0001 mm) is precise enough to detect measurable changes in one year. Due to signifcant improvement of the methodology, which allows to measure corrosion in streams with velocities over 3 m/s, in the period from 2005 to 2006 corrosion at 68 diferent sites was measured. Te average measured intensity was –0.02 mm/a. Gathered values are in accordance with measurements accomplished by dr. A. Mihevc, who measured corrosion on cave walls with micro-erosion meter. Measurements gave us insight into the activity of caves from the view of corrosion – some cave systems were recognized as corrosively active, while others have fossil cave rocky relief. On 29 locations the intensity of corrosion is greater than 0.005 mm/a. Tis means that at such places corrosion can be measured in 15-day intervals. Tis is so short period of time, that we can recognize the infuence of individual hydrological event, i.e. at low, medium or high water conditions. If the intensity of corrosion was almost the same through the year, in 6 separate cave systems we could measure it daily. On the basis of annual measurements we decided to place 78 limestone tablets in the following caves: Križna jama, Nova Križna jama, Lekinka, Pivka jama, Tkalca jama and Jelovička jama. In these caves we are measuring spatial and temporal diferences of corrosion in 15-, 30- and 50-day intervals. Although the caves have quite diferent recent speleogenesis our results show that in the period of low water levels corrosion does not occur at any experimental sites. Unexpectedly, at some places important fowstone deposition was observed. It seems that the 234 TIME in KARST – 2007 ABSTRACTS majority of corrosion is done during medium high water levels, which are relatively frequent. Te highest corrosion efcacy is certainly characteristic for high waters, but such hydrological events are quite rare. In the ponor cave Lekinka at medium high water level infuence of the 1 UMR 5125 du CNRS, Université Claude Bernard Lyon-1, Bât. France. Te largest part of the quercy regional area, Sw France, is made of a thick Jurassic limestone platform, which contains a dense net of karstic voids. Tese voids are flled with clay deposits, within which phosphatic crusts developed. Intense mining of this ore, in the last 19th century quarter, revealed the fossiliferous richness of the fllings, mostly with terrestrial vertebrate remains. Teir study, renewed for the last forty years, produced a corpus of fossil data and their bearing into various felds: knowledge of groups from amphibians to mammals, evolution of lineages, biochronology, paleoecology, paleoen-vironments, and so forth. Te fossil documented period in the quercy phosphatic area now extends through 30 million years, from the early Eocene up the early Miocene. Studied mammal lineages allowed numerical dating (Escarguel et al., 1997). Te Laprade net (Tarn-et-Garonne dept.) is mostly an unroofed cave, this resulting from the strong plio-quaternary erosion of the platform. Since mining, infll-ing red clay still occur in restricted parts. At the N-w end, this clay yielded a vertebrate fauna, the age of which is Mid-Eocene, MP 14 of the European mammal scale (Sudre et al., 1990). Its numerical dating is – 41.35 M.a. ± 0.426. In the Nw underground gallery, the E wall at entrance consists of chaotic superposition of large limestone blocks, some of metric size. Tis results from a vault and/or wall local collapse. Te inflling red clay between the blocks simply extends the bearing-fauna one outside. distance from the cave entrance was very well observed, namely the corrosion intensity in 65 m water course fell from 0.003 mm in 15 days to 0.002 mm in the same period. Geode, 2 rue Raphaël Dubois, La Doua, F-69622, Villeurbanne, Its age postdates the collapse, without signifcant time between collapse and flling. Similar collapses ofen occur in active karst systems, most of time without clear cause. However, the Laprade flling dating sent back us to another paleokarstic place in quercy, the Prajoux one (Lot dept.). Tere, few mammal remains predate a fault movement, at ca – 41.5 M.a. (Astruc et al., 2000). Te closeness of the two datings is best understood as relating to the same intense tectogenic and seismic period of the pyre-nean orogen, efects of which being distally preserved. Te Laprade collapse together with the Prajoux fault support such interpretation. Here, the dating method relies on evolutionary data of mammal lineages. while other, chemical dating methods, will soon be settled and allow checking. REFERENCES Astruc, J. G., Escarguel, G., Marandat, B., Simon-Coin-çon, R. & Sigé, B., 2000: Geodinamica Acta, 13: 271-280. Escarguel, G., Marandat, B. & Legendre, S., 1997: Actes du Congres BiochroM’97, Mém. EPHE, Inst. Montpellier, 21: 443-460, Sudre, J., Sigé, B., Remy, J.A.., Marandat, B., Hartenberger, J.-L., Godinot, M. & J.