Plant Remains from the Polish Triassic. Present Knowledge and Future Prospects

Acta Palaeobotanica 54(1): 3–33, 2014 DOI: 10.2478/acpa-2014-0001

Plant remains from the Polish Triassic. Present knowledge and future prospects

GRZEGORZ PACYNA

Department of Palaeobotany and Palaeoherbarium, Institute of Botany, Jagiellonian University, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected]

Received 12 November 2013; accepted for publication 4 March 2014

ABSTRACT. The Triassic plant macrofossils of Poland are very poorly known. There are few Triassic rock exposures here, they contain very few plant specimens, there is little scientific interest in the subject, and the rare plant remains found in drill cores are of low stratigraphical significance. The Lower Triassic macroflora is surprisingly poorer taxonomically than coeval European floras, and only single specimens have been found. The flora of the Middle Triassic is even poorer as a result of the Muschelkalk sea transgression. Only the Upper Triassic floras contain many specimens and taxa. The Upper Triassic macrofloras from Polish territory are well known since the early 19th century. Pioneering descriptions of these floras were given by Goeppert and Raci- borski. From the Polish Triassic, the seed fern Lepidopteris ottonis (index species for the Rhaetian stage) and Neocalamites lehmannianus (sphenopsid species typical of almost all European Upper Triassic and Lower Juras- sic floras) were described for the first time ever. In the 20th century only single specimens were described from outcrops and drill cores. Barbacka revised Lepidopteris ottonis specimens from old collections and described some new material. Palynological research on Triassic strata in Poland intensified from the 1970s on. That work has produced spore-pollen and megaspore zonations for Triassic strata in Poland, but the correlation of the dispersed spores and pollen grains with their parent plants is low. The Polish Triassic flora is comprised of ferns, lycopsids, sphenopsids, cycads, bennettitaleans, ginkgoaleans and conifers. This flora is taxonomically poorer than equally old and geographically close European floras. All available data about Polish Triassic plants fossils are critically summarised in this paper for the first time. The biostratigraphical and lithostratigraphical correlations of Pol- ish Triassic floras with other European Triassic floras are outlined. New macrofloral assemblages for the Lower and Middle Triassic and macrofloral assemblage zones for the Upper Triassic are proposed for Poland. Recent new finds of taxonomically rich, abundant and well-preserved floras accompanying vertebrate remains in Silesia provide an opportunity for comprehensive research on Polish Triassic floras. This should improve our perception of their taxonomy and allow them to be described in evolutionary and palaeoecological contexts.

KEYWORDS: fossil plants, biostratigraphy, Lepidopteris zone, Triassic, Krasiejów, Poland

INTRODUCTION

The Triassic was one of the most important (King 1996, Sues 2000). Yet the Triassic flo- periods because it witnessed dynamic evolu- ras of the Northern Hemisphere are rare and tionary changes and the formation of modern poorly known (Meyen 1987). In this circum- ecosystems for the first time in the history stance, each new site with plant macroremains of life on Earth (Fraser 2006, Sues & Fraser and each new piece of information may fill a gap 2010). It is also the period in which floras were in our understanding of Triassic palaeoecology restored after the most devastating extinction and evolution, locally and globally. The well- in Earth’s history, at the Permian-Triassic described localities of floras of this age are few boundary (Taylor et al. 2009). Floras that orig- (Kustatscher & van Konijnenburg-van Cittert inated during this period would occur through 2005, 2008, Roghi et al. 2006, van Konijnen- almost the entire Mesozoic era and would be burg-van Cittert et al. 2006, Kustatscher et al. the source of food for herbivorous dinosaurs 2012) but the information they provide forms 4 our knowledge of the taxonomic composition localities with well-preserved floral remains in of Triassic floras, particularly that of gym- the 1990s and early 21st century. Krasiejów, nosperms. Very intensive radiative processes Patoka and Lipie Śląskie-Lisowice are new fos- occurred in this group at that time, leading to sil plant sites found during excavation of the the appearance of new groups, possibly includ- bones of large land vertebrates. These locali- ing the angiosperms (Anderson & Anderson ties have yielded numerous plant remains. 1997, 2003). The Triassic is an exceptionally Besides expanding our knowledge of Triassic interesting period as it saw the evolution of floras in Poland, a full description of the mac- conifers (Miller 1977, Grauvogel-Stamm 1978, roflora from these localities will help - recon Rothwell et al. 2012). The oldest representa- struct the palaeoenvironment of the animals tives of some modern families appeared in the at those sites. Equipped with knowledge of Upper Triassic. The stages of transformation the flora that co-occurred with dinosaurs, we from primitive Votziales into evolutionarily can more fully characterise those ecosystems, advanced families of modern conifers in the including the plant-animal interactions. Triassic are not yet fully documented (Miller This paper gives an overview of the Triassic 1977, Rothwell et al. 2012). plant taxa of Poland, and summarises the data Problems of stratigraphy make it difficult on diversity reported in historical documents. to identify evolutionary events in the Triassic. Thick sequences of strata without good index fossils are difficult to correlate regionally and HISTORICAL SUMMARY OF RESEARCH globally, hampering the task of distinguishing ON TRIASSIC FLORAS IN POLAND evolutionary processes and sequences of terrestrial organisms (Hunt & Lucas 1991, Lucas Goeppert (1836, 1841–1846, 1844, 1845, 1998, Rayfield et al. 2005, Lucas et al. 2007b, 1846a, b) was the first to describe the plant Kozur & Bachmann 2008). Plant micro- and fossils of the Polish Triassic. His pioneering macrofossils have proved useful in Triassic works on the Upper Triassic flora of Silesia biostratigraphy, especially in palynological gave descriptions of new taxa later regarded as zonation (Dobruskina 1988, 1994, Fijałkowska- index or very common taxa in many European Mader 1999, Lucas 2006, 2007, Cirilli 2010, Triassic floras. Goeppert’s research was quite Kürschner & Herngreen 2010). Ash (1980, advanced for his time; for example, he described 1987) proposed a macrofossil plant biostratig- and illustrated anatomical details of the Lepi raphy for the Upper Triassic Chinle Group in dopteris ottonis cuticle (Goeppert 1846a). Such North America. Unfortunately not a single depth of approach was rare in early palaeobot- species is shared by the North American and any. Goeppert erroneously determined the age European Upper Triassic floras, so an exact of the flora as Middle Jurassic, though some correlation is almost impossible. specimens he described (wood fragments) are Little is known about the Triassic flo- indeed Middle Jurassic in age. Next, Roemer ras of Poland (Lilpop & Kostyniuk 1957, (1867, 1870), then the foremost expert on Orłowska-Zwolińska & Senkowiczowa 1970, Silesian geology, corrected the age given by Reymanówna 1986). There are only a very few Goeppert to Upper Triassic (Rhaetian). Roe- descriptions of macroremains (Goeppert 1844, mer compared the Silesian flora described 1846a, Raciborski 1890a), written mainly in by Goeppert with the German Keuper floras the 19th century. They require a modern revi- described by Schenk (1867). Roemer showed sion incorporating current nomenclature and Goeppert’s original specimens to Schenk, the meeting today’s standards of description for top expert on floras of the Triassic-Jurassic fossil plants (Tab. 1). Palynological research on boundary. Schenk studied Goeppert’s speci- Triassic strata in Poland intensified beginning mens and described new specimens collected in the 1970s. It produced a useful palynologi- by Roemer, including some new taxa (Schenk cal biostratigraphic scheme for the Polish Tri- 1867, Roemer 1867, 1870). Roemer also gave assic, proposed by Orłowska-Zwolińska (1985). the first complete description of the Triassic A comparable scheme based on plant macro- geological setting in Silesia (Roemer 1862, fossils is proposed here (Tab. 2). 1863, 1867, 1870). Roemer (1870) discovered New prospects for research on Triassic flo- another Upper Triassic flora in Upper Silesia, ras in Poland were opened by discoveries of preserved in crenogenic limestones known as 5 the Woźniki Limestone. Rich collections of fos- palynological data (Orłowska-Zwolińska 1967, sil plants have been assembled from this lime- Marcinkiewicz 1962, 1969, 1971, Fijałkowska stone since then (Różycki 1930, Reymanówna 1990, 1992a, 1994a, 1995, Pieńkowski et al. 1986, Szulc et al. 2006, Szulc & Becker 2007) 2012, Fijałkowska-Mader 2013). However, the but not described. Kunisch (1886) and Michael correlation between the parent plants and the (1895) described some plant remains from the dispersed spores and pollen grains requires Muschelkalk of Upper Silesia. In 1890, Raci- much more study. borski described the Rhaetian flora from the Recent finds of taxonomically rich, abun- Tatra Mountains at the Czerwone Żlebki local- dant and well-preserved floras accompanying ity (Raciborski 1890a, b, c). vertebrate remains in Silesia have opened an Then research on Triassic plants in Poland opportunity for new research on Polish Triassic came to a virtual halt. Some specimens dis- floras. The research on these localities is only covered by geologists studying the geology of just beginning (Dzik & Sulej 2007, Staneczko the Holy Cross Mountains at the beginning of 2007, Dzik et al. 2008a, b, Wawrzyniak & Ziaja the 20th century were noted in the literature 2009, 2010, Wawrzyniak 2010a, b, c, Barbacka (Czarnocki 1925, 1931, Samsonowicz 1929) but et al. 2012, Pacyna et al. 2013a, b). without descriptions or illustrations. After the Second World War, stratigraphical research by geologists produced new macroremains from COMPOSITION AND EVOLUTION drill cores and outcrops, only some of which OF PLANT ASSEMBLAGES IN THE have been described (Bocheński 1957, Piwocki POLISH TRIASSIC, AND THEIR 1970, Pawłowska 1979, Fuglewicz 1980b, STRATIGRAPHICAL VALUE Reymanówna 1986, Barbacka 1991, Brzyski & Heflik 1994, Barbacka & Wcisło-Luraniec LOWER AND MIDDLE TRIASSIC 2002, 2003, Ociepa et al. 2008, Barbacka (BUNTSANDSTEIN, MUSCHELKALK et al. 2009). Barbacka revised the Lepidopt AND LOWER KEUPER) eris ottonis specimens from old collections and described some new materials (Barbacka 1980, Lower Triassic floras worldwide are rare 1991, Reymanówna & Barbacka 1981). and taxonomically impoverished as a result Palynological research on Triassic strata of the greatest extinction in Earth’s history in Poland accelerated starting from the at the Permian-Triassic boundary (Looy et al. 1970s (Pautsch 1958, 1971, 1973, Orłowska- 1999, Grauvogel-Stamm & Ash 2005). This is Zwolińska 1967, 1971, 1976, 1977, 1983, 1984, also true for the Polish Triassic localities. The 1985, 1986, 1988, Marcinkiewicz 1962, 1969, Permian-Triassic boundary has been docu- 1971, 1976, 1978, 1981, 1992a, b, c, Fuglewicz mented palynologically from several Polish 1973, 1979a, b, 1980a, b, Dybova-Jachowicz boreholes (Fijałkowska 1990, 1992a, 1994a, & Laszko 1980, Fuglewicz & Śnieżek 1980, 1995, Fijałkowska-Mader 2013). Rdzanek 1982, Marcinkiewicz & Orłowska- Plant macrofossils from the Polish Lower Zwolińska 1985, 1994, Fuglewicz & Marcinkie- and Middle Triassic are exceptionally few wicz 1986, Fijałkowska 1989, 1990, 1992a, b, in number of specimens and number of taxa 1994a, b, 1995, Fijałkowska & Uchman 1993, (Tab. 1). Most recorded findings are from the Krupnik & Ziaja 2010, Pieńkowski et al. 2012, Holy Cross Mountains (Czarnocki 1925, 1931, Fijałkowska-Mader 2013). From the out- Samsonowicz 1929, Bocheński 1957, Lilpop set these studies focused on biostratigraphy. & Kostyniuk 1957). Some are from Silesia Palynological research became especially use- (Kunisch 1886, Michael 1895b), the Tatra ful in determining the age of nonmarine sedi- Mountains (Limanowski 1901), and boreholes ment sequences not bearing other index fossils. from various regions of Poland (Fuglewicz The investigations bore fruit in useful spore- 1980b). Only rare lycopsid, sphenopsid, conifer, pollen and megaspore zonations for Triassic and possibly seed fern remains have been iden- strata in Poland (Marcinkiewicz 1971, 1978, tified so far (Tab. 1). Intensive new research on 1992c, Fuglewicz 1980b, Orłowska-Zwolińska invertebrate and vertebrate tracks, which are 1983, 1985, Fijałkowska-Mader 1999). The very numerous and taxonomically diverse in Permian-Triassic and Triassic-Jurassic bound- the Holy Cross Mountains, has, surprisingly, aries have been well delineated based on provided only single new fossil plant specimens 6

