Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , 4 ˇ zek, 1958). Discussions , A. Petculescu ´ Biogeosciences ak, 1989), which is 1 ´ ak et al., 1989; Sasowsky ˇ ˇ zek, 1958). Two contrast- ce Quarry, Classical Karst, , I. Meleg 4 ˇ cek and Bos ˇ ´ a Crnoti ˇ zek, 1988; Bos ˇ a 31, 010986 Romania Bucures¸ti, ´ a 269, 165 00 Praha 6, Czech Republic 3404 3403 , S. Constantin 3 ´ o ˇ cek and Lo ´ a ˇ cina, and a cave in , L. Mik 2 ˇ ska pe ˇ ci includes fine-grained sediments transported from the vicinity of ˇ ´ zek, 1958; Hor ak, 1989) as well as of biological and anthropological information , A. Mihevc 1 2,5 ˇ ˇ a” Institute of Speleology, Clinicilor 5, 400006 Cluj-Napoca, Romania a” Institute of Speleology, Frumoas ´ ak ˇ cek and Bos entrance facies ´ a This discussion paper is/has been under review for the journal Biogeosciences (BG). Please refer to the corresponding final paper in BG if available. Institute of Geology AS CR, v. v. i., Rozvojov Karst Research Institute, SRC SASA,European Titov Commission, trg. DG 2, Environment, Postojna, Av. Slovenia de Beaulieu 5, 1160 Auderghem,“Emil Brussels, Racovit¸ “Emil Racovit¸ The the cave by wind, water and slope processes. It represents the most valuable section Received: 18 March 2011 – Accepted:Correspondence 23 to: March 2011 O. – T. Moldovan Published: ([email protected]) 30 MarchPublished 2011 by Copernicus Publications on behalf of the European Geosciences Union. Cave sediments represent conservers(Hor of the geological and(e.g., paleoenvironmental Kukla past and Lo and Mylroie, 2004). This iswhere of correlative special sediments importance are for mostlythe the terrestric missing case (continental) (Hor of history, theor studied are karst allochthonous region. ining facies origin Cave can sediments (Kyrle, be are 1923; distinguished formed Kukla among in and cave place Lo environments in (Kukla caves, and Lo groups as compared to mostthe of study the of surface cave habitats. sediments. This may open a new direction in 1 Introduction 5 the first pre-Quaternary invertebrateate fossils climate. found in The sediments Pliocene/Pleistoceneomagnetic of age dating continental of chronologically temper- the calibrated sediments bybrate was micromammal fossils determined biostratigraphy. could by Inverte- be pale- their validated suitability for as ecological new and paleogeographic proxycaves. correlations for in They caves also the and bring study outside additional the ofregime information in cave about the sediments cave area. formation due Even andthat if to karst climatic the hydraulic number conditions of in remains caves was allow very low, a it better represents preservation evidence of fossil remains of some Five samples of clasticrelic sediments caves (Trhlovca, from Ra interiorSW cave Slovenia) facies provided taken invertebratebut in fossil sparse remains. three individuals Slovenian Most of of Cladocera them and belong insects to were Oribatida also identified. They represent Abstract 2 3 Belgium 4 and P. Bos 1 Biogeosciences Discuss., 8, 3403–3434, 2011 www.biogeosciences-discuss.net/8/3403/2011/ doi:10.5194/bgd-8-3403-2011 © Author(s) 2011. CC Attribution 3.0 License. Invertebrate fossils from cave sediments: a new proxy for pre-Quaternary paleoenvironments O. T. Moldovan 5 20 25 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | develops interior facies ´ ak et al., 2003). Such sediments 3406 3405 ˇ zek, 1958; Bottrell, 1996; Courty and Vallverdu, ´ ak et al., 1998, 2003; Sasowsky, 2007). Although ´ ak, 2002, 2008; Bos erent kinds of proxies; e.g., charcoal, sediment pollen ff ´ on et al., 1992a, b, 1995, 1997, 2005; Finlayson et al., 2008), ˇ zek, 1958 and literature herein), the scientific interest in paleoclimatic ˇ zek, 1958), and (4) the long-lasting general evolution of the sub-surface erent types of information (Bos In karst areas, the evolution of the surface landscape is recorded inside the caves; ff Mexico, which were integrated in a paleoclimatic study. stable carbon isotopes ofrived organic from matter agricultural (Turney pollutiontype et (Bottrell, al., (Panno 1996), 2001), et the pesticide al.,2007), presence content magnetic 2004), of de- mineral a the properties certainor fulvic (e.g., mollusks vegetation acid in Elwood cave fraction et archeologicalsediments al., of sites, as etc. 1996; a the proxy Other Sroubek organic for studies the etto that matter environmental al., land-use changes focused (Polk (Polk on on 2001), et surface et recent were al., cave discovery specifically al., 2007). linked of Only a twelve single species paper of (Polyak et oribatid al., mites 2001) mentioned in the two Holocene stalagmites from New ble isotopes in combinationfeatures of with clastic numerical cave dating,Environmental sediments studies palynology of (Bastin, cave and sediments, 1978; especially sedimentological been Bastin those also of et carried the al., entrance out facies,and 1986 using have coprolite among di pollen (Carri others). through the caves aredi frequently preserved unalteredthe for recognition millions that of cave sediments years,(Kukla can providing and reflect Lo and conserverecord paleoclimatic from data is clastic old caveof deposits White, developed 2007). only in Mostlysediments the and fossil used past vertebrate to decades remains characterizeproxies. environmental (see have conditions, been summary or Speleothems identified used have as in been paleoclimate clastic broadly cave utilized as paleoclimatic proxies using sta- and surface karst (cf. Zupan Hajna et al., 2008b). lowering of the base level shiftslying active speleogenesis passages