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The Middle Miocene of the Fore-Carpathian Basin (Poland, Ukraine and Moldova)

Wysocka A., Radwański A., Górka M., Bąbel M., Radwańska U., Złotnik M.

Acta Geologica Polonica

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2016

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Vol. 66, no. 3

351--401

EN

Studies of Miocene sediments in the Fore-Carpathian Basin, conducted by geologists from the University of Warsaw have provided new insights on the distribution of the facies infilling the basin, particularly in the forebulge and back-bulge zones. The origin of the large-scale sand bodies, evaporitic deposits and large-scale organic buildups is discussed, described and verified. These deposits originated in variable, shallow marine settings, differing in their water chemistry and the dynamics of sedimentary processes, and are unique with regard to the fossil assemblages they yield. Many years of taxonomic, biostratigraphic, palaeoecologic and ecotaphonomic investigations have resulted in the identification of the fossil assemblages of these sediments, their age, sedimentary settings and post-mortem conditions. Detailed studies were focused on corals, polychaetes, most classes of molluscs, crustaceans, echinoderms, and fishes.

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Badenian (Middle Miocene) echinoids and starfish from western Ukraine, and their biogeographic and stratigraphic significance

Radwański A., Górka M., Wysocka A.

Acta Geologica Polonica

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2014

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Vol. 64, no. 2

207--247

EN

Echinoderms from the Badenian (Middle Miocene) of the Fore-Carpathian Basin of western Ukraine are facies restricted. The Mykolaiv Beds, stratigraphically older, yielded the starfish Astropecten forbesi (complete skeletons), two genera of sand dollars (Parascutella, Parmulechinus), and numerous other echinoids of the genera Psammechinus , Echinocyamus, Spatangus, Hemipatagus, Echinocardium, Clypeaster, Echinolampas, and Conolampas. The stratigraphically younger, calcareous Ternopil Beds yielded Eucidaris (complete coronae, isolated spines), Arbacina , Brissus, and Rhabdobrissus. Sixteen species of echinoids are distinguished and/or commented. A new brissid, Rhabdobrissus tarnopolensis sp. nov., is established. A mass occurrence of some species (Psammechinus dubius and Hemipatagus ocellatus) contrasts with that of mass aggregations (sand dollars and Echinocardium leopolitanum) by dynamic events in selected layers of proximal tempestites. Of special note is the occurrence of very small specimens, interpreted as juveniles (‘babies’) having been swept out of their restricted biotopes (‘nurseries’). Some species hitherto regarded as of Early Miocene age, and the problem of their persistence beyond the Fore-Carpathian Basin and/or migration into that basin during the Middle Miocene transgression are discussed.

3

Mykolaiv Sands in Opole Minor and beyond : sedimentary features and biotic content of Middle Miocene (Badenian) sand shoals of Western Ukraine

Wysocka A., Radwański A., Górka M.

Geological Quarterly

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2012

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Vol. 56, No 3

475--492

EN

The Mykolaiv Sands are a huge lithosome of Middle Miocene (Badenian) age, accommodated within the Fore-Carpathian Basin in the Western Ukraine. Typically developed in the area of Opole Minor, it spreads across adjacent regions of Opole to cover an area of about 1300 km2. The varied sedimentary structures and ubiquitous burrows, indicate their development as a stack of sand shoals or related bodies, up to a few tens of metres thick, some of which temporarily reached sea level. Amidst the shoals, storm scours intermittently formed channel-like infills, some with residual lags at the base. The reversed density stratification and/or an increasing gravity gradient involved mass movements, some of which may have been triggered by seismic shocks focused at the shore or the adjacent hinterland of Podolia and Volhynia. Special attention is paid to the diverse fossils, all taphonomically filtered (aragonite shells and chitinous carapaces being lost), but which locally are mass-aggregated. They typify particular sand sets/bodies, to form allochthonous assemblages, some members of which (the cirripedes Scalpellum and Creusia, the shark Hemipristis, the ray Myliobatis) are newly recognized in the Ukrainian part of the Fore-Carpathian Basin. The others enrich considerably the faunal content of the Middle Miocene (Badenian) Paratethyan basins, either in terms of taxonomic diversity, or the eco-taphonomy of selected taxa (the starfish Astropecten, diverse echinoids). The whole faunal content of the Mykolaiv Sands may owe its profuse development to the global Middle Miocene Climatic Optimum of early Badenian age.