-y. Crochet, 1990: Palaeover-tebrata 20 (1): 1-32. A MAMMAL-DATED MID-EOCENE CAVE COLLAPSE IN THE qUERCy PALEOKARST, Sw FRANCE Bernard SIGé1, J. G. ASTRUC, G. ESCARGUEL, S. LEGENDRE, T. PéLISSIé, R. SIMON-COINÇON TIME in KARST – 2007 235 ABSTRACTS VARIATIONS IN SPELEOTHEM TRACE ELEMENTS AND ?13C IN CENTRAL FLORIDA: POTENTIAL FOR PALEOCLIMATE RECONSTRUCTION Limaris R. SOTO1 1 Department of Geology, University of South Floriday, 4202 East Fowler Avenue, SCA 528, 33620 Tampa, Florida, USA; e-mail: isoto@cas.usf.edu A stalagmite collected from Briar Cave in central Florida, provided the frst Late Holocene Paleoclimatic record for Florida using speleothems. Te stalagmite was analyzed for trace elements Sr and Mg and for carbon isotopes. Termal ionization mass spectrometry of uranium-thorium isotopes indicate the speleothem was precipitated during the last 4200 years. when Sr/Ca and Mg/Ca time series data is combined with Uranium series dating techniques, the age of paleoclimate variations may be deduced. Using Sr/Ca and Mg/Ca ratios to learn past climatic data is under debate. Many academics believe temperature, efective rainfall, recharge rates, and vegetation patterns can be found from trace elements in spe-leothems, others argue that only the speleothem growth rate may be obtained from trace elements. Results showed that Sr concentrations are negatively correlated with ?13C, a relationship we inferred to record changes in soil productivity. Magnesium concentrations were not found to be signifcantly correlated with hemispheric temperature, however results suggest the residence time of percolation waters maybe the controlling factor. Coeval changes in the Sr content and ?13C signals, as induced by soil productivity, can only be explained by changing precipitation above the cave. Both proxies record a 170-180 year solar cycle that has also been found in the Gulf of Mexico marine records. Consequently, this result provides evidence of an extra-terrestrially driven modulator of precipitation in central Florida. DETERMINATION OF PAST FLOw REGIMES FROM SCALLOPS OF BEyyAyLA CAVE (BILECIK – TURKEy) By USING BOOTSTRAP HyPOTHESIS TESTING METHOD M. Evren SOyLU1, Haydar DEMIRHAN2 1 International Research and Application Center for Karst water Resources (UKAM), Hacettepe University, 06532 Beytepe, Ankara, Turkey. 2 Department of Statistics, Hacettepe University, 06530 Beytepe, Ankara, Turkey. Caves provide natural archives of records of the palaeo-hydrologic conditions. Scallops are among the corrosion – morphologies that may provide such records. In this paper scallops at the walls of the passage of the Beyyayla Cave in Turkey were measured and statistically analysed to infer about the past hydrologic conditions. Te cave is located within a dissected – relict karst, and it exhibits poly phase passage morphology. Relicts of scallops on the passage walls vary in shape and size. Data collected in the cave were analyzed by the bootstrap hypothesis test to determine the number of modes in the multimodal histogram, which is obtained over the collected data set. Te results of analyses revealed that the cave had been afected by at least two fow regimes. 236 URANIUM-SERIES DATING OF SPELEOTHEMS: ESTABLISHING TIME AND FORCING OF CARBONATE DEPOSITION IN KARST SySTEMS Christoph SPÖTL1 1 Institut für Geologie und Paläontologie, Leopold-Franzens-Universität Innsbruck, Austria; e-mail: christoph.spoetl@uibk.ac.at Over the last few years cave science has seen an strong in- as from the general public. Tis development is largely crease in interest from other academic disciplines as well due to the growing awareness of the importance of karst TIME in KARST – 2007 ABSTRACTS systems as groundwater resources in many regions of the world as well as the recognition that within these subsurface environments information about past climate and hydrology is reliably recorded in carbonate deposits. Paleoclimate scientists currently are very keen to study speleothems and some regard them even as valuable as ice cores from Polar regions. Although most speleothem studies are currently targeted toward providing high-quality records of climate and environmental change outside cave networks, these results are also undoubtedly of great signifcance for a better understanding of long-term dynamics within karst systems. Examples include the frequency of cave fooding recorded as detrital-rich layers in stalagmites, drought periods as indicated by narrowly spaced spe-leothem laminae and/or microhiatus, overgrowths of marine organisms on speleothems in near-coastal caves during sea-level highstands, and the occurrence of intermittent speleothem deposition in cold-climate caves prone to freezing. Tese studies therefore provide critical data for assessing the role of climate forcing of karst hydrological processes on timescales ranging from annual to millennial. Given the inherently complex nature of karst systems and the fact that these systems evolve over time, results obtained from individual speleothem samples, however, will require thorough validation to arrive at robust interpretations. KARST DEVELOPMENT AT THE RACISZyN AREA (KRAKÓw – wIELUŃ UPLAND, S POLAND) – PRELIMINARy RESULTS. Grzegorz SUJKA1 Institut of Geological Sciences, Polish Academy of Sciences, quaternary Geology