History of discovery and age of labyrinthodont remains in the Tatra Mts, Poland*

Prace Muzeum Ziemi Nr 43, 1996 PL ISSN 0032-6275 Prace paleozoologiczne ZBIGNIEW KOTAŃSKI History of discovery and age of labyrinthodont remains in the Tatra Mts, Poland* ABSTRACT. History of discovery and studies of labyrinthodont remains from the Western Tatra Mts in Poland is presented. The bony material was collected in 1959 from the Partnach Beds in the Wielkie Koryciska Ravine. The age of sediments yielding the bony remains and marine invertebrate fossils is precisely defined on the basis of foraminifers, Key words: labyrinthodonts, upper Anisian-lower Ladinian, Tatra Mts, Poland. STRESZCZENIE. Kości labiryntodontów zostały znalezione przez autora pracy w 1959 r. w dolince Wielkie Koryciska w Tatrach Zachodnich i przekazane do opracowania dr. Julianowi Kulczyckiemu z Muzeum Ziemi. Kulczycki po wypreparowaniu materiału stwierdził, że reprezentuje on nowy gatunek kapitozaurida. Po długiej przerwie, spowodowanej chorobą i śmiercią Kulczyckiego, opracowaniem tych materiałów, zgodnie z życzeniem Kulczyckiego, zajęła się dr T. Maryańska. Szczątki labiryntodontów i towarzyszące im okruchy drewna pochodzą z warstw złożonych z margli i wapieni, zaliczanych w czasie ich odkrycia do retyku. W warstwach tych zostały również znalezione amonity, daonelle i inne bezkręgowce morskie świadczące o póżnoanizyjskim-wczesnoladyńskim wieku tych osadów. Na podstawie fauny bezkręgowców i analogii litologicznych do AJp Wschodnich, warstwy te uznano za odpowiednik warstw z Partnach. Wiek ten jest pośrednio potwierdzony także tym, że ammonites and daonellas as Anisian and Ladinian boundary. The beds with labyrinthodonts belong to the Furkaska and Wielkie Koryciska units attributed to the Upper Subtatric (Strazov) nappe. Such tectonic position indicates paleogeographical situation of the beds with amphibian remains in the central part of the Tethys. leżące bezpośrednio niżej warstwy z Reifling zawierają konodonty charakterystyczne dla górnego anizyku, a leżący powyżej dolomit z Wetterstein jest wieku ladyńskiego. Pod względem tektonicznym warstwy z Partnach należą do jednostki Furkaski i jednostki Korycisk, zaliczanych przez Kotańskiego do płaszczowiny reglowej górnej (strażowskiej), która jest nasunięta na płaszczowinę reglową środkową (choczańską), ta zaś z kolei na płaszczowinę reglową dolną (kriżniańską). Taka pozycja tektoniczna wyznacza położenie paleogeograficzne warstw z labiryntodontami w środku Tetydy. Labiryntodonty musiały zapewne żyć na wyspie koralowej zbudowanej z wapieni i dolomitów z Wetterstein. Kilka osobników wraz z okruchami drewna, być może na większych jego kłodach, dostało się podczas burzy tropikalnej do basenu warstw z Partnach. Burze takie musiały być dość częste w środkowej części oceanu Tetydy. DISCOVERY OF LABYRINTHODONTS IN THE TATRA MTS The history of investigations of labyrinthodonts from the Tatra Mts spans several tens of years. The discovery of the bony remains was connected with the preparation for the 32nd Meeting of the Polish Geological Society which took place in Zakopane in 1959. At that time I followed the route G2 leading through the Subtatric Series of the Chochołowska Valley to the side Wielkie Koryciska Ravine. In the Chochołowska Valley Guzik (1959, 1961) described the contact of the Koryciska and Furkaska tectonic units which were, at that time, included into the Choć nappe. According to him the contact is found in conjunction with the Rhaetic synclinal belt of the Furkaska unit, which beginning.from the alp in the * Text edited by Teresa Maryańska Wielkie Koryciska is divided into two Rhaetic synclines. According to Guzik in the spectacular outcrops of this region the facies differentiation of the Rhaetian (and Hettangian?) in the Furkaska unit can be observed. Walking southwards from the alp (1050 m above sea level) towards the so-called Drwal's Gully, I observed the remains of rocks sliding into this gully forming a clearly visible debris cone. On some surfaces of blocks of the thick-bedded limestones the broad stems of crinoids, sections of ammonites and numerous large pelecypods were observed. The discovery of ammonites in the limestones previously considered as Rhaetian was very interesting. Ammonites are extremely rare in the Rhaetian sediments, their relative abundance suggested that the limestones were not Rhaetian but Middle Triassic. The first publica-