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Miocene burrows of the Ghost Crab Ocypode and their environmental significance (Mykolaiv Sands, Fore-Carpathian Basin, Ukraine)

Radwański A., Wysocka A., Górka M.

Acta Geologica Polonica

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2012

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Vol. 62, no. 2

217-229

EN

The huge lithosome of the Middle Miocene (Early Badenian) Mykolaiv Sands, developed at the external margin of the Fore-Carpathian Basin in western Ukraine, is recognized to represent a shallowing-up sequence. Special attention is paid to burrows of the Ghost Crab Ocypode which are pantropical in present-day littoral habitats. In the Stratyn section, burrows of this type become a crucial tool in the interpretation of basin bathymetry, which starts from distal offshore depths, through the foreshore, to the backshore where the Ocypode burrows record a temporary break in sedimentation. Lithification of the sand layers and the Ocypode burrows subsequently progressed in beachrock mode. The Stratyn section demonstrates that the development of submerged shoals and/or emergent parts, throughout the huge mass of the Mykolaiv Sands, is probably responsible for their great variation in thickness in western Ukraine, which has long proved difficult to explain.

5

'Entobia balls' in the Medobory Biohermal Complex (Middle Miocene, Badenian; western Ukraine)

Radwański A., Wysocka A., Górka M.

Acta Geologica Polonica

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2011

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Vol. 61, no. 3

265-376

EN

The peculiarly shaped 'Entobia balls', from the Middle Miocene (Badenian) Medobory Biohermal Complex, western Ukraine, are a maze of moulds of clionid sponge borings belonging to the ichnogenus Entobia Bronn. The ichnospecies recognized (Entobia geometrica, E. paradoxa, E. cateniformis, E. laquea) are ascribed to the activity of two extant zoospecies, Cliona vastifica Hancock and C. celata Grant. Their habitat was provided by thick-walled shells of the bivalve Chama gryphoides garmella De Gregorio, the shells of which were drilled through completely. Some small patches of borings are compatible with those of the extant zoospecies Cliona viridis (O. Schmidt).

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Queues of blind phacopid trilobites Trimerocephalus: A case of frozen behaviour of Early Famennian age from the Holy Cross Mountains, Central Poland

Radwański A., Kin A., Radwańska U.

Acta Geologica Polonica

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2009

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Vol. 59, no. 4

459-481

EN

An assemblage of blind phacopid trilobites of the genus Trimerocephalus McCoy, 1849, representing either the species Trimerocephalus mastophthalmus (Reinhard Richter, 1856) or its allies (possibly, a new species), from an Early Famennian (Early Marginifera Zone) marly sequence of the Holy Cross Mountains, Central Poland, is composed of well organized single-file queues. The trilobites in the queues appear almost intact, having been preserved in the position they kept when forming the queues, and are interpreted showing migratory behaviour known in various present-day arthropods, but unreported from the fossil state. This queuing was induced by environmental stress caused by a dramatic drop in sea level, temporarily leading to emersion. The preservation of the queues at omission horizons is thus ascribed to a mass mortality event, caused by subaerial exposure. The trilobites were suffocated and fossilized in a mortal spasm, and finally blanketed by calcareous ooze when inundated at a highstand. The assemblage of trilobite queues represents a unique example of frozen behaviour and a snapshot of the geological past.

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A unique habitat of endolithic biota: hurricane-induced limestone rubble in an Albian sand-mass of the Cracow Upland, southern Poland

Marcinowski R., Radwański A.

Acta Geologica Polonica

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2009

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Vol. 59, no. 4

505-521

EN

Peculiarly shaped, relatively large (up to 30 cm in diameter) concretions of quartzitic sandstone occur in a single horizon of Upper Albian loose sands in the Cracow Upland, southern Poland. They are characterized by hollow interiors adorned with mass-aggregated moulds of the borings of diverse sponges, polychaetes and bivalves. These moulds represent the siliceous filling of borings in limestone clasts that had been subject to dissolution, leaving a hollow within the concretion that had formed around them. Synsedimentary block-faulting and jointing affected the Jurassic limestone-basement, causing the uplift of a local horst (the Glanow Horst), to within the littoral zone so that it became exposed to abrasion. It is inferred that a hurricane or catastrophic storm surge swept limestone debris fallen from the cliff out to the sandy offshore, where nucleation of soluble silica was presumably favoured by the decay of the soft tissue of live or dead rock-borers. After filling the emptyo borings and solution of the limestone clasts, the nucleation progressed intensely, finally completed by precipitation of siliceous sinter in the hollow interiors of some of the concretions during subsequent diagenesis and/or epigenesis.