Department, Twarda 51/55, 00-818 warszawa, Poland; e-mail: gsujka@twarda.pan.pl In September 2005 in a quarry near Raciszyn (Kraków-wieluń Upland, S Poland) we discovered a new small cave. Te entrance was placed on the bottom of the lowest exploitation level. Both geographical and geological position of the cave allows us to expect new data on evolution and development of karst in this part of Poland. Te cave is a part of larger geological structure i.e. a large slit that runs along the whole quarry. During the exploration about 20 samples of speleothems from the cave and other parts of quarry have been collected to the further T/U analysis. we have found at least three generations of speleothems separated by erosion surfaces and clastic sediments (mostly clays). Tis indicates complicated history of the cave and radical changes of hydrogelogical conditions during Pleistocene in this area. Te oldest discovered speleothems are older than 350 ka, and probably older then 1.2 Ma, and exceeds the method’s range. Next generation of speleothems developed between 233 and 118 ka. Te youngest speleothems were Holocene age. A COMPARATIVE REVIEw OF FOREST MANAGEMENT HISTORy IN SOME HUNGARIAN KARST AREAS Eszter TANACS1 Department of Climatology and Landscape Ecology, University of Szeged, Hungary; e-mail: nadragulya@geo.u-szeged.hu Vegetation is one of the karstecological factors. As such it directly afects microclimate and soil and thus indirectly the whole system. Due to their geographical position, the potential vegetation of Hungarian karsts is mixed-stand deciduous forest so forest management methods in the past and present are a key issue in today’s karst surface development. On the other hand karst is a special environment with characteristic surface features and special water balance, which needs special considerations in management. Tis study aims to reveal the similarities and diferences of past forest management in the difer-ent karst regions of Hungary. TIME in KARST – 2007 237 ABSTRACTS THE KARSTIC REJUVENATION PROCESSES ON THE ALADAG MOUNTAIN (EASTERN OF CENTRAL TAURUS MOUNTAINS, TURKEy) SINCE THE LAST GLACIAL MAxIMUM Koray TÖRK1, Alexander KLIMCHOUK2, Lütf NAZIK1, Serdar BAyARI3 General Directorate of Mineral Research and Exploration of Turkey (MTA), Turkey; e-mail: cave@mta.gov.tr Ukrainian Institute of Speleology and Karstology, Ukraine; e-mail: klim@speleogenesis.info Geological Engineering Department of University of Hacettepe, Turkey; e-mail: serdar@hacettepe.edu.tr Te karstic processes had been ruled in Aladag region with the continental conditions for the beginning of the Paleocene-Eocene era in the area. Te trace of this term and the later karstifcation as Later Miocene can be seen on the paleokarstic ore deposits at Aladag region. Te morphologic development surfaces can be seen since the Upper Miocene on the area. Te Aladag Karst Plateau that bounded with Zamanti River and Ecemiş strike slipped fault zone is also an uplif area on the border of the Central and Eastern Taurus Belts. Te plateau consists of two main diferent parts which are as high altitude karst area and lower fuviokarstic area. Te high altitude karst area has been intensively afected from the quater- nary glacial processes. Te most of glacial were as valley type but also traces of ice cap glacial have been found on the mountain. Te resignation of the last glacial was continued till the 7000 year B P. Te landforms of epikarst were scraped and plugged through the last glacial period while the glacial were using the mature karstic basin. Tis processes afected the karst hydrological system and morphological structure as hanged karst springs, un-walled shafs, unroofed caves and polycyclic dolines on the area. Te rapid uplif of the plateau according to close region and the glacial fow also caused to deep incision of the Aladag Mountain. KARST IN THE CORDISBURGO REGION, MINAS GERAIS STATE, BRAZIL Luiz Eduardo PANISSET TRAVASSOS1 & Heinz C. KOHLER2 1 Scholarship provided by CAPES, Master’s Degree Student in the Post-Graduation Program on Geography – Spatial Analysis/PUC Minas - Pontifcia Universidade Catolica de Minas Gerais, Av. Itau 505 - Predio Emaus. - Belo Horizonte, Brasil; e-mail addresses: luizpanisset@uol.com.br & luizpanisset@gmail.com 2 Environmental Studies Laboratory Coordinator – Post-Graduation Program on Geography – Spatial Analysis/PUC Minas, Brasil; e-mail addresses: heinzkohler@gmail.com & charleskohler@uol.com.br Te Karst in Cordisburgo is considered to be one of the most signifcant karst regions in Minas Gerais State, Brazil. It is distinguished for both the magnifcent potential of its endokarst and the important archaeological, paleontological, historical and tourist sites it ofers. Minas Gerais, the third most important industrial area in Brazil, has intensive farming and mining activities as well. According to the Instituto Brasileiro de Geografa e Estatística (2001), about 35.83% of the economically active population is employed in farming, cattle raising, planting and fshing; 40.70% in the general services; 15.07% in