16oE 20oE 24oE

GDAŃSK

54oN 54oN

OLSZTYN

SZCZECIN

POZNAŃ WARSZAWA 52oN 52oN

3 ŁÓDŹ 4 6 5 7 LUBLIN WROCŁAW 8 9 10 11 12 13 16 14 15 17 19 18 20 22 21 24 23 KRAKÓW 0 100 km o 50oN 50 N

16oE 20oE 24oE

Fig. 1. Macrofossil and selected microfossil plant localities from the Polish Triassic. 1 – Chabowo 2 borehole; 2 – Gradzanowo 1 and 3 boreholes; 3 – Gostyń 46 G borehole; 4 – Kołaczkowice near Rawicz; 5 – Otyń IG-1 borehole; 6 – Studzianna borehole; 7 – Radoszyce 3 borehole; 8 – Wieluń; 9 – Pałęgi; 10 – Ratajów; 11 – Gorzów Śląski area (Lofkowitz, Sumpen); 12 – Majków area; 13 – Kluczbork area (Biadacz, Dobiercice, Gosław, Maciejów); 14 – Gacki 1, 3 and 4 boreholes; 15 – Krasiejów; 16 – Patoka; 17 – Lipie Śląskie-Lisowice; 18 – Krapkowice; 19 – Ligota near Woźniki; 20 – Woźniki; 21 – Myszków-Nowa Wieś; 22 – Kamie- nica Polska; 23 – Tarnowskie Góry; 24 – Poręba and Marciszów near Zawiercie; 25 – Czerwone Żlebki

(Karaszewski 1966, 1976, Gradziński et al. 1979a, b, 1980a, b, Marcinkiewicz 1976, 1992c, 1979, Fuglewicz et al. 1981, Senkowiczowa Orłowska-Zwolińska 1977, 1984, 1985, 1988, 1982, Mader & Rdzanek 1985, Fuglewicz et al. Rdzanek 1982, Fijałkowska 1994b, Fijałkowska 1990, Gradziński & Uchman 1994, Machalski & Uchman 1993, Fuglewicz & Marcinkie- & Machalska 1994, Ptaszyński 1996, 2000a, wicz 1986). A biostratigraphical scheme was b, Kuleta et al. 2001, 2005, 2006, Ptaszyński proposed based on palynology (Fuglewicz & Niedźwiedzki 2002, 2004, 2006, Niedźwiedzki 1980b, Orłowska-Zwolińska 1985, Marcin- et al. 2007, 2013, Niedźwiedzki & Ptaszyński kiewicz 1992c, Fijałkowska-Mader 1999) but 2007, Brusatte et al. 2011). In France, by con- the correlation of the dispersed spores and trast, well-preserved and taxonomically rich pollen grains with their parent plants is low. flora and fauna are known from coeval and Lower Triassic vertebrate bone fossils are similarly geologically structured Anisian strata very rare in Poland (Borsuk-Białynicka et al. – Grés a Voltzia (Grauvogel-Stamm 1978). 1999, Borsuk-Białynicka & Evans 2009, Sulej The palynological record of Lower Tri- & Niedźwiedzki 2013). assic plants in Poland is better than the Buntsandstein-type facies (mostly terres- record from macroremains (Fuglewicz 1973, trial) were replaced by fully marine facies in 7 the Anisian as a result of the Muschelkalk sea UPPER TRIASSIC transgression (Eck 1865, Marek & Pajchlowa (MIDDLE AND UPPER KEUPER) 1997, Nawrocki & Szulc 2000, Szulc & Becker The Upper Triassic stratigraphy of Poland is 2007, Kowal-Linka & Bodzioch 2012). Because confused, as it uses local stratigraphic divisions of this, the flora of the Muschelkalk in Poland, poorly correlated with the Germanic Basin as all over Europe, is even poorer than for the stratigraphy accepted in Europe (Znosko 1955, Lower Triassic. Palaeontological research on Szyperko-Śliwczyńska 1960, Dadlez & Kopik the Muschelkalk strata in Poland has a long 1963, Senkowiczowa 1966, 1970, Marcinkiewicz history (Eck 1865), and rich marine verte- 1969, 1971, 1978, Grodzicka-Szymanko 1971, brate fauna have been described (Roemer Grodzicka-Szymanko & Orłowska-Zwolińska 1870, Gürich 1884, Tarlo 1959, Lis & Wójcik 1972, Kotlicki 1974, Gajewska 1978, Czapowski 1960, Rieppel & Hagdorn 1997, Chrząstek & Romanek 1986, Pieńkowski 1988, Marek & Niedźwiedzki 1998, Rieppel 2000, Bardziński & Pajchlowa 1997). The first descriptions of et al. 2008, Chrząstek 2008). Macroscopic Upper Triassic strata in Poland, given by Romer plant remains are almost entirely absent, how- (1862, 1863, 1867, 1870), were based on the ever (Goeppert 1846b, Kunisch 1886, Michael German geological tradition and correspond to 1895), and the palynological record is meager the rest of the Germanic Basin. Unfortunately, (Orłowska-Zwolińska 1977, 1985, 1986, 1988, Samsonowicz (1924, 1929) then proposed a local Marcinkiewicz 1992a). Only calcified algae are stratigraphical division for the Polish Upper Tri- well known from these strata (Raciborski 1892, assic, a division somewhat similar to that used 1927, Szulc & Becker 2007), some even rock- in France. Polish geologists used stratigraphi- forming (Diplopora Beds). From the Muschel- cal nomenclature typical of the German part of kalk sea regression in the Lower Keuper the Germanic Basin but differing in meaning (Ladinian), rich swamp vegetation is known from that used in Germany: for example, Rhae- in Germany (Mader 1995), but in Poland only tian sensu polonico (Znosko 1955, Mader 1997, isolated horsetail remains have been described Marek & Pajchlowa 1997, Kozur & Bachmann (Pawłowska 1979, Barbacka et al. 2009, Sulej 2008, Franz 2009). This is the source of so much et al. 2011b). Lilpop (in Raciborski 1927) noted confusion in the geological literature, especially the discovery of the Lettenkohle flora from that created by foreign authors unfamiliar with Piekoszów in the Holy Cross Mountains. This the Polish local tradition. It is clear that the flora consisted of cycadaleans and conifers and Polish Triassic sedimentary basin is part of the was never illustrated or even preliminarily Germanic Basin (Mader 1997, Franz 2009), but described. because it is a marginal part of this basin, and Recently an interesting new locality was because of local variability in the facies typical found at the Pałęgi claypit in the Holy Cross of the Germanic Basin, the stratigraphical cor- Mountains (Kuleta et al. 2006, Żyła et al. 2013). relations are difficult (Racki 2010). This is the There, Lower/Middle Triassic sediments (latest Polish share in the global problems of Triassic Olenekian–early Anisian, Upper Buntsand- stratigraphy as discussed above (Szulc et al. stein) crop out with numerous well-preserved 2006, Szulc & Becker 2007). remains of crustaceans and insects, and inver- The Upper Triassic macro- and microflora tebrate and vertebrate tracks. Miospores from from Poland are well known but many descrip- the fossiliferous level are abundant and typical tions are outdated (Goeppert 1836, 1841–1846, of the Voltziacaesporites heteromorpha zone 1844, 1845, 1846a, b, Raciborski 1890a, b, c, (Żyła et al. 2013). Macrofossil plant remains Orłowska-Zwolińska & Senkowiczowa 1970, were also found there but they are not very Reymanówna 1986) and new discoveries have well-preserved; some remains were recognised yet to be described (Dzik & Sulej 2007, Dzik as horsetail stems (Kuleta et al. 2006, Żyła et al. 2008a, b, Niedźwiedzki & Sulej 2008, et al. 2013) and others were indeterminable. Ociepa et al. 2008, Wawrzyniak & Ziaja 2009, In the collection from this locality held in the 2010, Wawrzyniak 2010a, b). A comparison of W. Szafer Institute of Botany, Polish Academy the available data with floras described thus of Sciences, none of the plant remains could be far from Germany (Linck 1950, Kräusel 1952, determined even approximately (pers. observ.). Mägdefrau 1953, 1956, 1963, Kelber & Hansch This fossil assemblage is similar in age to the 1996, Axsmith & Taylor 1997, Kelber & van famous Grés à Voltzia. Konijnenburg-van Cittert 1997, Kelber 1998, 8