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Eco-taphonomy of mass-aggregated giant balanids Concavus (Concavus) concavus (DARWIN, 1854) from the Lower Pliocene (Zanclean) of Rafina near Pikermi (Attica, Greece)

Radwańska U., Radwański A.

Acta Geologica Polonica

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2008

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Vol. 58, no. 1

87-103

EN

The large- to giant-sized balanids, mass-aggregated in a tempestite of the Lower Pliocene (Zanclean) section at Rafina near Pikermi, Greece, represent a single species, Concavus (Concavus) concavus (DARWIN, 1854) [non BRONN, 1831], the taxonomy of which is revised. The peculiarly shaped forms .raphanoides. and .scutorum. are ecophenotypes of this species; the same applies to the .tulipiformis. specimens reported previously from this section. Discussed are dynamic events controlling the life and death conditions in the nearshore (offshore) environment of Rafina, where the giant specimens of Concavus (Concavus) concavus (DARWIN, 1854) flourished through several successive generations. As opportunistic species they adopted the r-selection reproduction strategy in order to dominate over other biota. The intermittent action of high-energy agents was responsible for the production of balanid-shell hash that involved taphonomic feedback. This enabled further colonization of the biotope and the growth of multiphased (bouquet-like and pyramidal) clusters. The demise of the monospecific balanid communities is ascribed to a heavy storm which stirred-up the whole balanid-bearing sequence, to produce a proximal tempestite. This final burial and subsequent depositional lull in the Rafina environment was favoured by a temporary deepening of the whole Lower Pliocene (Zanclean) basin beyond the bathymetric range in which the balanid population could survive.

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Middle Miocene coralgal facies at Maksymivka near Ternopil (Ukraine) : a preliminary account

Radwański A., Górka M., Wysocka A.

Acta Geologica Polonica

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2006

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Vol. 56, no. 1

89-103

EN

A peculiar coralgal facies is recognized in the Lviv-Ternopil region, Ukraine, from the northern shores of the Middle Miocene (Badenian) Fore-Carpathian Basin. Its complex structure is dominated by algal buildups composed of interfingering red-algal (lithothamnian) colonies and blue-green-algal crusts, associated locally with numerous hermatypic corals (Tarbellastraea reussiana, Porites vindobonarum prima), either isolated, or overlapping each other. The holes amidst, and the crevices in, the buildups are filled with coarse bioclastic sediment (shell-grit), burrowed commonly by crustacean decapods (alpheid shrimps). The alpheid burrows, filled with coarser or finer shell-grit, served frequently as taphonomic traps for crustacean decapods (squat lobsters and crabs) and echinoids.Special attention is paid to the activity of rock-boring bivalves (Jouannetia semicaudata, Lithophaga lithophaga) in coralgal buildups and/or in particular coral colonies, some of which are redeposited, and riddled densely by bivalve borings. Emphasis is given to the environmental significance of alpheid shrimps, the tiered burrows of which are recorded in the Fore-Carpathian Basin for the first time. Crustacean decapods and echinoids are systematically studied. A comparison of the studied coralgal facies with others of the Lviv-Ternopil region, and those from the territory of Poland, indicates their faunistic and biogeographic identity.

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Kolonie korali z muszlowców ostrygowych i oolitów dolnego kimerydu Małogoszcza

Radwański A., Roniewicz E.