the industrial sector and 8,40% in small commerce sectors, which is basically primary needs-oriented. Although there are some exceptions, this scenario leads to problems with natural resources management, especially in karst areas. Cordisburgo is located at about 110 km north of Belo Horizonte, the Minas Gerais State capital city, on the Late Proterozoic metasedimentary carbonate rocks of Grupo Bambuí (600-500 Ma B.P.). Te predominant climate is typical of tropical regions: two well defned seasons, a characteristic of the Cer-rado. Te annual temperature average is 22o C and the annual rainfall average is 1230 mm. Tis particular karst area is characterized by elongated massive limestone reliefs in the E-w direction (maximum height of 1055 m), intercalated by poljés (minimum height of 715 m) with temporary lagoons. Te endokarst is developed in four main caves with horizontal development equal or superior to 1000 m (Gruta da Morena - 4620 m; Lapa Nova do Maquiné - 1312 m; Gruta do Salitre - 1098 m; Gruta do Toboga - 1000 m). Tere are also 10 other caves that are developed in limestone. Te city of Cordisburgo was frst discovered by the 17th century explorers called bandei-rantes, and later became a settlement which was ofcially 238 TIME in KARST – 2007 ABSTRACTS established by a priest in 1883. Te toponym reinforces the sacred meaning of the place: Cordis = heart; burgo = city, an allusion to the Sacred Heart of Jesus. In the 19th century, this area was studied by Peter wilhelm Lund, who found important paleontological sites there, such as the Lapa Nova do Maquiné cave (1834). It is important to notice that Lund discovered important evidences of the pre-historic South American fauna from the Pleistocene in this cave. Tis cave is also known as the frst Brazilian touristic cave. In the 20th century (1956) the area was described in famous novels by an important Brazilian writer (Guimaraes Rosa), when he wrote about the relationship between farmers and the sertanejos, the ‘cerrado cowboys’ and the landscape. Traditionally the population used the karst waters as their primary resource for agriculture, farming and domestic usage. According to COPASA, the company that provides for water distribution and sewage treatment in the county, the supply comes from a 20.0 l/s outfow well. As a result of the intensive agricultural and farming activities, the drinking water supply of the municipality may be signifcantly exploited and contaminated. Nowadays, the municipality is investing much more in tourism. Besides all this, there is another important element that points out to the importance of that area conservation: the quarry expansion towards it, which is about to become a reality. Due to these threats, the conservation of this speleological and natural patrimony is urgent. Now the region is being studied by Travassos & Kohler, who are developing a map of the karst phenomena, intended to be the basis for the geoen-vironmental compartmentation of that region, ofering important subsides for the sustainable management of that environmental scenario, especially for the susceptibility of the karst aquifer to contamination. Processes that are similar to those mentioned by Kovačič and Rav- bar (2005) take place at Cordisburgo. Te presence of a thin soil cover leads to rapid water percolation in plantations and pastures, accelerating the contamination of the aquifer. Other distant areas of non-carbonated rocks may act as allogenic recharge areas that concentrate pollutants from diferent human activities as well. Tere are no sanitary landflls to protect groundwater from the leach-ates in that area. Te only solid waste disposal process is an inappropriate site. So, it is important to establish systematic studies in the area and, according to Kranjc (2000, p.123), also invest in the education of all inhabitants, from politicians and experts to pupils who may be of infuence to the threat and the protection of karst areas and the water supply. As a starting move “the most important task is to fll the gap between the karst specialists with their knowledge and “non-karst” specialists, also highly educated, with sometimes striking ignorance of karst and karst water.” (KRANJC, 2000, p.123). REFERENCES Kohler, H.C., 1989: Geomorfologia cárstica na Regiao de Lagoa Santa-mG. Doctorate Dissertation. p.113, Sao Paulo. Kovačič, G.; Ravbar, N., 2005: Mapping of hazards to karst groundwater on the Velika Planina Plateau (Slovenia). Acta Carsologica, 34/1, p.74-85, Ljubljana. Kranjc, A., 2000: Karst water research in Slovenia. Acta Carsologica, 29/1, p.117-125, Ljubljana. Palmer, A.N., 2001: Dynamics of cave development by allogenic water. Acta Carsologica, 30/2, p.13-32, Ljubljana. Ravbar, N., 2004: Drinking water supply from karst water resources (Te example of the Kras Plateau, Sw Slovenia). Acta Carsologica, 33/1, p.73-84, Ljubljana COMPARATIVE ANALySIS OF CAVE-PLANTHOPPER RADIATIONS IN AUSTRALIA AND HAwAI’I — PRELIMINARy RESULTS (HEMIPTERA: FULGOROMORPHA: CIxIIDAE) Andreas wESSEL1, Petra ERBE2 & Hannelore HOCH Museum für Naturkunde der