to lower are elevations, and thements, subsequently above sometimes abandoned intercalated with and chemical precipitates, filled transported from by the sediments. surface Clastic sedi- (Kukla and Lo 2001; Panno et al., 2004; Polk et al., 2007), (3) the sub-surface depositional processes quences of clastic sedimentsand usually velocity, reflect sediment conditions source, andtions surface changes weathering that in conditions are water and aof volume depositional the function condi- cave of environments delay climateand the change. destruction Mylroie, processes 2004; Moreover, of fossil specific Polkwell remains topoclimatic preserved et (Sasowsky in features al., the 2007). underground,no such longer as: Therefore, be many (1) foundincluding the types above-ground on information of source on the information (which past surface), may changing can vegetation (2) surface be and soils the land (Kukla environmental use and which evolution Lo is on not the preserved surface, in fast- represent a series ofare depositional separated and by erosional unconformities eventsof (breaks (sedimentary substantial in duration, cycles). deposition), and which the These the may erosional depositional represent phases events periods (e.g., may Bos berepresent of remnants much of longer past duration fluvial that or lacustrine conditions within the cave. Distinct se- tary sequences here areflood extensive, consisting or injecta of deposits fluvial of gravelsThey laminated and may silts also sands and contain overlain clays dejecta, often by colluvialmarine intercalated material by ones), and speleothems. outer often clastic re-deposited sediments(cf. (including and/or Ford injected and for Williams, longerried 1989, by distances 2007). (1) within sinking streams the Mostand and cave of (2) Williams, gravity-driven the 1989; infiltration sediments from Brinkman theenvironment result and surface of from (e.g., Reeder, cave Ford material 1995; interiors car- White, and 2007). periodicity of Due events, to sedimentary the sequences dynamic often and paleontological remains thatbiochemical are alteration protected (cf. Ford from and surface Williams,in 1989, erosion, those 2007). weathering parts The and ofsilts the and cave clays thatpart are are of more relatively sediments remote rare belonging from to and the the they interior surface. facies typically form Relics contain in of vadose no phreatic conditions. fossils. Sedimen- A dominant of the cave from a stratigraphic point of view as it may contain datable archeological 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | cult to ffi ´ ˇ ska Cave Sys- ak et al., 1998, ˇ ska Cave (Bos E) belongs to the Diva ˇ ce Quarry (Fig. 1) – also due to the fact 00 45 0 ˇ 56 Crnoti ◦ 3408 3407 N; 13 00 18.8 0 40 ◦ ˇ cina, and a cave in ˇ cek et al., 2007). ˇ cek et al., 2007; Zupan Hajna et al., 2008b, 2010). The selected ´ a ´ ˇ a ska pe ˇ ci The Trhlovca Cave (45 Clastic sedimentary sequences belonging to the interior cave facies were carefully Our study represents the first attempt to identify and study fossil invertebrates in than 1.77 Ma (Zupan Hajna et al., 2008b). 2000). The preserved fluvialtions, sediments are and located in speleothems, stratigraphically deposited relevantin position in (Fig. thickness, vadose 2a). was condi- A described vertical inwell section, 4.5 detail stratified m by and Zupan starts Hajnations from of et the light-greyish al. and top (2008b). yellowish-brown with sands.by Its a The multi-colored central brownish-red, rest clays part of in clayey the is sand theinterval, section upper with is silty half intercala- represented to and very chocolate-brownarrangement fine-grained clays of below. sandy R In admixtures (reverse) and the occur N basal in (normal) bands polarized and magnetozones shows laminae. ages older The two of them (Hor tem. The cave represents achoked by part sediments. of an The ancientas cave was a and consequence later more of partly extensive the system rejuvenated evolution completely and of sediments the lower-lying exhumed Diva 2 Materials and methods 2.1 The studied sites Three relic caves, all located in– the Trhlovca, Ra Dinaric Classical Karst,that were selected detailed for paleomagnetic this dating study was calibrated by micromammal biostratigraphy in karst evolved duringfollowed a by single cave infilling post-Eoceneready processes karst at (partial the period. ofpaleontologically-calibrated Oligocene/Miocene complete paleomagnetic boundary data fossilization) Speleogenesis from as that some wasHajna of started indicated et later the al- by studied al., sites 2010). fission (Zupan track (AFTA) and or terrestric deposits are preserved on the surface now. Surface morphology and thrust in tectonically complicated structures (Placer, 1999). No post-Eocene marine mostly been studied in lake(Gandouin et sediments al., and 2006, only 2007; few Engels of et them al., in 2008; fluviatileselected Howard environments et for al., a 2009). firstwell attempt dated to by find magnetostratigraphy fossil andtratigraphy invertebrates paleomagnetic (Hor in dates caves. werecaves Sedimentary calibrated are sections by located bios- in the(southeastern Classical Europe). Karst, the part Shallowsic of marine the to Dinaric Slovene Carbonate Dinaric Paleogene) region Platform are of deposits Kras covered (Juras- by Eocene flysch siliciclastics. Both units are over- chironomids (Diptera; Walker, 2001),the and most commonly ostracods used (Crustacea;identify invertebrate Holmes, them remains 2001) to in are aOther paleolimnology. remains, species It such level, is as andmollusks, not