2005, Arndt 2002, Kelber & Nitsch 2005) can insects (Dzik & Sulej 2007, Dzik 2008). The yield useful stratigraphical conclusions. I pro- site’s superbly preserved vertebrates include pose such a correlation, for the first time in the fish (Dzik & Sulej 2007), temnospondyls Polish palaeontological literature, in Table 2. (Sulej 2002, 2007, Sulej & Majer 2005, Bary- Many data on Polish Upper Triassic palynol- cka 2007, Konietzko-Meier & Wawro 2007, ogy have been gathered (Pautsch 1958, 1971, Konietzko-Meier & Klein 2013, Konietzko- 1973, Orłowska-Zwolińska 1967, 1971, 1976, Meier & Sander 2013, Konietzko-Meier et al. 1983, 1984, 1985, 1986, 1988, Marcinkiewicz 2013), a phytosaur (Butler et al. 2014), a raui- 1969, 1971, 1978, 1981, 1992b, Fuglewicz suchian (Sulej 2005, Brusatte et al. 2009), an 1980a, Fuglewicz & Śnieżek 1980, Marcinkie- aetosaur (Sulej 2010, Desojo et al. 2013) and wicz & Orłowska-Zwolińska 1985, 1994, Fugle- a dinosauromorph (Dzik 2003a, Dzik & Sulej wicz & Marcinkiewicz 1986, Fijałkowska 1989, 2004, Mazurek & Słowiak 2009, Fostowicz- 1992b, Pieńkowski et al. 2012). Well-grounded Frelik & Sulej 2010, Piechowski & Dzik 2010). biostratigraphical schemes have been proposed Numerous charophyte oogonia have been (Marcinkiewicz 1971, 1978, Orłowska-Zwolińska described (Zatoń & Piechota 2003, Zatoń et al. 1983, 1985, Fijałkowska-Mader 1999). The 2005) but unfortunately this locality contains Triassic-Jurassic boundary has been identi- no palynological record. This prevents the use fied based on palynological data (Orłowska- of palynological biostratigraphy here. The mac- Zwolińska 1967, Marcinkiewicz 1969, 1971, roflora of this site has not yet been described Pieńkowski et al. 2012, Pacyna 2013). in detail. Preliminary information has been given (Dzik 2003b, Dzik & Sulej 2004, 2007) The Carnian flora and a detailed report is in preparation (Pacyna – Voltzia floral assemblage zone & Zdebska in prep.). The macroflora assem- One of the most significant discoveries in blage from Krasiejów consists predominantly Polish palaeontology was the Krasiejów local- of conifers (Pseudohirmerella sp., Glyptolepis ity, found in the 1990s (Dzik et al. 2000, keuperiana, Voltzia spp., Pachylepis quinquies, Dzik 2001, 2003a, b, Sulej 2003, 2004). This Desmiophyllum spp., new taxa), and rare ferns is the most important Triassic Polish verte- (Sphenopteris schoenleiniana) and bennet- brate locality, of international repute (Sues titaleans (Pterophyllum sp.). This taxonomi- & Fraser 2010, Jagt-Yazykova et al. 2012). For cally poor flora is not well-preserved (cuticular our understanding of the evolution of Triassic details not recognisable). Xeromorphic features ecosystems the Krasiejów locality is among the of the plants indicate rather dry climatic con- world’s key sites, as seen in publications about ditions. The most taxonomically similar flora its fauna (Dzik 2001, 2003a, b, 2008, Sulej 2002, was described by Mägdefrau (1953, 1956, 1963) 2005, 2007, 2010, Dzik & Sulej 2004, 2007, from the Coburger Sandstein (=Semionotus Lucas et al. 2007a, Fostowicz-Frelik & Sulej Sandstein) at Haßfurt am Main. It could be 2010, Mazurek & Słowiak 2009, Olempska useful in determining the age of the Krasiejów 2004, 2011, Piechowski & Dzik 2010, Skawina fossil assemblage. The Coburger Sandstein cor- & Dzik 2011). Taxonomically, the fossil assem- responds to the Kieselsandstein and Blasen- blage of this locality is the richest in the Polish sandstein (Mader 1990), now referred to as the Triassic and one of the richest in the European Hassberge Formation (Kozur & Weems 2010), Triassic (Sulej 2003, 2004, Dzik & Sulej 2004, whose age is estimated as Late Carnian (Mid- 2007, Lucas et al. 2007a, Sues & Fraser 2010). dle Tuvalian). Thus, Late Carnian age is pro- The precise age, stratigraphy and taphonomy posed here for the Krasiejów fossil assemblage. of this locality are the subject of debate but A full description of the Krasiejów macroflora plausible solutions to these issues have been will improve our knowledge of the Triassic flo- offered (Kotlicki & Kubicz 1974, Bilan 1975, ras of Poland and the palaeoenvironment of the Kłapciński 1993, Dzik et al. 2000, Dzik 2001, animals at this site. Szulc 2005, Szulc & Becker 2007, Gruszka The fossil assemblage from the Woźniki & Zieliński 2008, Bodzioch & Kowal-Linka claypit may be similar in age to the Krasiejów 2012). Its rich invertebrate fauna consists of assemblage. Norian Woźniki Limestone has conchostracans (Olempska 2004, 2011, Kozur been identified without doubt above the fossil- & Weems 2010), bivalves (Skawina 2010, iferous level (Sulej et al. 2011a). Besides inver- 2013, Skawina & Dzik 2011), crustaceans and tebrate and vertebrate remains, characean 9 gyrogonites and conifer shoots similar to those There is also another new type of ovulate discovered in Krasiejów have also been found scale with two oval lobes – probably the place (Sulej et al. 2011a, T. Sulej pers. comm. 2011). of ovule attachment – and one reduced lobe between them, in one plane. The scales also The Norian flora – Brachyphyllum floral show cuticular structure similar to the mass of assemblage zone Brachyphyllum leaves in the same rock slabs. There are some Upper Triassic localities Together with the branches and scales was in Silesia with taxonomically very poor floras one ovule whose shape and size suggest that it whose most characteristic feature is the occur- might be associated with the scales. The ovule rence of Brachyphyllum-like shoots. These flo- is oval and has an exposed micropylar beak. ras are distinguished here as Norian (=Lower Pollen grains were found inside the ovule. The Rhaetian sensu polonico) in age, based on same type of pollen grain has been found in palynological and geological data (Szulc et al. well-preserved male cones. The Brachyphyl 2006, Szulc & Becker 2007). Floras of the lum-type conifers are accompanied by frag- Norian stage are seldom preserved and are ments of cf. Czekanowskia leaves, isolated extremely rare all over the world. The taxo- seeds of unknown affiliation, and gymnosperm nomic composition of the younger (Rhaetian) wood. Adaptations to dry climate are obvious and older (Carnian) floras indicate that rapid in the cuticle structure of all the plants from evolutionary changes took place just then, the Patoka locality (compare Preto et al. 2010). especially within the conifers. Modern conifer Yet another locality, Poręba, was recently families probably originated during this time. discovered; it may represent the same floral The first such flora was identified by assemblage (Sulej et al. 2012). A very interest- Roemer (1870) in crenogenic limestones in ing assemblage of turtles, aetosaurs, and coe- the northern part of Upper Silesia (Ligota, lophysoid dinosaurs was found at this locality. Myszków, Poręba), called the Woźniki Lime- In the flora, Brachyphyllum shoots and coni- stone. Rich collections of fossil plants have fer trunks, ginkgoalean (?) leaves and sporo- since been assembled from it (Różycki 1930, morphs were identified (Sulej et al. 2012). The Reymanówna 1986) but unfortunately not Marciszów-Zawiercie locality could also be described. The plants are preserved as voids or Norian in age; besides, the remains of freshwa- casts in the limestone (Szulc et al. 2006, Szulc ter bivalves, vertebrate (dicynodontid) bones, & Becker 2007). Besides the most numer- and tracks assignable to archosaurs and dicy- ous Brachyphyllum shoots along with seed nodonts, only palynomorphs and equisetalean scales and male cones probably belonging to remains are known to derive from that site the same plant, rare fern and bennetittalean (Szulc et al. 2006, Budziszewska-Karwowska leaves and sphenopsid shoots have been found et al. 2010, Racki 2010, Skawina & Dzik 2011, in this flora (Tab. 3). During his examination Sadlok & Wawrzyniak 2013). Also Norian in of the Roemer collection, Raciborski found age are rocks exposed in a claypit in Wyszyna some liverwort remains which he described as Machorowska near Radoszyce in the north- the new species Palaeohepatica roemeri (Raci- western part of the Holy Cross Mountains. borski 1892, 1893, Wcisło-Luraniec 1984) but At that site, apart from a vertebrate burrow this species was never illustrated and requires system some plant remains were recognised revision. (Tałanda et al. 2011). Patoka is an Upper Silesian locality newly discovered by T. Sulej and G. Niedźwiedzki, The Rhaetian flora – Lepidopteris ottonis with a rather homogeneous but very interest- floral assemblage zone ing flora of the same type as in the Woźniki It was from the Upper Silesian Kluczbork Limestone (Barbacka et al. 2012). The assem- area (Biadacz, Dobiercice, Gosław, Macie- blage from Patoka is also dominated by jów) that Goeppert (1836, 1841–1846, 1844, branches of Brachyphyllum, but unlike the 1845, 1846a) described the first Polish Trias- Woźniki Limestone the flora from Patoka is sic macroflora. It was also the first Rhaetian very well-preserved, with anatomical details. flora described in Europe. Then Schenk (1867) Besides Brachyphyllum branches and leaves, and Roemer (1867, 1870) published new data Pachylepis seed scales whose cuticle matches for this flora, based mostly on new specimens. some of the Brachyphyllum leaves were found. Plant remains were preserved in sideritic 10 nodules along with fish (Michael 1893, 1894, of this fossil assemblage. Cycad leaves and 1895a, Gallinek 1894). Goeppert’s papers (1836, cones, ginkgoalean leaves, conifer wood, twigs, 1846a) described for the first time the seed fern seed scales, male cones, and seeds have also Lepidopteris ottonis (the index species for the been preliminarily described (Dzik et al. 2008a, Rhaetic stage) and Neocalamites lehmanni b, Marynowski & Simoneit 2009, Wawrzyniak anus, a sphenopsid species typical of almost all & Ziaja 2009, 2010, Wawrzyniak 2010a, b, c, European Upper Triassic and Lower Jurassic d, e, 2011, Jarzynka & Wawrzyniak 2012). The floras. This flora is not very diversified; besides macroflora from Lipie Śląskie-Lisowice is sur- the taxa listed earlier, fern and bennettitalean prisingly advanced, with many taxa, mainly of or cycad leaves have been described (Tab. 3). cycads and conifers similar to younger Juras- The flora requires reworking in the context of sic taxa (Harris 1932, 1935, 1937), but there current taxonomy, nomenclature rules, and are also some taxa, especially of conifers, standards of description. As is already clear with obvious evolutionary roots in older Tri- from Schenk’s (1867) revision, some species assic Silesian floras, especially at Krasiejów described by Goeppert are junior synonyms of and Patoka. These facts and the richness of previously described species, some new species the well-preserved taxa place this flora among were described from small, badly preserved the most interesting Rhaetian floras of Europe remains, and species in some genera are over- (compare Harris 1926, Kelber & Hansch 1996, split (e.g. Pterophyllum). Needed is a compari- Kelber & van Konijnenburg-van Cittert 1997, son of the revised taxa from this flora with taxa Kelber 1998). from the new Silesian Rhaetian floral localities. In the Tatra Mountains the Upper Triassic Gothan (1909a) and Barbacka (Barbacka 1980, strata are developed in Keuper facies typical 1991, Reymanówna & Barbacka 1981) revised of the Germanic Basin. In Czerwone Żlebki the Lepidopteris ottonis specimens from the (West Tatras, Czerwone Wierchy Massif), Goeppert and Roemer collections and described Raciborski (1890a, b, c) discovered and then some new material. In the old collection, Bar- described an interesting Rhaetian flora. The backa (1991) recognised a Peltaspermum rotula plant remains are not well-preserved and the cupulate disc and several isolated seeds that cuticular details are not suitable for prepara- belong to Lepidopteris ottonis. tion. The occurrence of the endemic horsetail The Lipie Śląskie-Lisowice locality was species Equisetum chalubinskii is this flora’s discovered at the beginning of the 21st cen- most characteristic feature. Other horsetails, tury in an active claypit (Dzik et al. 2008a, b, ferns and conifers are rare in this flora (Tab. 1). Niedźwiedzki & Sulej 2008). Remains of thero- Later researchers (Reymanówna 1984, 1986, pods and dicynodonts were found together Michalík et al. 1976, 1988) attempted to find there, an interesting finding from biogeo- new plant remains at this locality but with- graphical and evolutionary points of view (Dzik out success. In the M. Reymanówna collection et al. 2008a, b, Niedźwiedzki & Sulej 2008, (W. Szafer Institute of Botany, Polish Acad- Niedźwiedzki et al. 2011, 2012). Fossils of emy of Sciences) from Czerwone Żlebki there other archosaurs, temnospondyls, fish, a mam- are no determinable plant remains. Nor has mal-like tooth, conchostracans and bivalves, the flora of Czerwone Żlebki been revised since and invertebrate and vertebrate tracks were its original description. Geologists and palaeo- also identified (Dzik et al. 2008a, b, Gorzelak botanists have long awaited a modern revision et al. 2010, Świło 2010, Skawina & Dzik 2011, of this flora (Reymanówna 1984, Niedźwiedzki Świło et al. 2014). Plant remains from the Lipie 2011, Lintnerová et al. 2013). Such a revision Śląskie-Lisowice claypit were first noted in the would improve our grasp of Tatra Upper Trias- early 1980s (Fuglewicz & Śnieżek 1980, Mar- sic palaeoecology, a key element of the complex cinkiewicz 1981) but only megaspores were geological history of the Tatra Mountains and preliminarily described. Current research on the palaeogeographic connections between the this locality has yielded many different well- Germanic Basin and the Alpine Tethys Ocean. preserved plant macroremains (Dzik et al. Those relationships can be determined from 2008a, b). Staneczko’s finding of cf. Lepidop the similarities or differences in the taxonomic teris ottonis isolated cuticles (Staneczko 2007, composition of the flora. Important dinosaur Ociepa et al. 2008) and also palynological data ichnotaxa were discovered at this site and (Dzik et al. 2008a, b) confirm the Rhaetian age described recently (Niedźwiedzki 2005, 2008, 11