Humboldt-Universität zu Berlin, Department of Research, Biosystematics Group, Invalidenstrasse 43, D-10115 Berlin, Germany; e-mail: andreas.wessel@museum.hu-berlin.de Uplands Program, Faculty of Agriculture, Chiang Mai University, 50200 Chiang Mai, Tailand; petra.erbe@gmx.net Ongoing studies have revealed numerous cases of parallel evolution of cavernicolous planthoppers all over the world in tropical and subtropical caves. Given that concepts of genetic change and speciation dynamics are subject to review by natural model systems, the comparative study of these independent evolutionary lineages and radiations can provide critical information. we survey species complexes from Australian and Hawai’ian caves exhibiting diferent degrees of troglo-morphy. In Australia (qld), Solonaima and Undarana TIME in KARST – 2007 239 ABSTRACTS species have colonized old karst caves as well as young lava tubes. On the Hawai’ian Islands, the cave-dwelling Oliarus species represent at least seven independent cave colonizations on three islands of diferent age, including the Oliarus polyphemus species complex from various, partly still active volcanic systems on Hawai’i Island . Karst cave development in carbonate rock is predicated by the existence of such rock, so the earliest a cave can form is synchronous with carbonate deposition. A hierarchy of cave development conditions can be established based on the postdepositional evolution of carbonate porosity into three time-porosity stages that conform to the rock cycle as stated by Choquette and Pray (1970, p. 215): “the time of early burial as eogenetic, the time of deeper burial as mesogenetic, and the late stage associated with erosion of long-buried carbonates as telogenetic”. In dia-genetic order, from immature to mature host rock, a progression can be established: A) Eogenetic Caves - Te carbonate rock is in its environment of deposition; cave development can be presented as three progressive stages: 1) Constructional - Caves formed by deposition of a soluble rock at the instant the rock is deposited. Example: Tufa caves; the rock is the end product of earlier carbonate dissolution but the void itself not produced by dissolution. 2) Syndepositional – Caves formed by dissolution while rock deposition is occurring. Examples: Caves formed in lagoonal carbonate sands when beach sands prograde over them, or eolian calcarenites deposited across existing surface stream channels. Te diversity of external factors (e.g., ressource availability and stability, migration possibilities, macro-and microclimatic changes, predators) as well as internal factors (e.g., genetic variability, population structure and density) allows us to incorporate these into concepts of time and mode of evolutionary change in cave organisms. 3) Syngenetic – Caves formed subsequent to deposition by dissolution while rock lithifcation is occurring. Examples: Eolian calcarenites invaded by a fresh-water lens and mixing zone, or subtidal carbonates exposed to meteoric water by uplif associated with tectonics and/or glacioeustasy. B) Mesogenetic Caves – Te carbonate rock is buried, isolated from surfcial processes, and undergoing diagenetic maturation. By defnition, all karst caves developed in mesogenetic rocks are hypogenic. C) Telogenetic Caves – Te carbonate rock is diage-netically mature and exposed to the surfcial weathering environment. A fourth category, metamorphic, could address caves formed in carbonates that have been metamorphosed into marbles. Te hierarchy does not require a fxed chronology. Geologic processes operate at diferent rates in diferent areas. For example, some New Zealand Oligocene carbonates are telogenetic as a result of the vigorous tectonic burial and uplif, while rocks of similar age in Florida are still eogenetic. Te age of the rock can be misleading, what is important is the diagenetic character of the rock at the time of cave genesis. THE EARLIEST TIME OF KARST CAVE FORMATION Susan q.wHITE1, Ken GRIMES2, John E. MyLROIE3 & Joan R. MyLROIE3 Environmental Geoscience, Latrobe University, Victoria 3086, Australia; e-mail: susanqwhite@netspace.net.au Regolith Mapping, RRN 795 Morgiana Road, Hamilton, Victoria 3300, Australia; e-mail: regmap1@ozmail.com.au Department of Geosciences, Mississippi State University, Mississippi State, MS 39762 USA; e-mail: mylroie@geosci.msstate.edu 240 TIME in KARST – 2007 ABSTRACTS THE DINARIC KARST EVOLUTION – THE CAVE SHRIMPS’ MOLECULAR AND GEOLOGICAL POINT OF VIEw Valerija ZAKŠEK1, Peter TRONTELJ2 & Boris SKET2 Department of Biology, University of Ljubljana, Biotechnical Faculty, Večna pot 111, SI-1000 Ljubljana, Slovenia; e-mail: valerija.zaksek@bf.uni-lj.si Department of Biology, University of Ljubljana, Biotechnical Faculty, Večna pot 111, SI-1000 Ljubljana, Slovenia. Te cave dwelling shrimps of the genus troglocaris show a disjunct distribution in European karstic groundwa-ters: in the Dinaric Karst on the Balkan Peninsula, on the west Caucasus and in South France. Te phylogenetic relationships within the genus based on the sequences of two mitochondrial