48 ZBIGNIEW KOTAŃSKI 1АК1АЛ PAUOZOOiOOłl А Им» «ми m m Warszawa,dn.9.V. 1960. Do: Pana Dr.Zbigniewa Kotarskiego Zakład Geologii Dynamicznej U.W. Warszawa,ul.Oboźna 8 Otreymane okazy z Tatr okazały się, zgodnie z przypuszozenlem Pana Doktora,szozątkami płazów tarczogłowych. Jest to bardzo oiekawe i na terenie Polski rzadkie znalezisko. Pożądanym byłoby zebranie większej ilośoi materiału i szczegółowe opracowanie paleontologiczne. Zakład Paleozoologii zwraoa się do Pana Doktora z uprzejmą prośbą o wskazanie punktu znalezienia,aby umożliwić naszym pracownikom przeprowadzenie eksploatacji wyżej wspomnianych szczątków kostnyoh. Fig.l The letter of Professor Roman Kozłowski confirming supposition concerning labyrinthodont bony remains. Fig. 2 Carbonized wood fragments transported together with labyrinthodonts from land to the sea presumably during a tropical storm. tion mentioned this fact was that by Kotański (1961). In 1959 I gathered much more paleontological material in Wielkie Koryciska Ravine. In addition to ammonites, nautiloids, echinoderms and pelecypods I found black bony remains with a characteristic wavy surface and some fragments of jaws with tiny teeth which I recognized as the remains of labyrinthodonts. In autumn 1959 I delivered all the specimens to Professor Roman Kozłowski who was at that time the Director of the Institute of Paleozoology of the Polish Academy of Sciences in Warsaw. In a letter dated the 9th of May 1960 (Fig. 1) Professor Kozłowski confirmed my view that we were dealing with the remains of labyrinthodont amphibians. The paleontological investigations of labyrinthodont material was charged to Julian Kulczycki, who was a well-known researcher of fossil vertebrates at the Museum of the Earth, Polish Academy of Sciences in Warsaw. Kulczycki began preparation works in November 1959 and continued them in 1960-1963. The preparation of bony remains involved etching the rocks using acetic and hydrofluoric acids, and then coating the bones with paraffins. It was soon realized that the prepared bony remains were precious paleontological material. In the unpublished account on the studies carried out in 1961 Kulczycki stated that the bony remains represented a new species of the capitosaurid labyrinthodonts. Informing me about the results of his studies he stated that up till now all bony remains found "fit in the same head". In autumn 1963 Kulczycki conducted the complementary field work at the Wielkie Koryciska Ravine and found more bony remains. Some of them fitted into the specimen found by me earlier. We wonder in which way the terrestrial amphibians got into the marine pelagic sediments. Because of presence of the carbonized remnants of wood in the limestones (Fig. 2) we can suppose that amphibians together with pieces of wood where pushed towards sea from land or an island after heavy storm. In 19631 carried out comparative studies in the Eastern Alps, paying attention to the lithological and paleontological analogies between the Triassic of the Tatra Mts and the East-Alpine Triassic (Kotański 1965) and I noticed that marls with intercalation of nodular limestones and cherts occurring in the Chochołowska Valley showed striking resemblance to the Partnach Beds. Their stratigraphical position also corresponds to the position of the Partnach Beds which overlie the Reifling Limestone and are underlain by the Wetterstein Dolomite. Thus, their age should be precisely determined as upper Anisian - lower Ladinian. We can draw conclusion that the labyrinthodont amphibians from the Tatra Mts should be of the same age. This suggestion has been proved by later stratigraphical and paleontological studies. Kulczycki had scheduled further field works for 1964, but in spring of that year he became seriously ill. At that time he told me that he did not intend to continue his studies on the labyrinthodont from the Tatra Mts and asked me to pass the bony material to dr. Teresa Maryańska for further studies. After the death of Kulczycki I asked Maryańska to derive a species name of a new labyrinthodont from Kulczycki's name, we discussed also the derivation of generic name from the Tatra Mts. After a long pause, the mentioned material was studied by Maryańska & Shishkin and described by these authors as a new genus and species - Tatrasuchus kulczyckii - of the cyclotosaurid capitosauroids ( Maryańska & Shishkin 1996, this volume, p. 53-83).

