(South Ern Po Land)? a Crit I Cal Over View of Stra Tig Ra Phy and Fa Cies

Home , Paleorhinus, Parasuchus, Rhynchosaur, Rutiodon

Annales Societatis Geologorum Poloniae (2015), vol. 85: 587–626. doi: http://dx.doi.org/10.14241/asgp.2015.037

HOW MANY UPPER TRIAS SIC BONE-BEARING LEVELS ARE THERE IN UP PER SILESIA (SOUTHERN POLAND)? A CRIT I CAL OVER VIEW OF STRA TIG RA PHY AND FACIES

Joachim SZULC1, Grzegorz RACKI2, 3, Karol JEWU£A1 & Jan ŒRODOÑ2

1 Insti tute of Geolog i cal Sciences, Jagiellonian Univer sity, Oleandry 2a, 30-063 Kraków, Poland; e-mail: [email protected] 2 Insti tute of Geolog i cal Sciences, Polish Academy of Sciences, Research Centre in Kraków, Senacka 1, Kraków, Poland; e-mail: [email protected] 3 Faculty of Earth Sciences, Univer sity of Silesia, Bêdziñska 60, 41-200 Sosnowiec, Poland; e-mail: [email protected]

Szulc, J., Racki, G., Jewu³a, K. & Œrodoñ, J., 2015. How many Upper Tri assic bone-bear ing lev els are there in Up per Silesia (south ern Po land)? A crit i cal over view of stra tig ra phy and fa cies. Annales Societatis Geologorum Poloniae, 85: 587–626.

Abstract: At least three widely sep a rated bone-bear ing in ter vals in the Up per Tri as sic suc ces sion of Upper Silesia, ranging in age from the Carnian to Rhaetian (i.e., in the in ter val of 25 Ma), are pre sented in pa pers by the Warsaw re search group, led mainly by Jerzy Dzik and/or Grzegorz NiedŸwiedzki. The stratigraphic ar guments are re viewed for the verte brate lo cal i ties stud ied so far, in partic ular for the well-known middle Keuper sites at Krasiejów and Lipie Œl¹skie, to show that the pre vi ously pro posed age as signments are still in ad e quately doc u mented and ques tion able. This un re li abil ity is ex em pli fied by the evolv ing strati graphic cor re la tion of the frag men tary Silesian sec tions (8–18 m thick) with in for mal subsurface units from cen tral-west ern Po land and with the Ger man stan dard suc ces sion, ul ti mately not cor rob o rated by com par i son with the com pos ite ref er ence suc ces sion of the Up per Silesian Keuper, in clud ing new pro files (ca. 260 m thick) from the WoŸniki K1 and Patoka 1 wells. Based on a multidisciplinary strati graphic study cov er ing con sistent litho-, bio-, climato- and chemostratigraphic pre mises, focused on the re gional ref er ence sec tion, two bone-bed lev els only are rec og nized in the Patoka Marly Mudstone-Sand stone Member (= Steinmergelkeuper) of the Grabowa Forma tion, not very dif fer ent in age (Classopollis meyeriana Palynozone; prob a bly IVb Subzone): (1) the lo cal ized Krasiejów bone brec cia level (early Norian in age) in the Opole re gion, and (2) the far more widely distrib uted Lisowice bone-bear ing level (middle Norian) in a vast al lu vial plain (braided to anastomosing river system) dur ing the Eo-Cim mer ian tec tonic-plu vial ep i sode. As a con se quence of the prin ci pal un cer tain ties and con tro ver sies in Up per Tri as sic ter res trial stra tig ra phy, this is still a some what pre lim i nary in fer ence. Typ i cal skel e tal con cen tra - tions of a com bined hy drau lic/sedimentologic type, re lated to flu vial pro cesses, are common in the Up per Silesian Fos sil-Lagerstätten, al though fac tors gov ern ing pres er va tion are prob a bly im por tant, as well.

Key words: Bone beds, lithostratigraphy, palynostratigraphy, chemostratigraphy, taphonomy, Grabowa For mation, mid dle Keuper, Upper Silesia.

Manu script re ceived 25 Au gust 2015, ac cepted 9 De cem ber 2015

IN TRO DUC TION

The Tri as sic, as the pe riod between the two great ex tinc - 2010; com pare Szulc, 2007c). The tetrapod local ities have tions, was char ac ter ized by the sub stan tial re con struc tion of major cogni tive impor tance for the evolu tion of many im - con ti nen tal tetrapod com mu ni ties (Benton, 2004). Af ter the portant groups of amphib i ans and reptiles (Dzik et al., 2000, discov ery of very rich bone accu mu la tions in southern Po- 2008a; Dzik and Sulej, 2007; Sulej and NiedŸwiedzki, land at Krasiejów vil lage, near the town of Ozimek, and at 2010; NiedŸwiedzki, 2012; NiedŸwiedzki et al., 2014), and Lisowice village (Lipie Œl¹skie clay pit), near the town of even mammals (Œwi³o et al., 2014). Accord ingly, it seems Lubliniec (Dzik et al., 2000, 2008a, 2008b), a new and at- hard to overes timate the impor tance of the Silesian ‘grave- trac tive sci en tific per spec tive sur pris ingly emerged for pre - yards’ with refer ence to the study of Late Trias sic evo lu - viously abandoned studies of the Polish Keuper (Racki, tionary patterns, exem pli fied by the still obscure dawn of di- 588 J. SZULC ET AL.

A more com pre hen sive pre sen ta tion of var i ous geo log i cal, sedimentological, geo chem i cal and strati graphic as pects of the middle Keuper tetrapod local i ties is given in a regional context by Œrodoñ et al. (2014), Szulc and Racki (2015), Szulc et al. (2015) and Fija³kowska-Mader et al. (2015).

STRATI GRAPHIC AND FA CIES BACKGROUND

The Upper Tri assic succes sion of Silesia belongs to the Keuper Group (Fig. 2), and, in palaeogeographic terms, cor- Fig. 1. A. Sche matic geo log i cal map (af ter Bardziñski and responds to the marginal part of the Germanic Basin (e.g., Chybiorz, 2013), showing the lo ca tions of the lo cal i ties stud ied in Beutler and Nitsch, 2005; Feist-Burkhardt et al., 2008; Upper Silesian region (out crops and bore holes; for de tail see Szulc Bachmann et al., 2010). Varie gated, fine-grained clastics et al., 2015), and B. lo ca tion of the stud ied re gion and the key and locally evaporites (mostly gypsum pseudomorphs) and bore hole in Po land. carbon ates are the most common deposit type and the records of ephem eral-lake and evolving flu vial sys tems devel - nosaur devel op ment (see Dzik, 2003a; Dzik et al., 2008a; oped under mostly arid to semi-arid cli mate condi tions with Brusatte et al., 2010; Nesbitt et al., 2010; Irmis, 2011; Nie- some pluvial inter ludes (Pieñkowski, 1988; Szulc et al., 2006, dŸwiedzki et al., 2012, 2014; Benton et al., 2014), as well as 2015; Szulc, 2007a). the sup posed bi otic turn overs and biodiversity cri ses, espe - The largely monot o nous and fossil-poor mudstone- cially at the Carnian-Norian boundary (Benton, 2004; Pieñ- claystone succes sion, up to 400 m thick in northern Silesia kowski et al., 2014). and almost without more distinc tive marker hori zons, in its Dur ing 15 years of in ten sive in ves ti ga tion, five ver te- facies de vel op ment dif fers some what from the clas si cal Ger - brate sites in to tal have been de scribed across Silesia in the manic Keuper, in par ticu lar in the oc cur rence of the palu- Up per Tri as sic suc ces sion (Fig. 1), sup ple mented by one strine carbon ates of the WoŸniki Limestone (see Fig. 2), basal Keuper, though still Middle Trias sic lo cal ity (Mie- a unique Keuper facies linked to a spring zone of deeply cir - dary; Sulej et al., 2011b; Fig. 2). Next, the bone-rich site re- culat ing groundwa ter that surfaces along the Kraków–Lubli- cently discov ered in 2014 in Zawiercie town was reported niec master frac ture (Szulc et al., 2006). Since the 19th by Rafa³ Piechowski (in Malinowska, 2015). Known since century, the mudstone-claystone rocks were in many places Römer’s (1867, 1870) papers, these rich fossil assem blages in small, local brickyards and thus they crop out in several of var ied tax o nomic com po si tion in clude both di verse scat tered clay pits. Strati graphic corre la tions of the profiles aquatic and ter res trial an i mals (?mam mals, rep tiles, amphi- in Up per Silesia are still un cer tain be cause of the pau city of bians, fishes; also bivalves and other vari ous inver te brates), available biostratigraphic data (see review in Bilan, 1991) as so ciated with al gae and vas cu lar plants (see Dzik and and the lack of any consis tent regional scheme of lithostrati- Sulej, 2007; Sulej et al., 2011a, 2012; NiedŸwiedzki et al., graphy with formally defined units (see Becker et al., 2008). 2014; Pieñkowski et al., 2014). The lithostratigraphic as- Moreover, the varied palaeotopography of the Silesian Keu- signment, age and, partly, depositional setting of the fossil- per basin, controlled mostly by synsedimentary tecton ics, bearing depos its were more or less con jectural at most of the re sulted in extreme lateral vari abil ity in facies across the ba - sites and quickly be came the subject of debate, ex empli fied sin, which ad di tion ally hin ders cor re la tion. by the al ready fa mous Krasiejów lo cality (see summary in To date, some fragmen tary suc ces sions, less than 20 m Dzik and Sulej, 2007 and Bodzioch, 2012). Thus, the corre - thick, have been corre lated with tra ditional or mod ern units lation and precise dat ing of the bone-bearing strata became of the Ger manic Keuper (af ter “Stratigraphische Tabelle a key re search challenge in in ves tiga tions of the Silesian von Deutschland”, STD 2002; Franz, 2008; Menning et al., Keuper, for exam ple, in the context of evolu tion ary infer - 2012) and/or with infor mal lithostratigraphic units in west- ences for the early dino saurs (Racki, 2010). ern Poland (Dadlez and Kopik, 1963) or likewise infor mal This ar ticle takes as its fo cus a crit ical re view of the local units (e.g., Bilan, 1976). Litho-, allo- and chronostrati- stratigraphic ideas previ ously proposed by the authors from graphic aspects of the several units and corre la tions pro- the In sti tute of Paleobiology, Polish Academy of Sciences posed are frequently mixed to gether (see Szulc et al., 2015). and Warsaw Uni versity, led mainly by Jerzy Dzik and/or An in te grated, re gional event-strati graphic ap proach Grzegorz NiedŸwiedzki. Ac cordingly, the views presented en ables an ex pla na tion of the climate-driven and fa cies- be low are con sid ered to be rep re sen tative for the War saw tem po ral re la tion ships of the bone-bear ing de pos its and re- research group. Further more, the authors present herein a fers them to the revised lithostratigraphic scheme of the new strati graphic per spec tive for all of the verte brate lo cali - middle Keuper (Szulc and Racki, 2015; Szulc et al., 2015). ties, on the ba sis of inte grated lithostratigraphy, biostrati- Within the Grabowa Varie gated Mudstone-Carbon ate For- graphy and chemostratigraphy. The data are mostly the re- mation (Fig. 2), the WoŸniki Lime stone Mem ber (here af ter sults of the study, supported by the grant, “The evo lu tion of WLM) is formally defined, along with two members that re- ter restrial en viron ments of the Upper Silesian Keuper as place tradi tional Keuper units: the Ozimek Mudstone-Eva- biotopes of verte brates” (N N307 11703; Racki, 2010). porite Mem ber (= the Upper Gyp sum Keuper) and the UP PER TRI AS SIC BONE-BEAR ING LEV ELS 589

Fig. 2. Schematic section of the Upper Trias sic of the Upper Silesia, and its partly formal (in higher part) lithostratigraphic subdi vi sion af ter Szulc and Racki (2015; changed af ter Jewu³a, 2010, fig. 4), with a fo cus on cor re la tion with the Ger manic Ba sin ref er ence suc ces sion (“Stratigraphische Tabelle von Deutsch land” 2002, Franz, 2008); note the oc cur rence of three bone-en riched levels in the Silesian Keuper, as shown here, but only two Upper Tri as sic bone beds are defined and dis cussed in this pa per (cf. Szulc and Racki, 2015; Szulc et al., 2015); the Ladinian bone-bed level cor re sponds to the Miedary Beds of Kotlicki (1974), and the “Miedary sand stone-mudstone member” of Kotlicki (1995; see also Szulc, 2007b; Sulej et al., 2011b).

Patoka Marly Mudstone-Sandstone Member (= the Stein- brick pro duc tion. The ex ca va tions even tu ally in ten si fied mergelkeuper). The ter res trial ver te brate lo cal i ties are re - between 1974 and 2002, when the mine was in cluded in a viewed be low to show their strati graphic set ting in re la tion cement plant. How ever, the mass oc cur rence of verte brate to hitherto proposed age desig na tions and to revised or new bones was discov ered formally only in 1993 by Robert data collected dur ing the course of this study. Thus, the mar- NiedŸwiedzki, although bone-bearing depos its were proba - ginal-marine succes sion at Miedary (Sulej et al., 2011b) is bly exposed in about 1980 and even were exploited in late omitted from the present account. 1980s by ama teur collec tors, such as Krzysztof Spa³ek (Bodzioch et al., 2010; see the complete case study in R. NiedŸwiedzki, 2012, 2015). KRASIEJÓW Strati graphic in ter pre ta tions: the state of de bate The middle Keuper strata in the Ozimek-Krasiejów (for merly Hüttendorf-Krascheow; Fig. 3) area have been As a re sult of the inten sive exploi tation bone mate rial known since Römer’s (1870) carto graphic work and be- and the stud ies ini tiated by Jerzy Dzik, the great sci en tific came more ac ces sible since 1905 in the clay pit for lo cal sig nif i cance of the Krasiejów ‘grave yard’ for the study of 590 J. SZULC ET AL.

Fig. 3. Simpli fied lithological column of the Krasiejów succes sion (af ter Szulc 2005, fig. 2, mod i fied; see also Bilan, 1975), as the stratotype of the bound ary be tween the Ozimek Mudstone-Evaporite Mem ber with the Patoka Marly Mudstone-Sandstone Member (Szulc and Racki, 2015), and pho to graphs of se lected sed i mentary fea tures of the Keuper suc ces sion in Krasiejów. A. Gyp sum-bear ing (ar rowed) red mudstones from Ozimek Mbr. B. Celest ite crys tals from Krasiejów clay pit. Pho to graph cour tesy of Eligiusz Sze³êg. C. Typ i cal var iegated mudstones and claystones of the Patoka Mem ber, com posed of al lu vial sed i ments interlayered with regolith ho ri zons (r). D. Two lay ers of re worked palaeosoil grains (c) in cluded in al lu vial de posits. E. Con glomer ates com posed of sieved pedogenic nod ules. Note the ero sive, rounded scour casts at the base (ar rowed). F. Complex palaeosoil sec tion com posed of lower spotty in ter val, fol lowed by vertisol with slick en sides and cov ered by blu ish al lu vial sed i ments. G. Car bon ate nod u lar palaeosoil. H. Gray flu vial-dom i nated sed i ments fea tur - ing the up per part of the sec tion in Krasiejów pit. I. Metaposaurid skulls from the main bone-bear ing ho ri zon. Pho to graphs C and E by Micha³ Matysik.

Late Tri as sic ter res trial tetra pods be came ob vi ous (e.g., starting point for the proposal of the new avian-line clade Sues and Fraser, 2010), already after the first publi cations Silesauridae, the closest sis ter group to di no saurs, is partic u - by Dzik et al. (2000) and Dzik (2001, 2003a). The excel - larly note wor thy (e.g., Nesbitt et al., 2010; Benton et al., lently pre served and diverse as semblage, called the Paleo- 2014). On the other hand, Bodzioch (2012, p. 33) stressed: rhinus fauna, in cludes nu mer ous, par tially ar tic u lated skel e- “(…) lots of uniden ti fied remains are in collec tions, numer - tons (see sum mary in Dzik and Sulej, 2007). The presence ous problems remain unre solved, and quite a lot of inter pre- of the dinosauriform Silesaurus opolensis Dzik, 2003, a ta tions are con tro ver sial (…). Studies of a geolog i cal nature UP PER TRI AS SIC BONE-BEAR ING LEV ELS 591

mainly revolve around inter pre ta tions of the exact age of the Rote Wand”. Dzik (2003a, p. 557) indi cated a more broad bone-bear ing de pos its, of the sed i men tary en vi ron ment, of strati graphic dis tri bu tion of the bone mate rial, and first diagenesis and of the ori gin of bone accu mu la tions” . clearly dis tin guished two Fossillagerstätte ho ri zons “of vir- The con troversy concerns the age of the bone bearing tu ally the same geo log i cal age” at this lo cal ity, where “the ho rizons. In the first pa pers of the Warsaw group (Dzik et upper fossiliferous hori zon was identi fied about 7 m above al., 2000; Dzik, 2001, 2003a, 2003b; Sulej, 2002, 2005; but the lower one during exca va tions in 2000”, and char ac ter - see also Majer and Lubka, 2003), the late Carnian timing of ized by terres trial reptiles and capitosauroid am phibi ans in “red-col ored middle Keuper marly claystone” was as sumed, flood depos its (see also Majer and Lubka, 2003; Dzik and mainly on the basis of consid er ations of the verte brates. Sulej, 2007; Skawina, 2013). This approach allowed a corre la tion with the standard Ger- Szulc (2005, 2007a) presented another approach to the man Keuper profile of Baden-Württemberg (for the Ger man question of the age of the Krasiejów succes sion, and thus unit ter minol ogy see Fig. 4), even though this is not a very ini ti ated a ba sic con tro versy be tween sedimentologic and appro pri ate succes sion for such a purpose (see the chap ter fa cies ver sus palaeontological “schools”, as de fined by on climatostratigraphy below): Bodzioch (2012; see also Bodzioch et al., 2010). The author 1. The dom inant semiaquatic labyrinthodont am phibian stressed that the succes sion about 18 m thick was not stud- Metoposaurus diagnosticus krasiejowensis Sulej (= Metopo- ied in de tail with re gard to age and fa cies, and “the pro posed saurus krasiejowensis af ter Brusatte et al., 2015) is known evo lu tion ary in fer ences (Dzik et al., 2000; Dzik, 2003) from the Lehrberg Beds of the middle Weser Forma tion. could be prema ture” (Szulc, 2005, p. 161). In the context of 2. Cor re spond ingly, the rel a tively short-snouted phy- dating, Szulc (2005, p. 165) indi cated: “Lack of age-diag - tosaur Paleorhinus from Krasiejów pos si bly rep re sents “an nostic fossils, includ ing palynomorphs, makes stratigraphic earlier stage in the evo lution of the lin eage than the popu la- state ments un cer tain. None the less, the lithostratigraphic tions from Blasensandstein in Franconia and other known framework (which re flects climatostratigraphic con straints) Paleorhinus fau nas” (Dzik et al., 2000, p. 226) and therefore is quite clear and allows one to recog nize the gypsum-bear - indi cates an equiva lent of the lower Bunte Mergel, under ly - ing, lower part of the profile from Krasiejów as the upper - ing the Blasensandstein (= a ba sin margin cor rela tive of the most Upper Gipskeuper (…)”. Thus, this sec tion is in ter - mid dle Weser Fm in the cen tral ba sin fa cies, see Fig. 4). preted as the tran si tion strata from the Weser Fm to the 3. The joint occur rence of Paleorhinus and Metopo- overlying, bone-bearing Steinmergelkeuper (Arnstadt For- saurus assigns the Krasiejów assem blage to the worldwide mation, STD 2002). Further more, Szulc (2005, fig. 2) as- and oldest dino saur-bearing Paleorhinus biochron (Otis- signed bone-bearing hori zons to the Norian stage, as sug- chalkian land-verte brate faunal chron, LVF; Lucas, 1998), gested by palynostratigraphic dating of the upper most Up- thought at that time as being dated in a marine context as per Gipskeuper by Or³owska-Zwoliñska (1983). A simi lar late Carnian (more ex actly, late Julian–early Tuvalian view point is pre sented by Gruszka and Zieliñski (2008, subages; see re vised view in Butler et al., 2014). fig. 2), who, with refer ence to Bilan (1975) and Deczkowski In terms of the local geol ogy, the tetrapod-bearing cal- et al. (1997), pointed out a discrep ancy between the pala- care ous hori zon is thought to be about 80 m above the eontological and lithological premises; they favored at least Schilfsandstein (Reed Sandstone) marker, recog nized in the partial assign ment to the Norian stage. Gruszka and Zieliñ- Ozimek 1a well (situ ated 5 km west of Krasiejów; see K³ap- ski (2008) ad vo cated also the Steinmergel (= Drawno to ciñski, 1993), but also about 20 m above the gyp sum-bear- Zb¹szynek Beds) tem po ral equiv a lency, in di cated by the ing unit (after the doc u mented borehole data at this clay pit: lack of sandstone and gyp sum inter cala tions in the profile Bilan, 1975; Dzik et al., 2000, p. 228). In summary, “based un der debate. Bodzioch and Kowal-Linka (2012) also on the posi tion of the Krasiejów fossiliferous hori zon in the agreed that the prob abil ity of a middle to late Norian age is lithostratigraphic column, it seems to be coeval to the Lehr- in di cated by the over all fa cies de vel op ment. berg Beds in Germany, which proba bly termi nate the Car- The age infer ence of Szulc (2005, 2007a) has been re- nian sed i men tary cy cle” (Dzik, 2001, p. 625). In the sub- jected by Dzik and Sulej (2007, pp. 22–23) because of the surface lithostratigraphic terms of Dadlez and Kopik conchostracan argu ment alone: “The identity of the rock (1963), the Krasiejów succes sion was thought to corre - unit exposed at Krasiejów with the Arnstadt Fm can be im- spond to the Drawno Beds and (in the upper most part) to the me di ately dismissed be cause of the conchostracan ev i- Jarkowo Beds (Dzik et al., 2000, p. 228). Dzik (2001, p. dence. Olempska (2004) identi fied an abundant conchostra- 625) noted a proba ble corre spon dence of these highest can fauna with Laxitextella laxitexta (Jones) in the main strata at Krasiejów to the Blasensandstein hori zon. Olem- fossiliferous hori zon of Krasiejów. Accord ing to Bachmann pska (2004, p. 432) concluded more carefully from the and Kozur (2004, p. 49), this species occurs also in Germany conchostracan cor re la tion with the Ger man suc ces sion that in the Coburg Sandstone (Hassberge Forma tion), imme di - this fauna “proba bly lies within the middle late Carnian time ately above the Lehrberg Beds (Steigerwald Forma tion). The span”. However, some authors re cently re gard all the con- Coburg Sandstone is corre lated with the middle part of the chostracan-based dates and corre la tions as highly conjec - late Carnian (Tuvalian) Weser Fm (formerly Oberer Gips- tural (Becker, 2015; Maron et al., 2015; see below). keuper)”. In their exten sive review, Dzik and Sulej (2007, In the abstract of the ar ticle only, Dzik et al. (2000) p. 3) care fully sum ma rized the sit u a tion: “The geo log i cal age noted: “(…) an incom plete skele ton has been found about of the Krasiejów strata can be deter mined, although with a 8 m below the most fossiliferous hori zon, in a calcar e ous in- rather low reso lu tion, based on posi tion of vari ous members ter cala tion within red clays compa ra ble with the Ger man of its fauna in their evolu tion ary lin eages. (…). The strata 592 J. SZULC ET AL.