and one nuclear gene showed a monophylum with two main lineages constituted of the Dinaric and Caucasian taxa. Both lineages have geographically largely separate ranges: one is extending from Italy to Bosnia and Herzegovina and the other from Bosnia and Herzegovina and Montenegro over Caucasus. Absolute dating of phylogenetic events is possible by using calibration points from the fossil record or vicari-ant events. In the atyid shrimps, or even more ideally in the troglocaris group, points like that are not available. Terefore the molecular clock approach has been used to estimate the divergence dates among the clades. Te divergence dates were estimated using substitution rate divergences previously defned in other decapods. Te split between t. inermis from France and the surface-living dugastella valentina from Spain is about 10 Myr old, but they had common predecessor with Dinaric and Caucasian troglocaris taxa in the middle Miocene, about 1 Department of Climatology and Landscape Ecology, University of zzboray@geo.u-szeged.hu & keveibar@earth.geo.u-szeged.hu Te most common form of land use in Hungarian karsts is woodland. An integrated forest management taking into account considerations of environmental protection as well could insure the conservation of karsts in a near-natural state. Before world war II, the forests of the Bükk Mountains were owned by the state. while earlier the forests were mainly cut down to increase the area suitable for grazing, afer the war deforestation followed due to a highly increased claim for wood production. Te exploitation of forests also had an impact on 15 Myr ago. Te most unexpected is a relatively young split between Caucasian and Dinaric Troglocaris populations that was estimated at 6-11 Myr. Te Dinarides represent a complex land formed during the Terciary and quaternary. Te tectonics strongly controlled the formation of many small Neogene basins within the Dinarides which were geological characterized by fresh-water sedimentation. Te analysis of faunal characteristics which occurred in sediments in the Neogene fresh-water basins within the Dinarides showed many similarities between North Croatian Basin and Northern Bosnia region. Te higher number of deep splits within the cave shrimps in the southern Dinaric Karst might be a result of the Miocene Dinaride lake system geographically comprising parts of Croatia, Bosnia-Herzegovina and Montenegro, where troglocaris lineages are taxonomically more diverse than in the Nw part of Dinaric Karst. younger lineages within troglocaris are congruent with hydro-graphically isolated areas in the Dinaric Karst. All age estimates are rough, calibration point within the Dinaric Karst would enable us to develop a more precise dating of a cladogentic events for troglocaris and other cave taxa in the area. of Szeged, 6722 Szeged, Egyetem u. 2. Pf. 653, Hungary; e-mails: their age composition. Te foundation of the Bükk National Park in 1977 meant a turning point in the history of the area’s forests, introducing the concept of sustainable management. In this study we compared time-series of aerial images in order to follow changes in the investigation area. Tese images were taken by the military cartography in the years 1956, 1965, 1975, 1987, 1988, 2004, related to the revision of earlier maps. we defne the horizontal and vertical changes of the area such as increase the net of the CHANGE DETECTION AND TIME-SERIES FROM AERIAL-, AND SATELLITE IMAGES ON THE BUKK-PLATEU (BÜKK-FENNSÍK), HUNGARy Zoltán ZBORAy1 & Ilona BÁRÁNy-KEVEI1 TIME in KARST – 2007 241 ABSTRACTS forestry roads which is close-textured that even before, and the height of forest based on digital surface models which contain the height of the trees above the digital elevation model. Te species composition of the stands was defned by the supervised classifcation of Landsat satellite imagery. TIME RECORDED IN CAVE DEPOSITS – 10 yEARS OF PALEOMAGNETIC RESEARCH IN SLOVENIAN CAVES Nadja ZUPAN HAJNA1, Andrej MIHEVC1, Petr PRUNER2 & Pavel BOSÁK1,2 Karst Research Institute, ZRC SAZU, Titov trg 2, 6230 Postojna, Slovenia. Institute of Geology, AS CR, Rozvojová 269, 165 02 Praha 6, Czech Republic. Te palaeomagnetic and magnetostratigraphic research of karst deposits has been carried out since October 1997 in 17 caves and 2 surface sites located in diferent geological units and geographical settings of Slovenia. Totally 33 sedimentary profles and more than 2,000 samples have been sampled and processed by the standard palaeomag-netic analyses (thermal and alternating feld demagnetisation, magnetic susceptibility measurements, etc.). Afer the last massive marine transgression during Eocene, when thick pile of fysch siliciclastics deposited, in the most of the area studied, there is no evidence of younger marine sediments, even if they have been expected by some authors (e.g., Rögl 1998). Te area was thus exposed to denudation. Correlate sediments preserved in isolated basins in vicinity, belonging to Miocene Paratethys deposits, are about 6-10 Ma old (e.g., Márton et al., 2002). Cave