2011, Niedźwiedzki & Sulej 2007). That dis- thus far been discovered anywhere. The genus covery gives a further reason for studying the Lepidopteris Schimper 1869 is known in the flora of that place. Germanic Basin from the beginning of the Some other plant macroremains have been Upper Triassic (Schimper 1869, Gothan 1909a, found in the Upper Triassic strata of other b, c, Antevs 1914). Lepidopteris stuttgardien Tatra localities but as a rule they are badly sis (Jäger 1827) Schimper 1869 and Lepido preserved (Raciborski 1892, Limanowski 1901, pteris brevipinnata Mägdefrau 1953 have been Uchman pers. comm. 2012) and have never described from the Schilfsandstein strata – been properly described or illustrated. Lower-Middle Carnian (Jäger 1827, Schimper There are other plant fossil localities origi- 1869, Gothan 1909b, Mägdefrau 1953). The nally proposed as Rhaetian in age but these cuticle structure of these species is not known. claims remain unconfirmed. Roemer (1866) Lepidopteris ottonis is known from Rhaetian identified a Lepidopteris ottonis specimen in strata but no species of Lepidopteris has been Mierzęcina (Miedzieczo in Roemer’s paper) described from the Norian. Because the cutic- near Kielce but as Barbacka (Reymanówna ular differences among Lepidopteris species & Barbacka 1981, Barbacka 1991) showed this from the Upper Triassic of Europe cannot be specimen belongs to an unidentified fern. Lil- compared at present, the question of extending pop (in Raciborski 1927) noted the discovery the Lepidopteris ottonis zone to the Norian on of a new Keuper or Rhaetian flora in iron ore the basis of isolated cuticle fragments cannot mines in Biała Góra near Bliżyn (Holy Cross be settled. The Norian specimens of Lepidopte Mountains). This flora, however, was never ris may belong to a species undescribed as yet. illustrated or even preliminarily described. The inter- and intraspecific variation of the The above mentioned plant remains probably cuticle in Lepidopteris is very poorly known. derive from Lower Jurassic strata widely out- Some authors have suggested that Lepidopteris cropping in this area (Pieńkowski 2004, Pacyna stuttgariensis and L. ottonis belong to the same 2013), but not from Keuper or Rhaetian strata, species (Kelber & van Konijnenburg-van Cittert as Roemer (1866) and Lilpop (Lilpop in Raci- 1997, Kiritchkova 2006, 2008) but this has not borski 1927) originally proposed. been shown with certainty. The type specimen A macrofloristic zonation of the strata of of Lepidopteris ottonis, described by Goeppert the Triassic-Jurassic boundary in Europe was (1836) from Wieluń, has not been found in Pol- proposed by Nathorst (1910) and developed by ish collections (Barbacka 1980, 1991, Reyman- Harris (1931, 1937). As currently understood, ówna & Barbacka 1981) and its whereabouts the occurrence of Lepidopteris ottonis marks are not known, so the typification proposed by the Rhaetian stage in nonmarine strata of the Kiritchkova (2006, 2008) is unwarranted. European Triassic. The known stratigraphic range of Lepidopteris ottonis coincides with the Riccisporites tuberculatus palynological zone. COMMENTS TO TABLES It is standard practice in Polish geology and palaeontology to refer the source strata to the Poland was partitioned by foreign powers Rhaetian when Lepidopteris ottonis remains three times in the second half of the 18th cen- are found in drill cores (Piwocki 1970, Bar- tury, ultimately ending Poland’s existence as backa 1980, 1991, Reymanówna & Barbacka an independent country for more than a cen- 1981, Staneczko 2007, Ociepa et al. 2008). Dur- tury. Poland was split between Prussia, the ing palynological examination of strata from Austro-Hungarian Empire and Russia. Mostly the Corollina meyerana zone (Norian), Marcin- German-speaking scientists were working in kiewicz and Orłowska-Zwolińska (1985, 1994) Silesia during that time, and the official place noted cuticle fragments which they determined names were in German. Below is a list of those to be cf. Lepidopteris ottonis. On this basis they localities in Silesia; their current names are suggested that the range of the Lepidopteris paired with their pre-1945 German names. It ottonis zone might overlap the Corollina meyer was not possible to establish the current Pol- ana and Riccisporites tuberculatus palynological ish names of some localities, especially when zones, thus the Norian and Rhaetian. They had the papers used vernacular German names of only small cuticle fragments, however; Norian localities in the Russian partition (e.g. in Goep- macroremains of Lepidopteris ottonis have not pert 1846a). Table 1 includes comments on the 12 nomenclatural status of taxa and the status of Panoszów (=Ponoszów) – Ponoschau (=Pon- specimens described from Poland. Synonyms noschau, Hegersfelde) are also given. The original published names Patoka – Klinkerwerk (=Patoka, Kreis Lublin- of boreholes, horizons and age determinations itz) are given in Table 1; Table 2 correlates the Sternalice – Sternalitz (=Ammern) strata. References for the main localities are Szubin – Schubin listed in Table 4. Tarnowskie Góry – Tarnowitz Woźniki – Woischnik Explanation of symbols and abbreviations Zakrzów – Sacrau in Table 1 Zborowskie – Zborowsky * – papers containing an illustration of a given taxon German – Polish ^ – papers in which a given taxon was described Ellguth – Ligota as new to science based on specimens from Pol- Goslau – Gosław ish territory Hellewald – Helenowo Landsberg – Gorzów Śląski MAIN LOCALITIES OF TRIASSIC GEOLOGICAL Lublinitz – Lubliniec OUTCROPS WITH FOSSIL PLANTS PRESERVED Ludwigsdorf – Biadacz IN UPPER SILESIA BEFORE 1945 (LOCALITY Klinkerwerk (=Patoka, Kreis Lublinitz) NAMES AFTER ROSPOND 1951) – Patoka Krappitz – Krapkowice Polish – German Kreuzburg, Kreu(t)zburg – Kluczbork Biadacz – Ludwigsdorf Matzdorf – Maciejów Bogdańczowice – Wüttendorf Neudorf – Nowa Wieś Oleska Byczyna – Pitschen Ober Kunzendorf – Kujanowice Dobiercice – Wilmsdorf Pitschen – Byczyna Gorzów Śląski – Landsberg Ponoschau (=Ponnoschau, Hegersfelde) Gosław – Goslau – Panoszów, Ponoszów Helenowo – Hellewald Sacrau – Zakrzów Kluczbork – Kreuzburg, Kreu(t)zburg Schubin – Szubin Krapkowice – Krappitz Sternalitz (=Ammern) – Sternalice Kujanowice – Ober Kunzendorf Tarnowitz – Tarnowskie Góry Ligota – Ellguth Wilmsdorf – Dobiercice Lubliniec – Lublinitz Woischnik – Woźniki Maciejów – Matzdorf Wüttendorf – Bogdańczowice Nowa Wieś Oleska – Neudorf Zborowsky – Zborowskie