Fig. 4. Three main corre la tion variants of the Krasiejów clay pit section with the updated refer ence German Keuper profile (in basin margin facies) of Baden-Württemberg (Nitsch, 2011) and Pol ish strati graphic schemes (Dadlez and Kopik, 1963; Becker et al., 2008; Szulc and Racki, 2015): (A) with the Lehrberg Beds (Steigerwald Fm), based on the orig i nal ver te brate ev i dence of Dzik et al. (2000), fa - voured, among oth ers, by the in fer ence from conchostracan dat ing (Kozur and Weems, 2010); (B) with the Blasensandstein or Co burg Sandstein of the Hassberge Fm, sup ported by re vised ver te brate data (Milner and Schoch, 2004; But ler et al., 2014, fig. 1B) and macrofloral ev i dence (Pacyna, 2014), re spec tively; (C) with the Arnstadt Fm, deduced ini tially from facies de vel op ment and palynostrati graphic dating (Szulc, 2005, 2007a; cf. also Gruszka and Zieliñski, 2008), con firmed by the pres ent, in te gra tive, strati graphic study (see Fig. 17); the Weser Fm is a lat eral equiv a lent in cen tral ba sin fa cies. Note more con tin u ous de po sition in Silesia re gion than as sumed in DST 2002 (Szulc et al., 2015), re corded in the tran si tion from the Up per Gipskeuper to Steinmergel fa cies at Krasiejów (= Ozimek and Patoka mbrs of Grabowa Fm, respec tively, Fig. 3; Szulc and Racki, 2015). UP PER TRI AS SIC BONE-BEAR ING LEV ELS 593

seem to corre spond with the upper part of the Weser Fm in A possi ble error in dating ranges from “a rather early part of Ger many, be lieved to be of Late Carnian age”. the Carnian” to “Early Norian”, accord ingly to the available De spite a somewhat dif fer ent ap proach to the same tax - cor re la tion vari ants with the Ger man Keuper ref er ence suc - o nom i cal ques tions, in clud ing the Metoposaurus sub spe - ces sion. Also Irmis (2011, p. 409 and fig. 2), in the context cies, Lucas et al. (2007) confirmed the corre la tion of the of the posi tion of Silesaurus, stressed that the Carnian age Krasiejów tetrapod site with the Lehrberg Beds, but the was “based on ‘stage of evolu tion’ biostratigraphic argu - Parasuchus (= Paleorhinus of Dzik, 2001) and Metoposa- ments, but an early Norian age is equally likely” on the ba sis urus co-occur rence is seen by Lucas (2010) as key indi ca - of lithostratigraphic cor re la tion. tors of the middle Carnian (Otischalkian) age. Kozur and In fact, the Norian alter native emerged in the light of re- Weems (2010, p. 383) formally defined the new conchostra- vised data on the tem po ral distri bu tion of Metoposaurus can Laxitextella seegisi Zone, encom pass ing both the Lehr- diagnosticus krasiejowensis in the west ern Ger manic Ba sin, berg Beds and the Krasiejów suc ces sion. Si mul ta neously extended upward to the middle Norian (middle Middle Stu- Kozur and Weems (2010, p. 383) stressed that Lucas (2010) bensandstein, Löwenstein Forma tion) by Milner and ex plained “the con tra dic tion be tween conchostracan cor re- Schoch (2004). However, Milner and Schoch (2004) sup- lations and the land-verte brate defi ni tion of the Otischal- posed that the prolif er a tion of Silesian subspe cies is the kian–Adamanian boundary” , be cause “the fa mous Krasie- tempo ral equiva lent to the Kieselsandstein fauna, whereas jów fauna” is assigned to the suc ceed ing Adamanian Chron the older Lehrberg Beds in fact are char acter ized by the LVF (Tuvalian; the Rutiodon Assem blage Zone of Lucas, most prim i tive chrono-sub spe cies M. diagnosticus diagnos- 1998); the fossiliferous strata were invari ably thought to be ticus Meyer. However, this asser tion is re jected by Sulej homotaxial with the German Lehrberg Schichten. This late (2007, pp. 127–128), who has tried to confirm the origi nal Carnian inter val is defined by the stratigraphic range of the cor re la tion of Dzik et al. (2000) with the under ly ing Lehr- aetosaur Stagonolepis, whilst the Metoposaurus-Parasu- berg Beds (see Fig. 4), and therefore consid ered the meta- chus as so ci a tion re mained a ver te brate in di ca tor of the pre - posaurid subspe cies as “rather geo graph ical races” (see an ceding mid-Carnian LVF in the biochronology scheme of updated discus sion in Lucas, 2015; compare Brusatte et al., Lucas (2010). 2015). Even though only “be lieved to be of Late Carnian age” In sum mary, dis en tan gling the cor re la tion di lemma of the by Dzik and Sulej (2007), this hypo thet i cal age assign ment Krasiejów suc ces sion with the Ger man ref er ence, re cent data was accepted in a long se ries of palaeontological studies on point to three differ ent hy pothe ses of corre la tion (Fig. 4). Both tetrapod and in ver te brate fau nal el e ments (e.g., Barycka, Carnian variants of the War saw group are stratigraphically 2007; Dzik, 2008; Brusatte et al., 2009; Piechowski and close and may be assigned to the upper Weser Fm (sensu Dzik, 2010; Sulej, 2010; Konietzko-Meier et al., 2013; see Franz, 2008) and late Carnian Tuvalian substage. the refer ence list in Bodzioch, 2012). Moreover, Dzik and Sulej (2007) and Skawina and Dzik (2011) even assigned a Fa cies and en vi ron men tal mod els 230 Ma age to this tetrapod graveyard (Dzik, 2003b, assumed 225 Ma). The late Carnian age is also desig nated by In the contro ver sial fa cies context, Bilan (1975) origi - the recent macrofloral corre la tion of Pacyna (2014), who nally in ter preted the Krasiejów mudstone succes sion as compared his “Voltzia flo ral as semblage zone” with the cross-strat i fied flu vial de pos its, suc ceeded up wards by mas- Coburger Sandstone flora. sive lithologies refer able to a brackish (mesohaline) basin. In the most re cent papers on the veterbrate problems Dzik et al. (2000) first consid ered the fine-grained, or- (But ler et al., 2014; Piechowski et al., 2014; Antczak, ganic-rich, dark and uniformly thick bone-bearing depos its 2016), this age is re tained, as well as the lithostratigraphic as the depositional re cord of a large lake-deltaic in ter lude affil i a tion to the Drawno Beds (Sulej, 2010; Desojo et al, within the cross-bedded fluvial mudstone succes sion, as- 2013; Skrzycki, 2015). In partic u lar, But ler et al. (2014) cribed to a mean der ing river. Dzik and Sulej (2007, p. 6) re-stud ied the phytosaur tax on omy and phylog eny, as well claimed that the profu sion of charophyte gyrogonites as the biochronological status of the ‘Paleorhinus Bio- (Zatoñ et al., 2005), bivalves, ostracodes, and artic u lated chron’, and concluded that the Krasiejów fauna is co eval fish skele tons is an addi tional argu ment for a lacus trine mi- with the late Carnian Bavar ian (Ebrach) as semblage of the lieu. More gener ally, this would be a ba sin-scale event of Blasensandstein. However, Butler et al. (2014, p. 205) rising ero sional base level in the highstand phase of the noted also “the ma jor dif fer ences be tween the as sem blages Carnian transgressive cy cle af ter the plu vial-type Schilf- are the pres ence of Ebrachosuchus neukami and a plagio- sandstein sed i men ta tion. Re mark ably, Dzik and Sulej (2007, saurid temnospondyl at Ebrach (…) and the presence of a p. 16) noted in addi tion to the lake com mu nity: “The most ‘rauisuchian’, Polonosuchus, and a silesaurid dinosauro- numer ous fossils of land tetrapods have been obtained from morph, Silesaurus, at Krasiejów (…); these dif fer ences may the upper fluvial hori zon, but virtu ally all data on coeval rep re sent palaeoenvironmental or sam pling bi ases, par tic u- flora and in ver tebrate fauna co mes from the basal layer of larly given the pre domi nance of aquatic taxa at Ebrach”. the lac us trine bed”. More advanced studies were not un der- Among the palaeontological papers, only Skawina taken by the Warsaw group to support this depositional (2013, pp. 70–71) high lights se ri ous dif fi cul ties in the in ter- model. preta tion of the age of the Krasiejów fauna. She stated that From a strictly sedimentologic viewpoint, Gruszka and “pre cise dat ing re mains a matter of dispute” and it is ex clu - Zieliñski (2008) de scribed the envi ron ment of the Krasie- sively “believed to corre spond to the Alpine Late Carnian”. jów suc ces sion as “a low-energy anastomosing to mean der - 594 J. SZULC ET AL.

ing silt-bed flu vial system” , evolving in a subtrop i cal, sea- Upper Gypsum Beds, and are obvi ous corre la tives for the sonally dry but increas ingly humid climate. However, “the Steinmergelkeuper (Arnstadt Fm; cf. Fig. 4). Further more, mas sive clayey mid dle unit” was thought to have been de- the strongly later ally variable Drawno Beds (as noted al- posited in an exten sive, long-lived lake basin. Szulc (2005) ready by Dadlez and Kopik, 1963) have been omit ted from high lighted the pro cess of bone con cen tra tion ex clu sively in many corre la tion schemes since Deczkowski (1977; see small, ephemeral lakes and ponds in an unsta ble clima tic Bilan, 1991, fig. 1) and included in the Jarkowo Beds (e.g., and tectonic setting. Likewise, Bodzioch and Kowal-Linka Or³owska-Zwoliñska, 1985; Becker et al., 2008; Marcinkie- (2012) dis cussed ep i sodic cli mate-con trolled flood events wicz et al., 2014; Fija³kowska-Mader, 2015). in the ori gin of the lower (= main) bone bed of the middle 4. The corre la tion of the Krasiejów lac us trine-al lu vial unit (i.e., lac us trine in ter val of Gruszka and Zieliñski, deltaic sed i menta tion with the late Carnian Lehrberg trans - 2008). The polem ics resulted in changing views on the ori - gression is an origi nal event-stratigraphic concept of Dzik et gin of the fossiliferous depos its and, for exam ple, Konie- al. (2000), but without com para ble ex am ples of such trans- tzko-Meier et al. (2013) already wrote about two bone-bear - gres sion-ini ti ated lac us trine sys tems in the strati graphic re - ing allu vial hori zons (see chapter on taphonomy). cord (see another lake inter pre ta tion in Gruszka and Zieliñ- ski, 2008). However, the main obsta cle is a sedimentologic flaw in the pro posal, be cause typ ical lacus trine fa cies are Dis cus sion char ac ter ized by fine hor i zon tal lam i na tion, re cord ing depo- As shown above, despite some uncer tain ties concern ing sitional rhythmicity, driven by wet/dry sea son al ity and tran - intra-Carnian corre la tion (Fig. 4), there is no biochronologi- sient chemocline devel op ment, paired with prodelta turbiti- cal ba sis for accep tance of the Norian age of the Krasiejów tes, or ganic-rich muds and carbon ate interlayers (see for fossil biota (compare Lucas, 2015). From multidisciplinary instance Bohacs et al., 2000; Co hen, 2003; Renaut and viewpoint of the present authors, this conclu sion of the Gierlowski-Kordesch, 2010). The di ag nos tic char ac ter is tics War saw group is seri ously chal lenged by the fol lowing set are mostly expressed in the Late Tri as sic bas ins (see papers of data: in Gierlowski-Kordesch and Kelts, 1994; also e.g., Demico 1. The ref er ence Ger man suc ces sion for the mid dle and Gierlowski-Kordesch, 1986, Clemensen et al., 1998, Keuper ver tebrates is in fact in complete (Fig. 4), if a great Vollmer et al., 2008), includ ing the adja cent Holy Cross re- early Norian hi atus is ac cepted in this part of Ger manic Ba - gion (lithofacies O of Czapowski and Romanek, 1986). In sin. Thus, it is pos si ble that sev eral ‘guide’ taxa are not this context, Dzik and Sulej (2007) explained the massive strictly limited to the Carnian, but contin ued into the Norian mudstone structure as an effect of pedogenic homog e ni za - (see also Lucas, 2015). tion of orig i nally lam i nated de pos its in con di tions of pro- 2. In strictly lithostratigraphic terms on a re gional scale, found dry wet sea sonal ity, but this is proba bly to a limited as noted already by Dzik et al. (2000) and stressed by Szulc de gree. As a matter of fact, the typ ical lacus trine de pos its (2005, 2007a) and Gruszka and Zieliñski (2008), the main encom pass a suite of facies from palustrine, through litto ral Krasiejów fossiliferous levels occur above evaporite-bear- to profundal (con trolled by wa ter col umn strat i fi ca tion) ing depos its. More precisely, in the revised scheme of Szulc ones. The con cept of to tal pedogenic ho mog e ni za tion of and Racki (2015, fig. 7), with refer ence to the succes sion pri mary lac us trine char ac ter is tics, in clud ing those from the from a borehole in the clay pit (Szulc, 2005, fig. 2; cf. also profundal facies, as assumed by Dzik and Sulej (2007), Bilan, 1975), the upper boundary of the Ozimek Mbr, a for- sounds irra tio nally even in terms of Walther’s Law. mal unit re plac ing the Upper Gyp sum Beds, is placed at the Gruszka and Zieliñski (2008) thought of this lith o logic ho- top of a celest ite-rich hori zon, above the gyp sum-bearing moge ne ity as the primary signa ture of quiet and stable depo - playa-type strata (see Szulc, 2005, plate 1A–B); this bound- sition from settling out of suspen sion. However, with regard ary also is placed directly above the oldest bone-bearing to the massive lacus trine facies, Szulc (2005, p. 165) ex- strata (Fig. 3). Thus, a straight forward, long-distance corre - plained that “the lithological prop er ties of the typ i cal Lehr- la tion of the Krasiejów sec tion with the Weser Fm of SW bergschichten, with the outstand ing 2-4 massive, dolomitic Germany, omit ting closer regions (Brandenburgia, Thurin- lac us trine ho ri zons, dif fer sig nif i cantly from those ob served gia and Sax ony-Anhalt) is very risky, since there are cru cial in the Krasiejów”. fa cies dif fer ences be tween the mid-Keuper in SW Ger many In sum mary, the Krasiejów section is not composed of and other Germanic basins, in cluding that of Upper Silesia. typ i cal lac us trine sed i ments. In stead, the de pos its rep re sent As a matter of fact, the typ ical Lehrberg Beds of Baden- a playa flat with small depres sions (of gilgai-type topog ra - Würtemberg, as un ques tion ably lac us trine car bon ate sed i - phy) form ing ephem eral ponds and strongly controlled by ments, are com pletely ab sent from the Silesian Keuper, as wet/dry cli matic season al ity (cf. Szulc 2005, 2007a; Bo- discussed below (see, for instance, the nearby drill sections; dzioch and Kowal-Linka, 2012; Gruntmejer et al., 2015). Assmann, 1926, 1929; Koeppen, 1997). 5. The au thors of the Warsaw group signif i cantly men- 3. In the inter-regional context of the Polish Basin, the tioned (Dzik et al., 2000, pp. 228–229; see also Dzik and cor re la tion of the Krasiejów succes sion with the Drawno Sulej, 2007, p. 22) that the subsurface equiva lents above the Beds and even in part with the Jarkowo Beds, proposed by gypsum-bear ing strata of the Krasiejów profile (Drawno Dzik et al. (2000), remains in distinct con tra diction to the and Jarkowo Beds) were palynostratigraphically dated by ac cepted late Carnian age of this sec tion and equiv a lence Or³owska-Zwoliñska (1983) as Norian (i.e., Classopolis with the evaporite Weser Fm (as given explic itly by Skrzycki, meyeriana Zone, poorly known ?IVa to lower IVb inter val; 2015). The Drawno and Jarkowo Beds in fact overlie the confirmed in Or³owska-Zwoliñska, 1985, Deczkowski et UP PER TRI AS SIC BONE-BEAR ING LEV ELS 595 al., 1997; Marcinkiewicz et al., 2014; Fija³kowska-Mader, LIPIE ŒL¥SKIE 2015). Of course, this is incon clu sive, owing to a lack of palynological mate rial for direct dating of the succes sion. The clay pit in Lipie Œl¹skie, at Lisowice village, near 6. The conclu sive argu ment on a regional scale is pro- Lubliniec, was active, with some inter rup tion, since 1928, vided by the miner al og i cal and chemostratigraphic data of and has been known in the lit era ture since 1980 (Fuglewicz Œrodoñ et al. (2014). The Krasiejów pro file, de spite its min- and Œnie¿ek, 1980). The bone mate rial was discov ered and er al og i cal dis tinc tive ness (oc cur rence of palygorskite), is initially exploited by the Warsaw research group in 2006, placed with certainty in the II “Cr/Ti Chemozone”, assigned af ter no ti fi ca tion from the am a teur min eral col lec tors, Piotr at least largely to the IVb Palynozone in the re gional ref er - Menducki and Rob ert Borzêcki. The first re ports, al ready ence sec tion Patoka 1 (see Szulc et al., 2015). with an as signment of Rhaetian age, were pub lished two Thus, the present authors support with the addi tional years later (Dzik et al., 2008a, 2008b; NiedŸwiedzki and evi dence the infer ence of Szulc (2005, 2007a) that the Kra- Sulej, 2008). Al most con cur rently, this ex ca va tion sec tion, siejów bone-bearing suc ces sion rep resents a mostly Silesian extended by two shal low bore holes, was studied by Szulc et counter part of the Steinmergel fa cies (Arnstadt Fm; Patoka al. (2006), who assumed a Norian age for the exposed Mbr) and therefore its Norian age is reason ably implied. strata, which were dated palynostratigraphically. This dating may be partly questioned because the Steinmer- gel fa cies oc curs also lo cally in the Carnian Weser Fm, be - The Rhaetian age and alleged atypi cal nature low the main Eo-Cimmer ian disconformity (see Nitsch et of the biota al., 2005, fig. 1; Feist-Burkhardt et al., 2008, fig. 13.13). The in ter preta tion reveals also the problem of diachroneity The stratig ra phy of the Lipie Œl¹skie section, located 25 of the facies tran sition be tween hypersaline playa and allu - km E of Krasiejów, in fact was es tab lished firmly in the first vial (gypsum-free) facies, known from the western Ger- pa pers. Dzik et al. (2008b, p. 733) described the succes sion manic Basin (Edgar Nitsch, pers. comm., 2015), which is (Fig. 5) as: “Bone bearing greenish, reddish, and grey flu- dif fi cult to ex plain with cer tainty in the Pol ish Ba sin (see vial mudstones and siltstones, interbedded with cross or hor- Szulc et al., 2015). izon tally strati fied greywacke sand stones (the whole strati- How ever, this prin ci pal Eo-Cim mer ian hi a tus sig na - graphic sec tion is aproximately 12 meters thick) (…)”. ture, such as a deeply weath ered sur face and/or an atyp i- Well-pre served Rhaetian ver tebrate re mains were found “in cally ma ture soil pro file, was not rec og nized in the Krasie- a len ticu lar body of grey mudstone and claystone”, partly jów–Ozimek area (e.g., Bilan, 1975; Szulc, 2005, 2007a). within lime stone con cre tions. Only a local scour in a low-energy distributary channel net- Un fortu nately, the most detailed data are presented in a work is docu mented by Gruszka and Zieliñski (2008). The pop u lar-sci ence ar ti cle in Pol ish, with out in di vid ual ci ta - consis tent sedimentologic and strati graphic data contra dict tions of refer ences (Dzik et al., 2008b). Even if no deci sive the hy pothe sis of a large-scale middle Keuper uncon formity stage as signment was proposed in this paper, the Rhaetian on a regional scale, under ly ing (or locally also partly over- tempo ral affin i ties were strongly favoured on the basis of flo- lying?) the Steinmergel-like strata (cf. Szulc and Racki, ral (co ni fers, twigs, seed-ferns, isoëtaleans) and fau nal (con- 2015; Szulc et al., 2015; but see Fig. 5). More over, the ob- chostracans, di no saur trace fos sils) ev i dence. Dzik et al. served re place ment of cal cium sulphates (from the Up per (2008a, p. 734) con sidered the macrofloral and palynolo- Gypsum Beds) by celest ite and then by carbon ate mineral gical, but also conchostracan data as in dic ative of a Rhae- phases (in Steinmergel depos its) is indic a tive for of playa tian age for this site. This age is accepted (e.g., Gorzelak et brine evo lu tion, formed un der chang ing cli ma tic con di tions al., 2010; Skawina and Dzik, 2011), and strengthened even – from very arid to more humid (Füchtbaeur and Müller, recently by the macrofloral study of Pacyna (2014). How- 1977). This, in turn, unequiv o cally provides evi dence of ever, the age of the Lipie Œl¹skie lo cal ity sub se quently also contin u ing fa cies changeover, and hence denies the erosive was de fined more widely, between the undi vided “late No- and time gap at between the Upper Gipskeuper and the rian–early Rhaetian” (NiedŸwiedzki et al., 2011; Sulej and Steinmergel succes sion, on subsid ing tectonic blocks (in a NiedŸwiedzki, 2010; Sulej et al., 2011a; see also e.g., Baj- graben?). dek et al., 2014; Œwi³o et al., 2014) and “lat est Norian and On the other hand, the early Norian age of the Krasie- early (possi ble also youn ger) Rhaetian” (Pieñkowski et al., jów suc ces sion implies a lon ger-last ing, re stricted playa flat 2014, p. 269; see Fig. 5 for this stratigraphic inter preta tion). in the eastern periph eral part of Germanic Basin during the A signif i cant contro versy concerns the age of the oddly late Carnian-early Norian regres sion, paired with the composed assem blage of verte brate fauna, a mixture of typ- Eo-Cimmerian re mod el ling of ba sin to pog ra phy. This com - i cally Tri as sic (‘con ser va tive’) and advanced, Juras sic bina tion of events resulted in a profound within-basin types. Also well-pre served flo ral con stit u ents are like wise biogeographical dif fer en ti a tion that fi nally led to a mis lead - surpris ingly diverse (cf. also Pacyna, 2014). Dzik et al. ing long-distance biostratigraphic corre la tion, as noted by (2008a) claimed that the discov ery of large theropod dino - Sulej (2007) for Metoposaurus pop u la tions af fected by the saurs (3 m in length) expands their fossil record from the pre ced ing Carnian (Schilfsandstein) fall in sea level. The Early–Mid dle Ju rassic into the Trias sic. However, the pred- strati graphic in ter pre ta tion of the pres ent au thors im plies a tory archosaur was de scribed by NiedŸwiedzki et al. that these amphib i ans in the Krasiejów area belong to the (2012) as a new spe cies, Smok wawelski, being “a mo saic of last survi vors, which is in agreement with the model of prim i tive archosaur, croc o dile-line archosaur, and di no saur Milner and Schoch (2004). char ac ters” (NiedŸwiedzki et al., 2012: 275; see “large 596 J. SZULC ET AL.