karst deposits (both clastic and chemogenic) ofer the record of processes, which evidences have not been preserved on the surface of most of karst regions of Slovenia and can help to decipher the younger geological and tectonic history. Te start of our paleomagnetic and magnetostratigraphic research was framed by chronostratigraphic models of cave inflling compiled by Rado Gospodarič (e.g., 1981, 1985, 1988) from diferent sources and applied methods. He dated cave deposits to period up to 350 ka and only some deposits were expected to be older (ca 600-900 ka; e.g. Gospodarič 1985). But later data of T/U dating of speleothems from diferent Slovenian caves (Zupan 1991, Mihevc 2001) indicate that a lot of speleothems are older than explained by Gospodarič. Even much greater age of caves and karst surfaces suggest the studies of unroofed caves. with assumption that the caves were developed deep below surface and considering recent denudation rate these caves and sediments in them can be 1 – 5 Ma years old (Mihevc 1996, 2001). Performed paleomagnetic and magnetostratigra-phy studies, combined by numerical dating methods, mineralogical, petrological, geochemical, paleontologi-242 TIME in KARST – 2007 cal and geomorphological analyses, ofered surprisingly new time frame for cave depositional processes – they proved that most of analyzed sediments can be up to several millions of years old, including sites studied by Rado Gospodarič. Te oldest paleontologically correlated ages have been found in the Račiška pečina Cave in the Classical Karst – about 3.5 Ma. Te obtained data of correlated- and numerical-ages of cave/karst deposits supported the new trends and ideas concerning the evolution of karst surfaces especially in the region of the Classical Karst, indicated that cave deposits occurring now in diferent altitudes can be of the same age, and proved that deposits in caves, now unroofed, are very ancient. REFERENCES Gospodarič, R., 1981: Sinter generations in Classical Karst of Slovenia. Acta Carsologica, 9, 87-110, Ljubljana. Gospodarič, R., 1985: On the spelogenesis of Divaška jama and Trhlovca Cave. Acta Carsologica, 13, 5-32, Ljubljana. Gospodarič, R., 1988: Paleoclimatic record of cave sediments from Postojna Karts. Ann. Soc. geol.. Belg., 111, 91-95. Márton, E., Fodor, L., Jelen, B., Márton, P. , Rifelj, H. & Kevric R., 2002: Miocene to quaternary deformation in NE Slovenia: complex paleomagnetic and structural study. J. Geodynam., 34, 627-651. Mihevc, A., 1996: Brezstropa jama pri Povirju. Naše jame, 38, 65-75, Ljubljana. Mihevc, A., 2001: Speleogenesis of the Divača Karst. Te Publishing House of the Scientifc Research Center of SASA, 180 pp, Ljubljana. Rögl, F., 1998: Palaeogeographic Considerations for Mediterranean and Paratethys Seaways (Oligocene to Miocene). Ann. Naturhist. Mus. wien, 99 a, 279-310, wien. Zupan, N., 1991: Flowstone datations in Slovenia. Acta Carsologica, 20, 187-204, Ljubljana. AUTHOR INDEx AUTHOR INDEx Prepared by Caitlin Nay, University of Akron Organized by last name of author or co-author, and indicating starting page of article or abstract. Audra, P. . ....................................... 53 Astrug, J.G. . ................................ 235 Backeljau, T. . ................................ 225 Bárány-Kevei, I. . ................ 207, 241 Baratti, M. . .................................. 226 Bayari, S. . .................................... 238 Bini, A. . ......................................... 53 Boch, R. . ...................................... 209 Bonacci, O. . ................................ 151 Bosák, P. . ............................. 207, 242 Broyer, C. . .................................... 224 Bruno, M. . ................................... 209 Campbell, B. . .............................. 212 Constantin, S. . ............................ 210 Culver, D. . ..................................... 87 Cunningham, K. . ........................ 216 Deharveng, L. . ............................. 212 Demirhan, H. . ............................ 236 Dittmar, K. . ......................... 173, 211 Dooley, T. . ................................... 226 Dragusin, V. . ............................... 210 Dreybrodt, w. . ............................... 25 Erbe, P. . .............. 199, 213, 214, 239 Escarguel, G. . .............................. 235 Fadem, C. . ................................... 215 Feier, I. . ........................................ 216 Fiers, F. . ............................... 224, 225 Florea, L. . ..................................... 216 Forte, J. . ....................................... 217 Fuller, L. . ..................................... 218 Gabrovšek, F. . ............................ 7, 53 Geest, P. . ....................................... 211 Giurginca, A. . ............................. 218 Gorički, Š. . .................................. 183 Grimes, K. . .................................. 240 Häuselmann, P. . ..................... 53, 93 Hendrickson, M. . ....................... 219 Hobléa, F. . ..................................... 