protozoans, their are di bryozoans, rather autoecology oribatid rarely is used mites well (Smol, (Acarina), 2002). known insects Until (Luoto, and recently, 2009). invertebrate fossils have imentary deposits, especiallyplant lacustrine macrofossils, ones, pollen, represent algaeoped paleoecological and as archives fossil a of invertebrates. multidisciplinarychemical science and Paleolimnology especially biological has in proxies devel- the preservedture in last and lake two the sediments composition decades (Luoto, of using 2009).by fossil physical, assemblages The Erickson (we struc- and prefer the Platt,vironment, term fossil, reflecting 2007, as past to proposed climate,or “subfossil”) ecological nutrient vary interactions conditions, in (Luoto, oxygen response 2009). content, to The pH, changes cladocerans pollution (Crustacea; in Rautio, the 2007), en- clastic sediments from the interiorological cave proxies facies in and paleoenvironmental tofrom studies. discuss other Similar their sedimentary methods possible deposits, use and ascan as yet proxies be studied bi- to applied in those also lakes,and seas to and paleoenvironmental cave lotic conditions sediments environments, in at order the to time obtain of information sediment on deposition. paleoclimate Surface sed- 5 5 20 25 15 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ). The atavus, ˇ cek et al., ˇ ˇ cek et al., cf. ce Quarry, ´ a ´ a E) uncovered 00 Potamon ˇ Crnoti . Fauna from the Marifugia cavatica 47.7 0 Apodemus 52 ◦ , N; 13 ´ ak et al., 2004; Hor E) represents a relic of an old 00 00 Ursus spelaeus sp.) that belong to MN15–MN16 56.3 0 1.56 ´ ak et al., 1999, 2004; Mihevc, 2001, 0 9 33 ◦ ◦ ) belong to middle to late MN17 (ca. 1.8– alcohol and each specimen was identified Cseria ◦ N; 14 Beremedia fissidens 3410 3409 00 sp., sp., Borsodia 12.5 0 ˇ ce Quarry (45 30 , cf. ◦ ˇ Crnoti Deinsdorfia Apodemus frequens, Glirulus ˇ cina Cave (45 cf. ˇ e, 1930 on the northern cave wall (Mihevc et al., 2001, 2002). The ˇ cek et al., 2007; Zupan Hajna et al., 2008b; Fig. 2b). Its lower part ´ a ˇ cek et al., 2007; position of R1 and R2 in Fig. 2). The clays are ponded ´ a ˇ ska pe ˇ ci ˇ cek et al., 2007). The top of the section represents flowstone layers with interca- ´ a An approximate quantity of 1 kg of sediment was taken from each sampling point Quarry operations in the The Ra the Olympus SZX2 stereomicroscope in 90 search, the samplesunits, were located selected in from such positions whatestablished that paleontological appeared would and to allow magnetic a besamples chronostratigraphy. clear were distinct, In correlation taken the with stratigraphic only theremained previously from intact the after a upper recent part collapse of of the the topmost originallyand sediments placed described in in the section sealed quarry plastic that face. in bags with 10% a KOH label. for Inand 30 the min, 40 laboratory, µm. the and samples washed were Sub-samples kept successively for through each sieves sieve of dimension 250 were µm, examined 125 µm separately under 2.2 Sampling protocols and invertebrate fossilA identification total of 11samples samples were were taken taken from frompaleomagnetic the the and selected exposed paleontological sites faces, studies. as which indicated have Given in been the Fig. already exploratory 2. sampled nature for The of our re- (ca. 3.0–4.1 Ma) were2007). found The in basal theoped 1 same m in is two horizon composed sequences asan of separated age the of multi-colored by serpulids the an laminated fillbly, (Hor the silts angular older fill than and unconformity. belongs 1.77 clays to Ma Thewithin the devel- (base the fauna Gauss of Gilbert chron indicated the chron (2.6–3.6 C2n (4.18–4.29 Ma)2007). or Olduvai or 4.48–4.62 subchron). the Ma; Bos other Most N-polarized proba- subchron cyclically/rhythmically arranged multi-colored fluvial sedimentscroconglomerates). (clays to intraclastic This mi- fillAbsolon rests et on Hrab sessile tubesmammal of remains the (with serpulid Rhagapodemus and a slight unconformity inside. The lowermost 7 m of the cave fill are composed of matrix. This issilts and underlain clays, by sometimes an sandy, overlying interval, the up rest to of 4.5 the thick, section with of deep light-colored, erosion laminated of C2n Olduvai subchronbe (1.770–1.950 Ma). correlated The with magnetozone theGauss alternation chron lower below (2.581–3.580 can Ma). part of the Matuyama chronmany caves (2.150–2.581 Ma) completely filled and with2007; the sediments Fig. (Bos 2c).sents The a studied relic section ofand is an a located extensive, height in sediment-filled of the passage more western with than quarry a 17 wall m. diameter and of Its repre- about top 10 is m filled by speleothem breccia with red clay (Hor lations of brown caveclay loams permitted with bone to fragments fixmagnetic of the Polarity Time arrangement Scale of (GPTS; the CandeR-polarized and interpreted magnetozone Kent, within magnetozones 1995). the with The interval boundary the of with Geo- N- fauna and was identified with the bottom micromammals (with 2.4 Ma; Hor and partly covered bysists several of collapse subhorizontally boulders. laminated,cient The porous rimstone upper and dams part light-colored andthe of flowstone flowstone interbedded the with layers section red resulted some from con- claysgrained an- successive allochthonous and sediments flooding This silts. that may deposited indicateponor either well-sorted Lutitic a of fine- distant interbeds the position far between surface from river the or an allogenic stream passing through a system of sumps cave