Fig. 5. Lithological suc cession of the clay pit at Lipie Œl¹skie and nearby bore holes (cf. Szulc et al., 2006, fig. 5), with marked re vised lithostratigraphy (for mal units af ter Szulc and Racki, 2015 and Szulc et al., 2015), bone- and macroflora-bear ing in ter vals (partly af ter Pieñkowski et al., 2014, fig. 4) and palyno logi cal ly dated level (PZ IVb; Fija³kowska-Mader et al., 2015; LS3 – sample stud ied by Anna Fija³kowska-Mader, see Fig. 6). Strati graphic cor re la tions of Pieñkowski et al. (2014, fig. 4) are pre sented for com par i son; note an er ro ne - ously esti mated thickness of the Lipie Œl¹skie sec tion (12 m in stead 8 m; marked by ar row) by the au thors of the War saw group since pa - per of Dzik et al. (2008a), as well as misin ter preted red beds as pri mary depositional units (see Fig. 9); true red beds oc cur in the Lipie 2 well below an erosional surface.

pred a tory di no saur” in Pieñkowski et al., 2014; Zatoñ et al., All biostratigraphic data from Lipie Œl¹skie and sup- 2015). Lastly, the im portance of the site was rein forced by posed corre la tions are still not docu mented in system atic the discov ery of a mammal-like tooth (Œwi³o et al., 2014). studies and even largely not illus trated, as ex em pli fied by UP PER TRI AS SIC BONE-BEAR ING LEV ELS 597

Fig. 6. Miospores from the Lipie Œl¹skie clay-pit (cour tesy of Anna Fija³kowska-Mader). For the lo ca tion of sample LS3 see Fig. 5. Scale 30 µm. A. Anapiculatisporites telephorus (Pautsch) Klaus. B. Enzonalasporites vigens Leschik. C. Ovalipollis cf. rarus Klaus. D. O. ovalis Krutzsch. E. Brachysaccus neomundanus (Leschik) Mädler. F. Duplicisporites granulatus Leschik. G. Praecirculina granifer (Leschik) Klaus. H. Granuloperculatipollis rudis Venktachala et Góczán. I, J. Classopollis meyeriana (Klaus) Venktachala et Góczán. K. Geopollis zwolinskae (Lund) Brenner. L. Monosulcites sp. the key palynological data of the Warsaw group. The Sub- Pieñkowski et al. (2014) and Œwi³o et al. (2014), though only zone IVb of the Classopollis meyeriana Zone and possi bly the pres ence of L. ottonis was con firmed by Pacyna (2014). higher ones (IVc to V, sensu Or³owska-Zwoliñska,1983) Twigs and as so ciated seed scales, described only as “simi lar were quoted by Dzik et al. (2008a) from the succes sion, al- to Stachyotaxus”, were il lus trated by Dzik et al. (2008b, pp. though the first one, deter mined earlier by the ex peri enced 8–9; com pare Dzik et al., 2008a) and most proba bly repre - palynologist Carmen Heunisch in Szulc et al. (2006) and by sent another genus (Elatocladus; Pacyna 2014, p. 17). Staneczko (2007), is clearly limited to the Norian (see be- The sec ond “flag” in dex Rhaetian spe cies was iden tified low). The dating was revised recently by Pieñkowski et al. as iso lated cu ticles merely as cf. [sic!] Lepidopteris ottonis (2014), who quoted only the Rhaetian IVc and V zones. The (Staneczko 2007), or as “very simi lar to Lepidopteris ottonis” palynological studies were supple mented recently by Anna (Zatoñ et al., 2015, fig. 17), so the occur rence of the species Fija³kowska-Mader and Subzone IV was recog nized finally is in fact not demon strated at Lipie Œl¹skie (Zuzanna Waw- in the organic-rich fossiliferous level (Figs 5, 6; see Fija³- rzyniak, pers. comm., 2015). Further more, the possi ble kowska-Mader et al., 2015). Therefore, there is no obvi ous Norian oc cur rences of L. ottonis in Po land are summa rized palynostratigraphic evi dence for the corre la tion of the (i.e., in the Steinmergelkeuper, IVb Subzone) in a recent bo- Lisowice bone-bearing hori zon with the higher Rhaetian tani cal study by Pacyna (2014, p. 11),: “Be cause the cu tic u - biozones. lar dif fer ences be tween Lepidopteris spe cies from the Up - The first, strongest argu ments for the presence of Rhae- per Tri assic of Europe can not be com pared at present, the tian strata at the Lipie Œl¹skie pit are de rived from macroflo- question of extend ing the Lepidopteris ottonis Zone to the ral ev i dence, in clud ing the ‘guide’ Rhaetian seed-fern Lepi- Norian on the ba sis of isolated cuti cle fragments cannot be dopteris ottonis (Goeppert) Schimper and the coni fer Sta- settled. The Norian spec imens of Lepidopteris may be long chyotaxus septentrionalis (Agardh) Nath, both quoted by to a spe cies un des cribed as yet. The inter- and intraspecific 598 J. SZULC ET AL.

Fig. 7. Sche matic geo log i cal sec tion across out crops in the Lisowice - Lipie Œl¹skie area (A; from Bardziñski and Chybiorz, 2013), and the WoŸniki Lime stone Mbr above the brown mudstones, mostly cov ered by a rub bish dump, as recently visi ble (No vember 2014) in an in ac tive quarry on the Lisowice Hill, sit uated some 400 m N of the pit (B; see Figs 5 and 8, compare Szulc et al., 2006, fig. 8B; see also G¹siorowski et al., 1986; cour tesy of Waldemar Bardziñski). UP PER TRI AS SIC BONE-BEAR ING LEV ELS 599

vari a tion of the cu ti cle in Lepidopteris is very poorly known” (see also Marcinkiewicz and Or³owska-Zwoliñska, 1994, ta ble 1). In contrast to the Krasiejów site, however, the evolu - tionary verte brate argu ments were never used. Foreign au - thors referred to the amazing Lipie Œl¹skie findings in dif- ferent ways. Lucas (2010, p. 465) remarked: “Dzik et al. (2008a) re cently reported a Tri assic dicynodont from Po- land in strata they deemed Rhaetian based on palaeobotany”, but does not try to compare this fauna with his LVF chrons. Kozur and Weems (2010, p. 386) noted: “a Norian (or possi bly Rhaetian) dicynodont recently has been reported from Poland”. Irmis (2011, p. 398) quoted “the unexpected co-occur rence of dicynodont synapsids and dino - saurs in the lat est Trias sic”, whilst Brusatte et al. (2010, p. 81 and ta ble 1) indi cated an unnamed and scarcely de scribed large theropod dino saur from Lipie Œl¹skie as “?Norian- Rhaetian” or “?Rhaetian” in age, but highlighted the fact that “Tri as sic theropods of the same gen eral size of the new Pol- ish mate rial are al ready known (Gojirasaurus, Lilienster- nus)”. In fact, both genera are reported from the Norian of Ger many and New Mex ico (Brusatte et al., 2010, table 1).

Geo log i cal and fa cies set ting Notewor thy, the lithostratigraphic assign ment of the bone-bearing level evolved upward from the Jarkowo and Zb¹szynek Beds (Staneczko, 2007) to the Zb¹szynek Beds (cf. NiedŸwiedzki et al., 2014), the upper most Zb¹szynek to lower Wielichowo Beds (Œwi³o et al., 2014), and lastly to Fig. 8. WoŸniki limestones at Lipie Œl¹skie, with post-evapo- the lower Wielichowo Beds by Pieñkowski et al. (2014; ritic cherts (A; ar rowed), photo taken in 1996 (compare Fig. 7B); and gyp sum pseudomorphs preserved in these cherts (B; thin sec- Fig. 5). In a lo cal strati graphic context, a corre la tion with tion). the in ter val 18.5–42.0 m of the nearby Lubliniec IG1 borehole (quoted as “Lesieniec IG 1”) and with the Krasiejów sec tion was of fered by Dzik et al. (2008a, 2008b). These cor re lations are rather specu lative, as the middle Keuper de - pres ently is re jected by Pacyna (2014), who accepted the posits from Lubliniec IG1 were described merely as Norian dating of the WoŸniki Limestone (as the Brachy- “crumby samples – marly claystones, spotty, var iegated” phyllum Floral Assem blage Zone, with Clathropteris). On (Siewniak-Madej, 1982, pp. 3–4), and re ferred to the upper - the other hand, Pacyna (2014) supported the Rhaetian age most “Lisów Beds”, 83.5 m thick, 60 m above the Upper (i.e., the Lepidopteris ottonis Zone) of the Lipie Œl¹skie suc - Gyp sum Keuper (= Weser Fm). Likewise, the re lation ship cession, which, if true, would imply a rever sal of the strati- to the Krasiejów succes sion (even ca. 150 m above the graphic succes sion (it would place the Norian above the playa-type strata – Dzik et al., 2008b, p. 7) is diffi cult to ac - Rhaetian!) in the area. cept, be cause Szulc et al. (2006, fig. 5) al ready docu mented As stressed above, the authors of the Warsaw group the occur rence of the evaporite-bearing depos its in the Lipie overlooked the data from Szulc et al. (2006), where the 2 borehole, just a few metres be low the exposed strata and Norian IVb subzone was identi fied at Lipie Œl¹skie, and below the erosional disconformity (Fig. 5). also the Norian age assigned to the WoŸniki Limestone (but On the other hand, the clastic depos its occur ring in the see Zatoñ et al., 2015). Pieñkowski et al. (2014) suggested Lisowice area pass gradu ally up-section into rel atively pure an al ter na tive: the lat eral re la tion ship be tween the Lipie limestone of crenogenic and palustrine ori gin, with abun- Œl¹skie succes sion and WoŸniki Limestone. However, these dant evaporite pseudomorphs (WLM, as rede fined by Szulc au thors mis in ter preted some dozen-cm-thick car bon ate con- and Racki, 2015; Figs 7, 8). Therefore, the key lithostrati- cre tions from the clay pit as “WoŸniki Limestone Beds”, graphic ar gu ment con cern ing the stra tig ra phy of the Lipie while this unit was clearly de fined by Szulc et al. (2006) as Œl¹skie sec tion is its po si tion be low the WLM (Figs 5, 7), a a carbon ate body of crenogenic, palustrine, and pedogenic marker unit in local stratig ra phy (Szulc et al., 2015). ori gin, several metres thick. Therefore, such thin and discon - Dzik et al. (2008b) es tab lished the su per po si tion of the tinu ous limestone occur rences are excluded from the WLM al leg edly Rhaetian fossiliferous strata at the pit and the defi ni tion in Szulc and Racki (2015; only contin u ous layers WLM in Lisowice, but they conse quently assigned a latest above 0.5 m thick are included). The true WoŸniki Limestone Rhaetian age to the WLM. The palaeobotanical ar gu ment of crops out in a small, inac tive quarry nearby (Figs 7, 8A), Dzik et al. (2008b), the presence of the fern Clathropteris, where it reaches above 5 metres in thick ness (see Fig. 5). 600 J. SZULC ET AL.

Fig. 9. Ap par ent “redbeds” from the Lipie Œl¹skie sec tion. A. Lat eral pinch ing of red col our (left side of the sec tion; ar rowed) in the “Lower red bed” of Pieñkowski et al. (2014). B. Complex strike-slip fault sur faces cov ered with a red Fe ox ide pa tina. C, D. Details from Fig. 9B. E. Top most part of the sec tion crop ping out in the clay pit. F. Sec ond ary red den ing (ar rowed) of the pri mary grey sed i ments from the “Upper red beds” of Pieñkowski et al. (2014). UP PER TRI AS SIC BONE-BEAR ING LEV ELS 601

In the same pa per, Pieñkowski et al. (2014, figs 4, 5) Dis cus sion mis in ter preted two “red in ter vals” vis i ble in the Lipie Œl¹skie exca va tion: one at the bottom (called “Lower red Sulej et al. (2011a, pp. 261–262), in fact, correctly no- beds”) and another in the topmost part of the section (“Up- ted two es sen tial stratigraphic at tributes of the WoŸniki sec - per red beds”) as pri mary redbeds. In fact, the reddish col- tion: (1) a sim ilar mid dle Keuper suc ces sion was pre sented by our is of second ary ori gin, since the red colour pinches out Grodzicka-Szymanko and Or³owska-Zwoliñska (1972, fig. 2) later ally (Fig. 9A). Due to a signif i cant contri bu tion of or- from the CW 62 well lo cated at WoŸniki, and the mid–late ganic matter and dis persed pyrite, the orig inal colour of the Norian age for this orig inally “Rhaetian” well sec tion is des - entire section cropping out in the clay pit is grey or even ig nated with ref er ence to Szulc et al. (2006), and, more sig- black. During postdepositional weather ing (during the Tri- nif i cantly, (2) “the fossiliferous strata at WoŸniki grade up- assic or even later) the Fe sulphides un derwent ox ida tion wards conform ably into the crenogenic–lac us trine de pos its and Fe ox ides were transported downsection by mete oric of the WoŸniki (…) consid ered to be of Norian age (Szulc et wa ters, per co lating along a dis tinc tive fault zone (see Fig. al., 2006)”. The authors did not re fer to the far more sim ilar 9B–F). This in ter pre ta tion in val i dates the con clu sion re- succes sion in the WoŸniki borehole, 45 m thick, docu - garding these “red” ho ri zons as evi dence of a semi-arid cli- mented by Szulc et al. (2006, fig. 4). The data alto gether are mate dur ing de po si tion and cor re la tion with the orig i nally suf fi cient to re ject the sur pris ing biochronological in fer ence red Wielichowo Beds (Pieñkowski et al., 2014). True of Sulej et al. (2011a) that this succes sion is coeval with the redbed sedi ments have been pene trated by the Lipie 2 late Carnian Krasiejów pro file, es pe cially since the con- drillhole, some 5 m deeper (Fig. 5). chostracan and bivalve basis for this corre la tion is poorly From a more refined sedimentological per spec tive, the docu mented and only the conchostracan Laxitextella cf. occur rence of gypsum pseudomorphs (Fig. 7B) within the laxitexta is il lus trated. In fact, the Krasiejów suc ces sion “Upper red beds” and WLM (see Szulc et al., 2006), and re- equiva lent occurs about 60 m below the WoŸniki clay-pit worked vadoids in the same stratigraphic posi tion (= Lisów succes sion, as in dicated by the data from nearby WoŸniki K1 Brec cia “Beds” of Pieñkowski et al., 2014), indi cate and Kozieg³owy WB3 wells (cf. Szulc et al., 2015, fig. 7). arid/semiarid cli ma tic con di tions (Szulc et al., 2006). This is another strong argu ment against assign ing the Lipie Œl¹skie suc ces sion to the per manently wet Rhaetian stage PORÊBA (see below), even though Pieñkowski et al. (2014, p. 283 and fig. 5) tried to explain this appar ent facies anomaly in a This ex ca va tion near the aban doned mu nic i pal waste hu mid set ting by an age cor re la tion with the late Rhaetian dump, located near the west ern limit of the town of Porêba cli mate re ver sal. (Fig. 11), was known to the Silesian geolog i cal commu nity since April 2008 (Krystyn Rubin, pers. comm., 2010; Fig. 12). This small dig was explored for bone mate rial after the WOZNIKI discov ery of the fossiliferous strata by Tomasz Sulej in Sep - tember 2008 (Urbañski, 2012). The local ity belongs to the histor i cally oldest area around the town of WoŸniki, where ver tebrate re mains (teeth, scales) Strati graphic in ter pre ta tion of fishes and reptiles were known already to Römer (1867, 1870). The pit was active for the produc tion of bricks since The Porêba site is described in pa pers of Sulej et al. 1920 and mining ended in the 1960s (Sulej et al., 2011a). (2012) and NiedŸwiedzki et al. (2014) as a source of excep - tion ally rich and di verse verte brate fauna from two dissim i- lar hab itats (ter res trial and aquatic). This lo cal ity is partic u - Strati graphic in ter pre ta tion larly signif i cant for the study of dino saur clade ori gins be- This exca va tion has been exploited palaeontologically cause basal dinosauriforms and theropod dino saurs co-oc- since 2007 by Tomasz Sulej and his collab o rators. This re- cur at the site, along with an “ar mored” reptile group, the search resulted in the dis covery of a dicynodont-archosaur aetosaurs (Desojo et al., 2013), and large temnospondyl am- assem blage (includ ing silesaurids?), both as bone mate rial phib i ans. Plen ti ful spec i mens of tur tles are es pe cially char - and tracks (Sulej et al., 2011a). The authors presented the acter istic, as well as the abun dance of coalified plants, in - section of about 10.5-m-thick red and grey mudstones (fig. cluding a woody stem, 14 m long (see also Kubik et al., 1A therein), with carbon ate (hard oolite-like grainstones) 2015), described as the coni fer Agathoxylon keuperianum and sand stone lay ers and inter cala tions (Fig. 10). The two (Unger) Philippe by Philippe et al. (2015). The overall fos sil-bear ing in ter vals in the east ern wall are lo cated about well-pre served bone ma terial was col lected from four in ter - 3 m (the main source of bone mate rial) and 10 m above its vals grouped in lower part of the alleg edly 11 m thick suc - bottom, assumed to be late Carnian in age. Sulej et al. cession, mostly from “yel low ish carbonatic con glom er ates (2011a, p. 261) gave the follow ing sum mary in their ab- (…) and grey fluvial carbonatic, organic-rich mudstones stract: “The WoŸniki ver te brate as sem blage is sim i lar to that and claystones with rare bones (mainly fish remains)” (Nie- of Lisowice-Lipie Œl¹skie (…), in the presence of dicyno- dŸwiedzki et al., 2014, p. 1122, see also Figs 13I, 14). donts, shark spines, plagiosaurs and a cyclotosaur, but Accord ing to the Warsaw group, the domi nantly grey conchostracans and bivalves are simi lar to those from the marly-mudstone strata exposed at Porêba, interbedded with Krasiejów site (late Carnian)”. hor i zon tally strat i fied yel low ish or red dish sand stones and 602 J. SZULC ET AL.

Fig. 10. Com pos ite sec tion (A) and field pho to graphs (B, C) of the WoŸniki clay pit suc ces sion. Strati graphic po sition of bone-bear ing ho ri zons of the Lisowice level are ap prox i mated from Sulej et al. (2011a, fig. 1A).

“Lisów Brec cia” carbon ates (Figs 12, 13), are assigned to Dis cus sion the Zb¹szynek Beds and WoŸniki Limestones sensu Szulc et al. (2006), corre la tive with the Steinmergelkeuper (= the In the opinion of the present authors, this middle Keu- up per Arnstadt Fm and low ermost Exter Fm). The strati - per lo cal ity is cor rectly dated, al though with out any at tempt graphic posi tion is evi denced by the palynological mate rial, to cor re late it with the nearby co eval sec tions, dated as the identi fied by Anna Fija³kowska-Mader (in NiedŸwiedzki et same IVb Subzone, namely with partly condensed strata al., 2014) as typi cal for the mid–late Norian IVb Subzone of from the Porêba well and the Zawiercie outcrop (Szulc et the C. meyeriana Zone. al., 2006, fig. 5). In the local scale, this bone-bearing profile belongs to the broad transi tion inter val be tween the Patoka Mbr and the WoŸniki Limestone Mbr that is widely dis trib - uted in the area. UP PER TRI AS SIC BONE-BEAR ING LEV ELS 603

ZAWIERCIE-MARCISZÓW

The lithological sec tion of this new ver te brate lo cal ity (Fig. 11), inter preted as coeval with the Lipie Œl¹skie bone-rich suc ces sion (Fig. 14), was presented by Szulc et al. (2006), as a result of obser va tions in 2000 by Joachim Szulc in an exca vated niche un der a new city dump. How- ever, the bone ma terial was first iden tified by Grzegorz Racki in May 2009 in the pile of Keuper depos its (Fig. 15), mined from the dump niche (now totally filled). The niche profile was extended by the Kobylarz 1 borehole, drilled in 2012, about 250 m east of the former expo sure (Figs 11, 14).

Strati graphic in ter pre ta tion The strati graphic posi tion of the bone-bearing in ter val was clearly deter mined by two obser va tions: (1) the bone- bearing strata occur just below the WoŸniki Limestone Fig. 11. Lo cal iza tion of the tetrapod lo cal i ties near mu nic i pal cover, sparsely outcrop ping in the Kobylarz hill top (Fig. waste dumps and the bore hole Kobylarz 1 in the Zawiercie-Porêba 11), and are asso ci ated with micro bial, coquinoid and lime bound ary area on a Google Earth sat el lite pho to graph. conglom eratic layers (Racki, 2010, figs 1, 2; Szulc et al., 2015, fig. 15C–E), and (2) palynostratigraphic dating of the strata as the IVb Subzone (Heunisch in Szulc et al., 2006; ska et al., 2010; NiedŸwiedzki and Budziszewska-Karwow- see also Sadlok and Wawrzyniak, 2013). As in the nearby ska, 2015), supple mented by a tetrapod track assem blage Porêba site, located 1.7 km NW, differ ently preserved, (Sadlok and Wawrzyniak, 2013) and partly charcoaled plant partly reworked bone mate rial is found in rare conglom er- ma te rial (Kubik et al., 2015; Philippe et al., 2015; Szulc et atic lay ers (Fig. 15B, C), whilst iso lated and well-pre served al., 2015, fig. 15B). remains occur in grey marly mudstones and claystones with A quite dif ferent strati graphic concept, presented lastly carbon ate concre tions (see Racki, 2010, fig. 1; Fig. 15D). by NiedŸwiedzki et al. (2014, p. 1122), is therefore aston - Large dicynodonts and preda tory archosaurs of the genus ishing, espe cially in that the previ ous data are not discussed Smok are es pe cially dis tinct el e ments iden ti fied in the col - by these authors. The Warsaw group suggests that the Mar- lected, not very rich bone mate rial (Budziszewska-Karwow- ciszów site lies about 20–30 m above the Norian Porêba

Fig. 12. Photo graphs of the Porêba lo cal ity, taken in April 2008. A. View of the western part of fresh exca va tion, adja cent to the inac tive munic i pal dump (see Fig. 11). B. Newly ex posed hor i zon tally lam i nated con glomer atic to sand stone layer in a mudstone se ries in the southern wall, in the up per part of the sec tion. Cour tesy of Krystyn Rubin. 604 J. SZULC ET AL. UP PER TRI AS SIC BONE-BEAR ING LEV ELS 605

Fig. 13. Setting of the most typ ical facies and microfacies charac teris tics from sec tions at Porêba (A, C, E, G, I) and Marciszów (B, D, F, H, J), to show a strict resem blance of the Lisowice bone-bearing level in the both local i ties (see also Szulc et al., 2015, fig. 16). A, B. Massive pedogenic nodules. C, D. Con glomer ates com posed of poorly sorted, sieved pedogenic nod ules (= “Lisów breccia” of NiedŸwiedzki et al., 2014; see Fig, 14). Note the erosive rounded scour casts at their base. E, F. Oncoidal coat ing de vel oped around lithoclasts, E – pol ished slab, (e – pseudomorphed gyp sum con cre tion with mi cro bial en ve lope – ar rows), F – thin sec tion; note densely packed small crystals of gyp sum, now pseudomorphed. G, H. Heavily sulphide en crusted oncoids. I, J. Thin-sec tion pho to graphs from finer-grained con glomer ates, com posed of typ i cal small pedogenic grains; bone frag ment (b) is ar rowed.

sec tion “ac cord ing to field ob ser va tions and lo cal cor re la- while the conglom er ates are mostly composed of reworked tions”, and conse quently should be attrib uted “to the lat est pedogenic nodules, and form a the bases of fining-up sed i- Norian–early Rhaetian fau nal as sem blage in clud ing dicyno- men tary rhythms. The nod ules are some times coated with donts and archosaurs from the Lisowice local ity” . The Za- cyanobacterial en ve lopes, sug gest ing ep i sodic and short wiercie area is no ticeably af fected by fault tecton ics in the wetting events. This all strongly suggests that the both sites Tri as sic strata (Fig. 16), as ev idenced also by re cent geo - en compass sed iments of the same age. physi cal study (Idziak, 2013), and any sim ple corre la tion of the surficial sections is therefore very risky. Con sid er ing the fa cies and microfacies char ac ter is tics CHEMOSTRATIGRAPHY of the main sedi ment types occur ring in the both nearby-sit- AND MIN ERAL IN DI CA TORS uated both outcrops (Marciszow and Porêba), they are uni - form (Figs 13, 14). The domi nant redbed facies abounds in This approach to the corre la tion of the Silesian Keuper evaporites (recently preserved as carbon ate pseudomorphs), profiles was elab o rated in detail by Œrodoñ et al. (2014),

Fig. 14. The lithological sec tion of the Zawiercie-Marciszów site (see Figs 11 and 16), as exposed in the mu nic i pal dump ex ca va tion in 2000 (re pro duced from Budziszewska-Karwowska et al., 2010, fig. 2, based on Szulc et al., 2006, fig. 5; modi fied) and in the Kobylarz 1 well (Szulc et al., 2015) and at the Porêba site (based on NiedŸwiedzki et al., 2014, fig. 1C and ob ser va tions of the pres ent au thors). Ar - rows indi cate al leged bone-rich ho ri zons of the hypo thet i cally-de fined Lisowice level, suspected as a source for ma te rial ex ploited since 2009 in the ad ja cent mound of the Keuper de posits de rived from the dig (see Figs 11, 15, 16). 606 J. SZULC ET AL.