53 Hoch, H. . ............ 199, 213, 214, 239 Horáček, I. . ................................. 219 Iepure, S. . .................................... 221 Iurilli, V. . ...................................... 221 Janžekovič, F. . ............................. 230 Jeannin, P. . ..................................... 53 Klimchouk, A. . ........................... 238 Kohler, H. . .................................. 238 Kramers, J. . .................................. 209 Kunaver, J. . .................................... 53 Lascu, C. . .................................... 210 Latella, L. . ...................................... 69 Lauritzen, S. . ............... 210, 223, 230 Legendre, S. . ............................... 235 Loucks, R. . .......... 121, 223, 224, 226 Luiszer, F. . ................................... 157 Lundberg, J. . ................................ 230 Martin, P. . .................................... 224 Mastronuzzi, G. . ......................... 221 Matthijs, S. . ................................. 225 McDonnell, A. . ........................... 226 Messana, G. . ............................... 226 Michel, G. . .................................. 224 Pichler, S. . .................................... 233 Pipan, T. . ............................... 87, 221 Polak, S. . .............................. 183, 230 Porter, M. . ................... 173, 212, 234 Prelovšek, M. . ............................. 234 Pruner, P. . .................................... 242 Rajka, G. . ....................................... 77 Reid, J. . ........................................ 209 Romanov, D. . ................................ 25 Sablon, R. . .................................... 224 Sanso, P. . ...................................... 221 Sasowsky, I. . ................................ 143 Sauro, U. . ........................................ 69 Selleri, G. . .................................... 221 Sigé, B. . ........................................ 235 Simon-Coinçon, R. . ................... 235 Sket, B. . ................................ 183, 241 L. . ....................................... 236 M. . .................................... 236 Soto, Soylu, Mihevc, A.....................35, 227, 242 Spöti, C................................209, 236 Mocochain, L. . ........................... 228 Moldovan, O. . ................................ 77 Monbaron, M. . ............................. 53 Munteanu, C. . ............................. 218 Mylroie, J.E. . ................................ 240 Mylroie, J.R. . ............................... 240 Nahill, N. . ................................... 228 Nassar, T. . .................................... 229 Nazik, L. . ..................................... 238 Novak, T. . .................................... 230 Onac, B. . ....................................... 230 Osborne, R. . ................................ 133 Otoničar, B. . ................................. 101 Paar, D. . ....................................... 233 Pace-Graczyk, K. . ...................... 231 Palmentola, G. . ........................... 221 Palmer, A. . .................................... 15 Papi, F. . ........................................ 232 Pélissié, T. . ................................... 235 Perez-Losada, M. . ...................... 173 Persoiu, A. . ................................. 232 Petculescu, A. . ............................ 233 Stringer, C. . ................................. 230 Sujka, G. . ..................................... 237 Summers Engel, A. . .................... 212 Šušteršič, F. . .................................. 53 Tanacs, E. . ................................... 237 Tognini, P. . .................................... 53 Törk, K. . ...................................... 238 Trajano, E. . .................................. 191 Travassos, L. . .............................. 238 Trimmel, H. . .................................. 53 Trontelj, P. . .......................... 183, 241 Vasileva Evtimova, V. . ................ 215 Verovnik, R. . ............................... 183 Viehmann, I. . .............................. 230 wessel, A. . .......... 199, 213, 214, 239 white, S. . ..................................... 240 white, w. . ..................................... 45 wildberger, A. . ............................. 53 wouters, K. . ................................ 224 Zakšek, V. . ........................... 183, 241 Zboray, Z. . ................................... 241 Zupan Hajna, N. . ......................... 242 TIME in KARST – 2007 243 Acta carsologica 36, 1 (2007) Izdala in založila Slovenska akademija znanosti in umetnosti in Znanstvenoraziskovalni center SAZU v Ljubljani Grafčna priprava in tisk Tiskarna Lotos Postojna Naklada 800 izvodov It is a great honor but also a big responsibility to protect the greatest Slovenian treasure. We are doing it for over 180 years and we'll do our best to keep it shining and in good health for generations to come. Postojnska jama turizem d.d. ActA cArsologicA issn 0583-6050 • letnik/ Volume 36 • Številka/Number 1 • ljubljana 2007 ¦r Contents