system. Theof 2 m 6.5 m thick (Hor verticalconsists section of has a a vaultedIt stalagmite composite is that stratigraphic includes overlain thickness several byand interbedded a thin layers thick calcite of interval crusts red and of clays. fossil red finds clays (vertebrates with and some invertebrate – silty and sandy intercalations 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ˇ ska ˇ ci are known ´ ocs and Hufnagel, sp. 2, two species in some characters Daphnia erent genus. Species of ff sp. (Suctobelbidae) of the ects via food limitation indirectly Suctobelbella ff Suctobelbella Oppiella (Rhinoppia) species in deep Quaternary lake sediments. (Microzetidae) was found in the sediments of 3412 3411 (Oribatulidae) found in the Trhlovca Cave is a (sensu lato) can be considered one of the most morphologically resembles a species known from erence is in the length of sensilla; the recent ones ff sp. 1 and Daphnia sp. of the same family (Opiidae) was represented in Miracarus Opiella Oppiella shore cladocerans. Korhola (1999) and Korhola et al. (2000) ff ˇ ska caves. Modern representatives of the genus are forest-litter cient numbers to be useful as proxies in investigations of Quater- ˇ Dissorhina ci ffi Zygoribatula frisiae sp. was identified in the Trhlovca Cave. The identified specimen is in a Oppiella (Rhinoppia) Daphnia A new species belonging to this family are common in litter and upper organic layer of forest soils, especially with the Trhlovca Cave by twoscience. specimens, probably Some belonging species to of the this(Seniczak genus same, et prefer new the species al., border to genus between 2006). forest to and drought. open Taylor areas andliving species, Wolters belonging (2005) to a mentioned2002). genus the often The found tolerance species in of more isand arid xero-tolerant, this lichens, settings today often (Shepherd known in et fromTrhlovca arboricol Cave al., resembles repeatedly microhabitats. species drying-out of the mosses recentbut genus displays some specific characters, which may define a di abundance in forest litter, alsoOne present of in the new shrublands, speciesmodern ecotone of cave zones environments. and The grasslands. di have very long ones. Thefauna species of in sediments Slovenia. is therefore an extinct element of cave 2001; Polyak et al., 2001).of Five species them were are identified new in to the science. studied samples and four the Trhlovca and Ra inhabitants. of the family Oppidaecaves. were Species identified of the incommon genus arthropod the groups sediments on Earth of (Norton the and Trhlovca Palmer, 1991) and with Ra high diversity and by a decrease ofNevertheless, some microbiota species in adapted soil alsoity. to or These xeric minute other conditions arthropods substratum or aresediments as high usually and values in preserved a of su well result humid- enoughnary of in paleoclimate, lacustrine lack paleoecology or of fluvial and water. stratigraphy (Solhøy and Solhøy, 2000; Solhøy, in humid habitats. Drought susceptibility exerts its e and van Straalen, 1995;2009). Behan-Pelletier, 1999; Almost Gulvik, all 2007; studies support Gerg the idea of a group with many representatives living found maximum lake depth totribution be in the Fennoscandian most lakes. importantclaws, mandibles factor and Jeppesen explaining ephippia et cladoceran of dis- al. (2001) found3.1.2 only post-abdominal Acarina Oribatida Mites of thecroarthropods, which suborder can Oribatida bebioindicators also (Acarina, of found the in Arachnida) ecological caves. conditions are Mites in are terrestric typical of and high aquatic soil-dwelling ecosystems potential (Lebrun mi- value as 3.1.1 Crustacea Cladocera One relatively good condition (Fig. 3),as but large-bodied completely pelagic flattened o laterally. The number of identifiableSome invertebrate of fossils the in samples were allments completely samples and invertebrate-sterile. was vegetal Unidentifiable veryspecimens animal fragments low frag- belong (Table were 1). to also theChironomida and groups found. Hymenoptera of (Insecta). Cladocera Theinvertebrate All (Crustacea), fossil these only assemblages Oribatida groups from are relatively (Acarina), lake commonly sediments well-preserved and identified (Elias, in 2007). 3.1 Faunal inventory following the specific methods for each group. 3 Results and discussion under the Olympus BX51 microscope. Identification of the individuals was carried out 5 5 25 15 20 10 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) Tetramo- Oppiella and and species, as an ˇ ce Quarry cave fill. Daphnia Daphnia Miracarus ˇ Crnoti ´ nska and Zawisza, 2005), was found in the sediments of the Trhlovca 3414 3413 was found in association with dry habitat taxa, such . Fossils from these three samples indicate a transi- Tetramorium Dissorhina Tetramorium ). Two taxa were identified at a genus level ( sp. in sample T5 suggests the presence of a lake or a low hydraulic and ). From the oldest samples, T4–T5, only T5 contained fossil remains. The Daphnia ) and one taxon as subfamily (Orthocladiinae), due to their poor conservation. The Zygoribatula Most of the invertebrate fossils were found in the Trhlovca Cave. Samples 1–3 belong All the analyzed samples were taken from cave sediments deposited during the Dissorhina Zygoribatula frisiae indicator of oligotrophic (low food) environments (Szeroczy a more humid and forestedence habitats of of the Lateregime Pliocene/Early of Pleistocene. the stream, The as pres- wellsupported as by the the proximity varved of sediments cave that entrance.ing indicate This single-flood continuous