Table 1. Triassic plant macroremains described so far from Poland

Taxon as originally Botanical affinity, described, selected References, taxon validity, taxo- Location Horizon, age nomenclatural comments nomical synonyms synonyms Alethopteris insignis Pterophytina, accord- Dobiercice Wilmsdorfer Schichten; Goeppert 1846a*, (Lindley & Hutton ing to Harris (1961) (Upper Silesia) Upper Triassic, Schenk 1867, Roemer 1833–1835) Goeppert Pecopteris insignis is Rhaetian 1867 1836 synonym of Clado (=Pecopteris insignis phlebis denticulata Lindley & Hutton (Brongniart 1834) 1833–1835, basonim) Schimper 1874; Goeppert’s (1846a) specimen is small and hardly determinable Araucarioxylon Coniferales Poręba near Zawiercie Upper Triassic – Lower Brzyski & Heflik 1994* Kraus 1870 (Upper Silesia) Jurassic, Rhaetico – Araucarioxylon sp. Liassic 13

Table 1. Continued

Taxon as originally Botanical affinity, described, selected taxon validity, taxo- Location Horizon, age References, comments nomenclatural nomical synonyms synonyms

Brachyphyllum Coniferales Patoka Jarkowo/Zbąszynek Barbacka et al. 2012 Brongniart 1828 (Upper Silesia) Beds; Upper Triassic, Brachyphyllum sp. Norian Lipie Śląskie-Lisowice Wielichowo Beds; Dzik et al. 2008a, (Upper Silesia) Upper Triassic, b (as Hirmeriella Rhaetian twigs), Wawrzyniak & Ziaja 2010, Jarzynka & Wawrzyniak 2012 Calamites Brongniart Equisetales Ratajów (north part of Upper Buntsandstein, Samsonowicz 1929 1828 Holy Cross Mts.) Röt; Lower Triassic (record without Calamites sp. description or illustration) Cladophlebis roes- Pterophytina, valid Dobiercice Wilmsdorfer Schichten; Roemer 1867, 1870, serti (Presl in Stern- species (Upper Silesia) Upper Triassic, Schenk 1867 berg 1838) Saporta Rhaetian 1872 (=Asplenites roes Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, sertii (Presl in Stern- (Tatra Mountains) Upper Triassic, c, Reymanówna 1986 berg 1838) Schenk Rhaetian 1867) Cladophlebis roes- Pterophytina, speci- Czerwone Żlebki Tomanová Formation; Raciborski 1890a*^, serti forma parvifo- mens need revision (Tatra Mountains) Upper Triassic, b, c lia Raciborski 1890 Rhaetian Calamites lehmann- Equisetales, valid Dobiercice Wilmsdorfer Schichten; Goeppert 1845, ianus Goeppert 1846a taxon, valid name (Upper Silesia) Upper Triassic, 1846a^*, Schenk 1867, (=Neocalamites hoeren – Neocalamites Rhaetian Roemer 1867 (as Equi sis (Schimper 1869) lehmannianus (Goep- setites arenaceus (?)), Halle 1908 sensu Halle pert 1846a) Weber 1870*, (older synonym 1908) 1968 of Neocalamites hoer (=Schizoneura hoeren ensis (Schimper 1869) sis (Hisinger 1840) Halle 1908 sensu Halle Schimper 1869) 1908 according to Weber 1968) Camptopteris juras- Pterophytina, Dipteri- Maciejów Wilmsdorfer Schichten; Goeppert 1846a^, sica Goeppert 1846a daceae, synonym of (Upper Silesia) Upper Triassic, 1841–1846*, Roemer (=Aspidium jurassi Clathropteris meni- Rhaetian 1867, Schenk 1867 cum (Goeppert 1846a) scoides (Brongniart (most probably synony- Ettingshausen 1865) 1824) Brongiart 1828 mous with Clathropt eris platyphylla = C. meniscoides) Carpolithes cardio- Gymnosperm seed Dobiercice Wilmsdorfer Schichten; Goeppert 1846a^* carpoides Goeppert (Upper Silesia) Upper Triassic, 1846a Rhaetian Clathropteris muen- Pterophytina, Dipteri- Dobiercice, Maciejów Wilmsdorfer Schichten; Schenk 1867, steriana (Presl in daceae, synonym of (Upper Silesia) Upper Triassic, Orłowska-Zwolińska Sternberg 1838) Clathropteris meni- Rhaetian & Senkowiczowa (1970) Schenk 1867 scoides (Brongniart erroneously listed (=Camptopteris mün 1824) Brongiart 1828 Clathropteris münsteri steriana Presl in (sic) based on Lilpop Sternberg 1838) & Kostyniuk (1957); using this species name they ascribed it to Goeppert (1845, 1846a), which is incorrect Ligota near Woźniki Woźniki Limestone; Roemer 1870* (as (Upper Silesia) Upper Triassic, Norian Clathropteris Münste riana) Clathropteris platy- Pterophytina, Dipteri- Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, phylla Goeppert 1846 daceae, synonym of (Tatra Mountains) Upper Triassic, c, Harris 1931, Rey- Clathropteris meni- Rhaetian manówna 1986 scoides (Brongniart 1824) Brongiart 1828 (Harris 1931) 14

Table 1. Continued

Taxon as originally Botanical affinity, described, selected taxon validity, taxo- Location Horizon, age References, comments nomenclatural nomical synonyms synonyms

Coniopteris lobata Pterophytina, probably Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, c (Oldham & Morris valid species referable (Tatra Mountains) Upper Triassic, (as Pecopteris lobata), 1863) Halle 1913 to genus Coniopteris Rhaetian Reymanówna 1986 (=Pecopteris lobata Old- (Harris 1961) ham & Morris 1863) Czekanowskia Heer Czekanowskiales Patoka Jarkowo/ Zbąszynek Barbacka et al. 2012 1876 (Upper Silesia) Beds; Upper Triassic, cf. Czekanowskia sp. Norian Desmiophyllum Les- Coniferales Krasiejów Drawno Beds?; Upper Dzik & Sulej 2007* quereux 1878 (Opole Silesia) Triassic, Desmiophyllum sp. Upper Carnian Dicranopteris roe- Pterophytina, species Dobiercice Wilmsdorfer Schichten; Schenk 1867^* (as meriana Schenk 1867 needs revision (Upper Silesia) Upper Triassic, Dicranopteris Römeri Rhaetian in text, as Dicrano pteris Römeriana in plate caption), Roemer 1870 (as Dicranopteris Roemeriana) Dictyophyllum aff. Pterophytina Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, dunkeri Nathorst (Tatra Mountains) Upper Triassic, c, Reymanówna 1986 1878 Rhaetian (poorly preserved small leaf fragment) Equisetites arena- Equisetales, valid Majków area Lower Keuper, Letten- Samsonowicz 1929 (as ceus (Jaeger 1827) taxon (north part of Holy kohle; Middle Triassic, Calamites arenaceus, Schenk 1864 Cross Mts.) Ladinian record without description or illustration) Equisetum Equisetales, valid Czerwone Żlebki Tomanová Formation; Raciborski 1890a^*, b, chałubińskii Racibor- taxon, proper taxon (Tatra Mountains) Upper Triassic, c, Reymanówna 1986. ski 1890 orthography Equise- Rhaetian Species very similar to tum chalubinskii Equisetites muensteri Raciborski 1890 Sternberg 1833; should properly be referred to genus Equisetites Equisetum an bun- Equisetales, specimens Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, c, buryanum? Zigno need revision (Tatra Mountains) Upper Triassic, Reymanówna 1986 1856 Rhaetian Equisetites Sternberg Equisetales, specimens Western Holy Cross Lower Buntsandstein; Czarnocki 1925 (only 1833 never illustrated or Mts. Lower Triassic record, without descrip- Equisetites sp./ Equi- revised tion or illustration) setum sp. Western Holy Cross Lower Keuper, Letten- Czarnocki 1925 (only Mts. kohle; Middle Triassic, record, without descrip- Ladinian tion or illustration) Nowa Wieś Woźniki Limestone; Różycki 1930, Znosko (Upper Silesia) Upper Triassic, Norian 1955 (only record, without description or illustration) Kołaczkowice near Wielichowo Beds; Piwocki 1970 (only Rawicz Upper Triassic, record, without descrip- (Lower Silesia) Rhaetian tion or illustration) Ginkgoites acosmia Ginkgoales, valid taxon Gradzanowo 3 borehole Upper Triassic, Barbacka & Wcisło- Harris 1935 (Central Poland) Rhaetian Luraniec 2002*, 2003* Glossopteridium Incertae sedis, origi- Radoszyce 3 borehole Middle Buntsandstein; Bocheński 1957^* (orig- czarnockii Bocheński nally stated as Glos- (Holy Cross Mts.) Lower Triassic inal proposed species 1957 sopteridales, specimen name Glossopteridium probably lost J. Czarnockii) Knorripteris Pterophytina – fern Krapkowice Lower Muschelkalk, Michael 1895b^*, mariana (Michael stem, older synonym (Upper Silesia) Chorzów beds; Middle Potonié 1897*, Hörich 1895) Potonié 1897 of Adelophyton jutieri Triassic 1910* (description of (=Knorria mariana Renault 1900 according specimen anatomy; Michael 1895) to Hörich (1910) Hörich questioned age determination of specimen by Michael) 15