Volumina Jurassica

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2005

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Vol. 3, no. 1

97--107

EN

Scleractinian corals occurring scarcely in the Lower Kimmeridgian Actinostl'eon (=Lopha, =Alectryonia) shellbeds at Malogoszcz in the Holy Cross Mountains, Central Poland, are represented by abraded colonies densely riddled by rock-boring bivalves (Lithophaga inclusa Phillips, GastTOchaena sp.) and polychaetes (potamilla sp.). The taxonomically recognised specimens include C01nplexastrea bUJ'gundiae (de Blainville, 1830), D 'imorphocoenia sp., Ovalastrea caryophylloides (Goldfuss, 1826), and Thamnasteria graeilis (Miinster, 1826). AlI colonies are preserved in the form of hollows, the wall s of which bear moulds of coral calyces, and of bivalve and polychaete borings. Taking into account the structural features of shellbeds and their faunistic content (uprooted crinoids Apioerinites, dug-out deeply-infaunal bivalves), stormy agitation is postulated as a basic agent responsible for damaging AeUnostreon communities, and their associates. The studied corals are thought to have lived aside the Aetinostreon gardens, up on a muddy bottom, from where they have been stirred-up during the storm cataclysm, having been then abraded and riddled by rock-borers repeatedly until laid down in a shellbed and transferred in to the fossil record. The extremely shallow-water conditions, under which the ostreid Actinostreon has lived, suggest the typically opportunistic nature or the associated corals, the same as of Ovalastrea caryophylloides (Goldfuss, 1826) from oolitic shoals, and the only colony of which completes the coral assemblage of Malogoszcz. The opportunism or ali these corals differs them from the habit of hermatypie forms from coeval and Oxfordian patch-reef communities of the Holy Cross Mountains (ef. Roniewicz & Roniewicz 1968, 1971).

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Gąbki wapienne Elasmostoma lomes W. Mliller, 1984, z dolnego kimerydu Karsów koło Korytnicy, ich ekofenotypia i tafonomia

Radwański A.

Volumina Jurassica

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2005

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Vol. 3, no. 1

109--121

EN

Calcareous sponges Elastostoma !'omes W MUller, 1984, from the Lower Kimmeridgian marly onkolitic limestones exposed at Karsy in the Holy Cross Mountains, Central Poland, are studied in terms of their settlement, growth, behaviour, and resulting phenotypy. The lichen-shaped auriculate specimens are interpreted as growing upwardly, at a higher angle to the bottom surface, in order to involve passive flow carrying suspended nutrients up on which they fed. Unstable bottom and hydrodynamie conditions, under which the growth of sponges progressed, we re basic prerequisites that controlled ecophenotypic variability of the studied specimens, A peculiar case of regeneration is discussed, to postulaie the skeleton parts having been lithified during the sponge's life, and healed after an accidental injury. Aviolent event that caused catastrophic burial of the studied sponge assemblage is ascribed to high-energy agents, most likely of storm origin.

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Myzostomid and copepod infestation of Jurassic echinoderms: A general approach, some new occurrences, and/or re-interpretation of previous reports

Radwańska U., Radwański A.

Acta Geologica Polonica

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2005

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Vol. 55, no. 2

109-130

EN

A study of diverse cysts developed on fossil echinoderms from Poland results in the recognition that these on Late Jurassic crinoid stems are attributable to the life activity of myzostomidan polychaetes, and those on Middle and Late Jurassic echinoids, to the activity of copepod arthropods. A review of formerly reported cysts of coeval age from Europe and western Asia permits the distinguishing of several types that differ in shape and/or location on the echinoderm skeleton. Although the studied cysts qualify as trace fossils (which require a separate ichnotaxonomy), their ethological and ecological characteristics are presented in terms of interspecific parasite-host relationships. The classical interpretation of VON GRAFF (1885) is affirmed for myzostomidan endocysts in crinoid stems, whereas for echinoid tests a new interpretation is offered for large exocysts ("Halloween pumpkin-mask" type) as having been induced by copepods,comparable in their ethology to those on present-day biota (hydrocorals) other than echinoderms. A copepod attribution by MERCIER (1936) of cysts (Castexia type) on some Middle Jurassic collyritid echinoids from France is fully accepted. This is supplemented by some new finds in Poland, a re-study of the Castexia cysts from France, and a re-interpretation of former reports from the literature. Eco-ethological consequences of the location of copepods in the ambulacral and peristomial parts of cidaroid and hemicidaroid echinoids are discussed; larval settling apparently took place at the tubefeet pores and gonopores, through which the copepod larvae reached the echinoid.s interior and began to parasitize it. Attribution of the discussed cysts to copepods yields, consequently, an extension of the stratigraphical range of the class Copepoda H. MILNE-EDWARDS, 1840, to the Early Jurassic.In POSTSCRIPT, suggested is the bald-sea-urchin disease to have caused some lesions in the collyritid echinoids (Middle Jurassic: Callovian) from France.