HISTORY OF DISCOVERY AND AGE OF LABYRINTHODONT REMAINS 49 STRATIGRAPHIC, TECTONIC AND PALEOGEOGRAPHIC POSITION OF THE LIMESTONES WITH LABYRINTHODONT REMAINS There was already a long history of opinions concerning the stratigraphic position of the strata in which the labyrinthodonts were found. The opinions changed many times. These strata make a part of the unit which had been generally called the Choć Dolomite. In view of the fact that in many places these strata overlie the Cretaceous marls, which is also seen in the Chochołowska Valley, the Choć Dolomite was thought to be of Cretaceous age (Uhlig 1897). It was not until after the victory of "the nappe revolution" in the Tatra Mts and acceptance of existence of great overthrusts that it become obvious that the Choć Dolomite is also overthrusted from south, consequently its age does not have to be Cretaceous. Based on lithological analogies and also the remains of calcareous algae found in few places in the Inner Carpathians, particularly so at Siwiańskie Turnie (Kuźniar 1913), Goetel (1915) stated that the Choć Dolomite is of Triassic age. Andrusov (1936) conducted research in the frontier range to the west of the Chochołowska Valley and in Orawice. He noticed, that the strata determined as Choć Dolomite were very differentiated and they showed great analogies to the lithostratigraphical members of the East- Alpine Triassic which have been well-known for a long time. He distinguished the Norian Hauptdolomite, Lunz Beds (in the Osobita Group), the Reifling Limestones and the associated Partnach Beds. Guzik (1936) strongly refuted these views on stratigraphy and stated that the Rhaetian age of the horizon regarded by Andrusov as the Reifling Limestones is unquestionable. Guzik's interpretation was presented on the map (1:10 000) of the Polish Tatra Mts (Guzik & Guzik 1958: sheet Furkaska). The apparent recurrence of marls and limestones in the Wielkie Koryciska was found to be due to fold tectonics with anticlines of the Rhaetic limestones and with synclines of the Liassic marls. Guzik's opinion was adopted by Andrusov (1959 a, b) and confirmed once more by Książkiewicz (1977). Guzik in 1959 described the stratigraphical position of two units - Furkaska and Koryciska. According to his interpretation the Rhaetian in the Furkaska unit is represented by two lithological facies: the Kóssen facies with coral and brachiopod fauna and by the limestones facies with cherts similar to the Reifling strata. However, it should be noticed that Guzik was not absolutely sure of his theory and he wrote: "[...] we cannot exclude the possibility that the belt of chert limestones in the Wielkie Koryciska-Kryta syncline is not of Rhaetian age, but may be older [...] It may be possible that Koryciska unit widens to the west from the Chochołowska Valley and contains under the limestones (»Choc Dolomite«) the known Reifling-Partnach elements, the platy dolomites and also the Gutenstein Dolomites of the Anisian-lower Ladinian age [...]" (Guzik 1959: p. 186). The fact that in Drwal's Gully in the limestones with intercalation of marls, ammonites and other fossils were found (Kotański 1961) suggested that the limestones did not belong to Rhaetian but to Middle Triassic. Kotański (1965) wrote on lithological analogies between the Triassic from the Tatra Mts and that from Eastern Alps. Further data concerning the discovered fauna and its Anisian-early Ladinian age were also reported by Kotański in 1967. Some descriptions and stratigraphical and tectonic information were included in the Geological guide book to the Tatra Mts (Kotański 1971). Zawidzka (1970) described a rich conodont