Fig. 15. Field (A, B), palaeontological (C, D) and microfacies (E, F) as pects of the Keuper de posits stored in the mound near munic i pal dump dig at Zawiercie-Marciszów (see Figs 11, 14; for fur ther de tail see in Szulc et al., 2015, figs 16, 17). A .View south ward from the muddy top of the waste pile, with vis i ble aban doned, over grown, and modern waste mu nic i pal dumps (see Fig. 11). B, C. Block of graded polymictic lime con glomer ate (with do lo mite clasts) in the heap, with visi ble bone cross sec tion (ar rowed in B, see also Racki, 2010, fig. 2B), and the recov ered frag ment of cracked, prob a bly dicynodont long bone in two views (C). D. An artic ulated tetrapod tail in lime stone concre tion (coll. Waldemar Bardziñski). E. Micritic-sandy microfacies of the co quina-oncoidal lime comglomerate, to show co-oc cur - rence of coalified wood de bris (W) and disarticulated ostracodes (Os, ar rowed). F. Finely-lam i nated cal car e ous mi cro bial en ve lope on large unionid shell, to show the zone with ver ti cally ori ented, fine tu bu lar struc tures, cor re sponding to cyanobacterial col o nies, in the stromatolite-like de posit. Pho to graphs taken by Maria Racka in May 2009 (A, B) and cour tesy of Ewa Budziszewska-Karwowska (C, D) and Józef KaŸmierczak (F). who studied the profiles from the Krasiejów and Lipie ported at the Kamienica gravel mine section, Fija³kowska- Œl¹skie pits and the WoŸniki K1, Patoka, Kozieg³owy WB3 Mader, 2015a). They found two geochem i cal indi ces: Cr/Ti and Kobylarz cores, encom pass ing the entire Silesian Keu- and Cr/Nb, which are the proxies for the ratio of basic/ per profile. This compos ite succes sion includes the bottom acidic source rocks, to de crease sys temat i cally in all longer of the Rhaetian, iden ti fied as the “Po³omia for ma tion” at Pa- profiles, thus offer ing a basis for the chemostratigraphic toka (this timing is ten ta tively palynostratigraphically sup - cor re la tion (Œrodoñ et al., 2014, fig. 19). Four zones were UP PER TRI AS SIC BONE-BEAR ING LEV ELS 607

Fig. 16. Geo log i cal sketch (A) and cross-sec tion (B) of the bor der land be tween Zawiercie and Porêba towns (af ter Bardziñski and Chybiorz, 2013, cf. Kotlicki, 1966; see Fig. 11), to show strati graphic set ting of the Porêba and Zawiercie-Marciszów ver te brate sites. 608 J. SZULC ET AL.

Ta ble 1 SUM MARY OF IN TE GRATED STRATI GRAPHIC IN TER PRE TA TION Zonation of the Silesian Keuper based on two geochem i cal in di ces. The aver age val ues are given in the table From the pa pers by the Warsaw group re viewed above, it may be con cluded that three widely sep a rated bone-bear - Average values of Cr/Ti ing in ter vals are pres ent in the Up per Trias sic Silesian suc - Patoka WoŸniki Kozieg³owy Kobylarz Lipie Krasiejów ces sion, ranging in age from the Carnian to Rhaetian. Be- Zone I 80 cause the intra-regional strati graphic rela tion ships of the Zone II 104 95 108 93 94 90 bone beds were never discussed in detail, the exact tempo ral Zone III 123 128 131 equiv alency of the Krasiejów and WoŸniki sec tions, as well Zone IV 156 151 as the Zawiercie and Lipie Œl¹skie sec tions, is rather ques- Average values of Cr/Nb tion able, and the pres ence of more than three bone-rich in- tervals would be worth consid er ing. From a geograph ical Patoka WoŸniki Kozieg³owy Kobylarz Lipie Krasiejów viewpoint, the Norian (Porêba-type) fos sil as sem blage of Zone I 3.6 the Warsaw group is known from one local ity only, while Zone II 4.7 4.9 5.4 5.1 4.6 4.7 the other two are re ported as hav ing poten tially wide dis tri- Zone III 7.5 7.3 7.5 bution in a regional scale. Zone IV 7.6 7.7 Lithostratigraphic aspect The present authors define two infor mal units, bone-en- spec ified, Zone I oc curs only at the top of the Patoka profile riched lev els, as the en tire strati graphic in ter val con tain ing and zone IV only at the bottom of WoŸniki and Kozie- bone ma te rial, even if the ver te brate-fos sil-rich ho ri zons are g³owy. Zone II is pres ent in all profiles, and both the Krasie- interlayers only in thick, (al most) barren depos its (cf. Szulc jów and Lipie Œl¹skie pits belong to this zone. The zonation and Racki, 2015). Pieñkowski et al. (2014, p. 272) showed dem on strated in fig. 19 of Œrodoñ et al. (2014) becomes that the main fossiliferous inter val at Lipie Œl¹skie “(…) has even more evi dent, when the aver age values for partic u lar four discrete bone-bearing beds, sepa rated by bone-barren zones are calcu lated (Ta ble 1). mudstone-siltstone levels with numer ous plant remains and The chemostratigraphic zonation is well corre lated with casts of bi valve shells”. At Porêba, four fossiliferous hori - lithostratigraphy. Zone I corre sponds to the Rhaetian (Po³o- zons were also dis tin guished (Fig. 14), while at least three mia gravels) and upper Steinmergelkeuper (Patoka Mbr), such inter vals oc cur at Krasiejów (Dzik and Sulej, 2007, zone II to the Steinmergelkeuper and/or upper most Upper p. 7; Gruszka and Zieliñski, 2008; compare Fig. 3). Gipskeuper (= lower Patoka Mbr), zone III to the Upper The proposed levels have distinctly differ ent lithostrati- Gipskeuper and Schilfsandstein and zone IV to the Schilf- graphic settings, and the lower one repre sents largely equiv- sandstein and Lower Gipskeuper. a lents of the Steinmergelkeuper at Krasiejów, in clud ing The quanti ta tive min eral data of Œrodoñ et al. (2014) also in its basal part a transi tion to the Upper Gypsum Beds support the dis tinction between the Rhaetian and the under - (re corded in the disap pear ance of the bar ite-bear ing celesti- lying Keuper strata, based on lithostratigraphy: the Rhaetian nes; see Szulc, 2005; Bzowska and Racka, 2006), that is, the rocks contain partic u larly abundant kaolinite and are free of broad pas sage between the Ozimek and Patoka Mbrs. This car bonates. By this cri terion, all the stud ied samples, ex cept rela tively thick unit (17 m; Bodzioch and Kowal-Linka, for the “Po³omia Fm” in the Patoka 1 well (and proba bly the 2012, fig. 2; Fig. 3) also is charac ter ized by a high density of kaolinite-rich upper most part of Patoka clay-pit sec tion; Szulc bone ma te rial ac cu mu la tion, cou pled with the rel a tive lat- et al., 2015), have to be classi fied as older than Rhaetian. eral con tinu ity of the main Fossillagerstätte de posit (= lower There is no rea son to spec u late that the chemostratigra- hori zon of Dzik, 2003; middle unit of Gruszka and Zie- phic zonation may be diachronic in such a small fragment of liñski, 2008), and therefore the term “Krasiejów bone-brec - one sedi men tary basin. If it is not, then it can be cali brated, cia level” is coined. On the other hand, com para ble fos- on the basis of available biostratigraphic data. Unfor tu - sil-rich depos its are totally absent from the appar ently con - nately, these are scarce (see Fija³kowska-Mader et al., tin u ous well suc ces sions (WoŸniki K1, Kozieg³owy WB3). 2015). The bot tom of the chemostratigraphic zone I at Pa- There fore, their dis tri bution is dem onstrated to be limited to toka and zone II at Lipie Œl¹skie were iden tified as the mid - the west ern Silesia re gion. Its re stric tion to the Opole re gion dle-late Norian Palynozone IVb and the upper part of che- might be partly controlled by the top o graphic reju vena tion of mostratigraphic zone IV at WoŸniki as the late Carnian bio- the ba sin and intraformational erosion fol lowing the Eo-Cim- zones II and III (cf. Fig. 17). The verti cal exten sions of merian movements in the Carnian-Norian transi tion. these biozones are not known, being lim ited in these pro- All other bone-bearing strata (i.e., Lipie Œl¹skie, WoŸ- files to their unoxidized sections. Thus, the chemostratigra- niki, Porêba-Zawiercie; Fig. 1B) are as so ciated with the phic zone II, contain ing both the Krasiejów and Lisowice tran sitional in ter val be tween the Patoka and WoŸniki Mbrs, bone hori zons, is – on the basis of this approach – most and this super po sition is di rectly ob served in the areas broadly dated as late Carnian to late Norian (see be low), but around the local i ties (Figs 5, 7, 14), with the excep tion of defi nitely not Rhaetian (com pare the magnetostratigraphic Porêba. How ever, several dis tinct lithologies at this site are in fer ences in Nawrocki et al., 2015). clearly cor re lative with the Lisowice level of the nearby UP PER TRI AS SIC BONE-BEAR ING LEV ELS 609 , ilc niam eerhT .)3C( lai re tam enob fo yr evo csid dna ,)2C( seicafonylap enir ts ucal ni )?( emca/noi ta vr eserp hpromonylap dez ila col yb dekram ,1 enozomehC rewol eht fo sla vre tni rehgih ylthgils rehgih tni vre sla fo eht rewol enozomehC ,1 dekram yb col ila dez hpromonylap eserp vr ta emca/noi )?( ni ucal ts enir seicafonylap ,)2C( dna csid evo yr fo enob tam re lai .)3C( eerhT niam ilc - eht fo noi ta le rroc eht fo stn airav eerht etoN .atad )4102 , )4102 .atad etoN eerht airav stn fo eht rroc le ta noi fo eht dna 1 akotaP ta spor ta akotaP 1 dna htiw dna ,)1C( se ci dni cihpargitartsomehc yb deta ci dni trap mo ttob sti htiw :1 akotaP ta st iso ped ra l imis htiw ,)3 .giF( el cyc lai vulf dedarg a gn imrof ,noi tces eiks tces ,noi imrof gn a dedarg vulf lai cyc el .giF( ,)3 htiw imis l ra ped iso st ta akotaP :1 htiw sti ttob mo trap dni ci deta yb cihpargitartsomehc dni ci se ,)1C( dna htiw .la te .la .snoi tces elo herob eht ni decalp ylno ylla c iteh to pyh ,level eciwosiL eht ot gn idnop se rroc edo s ipe cino tcet-lai vulf nairoN-dim eht gn idul cni ,llew sa dekram era stneve cino tcet ro/dna ci tam ci ro/dna tcet cino stneve era dekram sa ,llew cni idul gn eht nairoN-dim vulf tcet-lai cino ipe s edo rroc se idnop gn ot eht eciwosiL ,level pyh to iteh c ylla ylno decalp ni eht herob elo tces .snoi redaM-akswok ³ajiF( cihpargitartsonylap ,)5102 ,ikcaR dna cluzS( cihpargitartsohtil fo esu deni bmoc htiw ,)5102 ,6002 , ,6002 ,)5102 htiw bmoc deni esu fo cihpargitartsohtil cluzS( dna ,ikcaR ,)5102 cihpargitartsonylap ³ajiF( ctuo dna selo herob gn isir pmoc ,aiseliS re ppU rof noi tces repueK et iso pmoc ecn er efer eht htiw sei t ila col etar be trev eht fo noi ta le rroc cihpar gitarts detar ge tnI tnI ge detar gitarts cihpar rroc le ta noi fo eht trev be etar col ila t sei htiw eht efer er ecn pmoc iso et repueK tces noi rof ppU re ,aiseliS pmoc isir gn herob selo dna ctuo .la te .la oo ñodor Œ( cihpargitartsomehc dna la c ig ol ar enim evi sne her pmoc dna )3891 ,aks )3891 dna pmoc her sne evi enim ar ol ig c la dna cihpargitartsomehc Œ( .la te .la cluzS ni snoi sse ccus eht fo noi tpirc sed eht ees( 1K ikin 1K ees( eht sed tpirc noi fo eht ccus sse snoi ni cluzS ñilowZ-akswo ³rO re tfa sre bmun enozonylap ;5102 enozonylap bmun sre tfa re ³rO ¹l Œ eipiL morf level eciwosiL level morf eipiL Œ .71 .giF .71 ŸoW 610 J. SZULC ET AL.

Fig. 18. Dia gram showing the range of uncer tainty in tim ing of the Up per Tri as sic stages (af ter Ogg, 2012, figs 25.6 and 25.7; compare Lucas et al., 2012) and the range of the pos sible cor re la tion er ror for these bound aries with global palynozones (ac cord ing to Kürschner and Herngreen, 2010; adopted by Ogg, 2012, fig. 25.7, in “Geo log i cal Time Scale 2012”; GTS 2012) and the orig i nal and up dated Pol ish palynozones (see also Fija³kowska-Mader et al., 2015); ar rows show the strikingly dif fer ent po sitions of the palynozone bound aries on the ba sis of index spe cies ranges in Cirilli (2010, fig. 2), against the first ap pear ance da tum (FAD) of these species in Kürschner and Herngreen (2010, fig. 3; see also Barth and Kozur, 2011).

Marciszów site (see also Fig. 16B). In gen eral, the oc cur - marly set, under ly ing the WoŸniki Limestone, comprises rence of a thin (less than 0.5 m) and discon tin u ous hori zon cross-strat i fied sand stones and con glom er ate-oncolitic lay - of micro bial car bon ates (oncoids, stromatolites) is a distin - ers with macroflora debris. Therefore, a slight diachronism guishing feature of the greyish to locally black mudstone in- of the bone ac cumu lations is predicted in the later ally vari- ter vals, rich in coalified plants (note a coal part ing at Lipie able en vi ron ments, char ac ter iz ing the mid-Norian tec tonic- Œl¹skie) and even tree trunks many meters long (Lipie plu vial in ter val (see be low). Œl¹skie – Szulc, 2007a; Porêba – Sulej et al., 2012), as well Grey to blackish, sandstone-con glom erate to claystone as shelly bi valve mate rial (es pe cially at Marciszów; Racki, graded cyclic depos its, up to 20 m thick, in places with 2010, fig. 3; see also Szulc et al., 2015, figs 15, 16). It is abundant plant de bris, occur in the lower part of the succes - notewor thy that some 15 years ago, in a small outcrop at sion of the Patoka borehole (Szulc and Racki, 2015, figs 3D, Brudzowice village (some 13 km west from Porêba out- 4B), and allow the estab lish ment of the equiva lence of the crop), no lon ger in exis tence, a di rect contact was visi ble be- 6-m-thick flu vial cy cle in the Lipie Œl¹skie sec tion with this tween the oncoid- and bone-bearing conglom er ates with the part of the mudstone-sandstone Patoka Mbr in the regional overly ing, 3-m-thick carbon ates of the WoŸniki Limestone. refer ence section of the present authors (see Fig. 17). On a The thickness of the so delin eated Lisowice bone-bearing more refined scale, three fining-up cy cles may be taken into complex is surpris ingly constant, at least in the exposed suc - consid er ation, but only the oldest one (C1), incom plete at cessions, namely between 5 and 10 m. the bot tom of the drill core, is supported by chemostratigra- In light of the descrip tions of the borehole sections by phic corre la tion, whilst the real bone mate rial occurs about Bilan (1976, pp. 49–54), more broadly de fined Lisowice 80 m higher (C3). Thus, this attempt is a reso lu tion that is level proba bly contin ues at least 10 km southward from Za- far from conclu sive. wiercie, as gray-seledine mudstones with limestone-clayey con glom er ate sets and sand stone part ings, con tain ing macro- Age of ver te brate lo cal i ties flora detri tus and lignite fragments, up to a dozen cm at two or three lev els (up to 5.1 m thick), and mussels (see also The dating of the Krasiejów bone brec cia is somewhat Kotlicki, 1995 for ac count of his “Wojs³awice For ma tion”). un cer tain, ow ing to the con jec tural tem po ral re la tion ships The thickness of this charac ter istic bottom part of the be tween a lower seg ment of the Classopolis meyeriana “higher Rhaetian” unit of Bilan (1976) is proba bly about 20 Zone (i.e., its IVa Subzone) and the Carnian-Norian bound- m in two wells, but the macroflora- and coal-bearing in ter - ary (see e.g., Or³owska-Zwoliñska, 1985; Marcinkiewicz val is about 17.6 m and 13.7 m thick, respec tively. Also, in and Or³owska-Zwoliñska, 1994; Deczkowski et al., 1997; the Kozieg³owy WB 3 well section, a 14-m-thick grey Kürschner and Herngreen, 2010; Heunisch and Nitsch, UP PER TRI AS SIC BONE-BEAR ING LEV ELS 611

2011; Marcinkiewicz et al., 2014; Fija³kowska-Mader, 2015), com pli cated by barren in ter vals below and above this subzone in the scheme of Or³owska-Zwoliñska (1983). The initial assign ment of the upper part of Upper Gypsum Beds (more ex actly, grey claystones with an hyd rites) and the IVa Subzone to the Carnian-Norian stage transi tion by Or³owska-Zwoliñska (1983, tables 14, 15), narrowed by Or³owska-Zwoliñska (1985, table 1) to the Norian, was confirmed by Cirilli (2010, fig. 2), because the index spe - cies first ap pears ex actly at the Carnian-Norian bound ary (Fig. 19). On the other hand, the Drawno Beds (later included in the Jarkowo Beds), a supposed subsurface equiva - lent of the Krasiejów level (Dzik et al., 2000), are assigned to the basal IVb Subzone by Deczkowski et al. (1997). The Norian age of the Jarkowo Beds as well as the entire Patoka Mbr, is reason ably shown (see Fija³kowska-Mader et al., 2015), and supported also by miner al og ical and chemostratigraphic proxies (Fig. 17; Œrodoñ et al., 2014), viewed in the well-known Late Tri as sic climatostratigraphic con text (see be low). The Lisowice bone-bear ing level is there fore lo cated between 100 m and at least 180 m below the proba ble Rhaetian de pos its in the re gional ref er ence sec tion, marked Fig. 19. Di a gram show ing the im por tant dif fer ences in strati - by light kaolinite clays and sider ite-concretionary hori zons graphic dis tri bution of bone-bearing de posits in the Silesian Upper Tri as sic pro posed by the au thors of the War saw group (230 Ma for (see Fig. 17; Szulc and Racki, 2015, fig. 4D, E; Szulc et al., Krasiejów; Dzik and Sulej, 2007; 205 Ma for Lipie Œl¹skie; 2015, fig. 10). Thus, the Lisowice bone beds, as well as the Skawina and Dzik, 2011) and in the present pa per. Un cer tainty in over ly ing WLM (Szulc et al., 2006), are consid ered to be a ages of the Up per Tri as sic stages (taken from, Ogg 2012; see Fig. middle Norian inter val, corre spond ing to the lower part of 18) is marked as dot ted bound ary lines, as well as an ex tent of dat- the very lengthy IVb Subzone, because the three sites stud- ing am bi gu ity for par tic u lar bone-bear ing units (light grey ar - ied are assigned to this subzone (see Fija³kowska-Mader et rows), shown by the both re search groups. al., 2015). This conclu sion agrees well with the Norian dating of the WLM by macroflora by Pacyna (2014). Inde - pendent support is provided by the chemostratigraphic as - looks so surpris ingly young, if it is Rhaetian) is largely lost signment of the WLM in the Kozieg³owy WB3 well sec tion and the mid-Norian assem blage fits the global distri bu tion to zone II by Œrodoñ et al. (2014; see Fig. 17). It is also data much better (Lucas, 2015). notewor thy that in a core from a well drilled some 27 km E from Lisowice (Czarny Las), sands and conglom erates with Climatostratigraphic con text of the Keuper suc ces sion clasts of eroded WLM were dated (Fija³kowska-Mader et in Germany and in Up per Silesia al., 2015) by means of palynomorphs as Subzone IVb, which indi cates the Norian age of the WLM. The general lithostratigraphic pattern of the Upper Tri- In summary, in place of three widely sepa rated bone- as sic in the Ger manic Ba sin re flects long-term clima tic fluc - bearing inter vals in terms of stratig ra phy in the Upper Trias - tua tions between dry and hu mid in ter vals. The dry peri ods sic Silesian succes sion, in an inter val of about 25 Ma, the are rep resented by the Carnian evaporitic Lower and Upper authors propose a far narrower two-step tem po ral distri bu- Gypsum Beds, while the main humid in ter vals are in dicated tion. The bone-rich strata are assigned a Norian age, with a by the flu vial Schilfsandstein for the Carnian and the Rhae- range of less than 10 Ma (Fig. 19), but owing to the uncer - tian clastics for the up per most Tri as sic. Be cause of the su - tain ties and con tro ver sies in Up per Tri as sic ter res trial stra - praregional nature of the clima tic changes, the gen eral tig ra phy (see be low), this is still a somewhat pre limi nary in - climatostratigraphic framework of the Keuper may be reli - fer ence. This in ter pre ta tion is in di rectly sup ported by the ably cor re lated with the better dated Up per Trias sic of the con clu sion of Sulej et al. (2011a), who have corre lated the Tethys domain (Szulc, 2007a). This corre la tion is con- am phib ian-dom i nated Krasiejów fauna with the dicyno- firmed also by prelim i nary magnetostratigraphic studies dont-archosaur assem blage from WoŸniki. Of course, vari- (see Nawrocki et al., 2015). ability in local habi tat and/or taphonomical factors during With re gard to the in ter nal framework of the Stein- the de po sition of the Lisowice level is clearly re corded in mergelkeuper (Arnstadt Fm) from the closest German re- the sections under study and for exam ple the tur tle-dom i- gions, namely from Thuringia (Dockter, 1995), Lower Sax - nated commu nity at Porêba (NiedŸwiedzki et al., 2014) is ony (Arp et al., 2005), Saxony Anhalt and Brandenburgia strongly distin guished by such a wide repre sen ta tion of (Beutler, 2008) a con spic u ous tri par tite fa cies de vel op ment fresh wa ter rep tilian biota (see the co eval di ver sity data from becomes an ob vious fea ture of this succes sion. The Arn- Germany in Milner and Schoch, 2004). Con sequently, a stadt Fm is com posed of the so-called Lower Red (or Varie - par a dox of the pe cu liar ver tebrate fauna at Lipie Œl¹skie (it gated) Series, the Mid dle Grey Se ries and the Up per Red 612 J. SZULC ET AL.

Fig. 20. Strati graphic cor re la tion of Up per Silesian ver te brate-bear ing lev els with an ap pro pri ate Ger man mid dle Keuper ref er ence succes sion in Lower Sax ony (af ter Arp et al., 2005, fig. 2). Note that only major Krasiejów bone-rich ho ri zons are shown (see Fig. 3).