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Systemy piętrowych nor krewetek-alfeuszy i ich znaczenie środowiskowe w oksfordzie i kimerydzie Gór Świętokrzyskich

Radwańska U., Radwański A.

Volumina Jurassica

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2004

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Vol. 2, no. 1

113-130

EN

To the activity of alpheid shrimps genus Alpheus Weber, 1795) ascribed are the tiered burrows of a gridlike appearance from Lower Kimmeridgian oolitic shoals and Middle Oxfordian nearshore micritic limestones of the Holy Cross Mountains, Central Poland. The burrow networks are confined to beds of the soft or hard bottom type, the upper parts of which are more or less deeply truncated, to indicate erosional events of storm agitation. At low stand, the open burrows served as traps for solutions derived from the nearby hypersaline lagoons of the sabkha type, to cause precipitation either of dolomite, or of silica gel. At high stand, the open burrows, exemplified by the Małogoszcz section (Lower Kimmeridgian), became taphonomic traps and/or crevice habitats for diverse biota, the echinoderms in particular, to form their graveyards (EchinodermenlagerstŹtten). In these, represented are echinoids (tests, some spine-coated, all either empty, or sediment-filled; broken tests and their fragments, spines) stalkless crinoids (cusps, centrodorsals, radials, brachials, cirrals), stalked crinoids (columnals, pluricolumnals), starfish (marginalia, ambulacral plates), and ophiuroids (vertebrae, arm plates). Eco-taphonomic pathways for particular echinoderms (21 taxa taxonomically recognised) are interpreted since their death to burial in open burrows. Spine-coated echinoids were entrapped alive, others were swept into during successive storms which acted as a lethal agent. The storms, catastrophic for echinoderm communities, have prevailed through a longer timespan, when the alpheid-burrowed shoal evolved from the soft bottom to the hard ground colonized by a successive echinoderm community dominated by stalked crinoids.

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Chore i okaleczone zwierzęta jako ofiary pasożytów i drapieżców w jurze Polski - przegląd stanu wiedzy

Radwańska U., Radwański A.

Volumina Jurassica

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2004

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Vol. 2, no. 1

99-112

EN

The parasitism of, and predation upon, the diverse Middle and Late Jurassic invertebrates of Poland, personally recognised by the authors, are reviewed. All cases are discussed either in the biological (anatomical, ethological), or ecological terms, to show the mode of infection, or injury, and the relationship between the engaged taxa. The preys to parasites are exemplified by the prosopid crabs infested by bopyrid isopods, the crinoids infested by myzostomid polychaetes (both disk-shaped, and wormlike), and the echinoids attacked by copepod arthropods involving either swellings of spines, or gall-shaped cysts upon the test outerly. Of traumatic events, discussed is regeneration of injuries in the belemnite hooked guards, and in the ammonite shells of distorted ribbing. The pearl-like structures in belemnite guards (the “belemnite pearls”) are interpreted as caused by a tiny parasite encapsulated during further growth of the belemnite. Heaps of ammonite shell hash are thought to represent the ,“kitchen middens” of a larger predator which has feasted upon the fleshy tidbits alone, the beaten shell having been left. The post mortem damage of shells is remarked (taphonomic feedback and/or aftermath) to be distinguished from that one acted in shells of alive specimens of the Middle and Late Jurassic of Poland.

15

Farewell to Świniary sequence of mass-aggregated, spine-coated echinoids Psammechinus and their associates (Middle Miocene; Holy Cross Mountains, Central Poland)

Radwański A., Wysocka A.