Table 1. Continued

Taxon as originally Botanical affinity, described, selected taxon validity, taxo- Location Horizon, age References, comments nomenclatural nomical synonyms synonyms

Lepidopteris ottonis Pteridospermopsida, Biadacz, Dobiercice, Wilmsdorfer Schichten/ Goeppert 1846a*, (Goeppert 1836) Peltaspermales, valid Gosław, Maciejów Gorzów Beds; Upper, Schenk 1867, Roemer Schimper 1869 taxon (Upper Silesia) Triassic, Rhaetian 1867, 1870*, Gothan (=Alethopteris ottonis 1909a*, Harris 1932, Goeppert 1836, Marcinkiewicz 1969, basonim) Reymanówna & Bar- (=Pecopteris ottonis backa 1981*, Barbacka (Goeppert 1836) Goep- 1991* pert 1846a) Wieluń Upper Triassic, Goeppert 1836^* (=Aspidites ottonis (Upper Silesia) Rhaetian Schenk in Roemer Kołaczkowice borehole Wielichowo Beds; Piwocki 1970* 1867) near Rawicz Upper Triassic, Asplenites ottonis (= (Lower Silesia) Rhaetian (Goeppert 1836) Schenk 1867) Gostyń 46 G borehole Zbąszynek/Jarkowo Marcinkiewicz (Central Poland) Beds; Upper Trias- & Orłowska-Zwolińska sic, Rhaetian sensu 1985, 1994 (only iso- polonico, Norian? lated cuticules determined as cf. Lepidopte ris ottonis) Gradzanowo 1 borehole Wielichowo Beds; Barbacka 1980, Bar- (Central Poland) Upper Triassic, backa 1991* Rhaetian Gradzanowo 3 borehole Wielichowo Beds; Marcinkiewicz (Central Poland) Upper Triassic, & Orłowska-Zwolińska Rhaetian 1994 (only isolated cuticules determined as cf. Lepidopteris ottonis) Chabowo 2 borehole Upper Triassic, Ociepa et al. 2008 (Western Pomerania) Rhaetian Lipie Śląskie-Lisowice, Upper Triassic, Staneczko 2007*, Waw- (Upper Silesia) Rhaetian rzyniak & Ziaja 2009 (only isolated cuticules), Wawrzyniak 2010b Neocalamites Halle Equisetales Gacki 1 borehole Lower Keuper, Letten- Pawłowska 1979* 1908 (Holy Cross Mts.) kohle; Middle Triassic, Neocalamites sp. Ladinian Studzianna borehole Lower Keuper, Letten- Barbacka et al. 2009, (Holy Cross Mts.) kohle; Middle Triassic, Barbacka et al. in prep. Ladinian (as Neocalamites meria nii (Brongniart 1828) Halle 1908) Neuropteris sp. conf. Pterophytina, valid Ligota near Woźniki Woźniki Limestone; Roemer 1870* N. remota Presl in name Cladophlebis (Upper Silesia) Upper Triassic, Norian Sternberg 1838 remota (Presl in Stern- berg 1838) Zeiller 1911 Pachylepis quinquies Coniferales, valid spe- Krasiejów Drawno Beds?; Upper Dzik & Sulej 2007* (Linck 1951) Kräusel cies (Opole Silesia) Triassic, 1952 Upper Carnian (=Voltzia? quinquies Linck 1951) Pachylepis Kräusel Coniferales Patoka Jarkowo/Zbąszynek Barbacka et al. 2012 1952 (Upper Silesia) Beds; Upper Triassic, Pachylepis sp. Norian Pagiophyllum sp. Coniferales Lipie Śląskie-Lisowice Wielichowo Beds; Wawrzyniak & Ziaja (Upper Silesia) Upper Triassic, 2010 Rhaetian Palaeohepatica roe- Hepaticopsida, species Ligota near Woźniki Woźniki Limestone; Raciborski 1892, 1893 meri Raciborski 1892 never illustrated or (Upper Silesia) Upper Triassic, Norian (original Roemer label (=Hepaticites roemeri revised. Type speci- identification – Thau (Raciborski 1892) Oost- men found by M. Rey- matopteris Münsteri endorp 1987 manówna in Wrocław ß. longissima), Wcisło- Uniwersity collection Luraniec 1984, Oosten- (Wcisło-Luraniec 1984). dorp 1987 (she referred 16

Table 1. Continued

Taxon as originally Botanical affinity, described, selected taxon validity, taxo- Location Horizon, age References, comments nomenclatural nomical synonyms synonyms Harris (1942) referred species Palaeohepatica type species of genus roemeri to genus Hepa Paleohepatica Racibor- ticites and provided its ski 1889 – diagnosis in English P. rostafinskii Raci- based on Raciborski’s borski 1889 to genus description in Polish Thallites Walton 1925 (1892) as Thallites rostafin skii (Raciborski 1889) Harris 1942 Palissya braunii Coniferales, valid spe- Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, c, Endlicher 1847 cies (Tatra Mountains) Upper Triassic, Reymanówna 1986 Rhaetian Pecopteris (Brongni- Pterophytina or Pteri- Gacki 4 borehole Muschelkalk, transi- Pawłowska 1979* art 1822) Sternberg dospermopsida, genus (South-eastern Holy tional beds; Middle 1825 erroneously identified Cross Mts.) Triassic, Ladinian Pecopteris sp.? Peltaspermum rot- Pteridospermopsida, Dobiercice Gorzów Beds; Upper Barbacka 1991* (cupu- ula Harris 1932 Peltaspermales, valid (Upper Silesia) Triassic, Rhaetian late disc and isolated taxon seeds) Pinites jurassicus Coniferales, probably Kamienica Polska (in Upper Triassic (Rhae- Goeppert 1846a*^, Goeppert 1846a valid species, speci- original paper misspelt tian) or (Middle? – Mercklin 1852, 1855, (=Peuce jurassica mens need revision, as Kaminika Polska) Oolite) Jurassic Schenk 1867, Schimper (Goeppert 1846a) End- according to Seward (north part of Upper 1870–1872, Seward licher 1847) (1919) could be refera- Silesia) 1919, Jongmans ble to genus Cedroxylon & Dijkstra 1973 Pinites pertinax Coniferales, probably Sumpen, Gorzów Upper Triassic (Rhae- Goeppert 1846a^*, Goeppert 1846a valid species, specimen Śląski (?) tian) or (Middle?) Endlicher 1847, Merck- (=Peuce pertinax needs revision (north part of Upper Jurassic lin 1855, Schenk 1867, (Goeppert 1846a) End- Silesia) Schimper 1870–1872, licher 1847) Seward 1919, Jong- (=Cedroxylon perti mans & Dijkstra 1973 nax (Goeppert 1846a) Kraus in Schimper 1870–1872) Pinites Lindley Coniferales Lofkowitz, Sumpen (?) Keuper; Upper Triassic Roemer 1867, 1870 & Hutton 1831 (Upper Silesia) Pinites sp. Pleuromeia cf. stern- Isoetales, specimens North-west part of Upper Middle Bunt- Czarnocki 1931 (only bergi (Münster 1839) never illustrated or Holy Cross Mts. sandstein; record without descrip- Corda 1852 revised Lower Triassic tion) Pleuromeia Corda Isoetales Otyń IG-1 borehole Supra-Oolitic Beds, Fuglewicz 1980b* 1852 (Fore-Sudetic Mono- Middle Buntsandstein; (strobile with mega- ?Pleuromeia cline) Lower Triassic spores Pusulosporites Fuglewicz 1973 in situ) Pseudohirmerella Coniferales Krasiejów Drawno Beds?; Upper Dzik & Sulej 2007* Arndt 2002 (Opole Silesia) Triassic, Pseudohirmerella Upper Carnian sp. n. Pterophyllum braun- Bennettitales Gosław Wilmsdorfer Schichten, Schenk 1867, Roemer ianum Goeppert 1844 (Upper Silesia) Keuper; Upper Trias- 1870* (as synonym of sic, Rhaetian Pterophyllum oeynhau sianum Goeppert 1844) Pterophyllum car- Bennettitales Biadacz, Dobiercice, Wilmsdorfer Schichten, Goeppert 1844^*, 1845, nallianum Goeppert Gosław Keuper; Upper Trias- 1846a, Schenk 1867*, 1844 (Upper Silesia) sic, Rhaetian Roemer 1867, 1870* (as synonym of Ptero phyllum propinquum Goeppert 1844 Pterophyllum com- Bennettitales Extra-Carpathian area Wielichowo beds; Marcinkiewicz 1969 pressum Lundblad of Poland Upper Triassic, (only isolated cuticules) 1950 Rhaetian 17

Table 1. Continued

Taxon as originally Botanical affinity, described, selected taxon validity, taxo- Location Horizon, age References, comments nomenclatural nomical synonyms synonyms