(Var ie gated) Se ries. The tri par tite sub di vi sion re flects lon - all the bone beds is the time and fa cies equiva lent of the ger cli ma tic fluc tu a tion (sev eral Ma for each) and re flects Mid dle Grey Se ries and re flects gradual climate pluvialisa- grad ual cli mate pluvialisation, reaching its max imum in the tion that reached its climax around the time of forma tion of Middle Grey Se ries (as evi denced by the in creased con tri- the Lisowice bone-bearing complex (see below). The next bu tion of al luvial sedi ments, rich in plant and bone debris), aridisation trend is re corded in the redden ing of the clastic replaced then by the next, drier inter val of the Up per Red de pos its and the sed i men ta tion of the evaporite-bear ing car- Se ries. How ever, the lat ter is richer in al lu via than the bonates of the WoŸniki Limestone that postdates the grey Lower Red Se ries. The cli mat i cally-driven tri par tite sub di - allu vial sedi ments and that is equiva lent to the Upper Red vision is evi dent and unequiv o cal also in the more marginal Se ries. fa cies (Löwenstein Fm) (see overview by Franz, 2008). In summary, the Steinmergel succes sion in southern When the Upper Silesian Patoka Mbr is com pared to its Po land (in Upper Silesia in partic u lar) dis plays a simi lar Ger man coun ter parts, the tri par tite sub di vi sion is also ap- suc ces sion of lithofacies to its equiv a lents in the east ern re - parent (Fig. 20). In the Krasiejów borehole, the redbed in- gions of Germany. It is also notewor thy that the most abun - ter val ly ing be tween the last gyp sum-bear ing ho ri zon and dant bone breccias in the well exposed and exten sively stud- the bone breccia reason ably could be corre lated with the ied (for more than 100 years) German Keuper are found Lower Red Se ries in Germany. The sec tion en compass ing within the Mid dle Grey Series or its equiva lents (Seegis, UP PER TRI AS SIC BONE-BEAR ING LEV ELS 613

2005), that is in the middle Norian. It disap proves the age The Trias sic conchostracan-based zonation is hoped to assump tions discussed above that were presented by the be a con sis tent alter native tool of palynostratigraphy (Kozur War saw group for the Silesian Fossillagerstätte. and Weems, 2010). Argu ments as to the vagueness of the Early Tri as sic zonation were pre sented by Becker (2015), and include (1) an uncer tain conchostracan taxon omy, VAGUE NESS OF THE TER RES TRIAL jointly with the in dex spe cies, (2) unre li able palynostrati- NORIAN STAGE graphic dating of the zones, augmented by (3) specu la tive ter res trial-ma rine cor re la tions and con se quently, chrono- As shown in Fig. 18, any attempt to clarify the abso lute stratigraphic in fer ences that also are bi ased, because they age of Upper Tri assic sec tions is strongly in fluenced by the are deter mined exclu sively from marine-based stage bound- contro versy concern ing the ages of the stage boundaries, in- aries. The last fun da men tal inad equacy of the Late Tri as sic cluding the disputed inter vals of many millions of years zonal scheme of Kozur and Weems (2010) was re cently ex - (Ogg, 2012). For exam ple, the age of the Lipie Œl¹skie suc - posed by Maron et al. (2015, p. 973): “(…) conchostracans ces sion was iden tified as “Rhaetian; about 205 Ma old” by from the Weser Fm of the Germanic Ba sin are as signed an Skawina and Dzik (2011, p. 864). This conclu sion certainly early Tuvalian age (late Carnian) be cause the Weser Fm is is correct only for “the long-Rhaetian option” (i.e., with the consid ered corre la tive with the Dolomie de Beau mont of Rhaetian age lasting above 8 Ma, begin ning from 209.5 France, which contains marine bivalves consid ered to be of Ma), given ten tatively for the “Geo log ical Time Scale such age (Lucas et al., 2012). As a further exam ple, the 2012” (GTS 2012) by Ogg (2012). More recently, Wotzlaw conchostracan fauna from the Co burg Sandstein of the Ger- et al. (2014) esti mated the age of the lower boundary of this manic Ba sin is consid ered late Carnian, seemingly be cause stage as 205.5 ± 0.39 Ma and the short dura tion of the it lies im medi ately below the begin ning of a sporomorph as- Rhaetian as 4.14 ± 0.39 Ma; simi lar data are given by so ci a tion con sid ered to be late Tuvalian”. This res er va tion Maron et al. (2015) as: 205.7 Ma and 4.4 Ma, re spec tively. is signif i cant also for the dating of the Krasiejów fauna, in An other ma jor un cer tainty is as so ci ated with the se lec tion the context of the Polish-Ger man corre la tions (Fig. 4). of global stratotype sections for the Upper Trias sic stage As to the possi ble lower Rhaetian dating through the boundaries in marine settings, still under discus sion (see the use of conchostracans, Gregoriusella polonica Zone was Carnian–Norian boundary candi date in Canada; Onoue et iden ti fied at Lipie Œl¹skie (as the type lo cal ity), but it is con - al., 2016), and con jec tural ma rine-ter res trial cor re la tions firmed elsewhere in the Tarnow (Mecklenburg) borehole (see e.g., Lucas et al., 2012; Maron et al., 2015; Tanner and “in the basal few meters of the Exter For ma tion” (Kozur and Lucas, 2015). The corre la tion of the Silesian bone-bearing Weems, 2010, p. 360). As noted by Edgar Nitsch (pers. level with the global strati graphic points will be a se ri ous comm., 2015): “The facies boundary from Arnstadt Fm chal lenge, (“Steinmergelkeuper”, var iegated) to Exter Fm (“Rät- This am bi gu ity es pe cially con cerns ter res trial suc ces - keuper”, grey) is diachronous, however, and ranges region - sions because of the weak nesses of the present palynozonal ally from Sevatian (for merly taken as part of the “Rhae- schemes, par ticu larly with re gard to the Norian dates, be - tian”) to Rhaetian sensu stricto. The “basal few meters” in cause of the tran si tional na ture of suc ces sive flo ras (Cirilli, the Schwerin–Rostock area are proba bly Sevatian in age. 2010). As well, Kürschner and Herngreen (2010, p. 267) The species could possi bly appear already in the Norian (as stressed: “The Norian is, at about 15 mil lion years, the lon- pointed out by Kozur him self in Kozur and Weems, 2010, p. gest Tri as sic stage, but the pol len and spore as semblages are 388), and its range still has to be assessed” (see also the insuf fi ciently known due to the scattered nature of the Lower Saxonian occur rence in Barth and Kozur, 2011). palynomorph record”. The Polish scheme of Or³owska- Con sid er ing the lat est Tri as sic (= Apachean) non-ma- Zwoliñska (1983) is based gener ally on acme zones, but rine tetrapod biochronology, Lucas (2010, p. 473) indi - even in the Polish Basin, Pieñkowski et al. (2014, p. 279) cated: “The Apachean is the most dif fi cult Trias sic LVF in di cated a dis sim i lar ity be tween the “Rhaetian” palyno- [land-verte brate faunachron] to corre late globally. This al- floras from Lipie Œl¹skie and Kamieñ Pomorski. Po ten tially most cer tainly re flects a provincialization of the global re liable range zones of Kürschner and Herngreen (2010), tetrapod fauna near the end of the Trias sic. Some of the ap- imple mented in the “Geo log ical Time Scale 2012", are still par ent endemism of Apachean land-ver tebrate assembla- char acter ized by unaware ness of the de tails of these ranges ges may also be due to facies, sampling and taphonomical (Fig. 18), exac er bated by “dis crep an cies in the tax on omy of bi ases”. In fact, this ex peri enced author even does not try to sporomorphs” (Cirilli, 2010, p. 305; compare Lucas, 2015). cor re late the Lipie Œl¹skie fauna with any of his two Norian- Not surpris ingly, Becker and Nawrocki (2014), using mag- Rhaetian chrones (see an expla na tion in Lucas, 2015). Dzik netostratigraphic cor re la tion, showed a sub stan tial diachro- et al. (2008b, p. 19) expressed a simi lar opinion: “Re place - nism of the macrospore-based palynozones in the Lower ments of one faunas to others in single geolog i cal profiles Triassic. are merely a record of lo cal envi ron men tal shifts but not of As shown above, a careful taxo nomic study of the Sile- global evo lu tion ary changes. The par tic u lar spe cies have sian flo ras and com par a tive for eign col lec tions is still changed their areas of distri bu tion, mov ing not un com- needed urgently (com pare Pacyna, 2014). In gen eral, mega- monly to hard-to-identify refu gee in remote regions of the fossil plant genera are more use ful in Upper Tri assic bio- world”. This warning also surely applies to the Krasiejów stra tig ra phy than mostly en demic spe cies (see dis cus sion in fauna, as well as to Silesian land flo ras (see discus sion in Lucas, 2013; compare Kelber, 2005). Lucas, 2013). The biogeographical differ ences between the 614 J. SZULC ET AL.

west ern and east ern parts of the Ger manic Ba sin, de ter - Sulej (2007, pp. 32–33), the bone bed was formed as fol- mined by “the unique nature” of the Silesian fau nas, are lows: “The Metoposaurus speci mens from the lower hori - mentioned by some authors (e.g., Sulej et al., 2011a; But ler zon were depos ited proba bly on the bot tom of a lake as par- et al., 2014). Sulej and NiedŸwiedzki (2010, p. 113) only tially decayed ca dav ers. This is sug gested by the common concluded cautiously in this con text: “the fossil record of asso ci a tion of skulls with mandi bles and bones of the pecto - tetrapod suc ces sion in the Late Trias sic was strongly con- ral girdle. Winnow ing was frequent and many disarticulated trolled by ecolog i cal factors and biased by uneven repre sen - bones oc cur in len tic u lar in ter ca la tions of cal car e ous grain- ta tion of par tic u lar en vi ron ments”. stone. The clavi cles usually occur with their convex side As reca pit u lated by Edgar Nitsch (pers. comm, 2015): down, in di cat ing de po si tion from sus pen sion. Closed shells “Biostratigraphy of the Keuper has always been a kind of a of unionid bivalves co oc cur, which proba bly died from suf- quiz game that has lost the answer cards (…) Up to now, all foca tion under the load of dysaerobic mud” (cf. also Ska- groups have their limi ta tions: macrophytes have typi cally wina, 2013). On the other hand, an actu ally allochthonous too broad age ranges to be very helpful, acme palynozones ter res trial Aetosaurus-Silesaurus assem blage occurs in an up- are prone to show some bias from ecol ogy, verte brates in- per fluvial fossiliferous level at Krasiejów, “1 m thick and clude some uncer tain ties due to endemism at low taxo nomic 15 m wide lens of red clay within fluvial cross-lami nated de- lev els, and the conchostracan zonation scheme is yet too posits” (Dzik and Sulej, 2007, p. 7). new to have overcome its teething com pletely”. In another Accord ing to Gruszka and Zieliñski (2008), the lower - ex treme perspec tive, Cesar Schultz (pers. comm., 2015) in - most Krasiejów bone-bear ing ho ri zon cor re sponds to the dicated in the context of the paper by Lucas (2015): “I con- basal layer of the graded channel suc ces sion, whereas the sider valid the (biostratigraphic) corre la tion between the highest inter val is located in the massive mudstone, overly - paleofaunas present in Polish bone beds with Lucas’ LVFs, ing the chan nel cy cle set. The main bone-brec cia level lies but I do not consider that the assign ment of a Carnian and/or within dark brown, massive, lacus trine claystones, capping Norian Age (ac cording to the Standard Global Chronostrati- the vertisol, but more detailed genetic expla na tion is not graphic Scale) for Polish bone beds be conclu sive in the ab- provided. Thus, two channel-hosted, bone-bearing levels sence of abso lute dating for the latter” . were in ter preted by Gruszka and Zieliñski (2008) as a hardpart burial record in moist niches of low-energy allu vial plains under season ally dry condi tions, such as flood basins, BONE CON CEN TRA TION PRO CESSES abandoned channels and resid ual ponds. The seasonal arid - AND TETRAPOD HAB I TATS ity in a dry to semi-dry cli mate was also stressed ear lier by Szulc (2005; see also Bilan, 1975), who linked the bone ac - Ge netic as pects of the bone beds, as well their sed imen - cumu la tion with ephemeral lakes and ponds. In a tectoni - tary settings, were dis cussed mostly for the ref er ence Kra- cally mo bile re gime paired with cat astrophic runoffs, the ac - siejów site, but no exhaus tive taphonomical study was un- cu mu lated skel etal parts were easily replaced through de - der taken. Dzik et al. (2000, pp. 228–229) mentioned the po- bris-flow mass movements. Szulc (2005, p. 164) claimed ssible current transport of cara paces and/or preda tor and that “the bone-bearing bed is a typi cal high-viscos ity debris scav en ger ac tiv ity in an al leg edly calm and ox y gen-de fi - flow de posit, encom passing a rela tively narrow sheet of re - cient lake milieu, but concluded: “This “grave yard” was not placed bone-bearing mudstones”. This fluc tu at ing cli ma tic the cat a strophic na ture and dif fer en ti a tion of size does not set ting was further confirmed by a histological study of the cor re spond to the dis tri bu tion in the liv ing pop u la tion there”. bone pattern of growth of metoposaurids (e.g., Konietzko- River-in duced al ka lin ity of the lake en vi ron ment was pre- Meier et al., 2013; Konietzko-Meier and Klein, 2013). dicted in the carbon ate-rich drainage area. This hydroche- Bodzioch and Kowal-Linka (2012) also em phasized a mical setting would be expected to have neutral ized humic trig ger role for ep i sodic cli mate-con trolled cat a strophic ac ids, re sult ing from the bac te rial de com po si tion of or ganic events in the ori gin of the main bone breccia. The palaeo- matter, and led to the conser va tion of apa tite and carbon ate histological, min er al og i cal and geo chem i cal study of diage- hardparts (also Dzik, 2003b), in addi tion to dimin ished scav - netic overprint, based on mineral types of infills and cal cite enging, preda tion and bioturbation in a hypoxic habi tat. ce ment and the chem i cal com po si tion of bone, con clu sively A more-or-less oppo site model of depositional events in di cates cru cial dif fer ences be tween the early diagenetic emerges from sedimentological stud ies (see re view in his to ries and the ini tial fos sil iza tion of aquatic and ter res - Bodzioch and Kowal-Linka, 2012). Szulc (2005) first pro- trial verte brate bone mate rial. Bodzioch and Kowal-Linka posed a sheet-flood fan model for the main bone-bed level (2012) deduced the fol lowing three-step or der pf events for (see below). In the opinion of Dzik and Sulej (2007, p. 6), bone-brec cia for ma tion: (1) an ep i sode of rapid de po si tion, this inter pre ta tion is in con flict with the “pres ence of sev eral indi cated by the erosional base directly above the vertisol such strictly hor i zon tal in ter ca la tions at var i ous lev els mas sive de pos its and eco log i cally mixed (aquatic-ter res - within the approx i mately one meter thick fossiliferous trial) fos sil assem blage, whereas (2) the in situ burial of bi- unit”, combined with “the common occur rence of thin hori - valve pop u la tions points to the sub se quent de vel op ment of a zontal concretionary layers with unionid bivalves having transient, shallow pond, that (3) quickly evapo rated fi nally their valves closed”. Thus, “a lo cal Metoposaurus–Paleo- dur ing a water less in ter val. Con se quently, the skel e tal re- rhinus biota with a suite of fish and inver te brate fauna and mains were cer tainly re worked post-mor tem and trans- al gal flora” is ad vo cated for the vast lac us trine marly ported from differ ent habi tats of the allu vial plain toward an claystone level (Dzik and Sulej, 2007, p. 7). In the words of ephem eral, shal low de pression in the course of high-energy UP PER TRI AS SIC BONE-BEAR ING LEV ELS 615

flood ing af ter pe ri odic cat a strophic rain fall. Ex treme-weather calam ity events in season ally rainy condi tions undoubt - edly influ enced the rapid tetrapod mortal ity and mode of fos sil iza tion. There fore, this con cen tra tion Lagerstätten (Konzentrat-Lagerstätten sensu Seilacher, 1970) is cer tainly not pro moted by an ex cep tional cat a strophic mass-mor tal ity burial event of a single living commu nity (obrution scenario of Seilacher et al., 1985), as stressed by Dzik et al. (2000). Re cur rent resedimentation phe nom ena and re lated can ni - balism of muddy de posits al ready were in dicated by Bilan (1975) for the Krasiejów suc ces sion (see also e.g., Bilan, 1976; Dzik et al., 2000; Szulc et al., 2006).

Fig. 21. Dia gram matic scheme of general factors leading to the Ge netic as pects of the Krasiejów bone-brec cia Upper Silesian bone con cen tra tions in the Norian Patoka Mbr. As reviewed in depth by Rogers and Kidwell (2007), Tetrapod pop u la tions, con trolled by pro nounced cli ma tic fluc tu a - tions, were preserved prefer en tially in an acme time during the bone beds can be grouped genet i cally in terms of either most prolonged wet pe riod, com bined with suit able flu vial depo- biogenic or physi cal (hydrau lic or sedimentologic) control sitional/burial and diagenetic set tings, and are partly re cov ered in processes, although more com plex taphonomical histo ries ran dom out crops to day. are frequent. Consid er ing these agents, the mixed forma tive scenario is evi denced conclu sively by Bodzioch and Ko- wal-Linka (2012; see also Gruntmejer et al., 2015), owing to dif fer en ti ated taphonomic path ways of jointly fos sil ized, tive effect of many small precur sor water bodies along the skel e tal parts. There fore, the spec tac u lar bone-brec cia level floodplain, and not a single large-scale lacus trine source (as at Krasiejów rep re sents a complex case. In fact, all Up per claimed by the Warsaw group). Silesian bone-en riched inter vals seem to cor respond to 2. Syndepositional and early diagenetic augmen ta tion phys i cal ver te brate skel e tal con cen tra tions of a com mon of fos sil iza tion po ten tial and re stric tion of de struc tive pro - mixed hydrau lic/sedimentologic type (see diver sity of alike cesses, primar ily owing to the high pH and low Eh condi - Cre ta ceous ex am ples in Wood et al., 1988; Ryan et al., tions (a con ser va tion as pect sensu Seilacher, 1967; Dzik et 2001; Bell and Campione, 2014 and Botfalvai et al., 2015; al., 2000; Dzik, 2003b; Dzik and Sulej, 2007; Bodzioch, also review in Behrensmeyer, 2007). 2015; see below). In the case of the main Krasiejów bone-brec cia level, 3. Proba ble partic i pa tion of a very rapid burial process, the ef fi cient cu mu la tive ac tion of wa ter trans port and trap - man i fested in ex traor di nary sed i ment dis charge, such as the ping mech a nisms in a lat er ally ex ten sive de pres sion seems high-viscos ity debris flows, proposed by Szulc (2005). As to be more im por tant than sed i men ta tion/ero sion ra tio (= the shown by Rog ers (2005), ex cep tion ally in ten sive rain falls flood/drowning causal cate gory of Behrensmeyer, 2007) to in the Late Cre taceous of Mad a gas car were re corded in ep i- an extraor di nary degree. However, Bodzioch and Kowal- sodes of cata strophic sedi ment-laden subaerial flow, paired Linka (2012) have shown an initial role for the re-concen - with the amazing ver tebrate burial of many ar ticu lated spec - tra tion of rich preex ist ing bone sources from the ad jacent al- imens in the resul tant, massive, muddy de posits (see also lu vial do main, char ac ter ized by a re duced sed i men ta tion Britt et al., 2009 and Flaig et al., 2014). In the case of not rate or even epi sodic sedi ment star vation, and the gradual, very prolonged transport (Bodzioch and Kowal-Linka, pas sive ac cu mu la tion of the tetrapod re mains (attritional 2012, p. 35), the as sem bled hardpart accu mu lation still mode; Behrensmeyer, 2007, p. 87). Never the less, signa tures could be eco log i cally co her ent and within-hab i tat time av er- of abrasion, weather ing and preda tion are rare. When com- aged only (i.e., parautochthonous). On the other hand, this bined with the abundance of partially artic u lated skele tons, para dox could have been enhanced by the truly autochtho- this in di cates short-dis tance post-mor tem trans port and nous nature of some biotic el ements (not only bi valves) in quick burial. The taphonomic circum stances led Dzik and the relict aquatic ba sin (in the event scenario of Bodzioch Sulej (2007) to the final conclu sion that the “lake-dwell ing and Kowal-Linka, 2012). Metoposaurus–Paleorhinus biota” is of a lo cal and – im- plic itly – of dom inantly autochthonous nature. However, as Bone beds of the Lisowice level stressed by Lucas et al. (2010; cf. also Brusatte et al., 2015), the Krasiejów bone brec cia is differ ent from most metopo- The multitaxic and largely low-den sity ver tebrate ac cu - saurid bone beds that are nearly monotaxial and certainly mu lations in the higher parts of Patoka Mbr are overall com - monodominant, and inter preted as a mass-death record of pa ra ble to sub or di nate tetrapod oc cur rences at Krasiejów, breed ing pop u la tions. There fore, a com bi na tion of three strongly dom inated by the hardparts of scat tered ter restrial factors should be consid ered: species (with the ex ception of Porêba) for depos its varying 1. Large density and fertil ity of living tetrapod popu lain extent. The sedi men tary con text of the Lisowice-type tions, even if not ex actly coex istent (i.e., a time-av er ag ing tetrapod oc cur rences is more con jec tural, how ever. Typ i cal fac tor). Cer tainly many thou sands of in di vid u als were killed for the Steinmergel facies, short-term clima tic rhythmicity by the disas trous flooding, but this was a uniquely cumu la- re cords no tice able dry-wet sea son al ity in a mon soon-like or 616 J. SZULC ET AL.

trade-wind-influ enced system (Reinhardt and Ricken, 2000; rensmeyer (1988) dis tin guished between Lagerstätten con- Szulc et al., 2006; Bodzioch and Kowal-Linka, 2012; see cen trated by ac tive drain age (chan nel-lag as sem blages) and the discus sion of orbital controls in Vollmer et al., 2008). those that ac cu mu lated in an in ac tive chan nel (chan nel-fill Vast, extremely flat, muddy lowlands in wet peri ods and the as semblages). The lag va ri ety oc curs in coarse-grained conse quent floral acme offered many niches for a di verse lithologies, depos ited at the erosional base of a channel suc- fauna, while the hab itable lo cali ties con tracted to the envi - cession, and contains – in ex treme cases – allochthonous, rons of water sources during arid inter vals (Fig. 21). The destroyed and mostly uniden ti fi able bone pebbles that expe - muddy de pos its were pe ri od i cally pedogenically mod i fied ri enced several cycles of re work ing. The fill mode is as so ci- in the Late Trias sic cli matic set ting (e.g., Sarkar, 1988; ated with fine-grained to mixed fa cies of chan nels, es sen - Therrien and Fastovsky, 2000; Szulc, 2005). Verte brate re- tially abandoned by flow and the resul tant Lagerstätten may mains, when subaerially exposed to weather ing and abra- in clude even com plete skel e tons. sion, were pro gres sively de stroyed, par ticu larly as a re sult A chan nel-lag as semblage is clearly ev idenced in the of these soil processes (e.g., organic acid ifi ca tion, vary ing lower most fossil level at Krasiejów (Gruszka and Zieliñski, oxi da tion states in wet/dry seasons). Thus, the accu mu lated 2008) and in the lower hori zon at Lipie Œl¹skie (Pieñkowski hardparts did not survive the prolonged deg rada tion process et al., 2014; see Fig. 5). This cate gory also may be applied (Dzik and Sulej, 2007, p. 7), espe cially in peri ods of sedi - ten ta tively to the oc cur rences of dif fer ently pre served hard- ment starva tion. A genetic link between hardpart concen tra - parts in the carbon ate (partly reworked vadoids, Lisów brec- tions and discon ti nu ity surfaces is not observed in Keuper cia-type) or polymictic con glom eratic depos its at Porêba (ho- suc ces sions (a typ ical case for the fos sil re cord; Rog ers and ri zons a, c and d, NiedŸwiedzki et al., 2014; Fig. 14) and Kidwell, 2000). On the other hand, even bone lags are re- Zawiercie (Racki, 2010, fig. 1B). Less clear is the in ter pre - ported from greenish- and bluish-gray palaeosoil hori zons tation of the fossil-bear ing grey grainstones with “ooid-like in varie gated muddy floodplain facies (Eocene Willwood struc tures”, also compa ra ble to the “Lisów breccias” , in the Forma tion of Wyo ming; Bown and Kraus, 1981; see Upper WoŸniki Lagerstätte (Sulej et al., 2011a; Fig. 10), as well as Trias sic exam ple in Therrien and Fastovsky, 2000). a bone occur rence in the gravelly sands of the Patoka 1 The skele tons also were exposed to inten sive scaveng - borehole section (depth 119.7 m). In partic u lar, the Porêba ing and cracking by large archosaur (?dino saur) carni vores local ity is distin guished by numer ous, highly eroded frag - prior to burial, as ev idenced at Marciszów (Budziszewska- ments of tur tle cara pace or plastron in coarse-grained Karwowska et al., 2010). At Lipie Œl¹skie, about 12% of the lithologies, and the mixing of ele ments from aquatic and dicynodont bones exhibit tooth bite or patholog i cal traces ter res trial hab i tats is ev i dent in “a high-energy fluvial sys - that clearly man ifest a sig nif icant scale of pred a tor-prey in - tem” (Sulej et al., 2012, p. 1039). However, a resid ual hy - ter ac tions in that time (NiedŸwiedzki et al., 2011; see also draulic lag of highly abraded, dura ble ele ments was not ob- Pieñkowski et al., 2014). served at the sites, although actualistic analy sis does not The more-or-less bi ased death as semblages, de rived show a sim ple as so ci a tion be tween me chan i cal re work ing from within-hab i tat sources, were ep i sod i cally win nowed, effects and the length of transport before burial (Aslan and transported and finally concen trated in sedi ment traps during Behrensmeyer, 1996). re cur rent overbank in un da tions on dif fer ent scales, al though The up per most Krasiejów Lagerstätte cor re sponds to ex treme rain falls re sulted in ma jor flood ca tas tro phes and the chan nel-fill as semblages, lo cated in the massive mud- mass mortal ity, as envi sioned above for the Krasiejów bone stone cap of the chan nel-fill cy cle (Gruszka and Zieliñski, brec cia. The muddy sed iments of the Patoka Mbr were exten - 2008), where the mostly transported Aetosaurus-Silesaurus sively eroded, reworked and rede pos ited (Bilan, 1975, 1976; as semblage is ev idenced, but char ac ter ized by the high ar - Szulc et al., 2006, 2015) and the sig nifi cant transfor ma tion of ticu lation of many speci mens (Dzik, 2003a; Dzik and Sulej, river-stream systems was therefore a fre quent phe nome non. 2007; Piechowski et al., 2014). The authors assume a sim i- In con se quence, the bone mate rial could es cape (near)sur face lar taphonomical cate gory for the rich as semblage of ter res - degra da tion, and when rapidly buried in flood or mudflow trial tetrapod remains from the main dark organic-rich sedi ments, could be found only in the wet in ter vals with more mudstone level at Lipie Œl¹skie (Fig. 5), and also for the in ten sive flu vial ac tiv ity (Szulc et al., 2006, p. 149; see Fig. fish-dom i nated as sem blage at Porêba, collected as well-pre - 20). Ob vi ously, at least partly permineralized (“prefossili- served, isolated bones in the organic-rich hori zon b of zed”) spec imens, pre served to dif fer ent de grees, were as sem- NiedŸwiedzki et al. (2014; Fig. 13). However, even in this bled, as shown for verte brate signa tures from simi lar set ting a post-mor tem mix ing was sig nifi cant, as implied by low-gra dient river sys tems by Mukherjee and Ray (2012) in the diagenetic pattern for dicynodont bones at Lipie Œl¹skie the Upper Trias sic Tiki Forma tion of India, and, for exam ple, by Bodzioch et al. (2015). In fact, the palustrine facies of re- by Botfalvai et al. (2015) in an Upper Creta ceous Iharkút sidual ponds, mires and marshes frequently may develop in ver te brate as sem blage (Hun gary). The skel e tal ma te rial and the abandoned channels and this would be an effec tive the car casses of an i mals were re dis trib uted down stream and attritional trap for well-pre served skel e tal mate rial (Dzik et two effi cient trapping mecha nisms are known: in-channel al, 2008a, 2008b; NiedŸwiedzki et al., 2012). However, a blockades (e.g., tree trunks) and rapid falls in hydrau lic com - distinc tion between the channel-fill and floodplain succes - petence, usually at sin u ous chan nel bends (Aslan and Beh- sion is equiv o cal with out more de tailed spa tial doc u men ta - rensmeyer, 1996; Rogers and Kidwell 2007, p. 24). tion of partic u lar lithosomes in the river fa cies archi tec ture. There fore, the chan nel-hosted bone beds include pre- Floodplain muds can also provide rarer, but well-preserved, cursor mate rial from a vari ety of floodplain settings. Beh- even partly ar ticu lated bone ma terial, as noted for the UP PER TRI AS SIC BONE-BEAR ING LEV ELS 617

Fig. 22. Stra tig ra phy-fa cies and climatostratigraphy schema of the mid dle and up per Keuper of Up per Silesia, to show over all strati - graphic po sition and re la tion ships of bone- and flora-rich ho ri zons (with ref er ence to four gener al ized suc ces sion types, see Fig. 17 and Szulc et al., 2015; mod i fied af ter Szulc and Racki, 2015, fig. 9); palyzonation scheme af ter Or³owska-Zwoliñska (1983; see Fija³kowska- Mader et al., 2015).