fauna, associated with fish remains, scolecodonts, holothurian sclerites, sponge spicules, foraminifers, ostracods and tiny gastropods, from the Reifling Limestones and concluded that such assemblage ranges from the upper Anisian through the lower Ladinian. Most of holothurian sclerites described by Zawidzka (1971) are known from upper Anisian. According to Zawidzka (1972) conodonts and holothurian sclerites occur in the platy limestones and nodular limestones with cherts which are present at the bottom of the marly-shale complex in the Wielkie Koryciska. They are found neither in the marly limestones nor in the brecciated limestones with the carbonized plant detritus of the organodetritic limestones which are intercalated with shales nor in the shales themselves which are represented by the Partnach Beds. Conodonts which were found at the bottom of Partnach Beds belong to the I, II and III Hirsch zone, which means that they belong to Protrachyceras reitzi ammonite zone of the upper Anisian (Illyrian) to lower Ladinian (Fassanian). This span of time fully corresponds with the age of the Reifling Limestones with which the Furkaska limestones were compared. The marly beds lying higher up intercalated with the organodetritic limestones are identical in respect of macrofacies environment with the Partnach Beds from the Bavarian Alps (Zawidzka 1972). The shales which coexist with the organodetritic limestones yielded an extremely rich foraminifer detritus of Frondicularia cf. woodwardi (Zawidzka 1972). The foraminifer assemblage of the upper Anisian age from the Wielkie Koryciska closely resembles those recorded in the different regions of the Alpine-Carpathian geosyncline (Gaździcki & Zawidzka 1973). The foraminifers from the marly Partnach Beds have been described by Alexandrowicz & Szewczyk (1981). The dominant forms there are represented by Nodosaria genus, which have also been recorded in both geosynclinal and epicontinental Triassic deposits. The assemblage comprising Nodosaria genus belongs to Illyrian and occurs below the zone of Glomospira densa (Pelsonian-lower Illyrian). The rich macrofauna derived from the Partnach Beds of the Tatra Mts was described by Kotański (1965, 1967, 1973 a, b, 1974 b). Among the Triassic ammonites Kotański determined the following species: Semiornites lennanus, Arcestes cf. bramantei, Flexoptychites sp., Ptychites sp., Gymnites (?) sp. Apart from ammonites the following nautiloids:

50 ZBIGNIEW KOTAŃSKI Nautilus cf. tintoretti and N. cf. salinańus, and daonellas: Daonella (Daonella) lommeli, D. (D.) cf. taramelli, and D.(D.) tyrolensisparthanensis were determined. Most of the above ammonites and daonellas suggest uppermost Anisian age (Paraceratites trinodosus and Aplococeras avisianus zones), however the occurrence of the Ladinian species Daonella lommeli and Gymnites (?) sp. may witness that the Partnach Beds are transitional between Anisian and Ladinian. The other macrofauna in these beds is composed of pelecypods, numerous small gastropods, brachiopods as Mentzelia cf. mentzeli, Spiriferina cf. fragilis, Aulacothyris angustaeformis, echinoids: Miocidaris sp. and Cidaris sp., numerous thick stems of crinoids, bryozoans, and ostracods. The above mentioned fauna, particularly brachiopods, witness upper Anisian age of the Partnach Beds at Wielkie Koryciska. Thus, the age of the Partnach Beds at Wielkie Koryciska is precisely defined on the basis of ammonites, daonellas, brachiopods, and foraminifers. So, the same age should be applied to the labyrinthodont remains. This age was also indirectly confirmed by the fact that the underlying Reifling strata contain conodonts