assumption deposition events is in also or theyears flood past, (Zupan dur- pulses Hajna that et al., probablyis 2008a). lasted also The indicative less relatively of than good adue a preservation slow to of few flow the this from thousand still cladoceran the epiphreatic2007) surface position down and of to to some the parts the deposition of place short-distance the possibly transport. subterranean system (Mihevc, The presence of or taxa which can( be found in the ecotoneecotone zone zones – or typical in open,as moderately humid areas tion from a warmer period with relatively drier vegetation (Early or Middle Pliocene) to Early–Middle Pliocene (ca. 5.3 MaPliocene–Early Pleistocene to (ca. 3.6 Ma) 3.6 to withto 1.77 the higher Ma) present-day temperatures, with climate lower and (Haywood, temperatures, 2009). the more Late similar to a more recent periodall the than studied the samples older are infillingsamples older where than T1 samples 1.77 and Ma 4–5 T2 (Zupan wereamount Hajna include collected, et of taxa but al., that dead 2008b, are 2010). organic typical Clay matter for in forest the habitats upper with horizon relatively high of soil ( Pliocene and Pleistocene periods,content dated (Fig. by 4). magnetostratigraphy andcooler During paleontological and the drier than Pliocene, during thepronounced the climate Pleistocene Miocene (Robinson is cooling. et globally al.,riod Broadly, known 2008), covered two followed to by by climatic have the the stages become deposition more are of clastic defined sediments in in the the pe- sampled caves (Fig. 5): the due to climatic conditionssubsequently. and sedimentation processes at the time of deposition and 3.2 Paleoenvironmental significance of the fossilFive invertebrates taxa from identified onesOnly are a new single to taxon science( as and yet their identified descriptionrium is at in a progress. species levelcave is richest in found fossils in wassamples. the the The Trhlovca modern Cave maximum with number fauna of six taxa identified belongs taxa to in the oldest three sample of (T5), the presumably five One individual of theCave. genus The genus islacks a the typical head, inhabitant but of isdistribution dry in otherwise landscapes the well with present preserved. shrubs. fauna. It The represents individual a genus with very large The absence of mentumfamily made Orthocladiinae is the a identification group of ofwith chironomid Diptera cold the whose and lower larvae well-oxygenated prefer taxon waters loticRepresentatives habitats impossible. (Dimitriadis of and this Sub- Cranston, family 2001; inhabittoms Walker, cold 2007). of and lakes, running but streams areto or generally unstable low adapted sandy oxygen to bot- levels lowronomids (Eggermont food and et (Walker, its 2007) al., representatives and are 2008). are frequently intolerant found It in is lake3.1.4 a sediments. widely distributed Insecta family Hymenoptera of chi- of the living species have been found in rotting3.1.3 wood and under Insecta bark Chironomida of dead trees. One representative of Orthocladiinae was identified in the large amount of decomposing organic matter and with abundant fungal hyphae. Some 5 5 25 15 20 10 25 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | My- and Glir- , , Cseria Deinsdorfia Myodes Glirulus , frequens cf. ) that inhabit forest erence is the pres- ˇ ff cina. These are rep- genera), is indicative of a Apodemus erent Pliocene/Pleistocene ˇ ska pe ff ˇ ci Suctobelbella ˇ ce Quarry, including Rhagapodemus , and Deinsdorfia ˇ Crnoti and atavus typical for temperate climate, and of 3416 3415 cf. Miracarus ˇ cek et al., 2007) as indicators of a colder and , ´ a Blarinoides ˇ cina Cave, the most important di (Hor , ˇ cek et al., 2007), is typical for a temperate and humid ´ a Oppiella Apodemus , Rhagapodemus ˇ ska pe , ˇ ci Borsodia sp. (Hor and that are water-related micro-mammals. Samples T1 and R4 include Ori- Cseria Beremendia Beremendia fissidens . The presence of invertebrate drought-tolerant taxa in T5 indicates a land- Myodes ˇ ce Quarry. Located in the upper half of the sediment section, the Orthocladi- genera), together with taxa that are widely distributed in temperate and humid sp. and Beremendia fissidens Mimomys The fossil vertebrate assemblage from the Only sample R4 provided fossil invertebrates in the Ra The only chironomid identified at a family level was found in sample C1 in the ˇ Pleistocene. presence of trees inBorsodia wet forests and steppicscape habitats, typical with habitats shrubs for forest and inhabitants grassland in during T1vertebrates the points and late to R4 a Early correlatedchronologically colder Pliocene. extreme climate with samples, during invertebrates the the were The presence alsoT2. Mid-Pleistocene. identified presence of Between in of Sample samples these cold/dry-associated C1 two C1 and bly corresponds higher to flow the ratesSample advent on T2 of the fits the surface, within which Late the explains Pliocene the picture cooling presence of and of forests proba- the mixed chironomid. with open areas during the Early indicator for cold/well-oxygenated waterpresence (Orthocladiinae), of which isand supported by the batida invertebrates (genera litter, shrublands, ecotone zones and grasslands, and vertebrates which indicate the odes climates ( mild climate. Atence the Ra of drier climate. Owing towith the the scarce equally presence scarce of vertebratesamples identifiable record (C1, is invertebrates, problematic. a T1 Only correlation andsils, three and R4) of correlations the come were invertebrate from made only the for same these periods