Krasiejów succes sion by Dzik and Sulej (2007, p. 7). The level bone beds were proba bly depos ited mostly in multi ple ver tebrate hardparts seem to be as so ciated with marly mud- channel belts by a mostly low-gradi ent river sys tem (see be- stones and/or limestone concre tions, as well known at Kra- low). In Behrensmeyer’s (1988) model, attritional chan- siejów and Lipie Œl¹skie (Dzik et al., 2000, 2008a), but also nel-fill assem blages should be more frequently gener ated in at Marciszów (Racki, 2010, fig. 1A; Fig. 15D) and in the aban doned chan nels by anal ogy to Car bon if er ous bone beds, Kobylarz 1 well core (?mudflow recorded in grey mudsto- asso ci ated with sapropels (Behrensmeyer, 2007, p. 89). nes; depth 41.0 m; Szulc et al., 2015, fig. 13). Inter est ingly, Behrensmeyer (1988) stressed that the lat- The mid-Norian pluvial prelude erally migrat ing river would be expected to form channel-lag and tec tonic-flu vial event bone beds, in con trast to avulsion-dom inated, braided to anastomosing networks. This general deduc tion is not sub- The Lisowice bone-rich level is thought to have re- stan tiated in the Krasiejów suc ces sion, where, for ex am ple, sulted from a combi na tion of several factors (Figs 21, 22): the fill concen tra tion mode is noted only in the mean der ing (1) a more prolonged in ter val of cli matic pluvialization, a river stage of Gruszka and Zieliñski (2008). The Lisowice- prelude to the Rhaetian humid climate (Feist-Burkhardt et 618 J. SZULC ET AL.

Fig. 23. Sketch di a gram of en vi ron mental evo lu tion and main fa cies dis tri bution of the south ern Ger man Keuper (from Kelber and Nitsch, 2005, fig. 2), with ref er ence to the Silesian lithostratigraphy scheme and strati graphic con text of bone-rich lev els. Tetrapod re con - struc tions are af ter Dmitry Bogdanov (https://pl.wikipedia.org/wiki/Dicynodonty#/me dia/File:Dicynodont_from_PolandDB.jpg; https://up load.wikimedia.org/wikipedia/com mons/6/6b/Metoposaurus_diagnosticus_kraselovi_1DB.jpg). al., 2008; Fija³kowska-Mader, 2015b), that re sulted in (2) a Lisów brec cia lev els; see the above quoted con cept of Dzik floral acme, as a trophic basis for (3) more abundant and di- et al., 2000). More likely for most sites of the Lisowice verse ver te brate life. Ad di tional cir cum stances were in flu - level, the burial places were fed by an exten sive, fault-boun- enced sig nif i cantly by (4) tec tonic re ju ve na tion of sed i men- ded spring system in its initial phase, just before the com - tary ba sin to pog ra phy, man i fested prob a bly in block move- mon palustrine depo sition of the WoŸniki Limestone (an ments indi cated by very vari able, but lo cal ized hia tuses impli ca tion of the model of Szulc et al., 2006). Micro bial (e.g., Bilan, 1976). In effect (5), a braided to anastomosing films and in duced car bon ate pre cip i ta tion, re corded in com - river network devel oped (= mid-Norian tectonic-plu vial mon oncolites and stromatolites (see Szulc et al., 2015, fig. event; Figs 17, 22), as indi cated by the rarity of sheet sand- 16; Fig. 15F), might be an effi cient protec tive agent (see stones and lat eral ac cre tion surfaces. This early Cimmer ian Seilacher et al., 1985; Seilacher, 2008). An addi tional, pos- event (see Szulc et al., 2015) was proba bly controlled by the sible source of calcium and bi car bon ate ions was a bac teri - overall up lift of source areas and subse quent progradation ally induced, early diagenetic alter ation of ‘fresh’ bone ma- of allu vial fans in the envi ron ments of exten sive muddy- te rial (with soft tis sues), es pe cially in that cal cite is the main sandy flats. In fact, a more ex ten sive drain age sys tem, evol- mineral in bone voids at Krasiejów and Lipie Œl¹skie (Bo- ving with progres sive peneplenisation toward multi ple dzioch, 2015; Bodzioch et al., 2015). channel belts (see e.g., Makaske, 2001), promoted not only more diverse moist niches within the gently undu lat ing mud Ver te brate eco sys tems flats, but also strengthened ex hu mation, burial and preser - vation poten tial in the proxi mal Keuper facies (see below). The lithofacies suc ces sion of the Upper Tri assic of the All the fac tors re cord rather or di nary eco log i cal and Ger manic Ba sin al most per fectly re flects cli ma tic fluc tu a - geo log i cal cir cum stances in the mid-Norian plu vial/tec - tions overwhelm ing the West ern Tethys domain dur ing tonic-fluvial inter val, but (6) an addi tional taphonomical en- Carnian–Rhaetian time (Feist-Burkhardt et al., 2008). Cer- hance ment of early fos sil iza tion pro cesses was prob a ble tainly, the fun da mental cli matic turn over in the Late Tri as - both in the organic-rich swamp muds or overbank conglom - sic, which took place at about the Norian-Rhaetian bound- eratic depos its (Fig. 21). As mani fested by a frequent link of ary, consisted of a shift from dry/semi-dry to perma nently bone oc cur rences with car bon ate con cre tions or marlstone wet cli ma tic con di tions, partly vol ca ni cally in duced and of lay ers, an oc cur rence of cal cium-rich in ter sti tial flu ids, an extreme greenhouse type, which culmi nated during the stim u lat ing also com mon mi cro bial lime pre cip i ta tion, in - end-Tri as sic eco log i cal ca tas tro phe (Feist-Burkhardt et al., deed might have been crucial. This hydrochemical factor 2008; Preto et al., 2010; Pieñkowski et al., 2014; Fija³- was controlled mostly by the karstification of uplifted kowska-Mader, 2015b), recorded in floral shift (see below). Muschelkalk blocks and/or caliche soil degra da tion (cf. This turnover is re corded as the replace ment of a lithofacies UP PER TRI AS SIC BONE-BEAR ING LEV ELS 619

Fig. 24. En vi ronmen tal model for the Lipie Œl¹skie tetrapod local ity (after Jewu³a, 2010, fig. 22); for est fires are shown af ter Marynowski and Simoneit (2009).

of dry condi tions (redbeds includ ing local evaporites) by occu pied the highest level in the food pyra mid of the eco - wet, pluvial kaolinite-enriched lithofacies, includ ing fluvial systems (compare Mukherjee and Ray, 2012). and pe ren nial, lac us trine de pos its. As high lighted above, The Lisowice level re cords an impor tant wet in ter val in tetrapod popu lation sizes were certainly controlled by the the mid-Norian, but also an overall facies-tec tonic remod el- clima tic fluctu a tions and they thrived mostly during the ling. There fore, vari able al lu vial to palustrine en vi ron ments wet ter pe ri ods and re lated veg e ta tion growth in wet land prob a bly char ac ter ized this plu vial in ter val (chan nels, muddy hab i tats (Figs 21–23). floodplains, marshy ponds to small lakes etc.; Szulc, 2007a). The likely lo cal na ture of the Krasiejów-type biotope in Pieñkowski et al. (2014, p. 272) cate go rized the fossiliferous west ern Upper Silesia sug gests the devel op ment of exten - fa cies of Lipie Œl¹skie as: “(…) a mosaic of al lu vial-plain en- sive wetlands inside mean der ing to anastomosing rivers, viron ments swamps with ephem eral ponds and later ally shift- flowing from the rather degraded Variscan massif of the ing river channels” (see Fig. 24). Sudetes, situ ated several dozen kilometres to the southwest In palynofacies terms, lacus trine (or more ac cu rately, (cf. Dzik et al., 2000; Dzik, 2003b; Gruszka and Zieliñski, stag nant wa ter) (Porêba, Lipie Œl¹skie), floodplain (Koby- 2008; Bodzioch and Kowal-Linka, 2012). The Sudety Land larz 1) and floodplain or lacus trine to playa (Patoka 1) habi - (sensu Pieñkowski et al., 2014) prove nance was ev idenced tats are rec og niz able (Fija³kowska-Mader et al., 2015). In recently by the geochem i cal signa tures of bulk mudstone the palynomorph spectra, the increas ing Norian clima tic hu- sam ples (Konieczna et al., 2015). As supposed by Edgar midity to ward the “late Norian pluvial event” is de tectable Nitsch (pers. comm., 2015), the verte brate acme might have by an expan sion of hygrophytic ele ments (lycopsids and re corded “a kind of ‘mid-Tuvalian pluvial event’ that hap- equisetales) within the eurytopic xero phyt ic biota dom i- pened in the Bohe mian Massif (and neighbor ing Sudety nated by cheirolepidacean co ni fers (Fija³kowska-Mader, Mas sif) and prop a gated al lu vial (gyp sum-free) fa cies to re- 2015b; see also Or³owska-Zwoliñska, 1983; Fija³kowska- lated ba sin-mar ginal set ting”. An in creas ing hu mid ity in Mader et al., 2015; compare Brachyphyllum and Lepidopte- fact is ev idenced by the sedimentological at trib utes of the ris floras of Pacyna, 2014). This conclu sion is supported by Krasiejów succes sion (Gruszka and Zieliñski, 2008). Vast, an a tom i cal char ac ters of the co ni fer wood of Agathoxylon prob a bly co ni fer-dom i nated for ests, with ferns and benne- keuperianum from Zawiercie area (Philippe et al., 2015). ttitaleans (Voltzia flora of Pacyna, 2014), and a pleth ora of Thus, Fija³kowska-Mader (2015b) char ac ter ized these dif fer ent-sized fresh wa ter bas ins oc cu pied by charophyte enviromments as: “Moist to wet in land floodplain was oc- al gae (Zatoñ et al., 2005), were a prereq ui site factor for the cupied by lycopsids and equisetales. Dry uplands were cov- rich fauna flourish ing in the river catchment draining Su- ered with coni fer forests consist ing mainly of cedars. Chei- dety Land. In par tic u lar, metaposaurid am phib i ans richly rolepidacean formed bushfields along river banks and la- popu lated the ponds and small open lakes. The ubiqui tous goon coastal zones (...)”. A facies tran sition from carbon ate am phib i ans and the semi-aquatic croc o dile-like parasuchids to fine siliciclastic depositional re gimes along the Tethys 620 J. SZULC ET AL.

shelf might have been, at least in part, a signa ture of this nian to Rhaetian, as sumed in pa pers of the War saw re search same clima tic shift (Berra et al., 2010; see also Preto lain et group, should not be accepted, as their datings and corre la - al., 2010). Flo ral turnover took place in far more humid set- tive infer ences are poorly docu mented, flawed and ig nore tings in the Rhaetian, when pteridophyte vege ta tion defi - the results of other studies. nitely out num bered co ni fer for ests (Or³owska-Zwoliñska, 2. On the basis of a gener ally con sis tent com pila tion of 1983, p. 65; see also Krupnik et al., 2014; Fija³kowska- litho-, bio-, climato- and chemostratigraphic premises (Fig. Mader, 2015b). The wet and warm condi tions also are re- 17), the present authors propose two Norian bone-bearing corded in the thriving nature of the Agathoxylon keuperia- lev els, in a time in ter val less than 10 Ma (Fig. 19). Both lev- num trees (found at Patoka; Philippe et al., 2015). els no ticeably cor re late with the lower part of C. meyeriana In the spa tially het er o ge neous eco sys tem, di verse rep - Palynozone (likely mostly with the same IVb Subzone) and til ian-dom i nated land com mu ni ties thrived, with un usu ally the same II Cr/Ti Chemozone. The authors stress the im pre- large, her bivo rous dicynodonts and the giant top preda tor, cise charac ter of this dating, due to the vagueness of Late the archosaur Smok wawelski, in a most spec tacu lar role in a Tri as sic bio- and chronostratigraphy in ter res trial settings, rather com plex trophic ar ray (see Pieñkowski et al., 2014, and admit to a feasi ble amount of in forma tion noise. Their fig. 9; Zatoñ et al., 2015). In the Porêba as semblage, the top conclu sion, however, is well grounded in terms of Ockh- car ni vores in clude also me dium-to-large-bod ied di no saurs am’s ra zor. (NiedŸwiedzki et al., 2014). When com pared with the 3. The Krasiejów succes sion is placed in the lower part Krasiejów biota, aquatic life seems to have been impov er - of the Grabowa Fm, includ ing in its basal part a transi tion ished, restricted largely to fishes and local pop u lations of from the evapo rate-bear ing Ozimek Mbr to the Patoka Mbr turtles and amphib i ans. However, a strong taphonomical (= equiv alents of tran sitional beds between the Up per Gyp- bias, evi dent in the more-or-less reworked as sem blages, sum Keuper and Steinmergelkeuper). The Krasiejów bone- precludes the pos si bil ity of fur ther con sid er ation. brec cia level, limited to the Opole re gion, rep re sents most As al ready highlighted, the bone-bearing grey flu vial proba bly the early Norian inter val. Of course, the authors depos its of the Lisowice level, when ana lyzed in terms of can not pre clude the pos si bil ity that fu ture cor re la tion of the the gen eral cli mat i cally-con trolled lithofacies suc ces sion of criti cal profile with an approved global stratotype for the the Central Euro pean Keuper, appear to be most likely the Carnian–Norian boundary will lead to its re-loca tion in the time and fa cies equiva lent of the middle Norian, Middle late Carnian. Grey Series and Middle Löwenstein Fm from Thuringia and 4. The more widely distrib uted Lisowice bone-bearing Brandenburgia (cf. Franz, 2008; see also Milner and Schoch, level (also the WoŸniki and Porêba-Zawiercie sites) cor re - 2004, Schoch and Wild, 1999, Seegis, 2005 and Sues and sponds to the mid-Norian middle part of the Grabowa Fm Fraser 2010, pp. 89–95; Fig. 23). As noted by Milner and (= lower part of Patoka Mbr), and re cords a niche mo saic in Schoch (2004), these differ ent Germanic facies contain dis- an exten sive allu vial plain (braided to anastomosing river sim i lar rep til ian fau nas, char ac ter ized by plagiosaurs, capito- sys tem) dur ing the Eo-Cim mer ian tec tonic-plu vial ep i sode saurs, crocodylomorphs, and di no saurs, while metoposaurid re corded in the entire Ger manic Ba sin. am phib i ans were an in fre quent com po nent of the phytosaur- 5. The re vised local ity tim ing highlights the very am- tur tle assem blage. So, one can only as sume a simi lar dif fer - bigu ous problem of the evolu tion ary uniqueness of the Sile- en ti a tion of Up per Silesian tetrapod com mu ni ties. sian biotas, jointly with the mostly lost refugium status of The flu vial-erosive events, dis cussed above, corre late the ‘Rhaetian’ Lipie Œl¹skie fauna, and elimi nated a surpris - very well with simi lar ep i sodes, iden ti fied in other pe riph - ing contrast be tween the Rhaetian ver tebrate as semblages, eral zones of the Central Eu rope Basin, repre sented in Ger - rich in Silesia and very poor in Germany (see Schoch and many by the coarser-clastic Löwenstein Forma tion and the Wild, 1999; Seegis, 2005; Sues and Fra ser, 2010). A Norian sim i lar Kågeröd Forma tion in Den mark and Sweden (Franz, age for the Lisowice as semblage fits the global dis tri bu tion 2008). It is also note worthy that the great est and the most pattern much better (cf. Lucas 2015). On the other hand, this im portant local ities of the bone breccia of the Upper Tri as- char ac ter istic now may be as signed in part to the Krasiejów sic in Central Europe are asso ci ated with proxi mal Keuper as sem blage, at least to the dom i nat ing am phib i ans, but this fa cies (e.g., Löwenstein Fm, Schoch and Wild, 1999; fossil record proba bly is affected by a basal Norian hia tus Seegis, 2005). The marginal basin zones were the most rich (the main Eo-Cimmer ian disconformity). in wa ter, and, there fore, there were the most fa vor able 6. The Upper Silesian bone beds corre spond largely to niches for tretrapod commu ni ties on land. Intense fluvial typ i cal skel e tal con cen tra tions of com bined hy drau lic/ pro cesses (in clud ing di sas trous floods) fa cil i tated the rapid sedimentologic type and of fluvial ori gin. The taphonomic burial of organic remains that allowed the forma tion of the en hance ment of pres er va tion by hydrochemical (al ka lin ity) many Upper Silesian bone beds. and/or microbially- or bac te ri ally-in duced (synsedimentary or diagenetic) car bon ate pre cip i ta tion can be as sumed as well. Thus, a mixed con cen tra tion/con ser va tion na ture of CON CLU SIONS the Fossilagerstätten is consid ered on a prelim inary basis in sev eral cases, but in par ticu lar for the main breccia level at 1. On the ba sis of a multidisciplinary strati graphic study Krasiejów, proba bly distin guished by a vari ety of obrution of the complete compos ite section of the Silesian Upper Tri- mecha nisms, such as viscous mudflow. assic, the authors have concluded that three widely sepa - rated bone-enriched inter vals, ranging in age from the Car- UP PER TRI AS SIC BONE-BEAR ING LEV ELS 621

Ac knowl edg ments tatis Geologorum Poloniae, 85: 697–701. Becker, A., Kuleta, M., Narkiewicz, K., Pieñkowski, G. & Szulc, The thor ough re view and com ments by Spencer Lucas and J., 2008. Trias. In: Wag ner, R. (ed.) Tabela stratygraficzna Adam Bodzioch sig nif i cantly improved the manu script. In par tic u - Polski, Polska pozakarpacka. Pañstwowy Instytut Geolo- lar, the manuscript benefited greatly from the construc tive remarks giczny, Warszawa. [In Polish.] and com ments of Ed gar Nitsch who also kindly per mitted the use Becker, A. & Nawrocki, J., 2014. Magnetostratigraphy of the of two fig ures. We are grate ful also to Mike Benton, Simonetta Buntsandstein (Lower Tri assic) in the Gorzów Wielkopolski Cirilli and Anna Fija³kowska-Mader for help ful com ments on ear - IG 1 bore hole, eastern Ger man Ba sin in Poland: ev i dence of lier ver sions of the pa per. We thank Krystyn Rubin for shar ing substan tial diachronism of palynostratigraphic macro spore drill core from the WoŸniki K1 bore hole, and Waldemar Bar- zones. Geo log i cal Quar terly, 58: 369–378. dziñski for help in field work. We thank also Gernot Arp and Behrensmeyer, A. K., 1988. Ver te brate pres er va tion in flu vial Dmitry Bogdanov who per mits to use their fig ures in our pa per. chan nels. Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol - The study was supported by the Pol ish Minis try of Sci ence and ogy, 63: 183–199. Higher Ed u ca tion (grant N307117037 for Grzegorz Racki). Behrensmeyer, A. K., 2007. Bonebeds through time. In: Rog ers, R. R., Eberth, D. A. & Fiorillo, A. R. (eds), Bonebeds: Gen e - sis, Anal y sis and Paleobiological Sig nif i cance. The Uni ver - REF ER ENCES sity of Chi cago Press, Chi cago, pp. 65–101. Bell, P. R. & Campione, N. E., 2014. Taphonomy of the Danek Antczak, M., 2016. Late Trias sic aetosaur (Archosauria) from Bonebed: a monodominant Edmontosaurus (Hadrosauridae) Krasiejów (SW Poland): new spe cies or an ex ample of in di - bonebed from the Horse shoe Can yon Forma tion, Al berta. Ca- vid ual vari a tion? Geo log i cal Jour nal (in press). DOI: na dian Jour nal of Earth Sci ences, 51: 992–1006. 10.1002/gj.2691. Benton, M. J., 2004. Ver te brate Palae on tol ogy, 3rd Edtion. Arp, G., Bielert, F., Hoffmann, V.-E., & Loeffler, T., 2005. Blackwell, Ox ford, 455 pp. Palaleoenvironmental sig nif i cance of lac us trine stromatolites Benton, M. J., Forth, J. & Langer, M. C., 2014. Models for the rise of the Arnstadt For mation (“Steinmergelkeuper”, Up per Tri - of the di no saurs. Cur rent Bi ol ogy, 24: R87–R95. as sic, N-Ger many). Fa cies, 51: 419–441. Berra, F., Jadoul, F. & Anelli, A., 2010. En vi ron mental con trol on Aslan, A. & Behrensmeyer, A. K., 1996. Taphonomy and time res - the end of the Dolomia Principale/Hauptdolomit depositional o lu tion of bone as sem blages in a con tem po rary flu vial sys - sys tem in the central Alps: Coupling sealevel and climate tem: The East Fork River, Wy o ming. Palaios, 11: 411–421. changes. Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol - Assmann, P., 1926. Die Tiefbohrung “Oppeln”. Jahrbuch der ogy, 290: 138–150. Preussischen Geologischen Landesanstalt, 48: 332–335. Beutler, G., 2008. Keuper. In: Beutler, G., Ehling, B.-C., Eichner, Assmann, P., 1929. Die Tiefbohrung Leschna und ihre Bedeutung R. & Schwab, M. (eds.), Geologie von Sachsen-Anhalt. fuer die Stratigraphie der oberschlesischen Trias. Jahrbuch Schweitzerbart’sche Verlag. Stuttgart, pp. 213–230. der Preussischen Geologischen Landesanstalt, 50: 155–185. Beutler, G. & Nitsch, E., 2005. Palaeogeographischer Ueberblick. Bachmann, G. & Kozur, H. W., 2004. The Ger manic Tri assic: cor - In: Beutler, G., Hauschke, N., Nitsch, E. & Vath, U. (eds.), re la tions with the in ter na tional chronostratigraphic scale, nu - Stratigraphie von Deutschland IV. Keuper. Cou rier For- meri cal ages and Milankovitch cyclicity. Hallesches Jahr- schungsinstitut Senkenberg, 253: 15–30. buch für Geowissenschaften Reihe B, 26: 17–62. Bilan, W., 1975. The Rha etic pro file in Krasiejów near Opole. Bachmann, G. H., Geluk, M. C., Warrington, G., Becker-Ro man, Zeszyty Naukowe AGH, Geologia, 1 (3): 13–19. [In Pol ish, A., Beutler, G., Hagdorn, H., Hounslow, M. W., Nitsch, E., with English summary.] Röhling, H. G., Simon, T. & Szulc, J., 2010. Trias sic. In: Bilan, W., 1976. The stra tig ra phy of the Up per Tri as sic de posits of Doornenbal, J. C. & Stevenson, A. G. (eds), Pe tro leum Geo- the east ern mar gin of the Up per Silesian Coal Ba sin. Zeszyty log i cal At las of the South ern Perm ian Ba sin Area. EAGE Naukowe AGH, Geologia, 2 (3): 4–73. [In Pol ish, with Eng - Publi ca tions b.v., Houten, pp. 149–173. lish summary.] Bajdek, P., Owocki, K. & NiedŸwiedzki, G., 2014. Pu ta tive dicy- Bilan, W., 1991. Biostratigraphy of Upper Tri assic sed i ments of nodont cop ro lites from the Up per Tri as sic of Poland. Palaeo - the east ern mar gin of Up per Silesian Coal Ba sin. Zeszyty ge ogra phy, Palaeoclimatology, Palaeo ec ol ogy, 411: 1–17. Naukowe AGH, Geologia, 17 (1): 5–17. [In Pol ish, with Eng - Bardziñski, W. & Chybiorz, R., 2013. Opracowanie kartogra- lish summary.] ficzno-geologiczne wybranych ods³oniêæ kajpru Górnego Œl¹- Bodzioch, A., 2012. Gen e sis and cur rent state of knowl edge of ska (w oparciu o materia³y kartograficzne, obserwacje tere- bone ac cu mu la tions at Krasiejów. In: Jagt-Yazykova, E., nowe i dane fotointerpretacyjne). Un published Grant Re port, Jagt, J. W. M., Bodzioch, A. & Konietzko-Meier, D. (eds), Faculty of Earth Sciences, Univer sity of Silesia, Sosnowiec, Krasiejów – paleontological in spi ra tions. Stowarzyszenie 12 pp. [In Pol ish.] „Dinopark” & Zak³ad Paleobiologii Katedry Biosystematyki Barth, G. & Kozur, H. W., 2011. A Lat est Norian age for in - Uniwersytetu Opolskiego, Opole, pp. 28–35. [In Pol ish, with sect-bear ing beds of the Fuchsberg and Langenberg near Eng lish sum mary.] Seinstedt, north ern fore land of the Harz Mountains (Lower Bodzioch, A., 2015. Ide al ized model of min eral in fill ings in bones Sax ony, Ger many). In: Sullivan, R., Lucas, S. & Spielmann, of fossil freshwa ter ani mals, on the exam ple of Late Trias sic J. (eds), Fossil Re cord 3. New Mexico Mu seum of Nat ural metoposaurs from Krasiejów (Poland). Aus tin Jour nal of His tory and Sci ence Bul le tin, 53: 157–165. Earth Sci ence, 2: 1008. Barycka, E., 2007. Mor phol ogy and on tog eny of the hu merus of Bodzioch, A. Konietzko-Meier, D. & Jagt-Yazykova, E., 2010. the Tri as sic temnospondyl am phib ian Metoposaurus diagno- Wycieczka I. Œl¹skie stanowiska paleontologiczne ze szcz¹t- sticus. Neues Jahrbuch für Geologie und Paläontologie, Ab- kami krêgowców triasowych: Stanowisko dokumentacyjne handlungen, 243: 351–361. „Trias”w Krasiejowie (punkt 3). In: Zatoñ, M., Krawczyñski, Becker, A., 2015. Am bi gu ities in conchostracan biostratigraphy: a W., Salamon, M. & Bodzioch, A. (eds), Kopalne Biocenozy w case study of the Perm ian–Tri as sic bound ary. Annales Socie- Czasie i Przestrzeni. XXI Konferencja Naukowa Sekcji 622 J. SZULC ET AL.