Acta Geologica Polonica

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2004

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Vol. 54, no. 3

381--399

EN

The Świniary sand-pit (Middle Miocene; southern slopes of the Holy Cross Mountains, Central Poland), which has yielded innumerable fossils of various kinds over the last fifty years, is revisited. Most significant there were mass accumulation of a single echinoid species, Psammechinus dubius (L. AGASSIZ, 1840), tests of which preserved spine canopies and Aristotle.s lanterns in position. Other echinoids represented were rare, e.g. Spatangus austriacus LAUBE, 1871, 'Giant Psammechinus sp.', Echinocardium peroni COTTEAU, 1877, and Echinocyamus pusillus (O.F. M LLER, 1776). The pit has since been abandoned and recultivated entirely; in view of this an updated review of, and supplement to, previous data on the Świniary biotic assemblages, and their environmental living conditions, is here presented. The mass accumulations of Psammechinus dubius are interpreted to have been of storm origin, i.e. mass transportation and burial of live, or freshly dead, specimens.Hinted at is a predatory activity of the starfish Astropecten forbesi HELLER, 1858, upon juveniles of Psammechinus dubius absent from the storm-related thanatocoenoses. Associated with echinoderms are, amongst other groups, verrucid barnacles Verruca sp. and inarticulate brachiopods Discinisca leopolitana (FRIEDBERG, 1921) derived from eulittoral habitats, as well as pieces of terrestrial amber swept out of the shore. A general shallowing-up trend in the Świniary sequence is recognised, as indicated by environmental conditions changing from deeper offshore to proximal sublittoral, possibly shallow subtidal.

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Wapienie pasiaste dolnego kimerydu Gór Świętokrzyskich: przykłady prolapsji, diastazy oraz systemu nor piętrowych

Radwański A.

Volumina Jurassica

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2003

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Vol. 1, no. 1

71-76

17

Siedlisko życia, miejsce pogrzebania oraz epibionty ramienionogów Sellithyris subsella (Leymerie, 1846) z muszlowców dolnego kimerydu Małogoszcza

Radwańska U., Radwański A.

Volumina Jurassica

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2003

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Vol. 1, no. 1

85-92

18

Rezidualne brykiety kolapsyjne w pakietach margli górnego kimerydu Małogoszcza

Radwański A.

Volumina Jurassica

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2003

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Vol. 1, no. 1

77-84

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The Jurassic crinoid genus Cyclocrinus D'ORBIGNY, 1850 : still an enigma

Radwańska U., Radwański A.

Acta Geologica Polonica

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2003

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Vol. 53, nr 4

301-320

EN

A rich collection of isolated columnals and fragmentary pluricolumnals, varying considerably in size, shape, and sculpture, from the Upper Oxfordian of the Couiavia region (northwestern Central Poland), is the basis for a critical discussion of the crinoid genus Cyclorinus and its species. These are all based on dissociated columnals with tuberculate articular faces, and all with similar if not identical arrays of tubercles. The collection studied contains tuberculate columnals, as well as those with plain articular faces, which acquire tuberculation when etched with acids. This indicates the corrosional nature of tuberculation in the newly established species, Cyclocrinus couiavianus sp.nov., and in all other Cyclocrinus material. The structure of fragmentary pluricolumnals, whose arching and branching are reminiscent of root systems of some bourgueticrinids, would suggest that all Cyclorinus material represents nothing else but modified radicular cirrals of unrecognizable members of the order Bourgueticrinida SIEVERTS-DORECK, 1953, rather than Cyrtocrinida or Millericrinida as previously assumed.

20

Triassic brittlestar beds of Poland : a case of Aspiduriella ludeni (v. HAGENOW, 1846) and Arenorbis squamosus (E. PICARD, 1858)

Radwański A.

Acta Geologica Polonica

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2002

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Vol. 52, nr 4

395-410

EN

Exemplified by a new occurrence site of mass-aggregated Aspiduriella ludeni (v.HAGENOW, 1846) from the Lower Muschelkalk (lowest Anisian) sequence of the Holy Cross Mountains, Central Poland, discussed are life conditions of ophiuroids in that region of Muschelkalk Basin (Middle Triassic) in Central Europe. Postulated is their fast deposition (burial) by the rip-current transport of alive, possibly torpid, and/or newly specimens captured by stormy agitation from their shallow subtidal and/or intertidal habitats. The palaeogeographical setting of the Holy Cross region is compared to that of the Silesia region in southern Poland, highly influenced till the early-Middle Triassic by oceanic waters of the Tethyan Realm. Shortly reviewed and/or discussed are the formerly known ophiuroid occurrences in Silesia, to be supplemented by a mass-aggregated case of Arenorbis squamosus (E.PICARD, 1858). Commented are also other ophiuroid aggregations reperted from the Muschelkalk Basin, and whose occurrence is considered in terms of their environmental conditions controlled both by physical (fluctuation of water parameters, and its dynamics) and biotic (low predation stress) agents. A biogeographical significance of the Aspiduriella during Triassic is outlined, to suggest its Muschelkalk-refugee provenance in the late Middle and Late Triassic of the Tethyan Realm.