samples. as Sample C1 the provides found a vertebrate single fos- climate and more humid forest (Table 2, Fig. 5). sp., ulus climate. The high number of forest and shrub inhabitants ( periods, indicating a transition from a dry or mild climate with mixed forest to a colder ecology of some taxa canas be with correlated with the the climate discovered and vertebrate corresponding remains vegetation as of well the di for the taxonomy oftal the group significance. of mites Speciesfauna (in can biodiversity this bring linked case), additional but tothe information is the Pliocene/Pleistocene, about also and paleoclimate, the of the paleogeography importance paleoenvironmen- soilas and of and a the vegetation surface potential mid-Pliocene during warm analogal., period 2009). for (3 The future Ma) Pliocene/Pleistocene climate cavetaxa, sediments has or show been nothing a at already very all lowremains emphasized in concentration makes (Salzmann more of further than et biostratigraphic half interpretation of impossible, the but analyzed the samples. presence and The scarcity of fauna value of environmental variables asment a proxy function data of (Birks, biological 1995). data,presence/absence the Such so-called and proxy data environ- ordinal can estimates beertheless, defined of quantitatively, the or abundance by use (Prentice the was of et emphasized less al., by frequently 1996). Luoto occurringThe (2009) Nev- zoological discovery as of remains an five in important new paleolimnology proxy species for (and environmental perhaps changes. a new genus) is important not only Crnoti inae dipteran can be usedindividual as was an brought indicator under forgests colder the a and ground colder well-oxygenated climate by waters. with a more The relatively intense precipitation fast towards3.3 flowing the river, end which of Correlations the sug- with Pliocene. other proxies The primary aim of the quantitative reconstructions in paleolimnology is to express the resented by three individuals ofof oribatid forest mites inhabitants. belonging Sample to R4that the belongs comes same to from ecological the the group Early upper Pleistocene half (termination of of the Olduvai). section of this cave supports the above mentioned hypothesis of a drier vegetation on the surface. 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ˇ cek ´ a erent ff in the drainage area or to ff erent areas nearby. Vegeta- ff 3418 3417 erent types of fine laminated clays and silts. They were deposited ff erent species. Oribatida is a group with a lower mobility than flying inver- ff ˇ zek, 1988). This fact is probably related to the runo No pollen was found in the studied samples and sections, but detailed studies on base level was connected withFodor, the 2006). change in This tectonic caused regime a at transition ca. 6 from Ma an (Vrabec and epiphreatic regime to a vadose one, and deposited together with the sediments.studied caves The are most di common clasticfrom sediments in floodwater the suspended loadparable under with conditions slackwater facies of ofcorresponds pulsed to Bosch flow more and or or White caveassemblages less (2004). lakes of regular This com- flooding the depositional of caveing process karst products deposits areas of were by Eocene flysch sinking derivedlimestones sediments rivers. (Zupan from and Hajna Mineral soil et highly al., cover 2008b). homogenized on Paleocene The weather- and tectonically-driven lowering Cretaceous of the regional of the cavities withdeposited sediment in during clastic cave theand sediments Tertiary. Lo of the The interior verythe facies low hydraulics is of abundance well the of subterranean known streams.ples fossils (e.g., Almost have Hor all at of least theare studied one surface invertebrate aquatic sam- invertebrates, component, which and were all brought (with inside a the single caves possible by exception) allogenic streams 3.4 Implications for the reconstruction ofFossil karst invertebrates evolution may yield valuable informationhydraulic concerning regime and the deposition origin rate, of etc.condition sediments, of According the to fossil Ericksen is (1988), known,processes if its the that endpoint standing can acted be on usedpresence to the of infer surface some body of animals during the indicative geological its for pre-Quaternary an transport interior intense facies to karst of the evolution cave of site sediments the is of Dinaric in- deposition. karst, including the The filling of a part cave sediments as a proxy,small even in scales more and recent must sediments, beof can associated the provide with information other same at proxies region andsignificant in preferably at with order regional more scale. to sites provide paleoenvironmental information that would be taken into account only at small or medium scales. The use of invertebrate fossils from layson et al. (2008) for more recent sediments, the environmental changes must be the slowly migrating vegetationspecies or can vegetal be associations. considered eitherin The members the identified of present an new fauna extinct Oribatida of faunalundergone southern group evolution with Slovenia, in no or the relatives ancestorstion present-day of succession habitats a in or living time in speciesclimates/biomes contributed which di in to have the the Mediterranean settlement zonefor (Bertini, of flora 2010). a is This complex supported situation mosaicthat documented of by provided di the most invertebrates ofmixtures the found of specimens, in forests in the with the open cave Trhlovca dry sediments. Cave, areas yielded in The taxa all section indicative