Paleontologicznej Polskiego Towarzystwa Geologicznego, Pub li ca tion, 334: 221–262. ¯arki-Letnisko 13–16 wrzeœnia 2010 r. Wydzia³ Nauk o Clemmensen, L. B., Kent. D. V. & Jenkins, F. A., 1998. A Late Ziemi UŒ, Sosnowiec, pp. 107–111. [In Pol ish.] Trias sic lake system in East Greenland: facies, depositional Bodzioch, A. & Kowal-Linka, M., 2012. Un rav el ing the or i gin of cy cles and palaeoclimate. Palaeo ge ogra phy, Palaeoclimato- the Late Trias sic multitaxic bone accu mu la tion at Krasiejów logy, Palaeo ec ol ogy, 140: 135–159. (S Poland) by diagenetic analy sis. Palaeo ge ogra phy, Palaeo- Cohen, A. S., 2003. Paleolimnology: The History and Evo lu tion of cli ma tol ogy, Palaeo ec ol ogy, 346–347: 25–36. Lake Systems. Ox ford Uni ver sity Press, Oxford, 500 pp. Bodzioch, A., Kowal-Linka, M., NiedŸwiedzki, G. & Owocki, K., Czapowski, G. & Romanek, A., 1986. Lithostratigraphy and sed i - 2015. Taphonomy of dicynodont bones from the Lipie Œl¹s- men tary en vi ron ment of Up per Tri as sic rocks from the kie clay pit at Lisowice (Upper Trias sic, Norian-Rhaetian; Chmielnik area. Przegl¹d Geologiczny, 34: 635–643. [In Pol - Upper Silesia, southern Poland). In: Jagt, J. W. M., Hebda, G., ish, with English summary.] Mitrus, S., Jagt-Yazykova, E. A., Bodzioch, A., Konietzko- Dadlez, R. & Kopik, J., 1963. Prob lem of the Rheatic in West ern Meier, D., Kardyna³, K. & Gruntmejer, K. (eds), 13th An nual Poland in the light of tbe pro file at Ksi¹¿ Wielkopolski. Meet ing of the Eu ro pean As so ci a tion of Ver te brate Palaeon- Kwartalnik Geologiczny, 7: 131–158. [In Pol ish, with Eng lish tologists Opole, Poland, 8–12 July 2015 – Abstracts . Opole sum mary.] Uni ver sity, Opole, p. 106. Deczkowski, Z., 1977. Ge ol ogy of the Perm-Me sozoic cover and Bohacs, K. M., Carroll, A. R., Neal, J. E. & Mankiewicz, P. J., its base ment in the eastern part of the Fore-Sudetic Monocline 2000. Lake-ba sin type, source po ten tial, and hy dro car bon (Kalisz–Czêstochowa are). Prace Instytutu Geologicznego, char ac ter: an in te grated-se quence-strati graphic–geo chem i cal 82: 3–63. [In Pol ish, with Eng lish summary.] framework. In: Gierlowski-Kordesch, E. H. & Kelts, K. R. Deczkowski, Z., Marcinkiewicz, T. & Maliszewska, A., 1997. (eds), Lake Basins through Space and Time. AAPG Studies in Norian and Rhaetian. Sed i menta tion, paleogeography and Ge ol ogy. 46: 3–34. paleotectonics. Prace Pañstwowego Instytutu Geologicznego, Botfalvai, G., Õsi, A. & Mindszenty, A., 2015. Taphonomic and 153: 150–194. [In Pol ish, with Eng lish summary.] paleoecologic in ves ti ga tions of the Late Cre ta ceous (Santo- Demico, R.V. & Gierlowski-Kordesch, E., 1986. Fa cies se quences nian) Iharkút ver te brate as semblage (Bakony Mts, North west - of a semi-arid closed basin: the Lower Juras sic East Berlin ern Hun gary). Palaeo ge ogra phy, Palaeoclimatology, Palaeo - Forma tion of the Hartford Ba sin, New Eng land, U.S.A. Sedi- ec ol ogy, 417: 379–405. mentology, 33: 107–118. Bown, T. M., & Kraus, M. J., 1981. Ver te brate fos sil-bear ing Desojo, J. B., Heckert, A. B., Martz, J. W., Parker, W. G., Schoch, paleosol units (Willwood For mation, Lower Eocene, North - R. R., Small, B. J. & Sulej, T., 2013. Aetosauria: A clade of west Wy o ming, U.S.A.): Im pli ca tions for taphonomy, ar moured pseudosuchians from the Upper Tri assic con ti nen - biostratigraphy, and as sem blage anal y sis. Palaeo ge ogra phy, tal beds. In: Nesbitt, S. J., Desojo, J. B. & Irmis, R. B. (eds), Palaeoclimatology, Palaeo ec ol ogy, 34: 31–56. Anat omy, Phy log eny and Palaeo bi ol ogy of Early Archosaurs Britt, B. B., Eberth, D. A., Scheetz, R. D., Greenhalgh, B. W. & and their Kin. Geo log i cal So ci ety, Lon don, Spe cial Pub li ca - Stadtman, K. L., 2009. Taphonomy of de bris-flow hosted di - tions, 379: 203–239. nosaur bonebeds at Dal ton Wells, Utah (Lower Cre ta ceous, Dockter, J., 1995. Keuper. In: Seidel, G. (ed.), Geologie von Thue- Ce dar Moun tain For ma tion, USA). Palaeo ge ogra phy, ringen. Schweitzerbart’sche Verlag, Stuttgart, p. 332–362. Palaeoclimatology, Palaeo ec ol ogy, 280: 1–22. Dzik, J., 2001. A new Paleorhinus fauna in the early Late Trias sic Brusatte, S. L., Butler, R .J., Mateus, O. & Steyer, J. S., 2015. A of Poland . Jour nal of Ver te brate Pa le on tol ogy, 21: 625–627. new species of Metoposaurus from the Late Trias sic of Portu - Dzik, J., 2003a. A beaked her biv o rous archosaur with di no saur af - gal and com ments on the system at ics and biogeography of fini ties from the early Late Trias sic of Poland. Jour nal of Ver - metaposaurid temnospondyls. Jour nal of Ver te brate Pa le on - te brate Pa le on tol ogy, 23: 556–574. tol ogy, 35: DOI: 10.1080/02724634.2014.912988. Dzik, J., 2003b. Krasiejów – u zarania ery dinozaurów.Wystawa w Brusatte, S. L., Butler, R .J., Sulej, T. & NiedŸwiedzki, G., 2009. Muzeum Ewolucji w Warszawie. Ewolucja, Biuletyn Muzeum The tax on omy and anat omy of rauisuchian archosaurs from Ewolucji Instytutu Paleobiologii PAN, 1: 2–13. [In Pol ish.] the Late Trias sic of Germany and Poland. Acta Palaeontolo- Dzik, J., 2008. Gill struc ture and re la tion ships of the Tri as sic gica Polonica, 54: 221–230. cycloid crus ta ceans. Jour nal of Mor phol ogy, 269: 1501–1519. Brusatte, S. L., Nesbitt, S. J., Irmis, R. B., Butler, R. J., Benton, M. Dzik, J., Niedzwiedzki, G. & Sulej, T., 2008a. Zaskakuj¹ce uwieñ- J. & Norell, M. A., 2010. The or i gin and early ra di a tion of di - czenie ery gadów ssakokszta³tnych. Ewolucja, Biuletyn Muze- no saurs. Earth Science Reviews , 101: 68–100. um Ewolucji Instytutu Paleobiologii PAN, 3: 1–21. [In Pol ish] Budziszewska-Karwowska, E., Bujok, A. & Sadlok, G., 2010. Dzik, J. & Sulej, T., 2007. A review of the early Late Tri assic Bite marks on an Up per Tri as sic dicynodontid tibia from Krasiejów biota from Silesia, Poland. Palaeontologia Polo- Zawiercie, Kraków-Czêstochowa Up land, south ern Po land. nica, 64: 3–27. Palaios, 25: 415–421. Dzik, J., Sulej, T., Kaim, A. & NiedŸwiedzki, R., 2000. A late Tri - But ler, R. J., Rauhut, O. W. M., Stocker, M. R. & Bronowicz, R., as sic tetrapod grave yard in the Opole Silesia (SW Poland). 2014. Redescription of the phytosaurs Paleorhinus (‘Franco- Przegl¹d Geologiczny, 48: 226–235. [In Pol ish, with Eng lish suchus’) angustifrons and Ebrachosuchus neukami from Ger- sum mary.] many, with impli ca tions for Late Trias sic biochronology. Dzik, J., Sulej, T. & Niedzwiedzki, G., 2008b. A dicynodont- Zoo log i cal Jour nal of the Lin nean So ci ety, 170: 155–208. theropod as so ci a tion in the lat est Tri as sic of Po land. Acta Bzowska, G. & Racka, M., 2006. Geo chem i cal and min er al og i cal Palaeontologica Polonica, 53: 733–738. look at the Keuper of Krasiejów. Gospodarka Surowcami Feist-Burkhardt, S., Götz, A. E., Szulc, J., Aigner, T., Borkha- Mineralnymi, 22, (zeszyt specjalny, 3): 355–358. [In Pol ish, taria, R., Geluk, M., Haas, J., Hornung, J., Jordan, P., Kempf, with English summary.] O., Michalík, J., Nawrocki, J., Reinhardt, L., Ricken, W., Cirilli, S., 2010. Up per Tri as sic - low er most Juras sic palynology Röhling, H. G., Rüffer, T., Török, A. & Zühlke, R., 2008. Tri - and palynostratigraphy: A re view. In: Lucas, S. G. (ed.), The assic. In: McCann, T. (ed.), The Ge ol ogy of Cen tral Eu rope. Trias sic Timescale. Geo log i cal So ci ety of Lon don Spe cial The Geo log i cal So ci ety, Lon don, Lon don, pp. 749–821. UP PER TRI AS SIC BONE-BEAR ING LEV ELS 623

Fija³kowska-Mader, A., 2015a. Palinostratygraficzne aspekty nych wykonanych w rejonie Krasiejowa, Lisowic i Zawiercia- sukcesji kajpru Górnego Œl¹ska. In: Racki, G. (ed.), V Sesja Marciszowa. Un pub lished Grant Re port, Fac ulty of Earth Sci - Popularnonaukowa “Œl¹skie Dinozaury (nie tylko) z Krasie- ences, Uni ver sity of Silesia, Sosnowiec, 27 pp. [In Pol ish.] jowa - Kraków 2015, p. 8. [In Pol ish.] Irmis, R. B., 2011. Eval u at ing hy poth e ses for the early di ver si fi ca - Fija³kowska-Mader, A., 2015b. Re cord of cli matic changes in the tion of di nosaurs. Earth and En vi ron men tal Sci ence Trans ac- Tri as sic palynological spec tra from Poland. Geo log i cal Quar- tions of the Royal So ci ety of Ed in burgh, 101: 397–426. terly, 59 (in press). DOI: 10.7306/gq.1239. Jewu³a, K., 2010. Skala magnetostratygraficzna dla utworów Fija³kowska-Mader, A., Heunisch, C. & Szulc, J., 2015. Palyno- górnego triasu z Górnego Œl¹ska. Un published MSc The sis, stra tig ra phy and palynofacies of the Upper Silesian Keuper In sti tute of Geo log i cal Sci ences, Jagiellonian Uni ver sity, (south ern Po land). Annales Societatis Geologorum Poloniae, Kraków, 107 pp. [In Pol ish.] 85: 637– 661. Kelber, K. P., 2005. Makroflora (Die Keuperfloren). In: Beutler, Flaig, P. P., Fiorillo, A. R. & Mc carthy, P. J., 2014. Dino saur- G., Hauschke, N., Nitsch, E. & Vath, U. (eds), Deut sche bear ing hyperconcentrated flows of Cre ta ceous Arc tic Stratigraphische Kommission, Stratigraphie von Deutsch land Alaska: Re cur ring cat a strophic event beds on a dis tal paleo- IV – Keuper. Courier Forschungsinstitut. Senckenberg, 253: polar coastal plain. Palaios, 29: 594–611. 32–38. Franz, M., 2008. Litho- und Leitflächenstratigraphie, Chrono- Kelber, K. P. & Nitsch, E., 2005. Paläoflora und Ablagerung- stratigraphie, Zyklo- und Sequenzstratigraphie des Keupers im sräume im unterfränkischen Keuper (Exkursion H am 1. April Östlichen Zentraleuropäischen Becken (Deutschland, Polen) 2005). Jahresberichte und Mitteilungen des Oberrheinischen und Dänischen Becken (Dänemark, Schweden). Un pub lished Geologischen Vereins, 87: 217–253. thesis (Dr. rer. nat.), Naturwissenschaftlichen Fakultät III der K³apciñski, J., 1993. Litostratygrafia profili g³êbokich otworów Martin-Lu ther-Universität Halle-Wittenberg, 198 pp. wiertniczych w regionie opolskim. Acta Universitatis Wrati- Füchtbauer, H. & Müller; G., 1977. Sed i ment-Petrologie, Teil II: slaviensis, 8: 1–129 [In Pol ish.] Sedimente und Sedimentgesteine. Schweitzerbart’sche Ver- Koeppen, A., 1997. Fazieentwicklung in den fruehen Obertrias lag, Stuttgart, 784 pp. Mitteeuropas – ein sequenzstratigraphischer Vergleich. Gea Fuglewicz, R. & Œnie¿ek, P., 1980. Upper Tri as sic megaspores Heidelbergensis, 2: 1–233 from Lipie Œl¹skie near Lubliniec. Przegl¹d Geologiczny, 28: Konieczna, N., Belka, Z. & Dopieralska, J., 2015. Nd and Sr iso to - 459–461. [In Pol ish.] pic evi dence for prove nance of clastic mate rial of the Upper G¹siorowski, S. M., Michalik, M., Paszkowski, M., Piekarska, E. Trias sic rocks of Silesia, Poland. Annales Societatis Geolo- & Szulc, J., 1986. Shal low-wa ter and subearial car bon ates of gorum Poloniae, 85: 675–684. the Czêstochowa-Cra cow Upland. Case histo ries from the Konietzko-Meier, D., Bodzioch, A. & Sander, P. M., 2013. Lower Car bon if er ous to Ho lo cene. Ex cur sion B-10. In: Histological char ac ter is tics of the ver te bral intercentra of Teisseyre, A. K. (ed.), IAS 7th Eu ropean Re gional Meet ing Metoposaurus diagnosticus (Temnospondyli) from the Up per (Kraków, May 23–25, 1986), Ex cur sion Guide book. Ossili- Tri as sic of Krasiejów (Upper Silesia, Poland). Earth and En - neum, Wroc³aw, pp. 179–201. vi ron men tal Sci ence Trans ac tions of the Royal So ci ety of Ed - Gierlowski-Kordesch, E. H. & Kelts, K. (eds), 1994. Global Geo - in burgh, 103: 1–14. log i cal Re cord of Lake Bas ins, Volume 1. Cam bridge Uni ver - Konietzko-Meier, D. & Klein, N., 2013. Growth pat tern of Meto- sity Press, Cambridge, 427 pp. posaurus diagnosticus krasiejowensis (Amphibia, Temno- Gorzelak, P., NiedŸwiedzki, G. & Skawina, A., 2010. Pa thol o gies spondyli) from the Late Trias sic of Krasiejów, Poland, as de - of non-marine bi valve shells from the Late Trias sic of Poland. duced from fe mur histol ogy. Palaeo ge ogra phy, Palaeoclima- Lethaia, 43: 285–289. tology, Palaeo ec ol ogy, 370: 145–157. Grodzicka-Szymanko, W. & Or³owska-Zwoliñska, T., 1972. Stra - Kotlicki, S., 1966. De tailed Geo log i cal Map of Po land, Zawiercie tig ra phy of the Up per Tri as sic in the NE mar gin of the Up per Sheet, 1: 50 000. Wydawnictwa Geologiczne, Warszawa. [In Silesian Coal Basin. Kwartalnik Geologiczny, 16: 216–232. Pol ish.] [In Pol ish, with Eng lish summary.] Kotlicki, S., 1974. Strati graphic po sition of the Tri as sic sed i ments Gruntmejer, K., Konietzko-Meierm, D. & Bodzioch, A., 2015. in the Up per Silesian region. Bul le tin de l’Académie Polo - The Trias sic world of Krasiejów. In: Jagt, J. W. M., Hebda, naise des Sciences , Sciences de la Terre, 22: 161–166. G., Mitrus, S., Jagt-Yazykova, E. A., Bodzioch, A., Ko- Kotlicki, S., 1995. Badania nad litostratygrafi¹ triasu Górnego nietzko-Meier, D., Kardyna³, K. & Gruntmejer, K. (eds), 13th Œl¹ska. Un pub lished re port. Na tional Geo log i cal Ar chive, An nual Meet ing of the Eu ro pean As so ci a tion of Ver te brate Pol ish Geo log i cal In sti tute, Up per Silesian Branch, Sosno- Palaeontologists Opole, Po land, 8–12 July: Field Guide. wiec, 157 pp. [In Pol ish.] Opole Uni ver sity, Opole, pp. 17–27. Kozur, H. W. & Weems, R. F., 2010. The biostratigraphic impor - Gruszka, B. & Zieliñski, T., 2008. Ev i dence for a very low-en ergy tance of conchostracans in the con ti nen tal Trias sic of the flu vial system: A case study from the di nosaur-bear ing Upper north ern hemi sphere. In: Lucas, S. G. (ed.), The Tri as sic Tri as sic rocks of South ern Po land. Geo log i cal Quar terly, 52: Timescale. Geo log i cal So ci ety of Lon don Spe cial Pub li ca - 239–252. tion, 334: 315–417. Haisig, J., Kotlicki, S., Wilanowski, S. & ¯urek, W., 1983. Krupnik, J., Ziaja, J., Barbacka, M., Feldman-Olszewska, A. & Objaœnienia do Szczegó³owej Mapy Geologicznej Polski, Jarzynka, A., 2014. A palaeoenvironmental re con struc tion Arkusz Lubliniec. Wydawnictwa Geologiczne, Warszawa, 46 based on palynological anal y ses of Upper Tri as sic and Lower pp. [In Pol ish.] Juras sic sed i ments from the Holy Cross Mountains re gion. Heunisch, C. & Nitsch, E., 2011. Eine seltene Mikroflora aus der Acta Palaeobotanica, 54: 35–65. Mainhardt-For mation (Keuper, Trias) von Baden-Württem- Kubik, R., Uhl, D. & Marynowski, L., 2015. Ev i dence of wildfires berg (Süddeutschland). Jahresberichte und Mitteilungen des dur ing de po si tion of the Up per Silesian Keuper suc ces sion, Oberrheinischen Geologischen Vereins, Neue Folge, 93: south ern Po land. Annales Societatis Geologorum Poloniae, 55–76. 85: 685–696. Idziak, A., 2013. Opis wyników i interpetacja pomiarów geofizycz- Kürschner, W. M. & Herngreen, G. F. W., 2010. Tri as sic paly- 624 J. SZULC ET AL.