of upper Anisian age (Zawidzka 1970), but the overlying Wetterstein Dolomite is of the Ladinian age, based on index-fossils as: Teutloporella herculea, T. aequalis and Diplopora annulata dolomitica (Kotański 1973 a). The sequence of these beds is illustrated on Fig. 3. The paleogeographical position of the Partnach Beds is another problem. All the paleogeographical considerations concerning the mountains which show nappe structure, as for example the Tatra Mts, depend on exact determination of tectonic position of the unit to which the beds belong. The Partnach Beds of the Furkaska unit (Guzik 1936,1959) together with the Koryciska unit were incorporated into the Choć nappe. The Rhaetic beds from the Chochołowska and Lejowa Valleys were also included into the same nappe. As a consequence of extensive studies the upper Anisian age of the Partnach Beds in the Wielkie Koryciska Ravine was confirmed. On the other hand the Rhaetian age of the limestones from the Chochołowska 1 Furkaska Fig. 3 Geological section across Furkaska and Wielkie Koryciska units. Triassic of the Upper Subtatric (Strazov) nappe thrusted over the Cretaceous of the Lower Subtatric (Kriżna) nappe. Lower Subtatric (Kriżna) nappe: 1 - Lower Cretaceous marls; Upper Subtatric (Strażov) nappe: 2 - Anisian saccharoid and piaty dolomites, 3 - upper Anisian Reifling Limestones. 4 - Partnach Beds with very rich upper Anisian marine fossils and labyrinthodont bones, 5 - Wetterstein Dolomite; 6 - transgressivc Eocene beds. Fig. 4 Schematic map showing the distribution of the Middle and Upper Subtatric units in the Western Tatra Mts. 1 - Hightatric unit - cristalline core; 2 - Lower Subtatric Kriżna group of nappes; 3 - Middle Subtatric Choć nappe; Upper Subtatric nappe: 4 - Furkaska unit, 5 - Koryciska unit; 6 - transgressive Eocene of the Innercarpathian Podhale basin. and Lejova Valleys is unquestionable and based on numerous index-fossils such as brachiopod Rhaetina gregaria (Guzik 1959), corals (Roniewicz 1974), foraminifers as Triasina hantkeni and Glomospirella cf.friedli (Gaździcki & Zawidzka 1973; Gaździcki 1984) and conodonts, e.g. Misikella posthemsteini (Gaździcki 1984). Taking into consideration the tectonic reasons, the attachment of both stratigraphical members to the same Furkaska unit was impossible. That is why the Partnach Beds were distinguished only in the Furkaska unit (Fig. 3), while the underlying Rhaetic beds together with the dolomites were incorporated into the newly distinguished Siwa Woda unit (Kotański 1973 a, b, c, 1974 a, b), which is overlain by the Furkaska and Koryciska units (Figs 4, 5). On the basis of the facies considerations only the Siwa Woda unit was included into the Middle Subtatric nappe (Choć nappe), whereas the Furkaska and Koryciska units into the Upper Subtatric-Strażov nappe (Kotański 1973 a, b, 1974 a, b, 1976,1977, 1979 a, b, c, 1985 a, b). The attachment of the Furkaska unit to the Strazov nappe, as well as the presence of the Upper Subtatric nappe in the Tatra Mts was questioned e.g. by Passendorfer (1978 a, b), Mahel (1979), Michalik & Gaździcki (1980), Gaździcki & Michalik (1980). The above authors included all these units into the Choć nappe. Recently, opinion concerning the attachment of the Furkaska and Koryciska units into the Upper Subtatric (Strazov) nappe has become almost commonly accepted (Lefeld 1979, 1984:

HISTORY OF DISCOVERY AND AGE OF LABYRINTHODONT REMAINS 51 sw Furkosko 14005 NE I WSW Wielkie Zgmcasko 1339,6 Mote Zomcrysko Wie k, Koryciska Mflfe н JEJ 2 3 Fig. 5 Koryciska Siwiańskje Turrne 1069,9 rtrrvt? Geological section along the frontier crest on the western side of the Chochołowska Valley. Lower Subtatric (Kriżna) group of nappes (pb - Bobrowiec partial nappe, Parządczak scale): 1 - Jurassic limestones and Neocomian marls; Middle Subtatric (Choć) group of nappes (psiw - Siwa Woda partial nappe): 2 - Upper Triassic dolomites and limestones belonging mainly to the Upper Triassic Hauptdolomite. including Rhaetian of Kossen facies, in normal position; Upper Subtatric (Straiov) group of nappes (Furkaska-Koryciska partial nappe, łf - Furkaska scale, łk - Koryciska scale): 3 - middle Anisian Ramsau Dolomite, 4 - upper Anisian Reifling Limestone, 5 - upper Anisian-lower Ladinian Partnach Beds, 6 - Ladinian-lower Carnian Wetterstein Dolomite; 7 - transgressive Eocene beds. Fig. 6 ENE FATRIC TATRIC VEPOR PIENINY Schematic palinspastic section through Internal Carpathians during the Middle Triassic time. The northern carbonate platform is separated from the southern one (North Gemer and Strazov) by basinal Choć sediments. Forereef Debris and Forereef Internbreccias Debris Flows Pro*imol Turbidltes Initial Reef Growth Vodose Pisolitic Crusts Backreef Sands ond Coral Heads Reef Knolls Reef Debris Fig. 7 S solenoporacean Corol Biocoenoses TY» Tubiphytes Zone С» Colcisponge Hydroioan Corol Community Environmental model of the platform-basin setting (Kotański 1985). T Dosyclodoceon Faciei Thecosmilia Thaumatoporella Biocoenoses Wieczorek 1984; Iwanow & Wieczorek 1987; Wieczorek 1989). Explanation of the tectonic-facies attachment of the Furkaska unit to the Upper Subtatric nappe is of primary importance for determination of the paleogeographical position of the beds with labyrinthodonts. The Strazov nappe has a distinctive southern character and occupied definite position in the Tethys geosyncline. It is a counterpart of the Upper Austro-Alpine (Tyrolian) nappes from the Eastern Alps and also some Transylvanian nappes of the Eastern Carpathians. The isopic zones are spread over great distances in the Tethys (Ryszkiewicz 1979). The Hightatric-Kriżna- Vepor zone made during the Middle Triassic the northern carbonate platform and was separated by basin deposits of the Choć zone from the southern carbonate platform, which was composed of the North-Gemer and Strazov zones (the latter was situated on the border of the Choć zone, Fig. 6). To the south from the Gemer zone in the Biikk Mts in Hungary there was the Triassic continental rift, which constituted a nucleus of the later oceanic rift. From there originated the volcanic ash which was found in the Partnach Beds in the Wielkie Koryciska Ravine (Kotański 1979 a, b, 1985 a, b; Lefeld 1979). An environmental model of the jointing between the carbonate platform of the Wetterstein limestones or dolomites and the basin of the Partnach Beds is illustrated on Fig. 7. Kotański wrote: "Findings of the labyrinthodont amphibians, living in the terrestial environment are very rare, but the discovery of very-well preserved skull of labyrinthodont in the marine sediments of the Upper Subtatric nappe in the Tatra Mts is due to an unusual coincidence" (Kotański 1977: p. 496). The labyrinthodonts had to live on coral island which was built of the Wetterstein limestones and dolomites - probably it might be an atoll (Kotański 1985 a: p. 551). A few individuals of them were drifted by ocean current on a piece of wood and they were deposited in the basin of the Partnach Beds. Probably, this situation was caused by a tropical storm; this kind of storms had to be frequent in the middle part of the Tethys ocean. Zbigniew Kotański Państwowy Instytut Geologiczny ul. Rakowiecka 4 02-517 Warszawa, Poland

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