samples. for As already documented by Fin- cave sediments of Slovenia.defined In mostly general, by faunal climate changesrelatively depend short and on duration, vegetal some the associations. ofevolve environment, into the di If taxa changes that aretebrate are unable species, frequent to and and adapt their may of migration get can extinct, or only be done over short distances, following general decrease of humidity andoccurred installation prior of drier to conditions 2.5steppes, Ma during with the (Bertoldi Piacenzian typical et open al.,tocene, vegetation 1989). during phases the characterized Alternations Gelasiancomposition the of and and following forest structure early the vegetation of Pleis- Calabrian and vegetationby of (Bertini, the the rare 2010). Pliocene-Pleistocene appearance are ofpearance In characterized new of general, species the the and same floral vegetation by can spatial taxon explain asynchronous (Bertini, the events 2010). of extinction of disap- some Frequent of changes the in identified both invertebrate climate taxa in and the the nearby Italian and peri-Mediterraneanrelated sites with given in the Bertini invertebrateDuring (2010) fossils can the found be Zanclean, in well the cor- the climatepending was Slovenian on warmer cave the and sediments drier geographical (Tableof or 2). latitude more humid and humid subtropical altitude than to today, (Faquette de- warm et al., temperate 2006); zones forest were taxa dominant (Bertini, 2010). The 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 31/2010 funded by CNCS- ID sp. and its relatively good preser- Daphnia 3420 3419 , serpulid polychaetes attached to walls at the ˇ c has kindly helped with the English revision of the manuscript. Marifugia cavatica The authors are grateful to Ilse Bartsch, Karina Battes, Steve Brook, Adamovi ´ ı ´ alint Marko, Laura Momeu for the help with taxa recognition and ecologi- ˇ r mpean, B ı ˆ Caves are known as systems with low-energy input from the surface and low- Pre-Quaternary invertebrate remains, abundant in marine settings and quite com- As already mentioned, the presence of Acknowledgements. Mirela C cal information. Ji This study was funded throughUEFISCDI the (Romania; KARSTHIVES granted Project to PCCE SC), Grantresearch Agency of AS CR karst No. sediments: IAA300130701 (Paleomagnetic and paleotectonic the and Institutional geomorphological Research implications; Plan granted No. to AV0Z30130516 PB) of the GLI AS CR, v. v. i. tebrate fauna, these findingsPliocene/Pleistocene support transition, the while also ideaevolution. bringing of new a insights colder on phase the associated regional with karst the okarst settings. energy in situcaves production. due toimens the This relatively transported may low into hamperlow chance caves. the of number use finding However, ofbiostratigraphy-calibrated of identifiable for invertebrate magnetostratigraphy cave fossil the back specimens or invertebrates threeacross to surface were the from described spec- Pliocene/Pleistocene ca. successfully boundary. sites, When 1.77 identified a cross-correlated to with relatively and the ca. coeval ver- dated 4.8 Ma, by i.e., of vertebrate fossils in pre-Quaternaryof inferred cave paleoenvironmental deposits; settings (ii) in theproxies; the (iii) need case the for of cave cross-validation incomplete conditions orinvertebrates that ambiguous when are coeval prone compared to to ainformation surface better on settings; preservation speleogenesis of (iv) and old its chitinous evolution added-value of as karst a hydraulic source regimes of even for pale- conditions. This new proxy may be important for a number of reasons: (i) the scarcity cave facies in temperateserve regions. yet These another finds proxy suggest for that the cave assessment sediments of can paleoclimatic pre- and paleoenvironmental slow transport to therate, and site (3) of subdued microbial deposition,entire and organisms. biochemical (2) processes a that could rapid have burial, altered the i.e., a high sedimentation 4 Conclusions Three sites of relic caves fromPliocene/Pleistocene the invertebrate Classical remains in Karst continental of clastic Slovenia sediments provided from first interior evidence of mechanisms and the low intensitygood of state of biochemical preservation processes of canArctic old explain lake invertebrate the sediments. remains relatively both AlthoughSlovenian in sites the the number is cave sediments of small,age and identified their of invertebrates state the at of sediments. the preservation This studied is may relatively suggests good a combination considering of: the (1) a relatively short and varved and/or cyclically arrangeda sediments. period The of chironomid relatively inat high the sample same C1 flow level indicates rate as air/karst-water in interface, a still living vadoseMihevc in regime. et the al., caves The 2001). of chironomid the was Dinaric found karst (formon summary in see amber, areonly very from rare and Arctic have environments as (see yet a been complete reported list in in continental Elias, sediments 2010). The depositional parts and a complete fill withHajna fluvial et sediments al., as a 2008b). consequence (Mihevc, 2007; Zupan vation in sample T5 indicatescave entrance the and existence multiple of flood a events that system contributed with to low the hydraulic continuous deposition head of at the followed by a decrease in hydraulic head in some cave systems or their higher-situated 5 5 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ´ es depui Marifugia ˇ zba ZRC, nidos, edited by: ı ´ ´ a, A.: Cave fill in the ˇ ce II site with ´ e d’approche des fluctu- ˇ Crnoti ˇ sek, F., Carsologica, Zalo ´ eriz, P., Finlayson, C., and Riquelme, ´ a, D., and Langrov ´ alez-Samp ˇ cek, I. (Eds.): Paleokarst. A Systematic and Re- 3422 3421 ´ a a, M. S., Gonz ı ´ azek, J., and Hor ł ´ e, M., Fumanal, M. 