nology of cen tral and north west ern Eu rope: A re view of of Ger many Compact 2012.] Zeitschrift der Deutschen palynofloral di ver sity pat terns and biostratigraphic sub di vi - Gesellschaft für Geowissenschaften, 163: 385–409. sions. In: Lucas, S. G. (ed.), The Tri as sic Timescale. Geo log i - Milner, A. R. & Schoch, R. R., 2004. The lat est metaposaurid am - cal So ci ety of Lon don Spe cial Pub li ca tion, 334: 263–283. phib i ans from Eu rope. Neues Jahrbuch für Geologie und Lucas, S. G., 1998. Global Tri as sic tetrapod biostratigraphy and Paläontologie Abhandlungen, 232: 231–252. biochronology. Palaeo ge ogra phy, Palaeoclimatology, Pala- Mukherjee, D. & Ray, S., 2012. Taphonomy of an Up per Tri as sic eoecology, 143: 347–384. ver te brate bonebed: A new rhynchosaur (Reptilia; Archosau- Lucas, S. G., 2010. The Trias sic timescale based on nonmarine romorpha) ac cu mu la tion from In dia. Palaeo ge ogra phy, Pala- tetrapod biostratigraphy and biochronology. In: Lucas, S. G. eoclimatology, Palaeo ec ol ogy, 333–334: 75–91. (ed.), The Tri as sic Timescale. Geo log i cal So ci ety of Lon don Nawrocki, J., Jewu³a, K., Stachowska, A. & Szulc, J., 2015. Mag- Spe cial Pub li ca tion, 334: 447–500. netic polar ity of Upper Trias sic sedi ments of the Germanic Lucas, S. G., 2013. Plant megafossil biostratigraphy and biochro - Ba sin in Poland. Annales Societatis Geologorum Poloniae, nol ogy, Up per Tri assic Chinle Group, West ern USA. In: Tan - 85: 663–674. ner, L. H., Spielmann, J. A. & Lucas, S. G. (eds), The Tri as sic Nesbitt, S. J., Sidor, C. A., Irmis, R. B., Angielczyk, K. D., Smith, Sys tem. New Mex ico Mu seum of Nat ural History and Sci ence, R. M. H. & Tsuji, L. A., 2010. Ecolog i cally distinct dinosaur - Bul le tin, 61: 354–364. ian sis ter group shows early di ver sifi ca tion of Ornithodira. Lucas, S. G., 2015. Age and cor re la tion of Late Tri as sic tetrapods Na ture, 464: 95–98. from southern Po land. Annales Societatis Geologorum Polo- NiedŸwiedzki, G., Brusatte, S. L., Sulej, T. & Butler, R. J., 2014. niae, 85: 627–635. Basal dinosauriform and theropod di nosaurs from the mid– Lucas, S. G., Rinehart, L. F., Krainer, K., Spielmann, J. A. & late Norian (Late Trias sic) of Poland: Im plica tions for Trias - Heckert, A. B., 2010. Taphonomy of the Lamy amphib ian sic di no saur evo lu tion and dis tri bu tion. Palae on tol ogy, 57: quarry: A Late Trias sic bonebed in New Mex ico, U.S.A. 1121–1142. Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol ogy, 298: NiedŸwiedzki, G. & Budziszewska-Karwowska, E., 2015. Smok 388–398. z Zawiercia-Marciszowa – wstêpne dane. In: Racki, G. (ed.), Lucas, S. G., Spielmann, J. A. & Hunt, A. P., 2007. Biochro- V Sesja Popularnonaukowa “Œl¹skie Dinozaury (nie tylko) nological signif i cance of Late Trias sic tetrapods from Kra- z Krasiejowa - Kraków 2015, p. 12 [In Pol ish.] siejów. In: Lucas, S. G. & Spielmann, J. A. (eds), The Global NiedŸwiedzki, G., Gorzelak, P. & Sulej, T., 2011. Bite traces on Tri as sic. Mu seum of Nat u ral His tory and Sci ence Bul le tin, dicynodont bones and the early evo lu tion of large ter res trial 41: 248–258. pred a tors. Lethaia, 44: 87–92. Lucas, S.G., Tanner, L.H., Kozur, H.W., Weems, R.E. & Heckert, NiedŸwiedzki, G. & Sulej, T., 2008. Lipie Œl¹skie off Lisowice – a A.B., 2012. The Late Trias sic timescale: Age and corre la tion window on the Late Trias sic con ti nen tal eco sys tem. Przegl¹d of the Carnian-Norian bound ary. Earth-Sci ence Re views, Geologiczny, 56: 821–822. [In Pol ish.] 114: 1–18, NiedŸwiedzki, G., Sulej, T. & Dzik, J., 2012. A large pred a tory Majer, D. & Lubka, M., 2003. An ac cu mula tion of bones of large archosaur from the Late Trias sic of Poland. Acta Palaeonto- Late Tri assic reptiles at Krasiejów, Opolean Silesia, Po land. logica Polonica, 57: 267–276. Opole Sci en tific So ci ety, Na ture Jour nal, 36: 69–76. NiedŸwiedzki, R., 2012. From iron to Silesaurus. The dis cov ery of Makaske, B., 2001. Anastomosing rivers: A re view of their clas si- a Tri as sic tetrapod grave yard and the history of mining at fi ca tion, or i gin and sed i men tary prod ucts. Earth-Sci ence Re - Krasiejów. In: Jagt-Yazykova, E., Jagt, J. W. M., Bodzioch, views, 53: 149–196. A. & Konietzko-Meier, D. (eds), Krasiejów – paleontological Malinowska, A., 2015. Zawiercie. Naukowcy nie mog¹ dobraæ siê in spi ra tions. Stowarzyszenie „Dinopark” & Zak³ad Paleobio- do cennych koœci. Gazeta Wyborcza, Katowice. http:// logii Katedry Biosystematyki Uniwersytetu Opolskiego, katowice.wyborcza.pl/katowice/1,35063,18980392,zawierci Opole, pp. 6–27. [In Pol ish, with Eng lish summary.] e-naukowcy-nie-moga-dobrac-sie-do-cennych-kosci.html [In NiedŸwiedzki, R., 2015. The sci en tific, geotouristic and eco nomic Pol ish; 15.10.2015]. signif i cance of the dis cov ery of the Tri as sic tetrapod La ger- Marcinkiewicz, T. & Or³owska-Zwoliñska, T., 1994. Miospores, stätte at Krasiejów (Opole Silesia, Po land). In: Jagt, J. W. M., megaspores and Lepidopteris ottonis (Goeppert) Schimper in Hebda, G., Mitrus, S., Jagt-Yazykova, E. A., Bodzioch, A., the up per most Tri as sic de pos its from Po land. Geo log i cal Konietzko-Meier, D., Kardyna³, K. & Gruntmejer, K. (eds), Quar terly, 38: 91–116. 13th An nual Meet ing of the Eu ro pean As so ci a tion of Ver te - Marcinkiewicz, T., Fija³kowska-Mader, A. & Pieñkowski, G., brate Palaeontologists Opole, Po land, 8–12 July 2015 – Ab- 2014. Megaspore zones of the epicontinental Tri as sic and Ju - stracts. Opole Uni ver sity, Opole, p. 107. ras sic de pos its in Po land – over view. Biuletyn Pañstwowego. Nitsch, E., 2011. Keuper im Baden-Württemberg. http://portale. Instytutu Geologicznego, 457: 15–42. [In Pol ish, with Eng lish wisutec.de/Portals/10/Stratigraphie/bw/tri/k_Nit_058.pdf. sum mary.] [11.02.2015]. Marynowski, L. & Simoneit, B. R. T., 2009. Wide spread Late Tri - Nitsch, E., Seegis, D., Vath, U. & Hauschke, N., 2005. Sedimente assic to Early Juras sic wildfire records from Poland: Evi dence und Sedimentationspausen im deutschen Keuper: Wie voll- from charcoal and pyrolytic polycyclic aro matic hy drocar - ständig ist die Überlieferung der späten Triaszeit? News let - bons. Palaios, 24: 785–798. ters on Stratig ra phy , 41: 225–251. Maron, M., Rigo, M., Bertinelli, A., Katz, M. E., Godfrey, L., Ogg, J. G., 2012. Tri as sic time scale. In: Gradstein, F. M., Ogg, J. Zaffani, M., & Muttoni, G., 2015. Magnetostratigraphy, bio- G., Schmitz, M. D. & Ogg, G. M. (eds), The Geo logic Time stra tig ra phy, and chemostratigraphy of the Pignola-Abriola Scale 2012. Vol. 2. Amster dam, Elsevier, pp. 700–730. sec tion: New con straints for the Norian-Rhaetian bound ary. Olempska, E., 2004. Late Trias sic spinicaudatan crusta ceans from Geo log i cal So ci ety of Amer ica Bul le tin, 127: 962–974. SW Poland. Acta Palaeontologica Polonica, 49: 429–442. Menning, M. & Deutsche Stratigraphische Kommission, 2012. Onoue, T., Zonneveld, J. P., Orchard, M. J., Yamashita, M., Ya- Erläuterung zur Stratigraphischen Tabelle von Deutsch land mashita, K., Sato, H., Kusaka, S., 2016. Palaeoenvironmental Kompakt 2012. [Ex plan a tory notes to the Strati graphic Table changes across the Carnian/Norian bound ary in the Black UP PER TRI AS SIC BONE-BEAR ING LEV ELS 625

Bear Ridge section, British Colum bia, Can ada, Palaeo ge - Gesellschaft, 19: 255–269 ogra phy, Palaeoclimatology, Palaeo ec ol ogy, 441: 721–733. Römer, F., 1870. Geologie von Oberschlesien. Breslau, Nisch- Or³owska-Zwoliñska, T., 1983. Palynostratigraphy of the Up per kowsky, 587 pp. part of Trias sic epicontinental sedi ments in Poland. Prace Ryan, M. J., Russell, A. P., Eberth, D. A. & Currie, P. J., 2001. Instytutu Geologicznego, 104: 1–89. [In Pol ish, with Eng lish The taphonomy of a Centrosaurus (Ornithischia: Ceratop- sum mary.] sidae) bone beds from the Di no saur Park Forma tion (Up per Or³owska-Zwoliñska, T., 1985. Palynological zones of the Pol ish Campanian), Al berta, Canada, with comments on cra nial on - epicontinental Tri as sic. Bul le tin Acad emy of Sci ences, Earth tog eny. Palaios, 16: 482–506. Sci ences, 33: 107–119. Sadlok, G. & Wawrzyniak, Z., 2013. Up per Tri as sic ver te brate Pacyna, G., 2014. Plant re mains from the Polish Tri as sic. Pres ent tracks from Kraków-Czêstochowa Up land, South ern Po land. knowledge and fu ture pros pects. Acta Palaeobotanica, 51: Annales Societatis Geologorum Poloniae, 83: 105–111. 3–33. Sarkar, S., 1988. Pe trol ogy of caliche-de rived peloidal cacirudite/ Philippe, M., Pacyna, G., Wawrzyniak, Z., Barbacka, M., Boka, calcarenite in the Late Trias sic Maleri Forma tion of the K., Filipiak, P. Marynowski, L., Thévenard, F. & Uhl, D., Pranhita–Godavari val ley, south In dia. Sed i men tary Ge ol ogy, 2015. News from an old wood – Agathoxylon keuperianum 55: 263–282. (Unger) nov. comb. in the Keuper of Po land and France. Re - Schoch, R. & Wild, R., 1999. Die Wirbeltier-Fauna im Keuper von view of Palaeobotany and Palynology, 221: 83–91. Süddeutschland. In: Hauschke, N. & Wilde, V. (eds), Trias. Piechowski, R. & Dzik, J., 2010. The ax ial skel e ton of Silesaurus Eine Ganz andere Welt. Mitteleuropa im Frühen Erdmittel- opolensis. Jour nal of Ver te brate Pa le on tol ogy, 30: 1127– al ter. Dr Pfeil Verlag, München, pp. 395–408. 1141. Seegis, D., 2005. Tetrapoden. In: Beutler, G., Hauschke, N., Piechowski, R., Ta³anda, M. & Dzik, J., 2014. Skel e tal vari a tion Nitsch, E. & Vath, U. (eds), Stratigraphie von Deutschland and on tog eny of the Late Tri as sic dinosauriform Silesaurus IV, Keuper. Courier Forschungsinstitut. Senckenberg, 253: opolensis. Jour nal of Ver te brate Pa le on tol ogy, 34, 1383– 50–54. 1393. Seilacher, A., 1970. Begriff und Bedeutung der Fossil-La gerstä- Pieñkowski, G., 1988. Fa cial anal ysis of the up per most Trias sic tten. Neues Jahrbuch für Geologie und Paläontologie Mo- and the Liassic of the Cracow-Wieluñ Upland and pros pects natshefte, 1970: 34–39. for occur rence of clay depos its. Przegl¹d Geologiczny, 36: Seilacher, A., 2008. Biomats, biofilms, and bioglue as preser- 449–456. [In Pol ish, with Eng lish summary.] vational agents for ar thro pod trackways. Palaeo ge ogra phy, Pieñkowski, G., NiedŸwiedzki, G. & Brañski P., 2014. Cli matic Palaeoclimatology, Palaeo ec ol ogy, 270: 252–257. re ver sals re lated to the Cen tral At lan tic mag matic prov ince Seilacher, A., Reif, F. & Westphal, R., 1985. Sedimentological, caused the end-Tri as sic bi otic crisis – Evi dence from con ti - eco log i cal and tem po ral pat terns of Fos sil Lagerstatten. nental strata in Poland. In: Keller,G. & Kerr, A. (eds), Vol ca- Philo soph i cal Trans ac tions of the Royal So ci ety of Lon don. nism, Impacts, and Mass Ex tinc tions: Causes and Ef fects. Se ries B, Bi o log i cal Sci ences, 311: 5–24. Geo log i cal So ci ety of Amer ica Spe cial Pa pers, 505: 263–286. Siewniak-Madej, A., (ed.), 1982. Lubliniec IG 1. Pro file G³êbo- Preto, N., Kustatscher, E., & Wignall, P. B., 2010. Trias sic cli- kich Otworów Wiertniczych Instytutu Geologicznego, 55. mates – State of the art and perspec tives. Palaeo ge ogra phy, Wyd. Geol., Warszawa: 67 pp. [In Pol ish.] Palaeoclimatology, Palaeo ec ol ogy, 290: 1–10. Skawina, A., 2013. Popu la tion dynam ics and taphonomy of the Racki, G., 2010. Ewolucja œrodowisk l¹dowych kajpru Górnego Late Trias sic (Carnian) freshwa ter bivalves from Krasiejów Œl¹ska jako biotopów krêgowców – nowy projekt badawczy. (Po land). Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol - Przegl¹d Geologiczny, 58: 124–126. [In Pol ish.] ogy, 379–380: 68–80. Reinhardt, L. & Ricken, W., 2000. The stratigraphic and geochem - Skawina, A. & Dzik, J., 2011. Umbonal mus cu la ture and re la tion - ical record of playa cycles: Moni tor ing a Pangaean mon- ships of the Late Tri as sic filibranch unionoid bi valves. Zoo- soon-like sys tem (Trias sic, Middle Keuper, S. Germany). log i cal Jour nal of the Lin nean So ci ety, 163: 863–883. Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol ogy, 161: Skrzycki, P., 2015. New spe cies of lungfish (Sarcopterygii, 205–227. Dipnoi) from the Late Trias sic Krasiejów site in Poland, with Renaut, R. W. & Gierlowski-Kordesch, E. H., 2010. Lakes. In: re marks on the on tog eny of Tri as sic dip no an tooth plates. Dalrym ple, R. & James, N. (eds), Fa cies Mod els, 4th Edition . Jour nal of Ver te brate Pa le on tol ogy, 35: DOI: 10.1080/ Geo log i cal As so ci a tion of Can ada, To ronto, pp. 541–575. 02724634.2015.964357 Rogers, R. R., 2005. Fine-grained de bris flow and ex traor di nary Œrodoñ, J., Szulc, J., Anczkiewicz, A., Jewu³a, K., Banaœ, M. & ver te brate buri als in the Late Cre ta ceous of Mad a gas car. Ge - Marynowski, L., 2014. Weath er ing, sed i mentary, and diage- ol ogy, 33: 297–300. netic con trols of min eral and geo chem i cal char ac ter is tics of Rogers, R. R. & Kidwell, S. M., 2000. Asso ci a tions of ver te brate the ver te brate-bear ing Silesian Keuper. Clay Min er als, 49: skel e tal con cen tra tions and dis con ti nu ity sur faces in ter res - 569–594. trial and shallow-ma rine records: A test in the Creta ceous of Staneczko, K., 2007. Nowe dane paleobotaniczne na temat gór- Montana. Jour nal of Ge ol ogy, 108: 131–154. nego triasu z Lipia Œl¹skiego ko³o Lubliñca (po³udniowa Rogers, R. R. & Kidwell, S. M., 2007. A con cep tual frame work Polska). In: Geo-Sympozjum M³odych Badaczy Silesia 2007. for the gen e sis and anal y sis of ver te brate skel e tal con cen tra- Uniwersytet Œl¹ski, Sosnowiec, pp. 157–170. [In Pol ish.] tions. In tro duc tion. In: Rog ers, R. R., Eberth, D. A. & Stratigraphische Tabelle von Deutsch land, 2002. Deut sche Strati- Fiorillo, A. R. (eds), Bonebeds: Gen e sis, Anal y sis and graphische Kommission, Potsdam (GeoForschungszen- Paleobiological Sig nif i cance. Uni ver sity of Chi cago Press, trum), Frank furt a. M. (Forschungsinstitut Senckenberg); Chi cago, pp. 1–63. http://www.stratigraphie.de/std2002/down load/STD2002_ Römer, F., 1867. Neuere Beobachtungen über die Gliederung des large.pdf [10. 03. 2015]. Keupers und der ihn zunächst überlagernden Abtheilung der Sues, H. D. & Fraser, N. C., 2010. Trias sic Life on Land: The Juraformation in Oberschlesien und in den angrenzenden Great Tran si tion. Co lumbia Uni ver sity, New York, 236 pp. Theilen von Polen. Zeitschrift der Deutschen Geologischen Sulej, T., 2002. Species discrim i nation of the Late Trias sic tem- 626 J. SZULC ET AL.

nospondyl am phib ian Metoposaurus diagnosticus. Acta Pa- Fieldtrip Guide. Pol ish Geo log i cal So ci ety, Pol ish Geo log i cal laeontologica Polonica, 47: 535–546. In sti tute, In sti tute of Geo log i cal Sci ences, Jagiellonian Uni- Sulej, T., 2005. A new rauisuchian rep tile (Diapsida: Archosauria) ver sity, Cra cow, p. 61. from the Late Trias sic of Poland. Jour nal of Ver te brate Pa le - Szulc, J., 2007c. Tatra and Up per-Silesian Trias sic. State of exam - on tol ogy, 25: 75–83. i na tions. Przegl¹d Geologiczny, 55: 947–950. [In Pol ish.] Sulej, T., 2007. Osteology, vari abil ity, and evo lu tion of Metopo- Szulc, J., Gradziñski, M., Lewandowska, A. & Heunisch, C., saurus, a temnospondyl from the Late Tri as sic of Po land. 2006. The Up per Tri as sic crenogenic lime stones in Up per Palaeontologia Polonica, 64: 29–139. Silesia (south ern Po land) and their paleoenvironmental con - Sulej, T., 2010. The skull of an early Late Tri as sic aetosaur and the text. In: Alonso Zarza, A. M. & Tanner, L. H. (eds), Paleo- evo lu tion of the stagonolepidid archosaurian reptiles. Zoo log- en vi ron men tal Re cord and Ap pli ca tions of Calcretes and i cal Jour nal of the Lin nean So ci ety, 158: 860–881. Palustrine Car bon ates. Geo log i cal So ci ety of Amer ica Spe - Sulej, T., Bronowicz, R., Ta³anda, M. & NiedŸwiedzki, G., 2011a. cial Paper , 416: 133–151. A new dicynodont-archosaur assem blage from the Late Tri as - Szulc, J. & Racki G., 2015. The Grabowa For mation – the ba sic sic (Carnian) of Po land. Earth & En vi ron men tal Sci ence: lithostratigraphic unit of the Keuper of Silesia. Przegl¹d Transac tions of the Royal So ci ety of Ed in burgh, 101: 261– Geologiczny, 63: 103–113. [In Pol ish, with Eng lish ab stract.] 269. Szulc, J., Racki, G. & Jewu³a, K., 2015. Key as pects of the stra tig - Sulej, T. & NiedŸwiedzki, G., 2010. Late Trias sic verte brates from ra phy of the Up per Silesian mid dle Keuper, southern Po land. Poland: short over view. In: Golubiev, W. K, & Sennikov, A. Annales Societatis Geologorum Poloniae, 85: 557–586. G. (eds), Perm ian and Tri as sic Pa le on tol ogy and Stra tig ra phy Tanner, L. H. & Lucas, S. G., 2015. The Trias sic-Ju ras sic strata of of North Eur asia. Ma te ri als of the 5th In ter na tional Con fer - the New ark Ba sin, USA: Ac cu rate as tro nom i cally-tuned ence De voted to 150 Anni ver sary of Vladi mir Prokhorovich timescale? Stra tig ra phy, 12: 47–65. Amalitzky (1860-1917). Moscow, 22–23 No vem ber 2010, Therrien, F. & Fastovsky, D. E., 2000. Palaeoenvironments of Borissyak Paleontological In sti tute of RAS, Mos cow, pp. early Theropods, Chinle Fm. (Late Trias sic), Petri fied Forest 112–114. Na tional Park, Ar i zona. Palaios, 15: 194–211. Sulej, T., NiedŸwiedzki, G. & Bronowicz, R., 2012. A new Late Urbañski, K., 2012. ¯ó³w z Porêby i „Smok Wawelski”. Uwa¿am Trias sic verte brate fauna from Poland with turtles, aetosaurs, Rze 39 (86): 92–93. [In Pol ish.] and coelophysoid di nosaurs. Jour nal of Ver te brate Pa le on tol - Vollmer, T., Ricken, W., Weber, M., Tougiannidis, N., Röhling. ogy, 32: 1033–1041. H. G. & Hambach, U., 2008. Or bital con trol on Up per Tri as - Sulej, T., NiedŸwiedzki, G., NiedŸwiedzki, R., Surmik, D. & sic playa cy cles of the Steinmergel-Keuper (Norian): A new Stachacz, M., 2011b. New verte brate assem blage from mar- concept for ancient playa cycles. Palaeo ge ogra phy, Palaeo- ginal-ma rine and land Lower Keuper strata (Ladinian, Mid dle cli ma tol ogy, Palaeo ec ol ogy, 267: 1–16. Trias sic) of Miedary, Silesia, SW Poland. Przegl¹d Geolo- Wood, J. M., Thomas, R. G. & Visser, J., 1988. Flu vial pro cesses giczny, 59: 426–430. [In Pol ish, with Eng lish summary.] and ver te brate taphonomy: the Up per Cre ta ceous Ju dith River Œwi³o, M., NiedŸwiedzki, G. & Sulej, T., 2014. Mam mal-like For ma tion, south-cen tral Di no saur Pro vin cial Park, Al berta, tooth from the Up per Tri as sic of Poland. Acta Palaeonto- Can ada. Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol - logica Polonica, 59: 815–820. ogy, 66: 127–143. Szulc, J., 2000. Mid dle Tri as sic evo lu tion of the north ern peri- Wotzlaw, J. F., Guex, J., Bartolini, A., Gallet, Y., Krystyn, L., Tethys area as in flu enced by early open ing of the Tethys McRoberts, C. A., Taylor, D., Schoene, B. & Schaltegger, U., ocean. Annales Societatis Geologorum Poloniae, 70: 1–48. 2014. To wards ac cu rate nu mer i cal cal i bra tion of the Late Tri- Szulc, J., 2005. Sed i men tary en vi ron ments of the ver te brate-bear - as sic: High-pre ci sion U-Pb geo chron ol ogy con straints on the ing Norian de posits from Krasiejów, Upper Silesia (Po land). du ra tion of the Rhaetian. Ge ol ogy, 42: 571–574. Hallesches Jahrbuch der Geowissenschaften, B19: 161–170. Zatoñ, M., NiedŸwiedzki, G., Marynowski, L., Benzerara, K., Szulc, J., 2007a. Keuper. In: Szulc, J. & Becker, A. (eds), In ter na - Pott, C., Cosmidis, J., Krzykawski, T. & Filipiak, P., 2015. tional Work shop on the Trias sic of south ern Po land, Sep tem - Cop ro lites of Late Tri as sic car niv o rous ver te brates from Po- ber 3–8, 2007. Fieldtrip Guide. Pol ish Geo log i cal So ci ety, land: An in te gra tive ap proach. Palaeo ge ogra phy, Palaeocli- Pol ish Geo log i cal In sti tute, In sti tute of Geo log i cal Sci ences, matology, Palaeo ec ol ogy, 430: 21–46. Jagiellonian Uni ver sity, Cra cow, pp. 33–41. Zatoñ, M., Piechota, A. & Sienkiewicz, E., 2005. Late Trias sic Szulc, J., 2007b. Stop. III. 1. Laryszów – commu nal waste de pot. charophytes around the bone-bear ing bed at Krasiejów (SW In: Szulc, J. & Becker, A. (eds), In ter na tional Work shop on Po land) – palaeo eco logi cal and en vi ron men tal re marks. Acta the Tri as sic of south ern Po land, Sep tem ber 3–8, 2007. Geologica Polonica, 55: 283–293.