Pterophyllum cf. jae- Bennettitales Gacki 3 borehole Lower Keuper, Letten- Pawłowska 1979* geri Brongniart 1828 (South-eastern Holy kohle; Middle Triassic, Cross Mts.) Ladinian Pterophyllum muen- Cycadales, synonym Dobiercice Wilmsdorfer Schichten; Schenk 1867, Roemer steri (Presl in Stern- of Nilsonia acuminata (Upper Silesia) Upper Triassic, 1870, Raciborski 1892 berg 1838) Goeppert (Presl in Sternberg Rhaetian (Raciborski referred 1844 1838) Goeppert 1844 this specimen to Ptilo (=Nilssonia muensteri zamites nilsonii) (Presl in Sternberg 1838) Schimper in Zit- tel 1880) Pterophyllum oeyn- Bennettitales Biadacz Wilmsdorfer Schichten; Goeppert 1844^*, 1845, hausianum Goeppert (Upper Silesia) Upper Triassic, 1846a, Schenk 1867, 1844 Rhaetian Roemer 1867, 1870 Pterophyllum pro- Bennettitales Biadacz Wilmsdorfer Schichten; Goeppert 1844^*, 1845, pinquum Goeppert (Upper Silesia) Upper Triassic, 1846a, Schenk 1867*, 1844 Rhaetian Roemer 186 Pterophyllum Bennettitales Krasiejów Drawno Beds?; Upper Dzik & Sulej 2007 Brongniart 1828 (Opole Silesia) Triassic, Upper Car- Pterophyllum sp. nian Nowa Wieś Woźniki Limestone; Znosko 1955 (only (Upper Silesia) Upper Triassic, Norian record, without description or illustration) Schizoneura hoer- Equisetales, synonym Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, c, ensis (Hisinger 1840) of Neocalamites (Tatra Mountains) Upper Triassic, Reymanówna 1986 Schimper 1869 lehmannianus (Goep- Rhaetian pert 1846a) Weber 1968 Sphaerococcites Nomen dubium, origi- Tarnowskie Góry Muschelkalk; Middle Goeppert 1845, blandowskianus nally stated as alga, (Upper Silesia) Triassic 1846b^*, Eck 1865 Goeppert 1846b most probably trace fossil Stachyotaxus sep- Coniferales, illustrated Lipie Śląskie-Lisowice Wielichowo Beds; Dzik el al. 2008a*, b, tentrionalis (Agardh by Dzik et al. (2008a), (Upper Silesia) Upper Triassic, Świło et al. 2014 1823) Nathorst 1886 remains most probably Rhaetian belong to genus Elato cladus Taeniopteris Cycadales Dobiercice Wilmsdorfer Schichten; Schenk 1867^*, Roe- gigantea Schenk 1867 (Upper Silesia) Upper Triassic, mer 1867, 1870* Rhaetian Voltzia krappitzen- Coniferales, species Krapkowice Muschelkak; Middle Kunisch 1886^* sis Kunisch 1886 needs revision (Upper Silesia) Triassic, Anisian Voltzia Brongniart Coniferales, specimens North-west part of Upper Middle Bunt- Czarnocki 1931 1828 never illustrated or Holy Cross Mts. sandstein; (only record, without Voltzia sp. revised Lower Triassic description) Widdringtonites End- Coniferales Czerwone Żlebki Tomanová Formation; Raciborski 1890a*, b, c, licher 1847 (Tatra Mountains) Upper Triassic, Reymanówna 1986 Widdringtonites sp. Rhaetian Xylomites irregula- Originally stated as Dobiercice Wilmsdorfer Schichten; Goeppert 1846a^*, ris Goeppert 1846a epiphyllous fungi on (Upper Silesia) Upper Triassic, Roemer 1870 (as fungi leaves Rhaetian on Pterophyllum car nallianum leaf fragments) 18

Table 2. Proposed approximate zonation and correlation of Triassic floras of Poland

Germanic Triassic divi- Polish Triassic division German/French Polish macrofloral Palynological zones Megaspore zona- Most typical plant sion (Kozur & Bach- (Orłowska-Zwolińska floral assemblages assemblages and (spores and pollen grains) tion in Poland Evolutionary events in Age macrofossil taxa in Localities in Poland mann 2008, Franz 2009, 1985, Mader 1997, (Dobruskina 1988, assemblage zones in Poland (Orłowska- (Marcinkiewicz Polish macroflora Poland Kozur & Weems 2010) Fijałkowska-Mader 1999) 1994, Kelber 1998) (here proposed) Zwolińska 1985) 1978, 1992) Rhaetian Upper Keuper/ Upper Rhaetian sensu Rhät-Flora/ Lepidopte Lepidopteris ottonis Riccisporites tuberculatus Trileites pingius Lepidopteris ottonis, Kluczbork area (Goep- During stage rise of Rhätkeuper/ polonico/ ris ottonis zone zone Neocalamites lehman pert’s sideritic nodules floral biodiversity, grow- Exter Formation Wielichowo Beds nianus, Pterophyllum flora) Lipie Śląskie- ing similarity to Lower and perhaps Upper (most) spp. Lisowice, Czerwone Jurassic floras Zbąszynek Beds (?) Żlebki Norian Middle Keuper/ Lower Rhaetian sensu Stubensandstein/ Burg- Brachyphyllum zone Corollina meyeriana Brachyphyllum, Ptero Poręba, Patoka, Woźniki Appearance of modern Steinmergelkeuper/ polonico/Zbąszynek Beds, sandstein Flora phyllum, Equisetites Limestone (Ligota, Mysz- conifer families (Brachy Arnstadt Formation Jarkowo Beds (Grabowa ków, Poręba), Marciszów- phyllum leaves appear- Formation), Drawno Beds Zawiercie ing) Carnian Middle Keuper/ Upper Keuper sensu polo- Coburger Sandstein (Younger) Voltzia Hiatus in pollen zones in Voltzia, Glyptolepis, Krasiejów, Woźniki Decrease of diversity and Upper Gipskeuper/ nico/Upper Gipskeuper Flora zone Poland, upper Camero Desmiophyllum, Pseu claypit extinction of Voltziales Weser Formation sporites secatus zone in dohirmerella, Pachy German part of Germanic lepis, Sphenopteris Basin schoenleiniana Middle Keuper/ Upper Keuper sensu polo- Schilfsandstein Flora Lack of macroscopic Aulisporites astigmosus Narkisporites har Schilfsandstein/ nico/Schilfsandstein plant remains in risi Stuttgart Formation Poland Middle Keuper/ Upper Keuper sensu polo- Gipskeuper Flora Lack of macroscopic Porcelispora longdonesis Lower Gipskeuper/ nico/Lower Gipskeuper plant remains in Upper Ladinian Grabfeld Formation and Grenzdolomit Poland Lower Keuper/ Letten- Lower Keuper/ Unterkeuper (Letten- Equisetales-flora Heliosaccus dimorphus Dijkstraisporites Neocalamites merianii Holy Cross Mountains Floras very rare and keuper/ Erfurt Formation Sulechów Beds kohle) Flora beutleri impoverished in Poland Middle and Lower Upper Muschelkalk Upper Muschelkalk Lack of macroscopic Lack of macroscopic Hiatus Ladinian plant remains in Ger- plant remains in Upper Anisian manic Basin Poland Middle Anisian Middle Muschelkalk Middle Muschelkalk Tsugaepollenites oriens Lower Muschelkalk Lower Muschelkalk Perotrilites minor Lower Anisian/ Upper Upper Buntsandstein- Upper Buntsandstein-Röt Grés à Voltzia flora (Older) Voltzia-flora Voltziacaesporites hetero Trileites validus Voltzia, ferns and Krapkowice, Pałęgi Floras very rare and Buntsandstein- Röt Röt morpha equisetalean remains impoverished in Poland Scytian/ Lower and Middle Buntsandstein Middle Buntsandstein Pleuromeia-flora Pleuromeia-flora Densoisporites nejburgii Hiatus Pleuromeia and equi- Holy Cross Mountains Floras very rare and Middle Buntsand- setalean remains only impoverished in Poland Talchirella daciae stein Trileites polonicus Hiatus Lower Buntsandstein Lower Buntsandstein Lundbladispora obsoleta- Otynisporites Protohaploxypinus pantii eotriassicus

CONCLUSIONS AND FUTURE These floras will also fill in the palaeoecologi- PROSPECTS cal and palaeoenvironmental background of the vertebrates, especially the dinosaurs at The Triassic floras of the Northern Hemi- Polish localities. A better understanding of sphere are rare and poorly known. Each new the dinosaurs’ environment will help expand site with plant macroremains fills a gap in our our knowledge about their biology and explain knowledge of plant evolution in this period their evolutionary success. Such interdiscipli- (Kustatscher & van Konijnenburg-van Cittert nary dinosaur research has been very intense 2005, Roghi et al. 2006, van Konijnenburg-van lately, particularly in North America (Samp- Cittert et al. 2006, Kustatscher & van Koni- son 2012), but has not included the dinosaurs jnenburg-van Cittert 2008, Kustatscher et al. of Poland; they have yet to take their rightful 2012). Good, complete descriptions of the flo- place in the European Triassic. ras of Upper Triassic localities in Poland will The very few descriptions of the Triassic flo- help researchers study Triassic ecosystems in ras of Poland come mainly from the 19th cen- Poland and worldwide, and will be useful in tury. They require modern revisions incorporat- analysing the evolution of floras in the Trias- ing current nomenclature and meeting current sic. Some taxa found in Poland may have large standards of fossil description. Expert investi- implications for reconstruction of the gymno- gations begun in the 19th century were not con- sperm phylogeny, particularly for conifers. tinued in the 20th century, except for a very few 19