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C.: The distribution, mechanisms, and evolutionary significance 5 5 30 25 20 15 30 10 25 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | incomplete specimen in poor condition Observation new species specimen in poor condition new species cosmopolite cosmopolite unidentified new species new species specimen in poor condition 1 1 individuals 1 1 1 1 1 1 1 4 2 1 ) n. sp. 1 ˇ zba ZRC, Ljubljana, 8, 266 pp., 2008b. ) n. sp. 2 ´ ak, P.: Palaeomagnetic research on karst sp. ? ´ ak, P.: Cave sediments from the Postojnska- sp. ´ ak, P.: Palaeomagnetism and Magnetostratig- Rhinoppia n. sp. n. sp. n. sp. 1n. new species sp. Rhinoppia sp. ( (cf. 3428 3427 Astigmatida? Identified taxaMiracarus Opiella Daphnia No. Dissorhina Zygoribatula frisiae Tetramorium Insecta larvae Oppiella Miracarus Suctobelbella Dissorhina of the sediment ˇ ce 1 (C1) yellow clay Orthocladiinae ˇ ska 4 (R4) red clay A list of invertebrate fossils in the samples of cave sediments in Slovenia (only samples ˇ ci ˇ Crnoti Site/Sample Short description raphy of Karst ediments in Slovenia, Carsologica, Zalo sediments in Slovenia, Int. J. Speleol., 39, 47–60, 2010. Planinska cave systemActa (Slovenia): Carsologica, 37, evidence 63–86, of 2008a. multiphase evolution in epiphreatic zone, Using Lake Sediments, Vol.and 4, Last, Zoological W. Indicators, M., Kluwer edited Academic by: Publishers, Dordrecht, Smol, 2001. J. P., Birks, H. J. B., 4 Ra 5 1 Trhlovca 1 (T1)2 red Trhlovca 2 clay (T2) beige clay 3 Trhlovca 5 (T5) red clay Table 1. with identifiable animal remains are shown). Zupan Hajna, N., Mihevc, A., Pruner, P., and Bos Zupan Hajna, N., Mihevc, A., Pruner, P., and Bos Zupan Hajna, N., Mihevc, A., Pruner, P., and Bos White, W. B.: Cave sediments and palaeoclimate, J. Cave Karst Stud., 69, 76–93, 2007. Walker, R.: Midges: Chironomidae and related Diptera, in: Tracking Environmental Change 5 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ˇ ska ˇ ci ), Ra ∗ = riverside habitats + ˇ ce Quarry (the section ˇ Crnoti river living in steppe + river fluvial + ecotone zones,grasslands fluvialwell oxygenated micromammals fluvial micromammals ecotone zones fluvial fossil cave in the ) water living in tree = 3430 3429 ∗ ˇ ce origin Marifugia ˇ ( cavatica Crnoti – forest, surface, (fish teeth ) and tree habitats ∗ ˇ ska ˇ ci Potamon Suctobelbella ( Miracarus – – forest, surface, – Oppiella ˇ car et al., 1969): C , shrubs , – – dry landscape with surface, – ˇ cek et al. (2007). Trhlovca Cave (the oldest part of the section with fluvial sediments), R ´ a = – – Orthocladiinae cold, surface, (fish teeth*), Miracarus Opiella Daphnia Dissorhina Zygoribatula Tetramorium Trhlovca Ra Dissorhina – Taxa found in cave sediments of Slovenia with the corresponding vegetation, sediment Location of the studied sites in Slovenia (see the geological map in the upper right ˇ cina Cave (the section is an alternation of flowstone and clay deposits; Photos by A. Mihevc). Upper Pliocene/ Lower Pliocene StageLower Pleistocene TAXA Environment Sediment Vertebrates mentioned in Hor is represented by anflowstone), unroofed T cave filled with yellowpe fluvial sediment covered by red clay with Fig. 1. corner; map after Pleni ∗ Table 2. type and origin, and vertebrate fossils. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ; sampling Oppiella = , 6 (h) large flowstone boul- = red clay, silt and sand, sp. ?; Zygoribatula frisiae = (d) Marifugia cavatica sampling points (T1–T5; modified speleothems on the section surface, = flowstone, 3 = n. sp.; clays lighter in color, with a higher sand = Suctobelbella = (g) limestone 2 Dissorhina = (c) a section in the filled horizontal cave passage cut by the ) n. sp. 1; flowstone mixed with reddish clay, 3 A section exposed in the Trhlovca Cave, several meters in = 3432 3431 (a) (b) stalagmites in flowstone, 6 wall and sediment with tubes of = = brownish to ochre sandy clay, 5 = Rhinoppia limestone, 2 ˇ cina Cave section: 1 Orthocladiinae; = (cf. (b) ´ sampling points (R1–R4; modified from Zupan Hajna et al., 2008b). ak et al., 2004). ˇ ska pe ˇ = ci sp.; Opiella Ra reddish younger fill of clay, silt and fine sand, 3 = (f) (c) Daphnia n. sp.; (a) yellowish brown sandy clay, 4 limestone, 2 ) n. sp. 2. = = Lithological sections at the studied sites: allogenic laminated fluvial sediment, 5 ´ ak et al., 2006); Light microscope photographs of the identified fossil invertebrates in the cave sediments = ˇ ce Quarry (the original section): 1 Miracarus brown clays with gravel and cave bear bones, 5 collapsed limestone blocks, 8 = = Rhinoppia ˇ (e) ( Fig. 3. in Slovenia: Fig. 2. Crnoti ders, 4 points (C1–C2; modifiedthickness: from 1 Bos proportion, 3 4 from Bos 7 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | fossil = ˇ ska Cave, C ˇ ci Ra = Trhlovca Cave, R = 3434 3433 ˇ cek et al., 2007), mammal zones, temperature estimates (after Lisiecki and ´ a ˇ ce Quarry and vertebrates (in blue). ˇ Crnoti Invertebrate fossils found in the studied caves, correlated with climate, vegetation and A correlation of the obtained magnetostratigraphic results with the standard paleomagnetic scale (GPTS; after Fig. 5. fossil vertebrates. Cande and Kent, 1995; modified after Hor Fig. 4. Raymo, 2005) and position of the samples with invertebrates (in red): T cave in the