Table 2. Proposed approximate zonation and correlation of Triassic floras of Poland Table 2. Continued

papers and excerpts in publications on stratig- a useful palynological biostratigraphic scheme raphy. There is a good record of Polish Lower for the Polish Triassic (Orłowska-Zwolińska and Middle Triassic floras in the palynological 1985). New palynological data gathered from data but not in data on macroremains. Upper Poland fits this scheme perfectly (Fijałkowska- Triassic macrofloras are better known but their Mader 1999). Orłowska-Zwolińska’s scheme is descriptions are outdated. Most determina- congruent with Western European palynologi- tions of macroremains from the Polish Trias- cal biostratigraphy (Fijałkowska-Mader 1999, sic should be revised. The plants described by Cirilli 2010, Kürschner & Herngreen 2010). Goeppert (1836, 1844, 1845, 1846a) were revised Correlating dispersed spores and pollen grains by Schenk alone (1867). Later only Lepidopteris with their parent plants in Triassic floras pre- ottonis was redescribed (Barbacka 1991). As is sents great difficulties not just in Poland but clear from Schenk’s (1867) revision, some spe- worldwide. The true botanical affiliation of cies Goeppert described are junior synonyms even index taxa such as Ricciisporites tuber of previously described species, some new spe- culatus is disputed (Mander et al. 2012). This cies were described from small, badly preserved presents problems, as broad palaeoecological remains, and species in some genera are over- conclusions are based on this palynological split (e.g. Pterophyllum). data (e.g. Dzik et al. 2008b). The newly dis- Palynological research on Triassic strata in covered floras, especially those in Krasiejów, Poland intensified in the 1970s and produced Patoka and Lipie Śląskie-Lisowice, contain 20

Table 3. Complete taxonomical composition of Upper Triassic macrofloral assemblage zones here proposed for main localities

Assemblage Formation/ Age zones (here Locality Floral assemblage composition Beds proposed) Rhaetian Lepidopteris Wilmsdorfer Kluczbork area (Goep- Neocalamites lehmannianus, Alethopteris insignis, Clado ottonis assem- Schichten pert’s sideritic nod- phlebis roesserti, Clathropteris meniscoides (=Camptopte blage zone ule flora) Biadacz, ris jurassica =Clathropteris muensteriana), Carpolithes Dobiercice, Gosław, cardiocarpoides, Dicranopteris roemeriana, Lepidopteris Maciejów ottonis, Peltaspermum rotula, Pterophyllum braunianum, (Upper Silesia) Pterophyllum carnallianum, Pterophyllum muensteri, Pte rophyllum oeynhausianum, Pterophyllum propinquum, Taeniopteris gigantea, Xylomites irregularis Wielichowo Lipie Śląskie-Lisowice Lepidopteris ottonis, Brachyphyllum sp., Pagiophyllum Beds (Upper Silesia) sp., Cheirolepidiaceae Tomanová Czerwone Żlebki Neocalamites lehmannianus, Equisetum chalubinskii, Formation (Tatra Mountains) Equisetum an bunburyanum, Cladophlebis roesserti, Cladophlebis roesserti forma parvifolia, Clathropteris meniscoides (=Clathropteris platyphylla), Dictyophyllum aff. dunkeri, Pecopteris lobata, Palissya braunii, Wid dringtonites sp. Norian Brachyphyl Woźniki Woźniki area (Ligota, Clathropteris meniscoides (=Clathropteris muensteriana), lum assem- Limestone Myszków, Nowa Wieś, Neuropteris sp. conf. N. remota, Palaeohepatica roemeri, blage zone Poręba) (Upper Silesia) Pterophyllum sp., Brachyphyllum sp., Equisetites muen steri Carnian Voltzia Upper Gip- Krasiejów Sphenopteris schoenleiniana, Pterophyllum sp., Desmio assemblage skeuper (Opole Silesia) phyllum spp., Glyptolepis keuperiana, Pachylepis quin zone quies, Pseudohirmerella sp., Voltzia spp. many new taxa yet to be described, and some Cretaceous conifers are accompanied by a new of them could have great evolutionary impor- type of seed scale which may have evolved from tance. A complete description of the Woźniki primitive Voltzia-type scales. This is a plant of limestone flora, which has been on record since evolutionary significance for modern conifers. the end of the 19th century, is urgently needed. Preliminary work indicates that it will be pos- The Triassic is an exceptionally interest- sible to fully reconstruct a conifer from Patoka ing period as it witnessed rapid evolution of with trunks, twigs, male and female cones – conifers. The oldest representatives of modern and pollen grains. Completely reconstructed conifer families occurred in the Upper Triassic extinct plants can be used in developing phy- (Rothwell et al. 2012). The stages of transfor- logenetic hypotheses. The discovery of a plant mation from the primitive Votziales to evolu- with primitive female cones accompanied by tionarily advanced families of modern conifers evolutionarily advanced shoots at the Patoka in the Triassic are insufficiently documented. site may form the basis for erecting a new fam- The newly discovered Silesian sites (Krasie- ily of conifers and revising current views on jów, Patoka, Lipie Śląskie-Lisowice), contain- the taxonomy of this group of plants. This find ing Upper Triassic floras rich in conifer taxa, could also have far-reaching implications for can shed new light on this poorly known stage the taxonomy of conifers. of conifer evolution. The flora from Krasiejów The task of correlating Triassic floras world- contains relatively primitive conifers: they wide is hampered by stratigraphical problems fall into the genus Voltzia based on twig mor- and regional differences in floral composition. phology and into the genus Glyptolepis based Some approximate correlations are possible on seed scales. Further research is needed to regionally on the continental scale, however. identify the Krasiejów conifers to species level Such biostratigraphical and lithostratigraphi- based upon twig, seed scale, and pollen cone cal correlations of the Polish Triassic floras morphology, because these specimens repre- from the main localities with other well-known sent genera and species new to science. The European Triassic floras are provided here Krasiejów conifers hold clues to the evolution (Tab. 2), and new macrofloral assemblages for of these plants in the early Upper Triassic. In the Lower and Middle Triassic and macroflo- the flora of Patoka, which is slightly younger ral assemblage zones for the Upper Triassic geologically, relatively advanced Brachyphyl in Poland (Tabs 2 and 3), corresponding to lum-type shoots characteristic of Jurassic or Orłowska-Zwolińska’s palynological zonation 21

Table 4. Main Polish Triassic macrofossil and selected microfossil plant localities with basic palaeobotanical, palaeozoological and geological references

Locality of fossil plant remains References Chabowo 2 borehole Ociepa et al. 2008, Barbacka et al. in prep. Czerwone Żlebki Raciborski 1890a, b, c, Michalík et al. 1976, 1988, Reymanówna 1984, 1986, Niedźwiedzki 2005, 2008, 2011, Niedźwiedzki & Sulej 2007 Gacki 1, 3 and 4 boreholes Pawłowska 1979 Gorzów Śląski area Goeppert 1846a, Endlicher 1847, Mercklin 1855, Roemer 1867, 1870, Schenk 1867, Schimper 1870–1872, Seward 1919, Znosko 1955, Jongmans & Dijkstra 1973 Gostyń 46 G borehole Marcinkiewicz & Orłowska-Zwolińska 1985, 1994 Gradzanowo 1 and 3 boreholes Barbacka 1980, Barbacka 1991, Marcinkiewicz & Orłowska-Zwolińska 1994, Barbacka & Wcisło-Luraniec 2002, 2003 Holy Cross Mts. Samsonowicz 1924, 1929, Czarnocki 1925, 1931, Fijałkowska 1989, 1992a, b, 1994a, b, Rdzanek 1982, Fijałkowska-Mader 1999 Kluczbork area (Biadacz, Goeppert 1836, 1841–1846, 1844, 1845, 1846a, b, Roemer 1867, 1870, Schenk 1867, Dobiercice, Gosław, Maciejów) Michael 1893, 1894, 1895a, Gallinek 1894, Gothan 1909a, Znosko 1955, Orłowska-Zwoliń- ska & Senkowiczowa 1970, Barbacka 1980, 1991, Reymanówna & Barbacka 1981, Mader 1997, Kiritchkova 2006, 2008 Kamienica Polska Goeppert 1846a, Mercklin 1852, 1855, Schenk 1867, Schimper 1870–1872, Seward 1919, Jongmans & Dijkstra 1973 Kamień Pomorski IG 1 borehole Pieńkowski 2004, Pieńkowski et al. 2012 Kołaczkowice near Rawicz Piwocki 1970 Krapkowice Kunisch 1886, Michael 1895b, Potonié 1897, Hörich 1910, Rieppel & Hagdorn 1997, Rieppel 2000 Krasiejów Dzik et al. 2000, Dzik 2001, 2002, 2003a, b, 2008, Sulej 2002, 2003, 2004, 2005, 2007, 2010, Zatoń & Piechota 2003, Dzik & Sulej 2004, 2007, Olempska 2004, 2011, Sulej & Majer 2005, Szulc 2005, Zatoń et al. 2005, Barycka 2007, Konietzko-Meier & Wawro 2007, Lucas et al. 2007, Szulc & Becker 2007, Gruszka & Zieliński 2008, Brusatte et al. 2009, Fosto- wicz-Frelik & Sulej 2009, Mazurek & Słowiak 2009, Kozur & Weems 2010, Piechowski & Dzik 2010, Racki 2010, Skawina 2010, 2013, Sues & Fraser 2010, Skawina & Dzik 2011, Bodzioch & Kowal-Linka 2012, Desojo et al. 2013, Konietzko-Meier & Klein 2013, Konietzko-Meier & Sander 2013, Konietzko-Meier et al. 2013 Lipie Śląskie-Lisowice Fuglewicz & Śnieżek 1980, Marcinkiewicz 1981, Staneczko 2007, Dzik et al. 2008a, b, Niedźwiedzki & Sulej 2008, Ociepa et al. 2008, Marynowski & Simoneit 2009, Gorzelak et al. 2010, Wawrzyniak & Ziaja 2009, 2010, Wawrzyniak 2010a, b, c, d, Racki 2010, Niedźwiedzki et al. 2011, 2012, Skawina & Dzik 2011, Świło et al. 2014 Majków Samsonowicz 1929 Mechowo IG I borehole Marcinkiewicz 1962 Otyń IG-1 borehole Fuglewicz 1980b Pałęgi Kuleta et al. 2006, Żyła et al. 2013 Patoka Znosko 1955, Barbacka et al. 2012 Poręba Znosko 1955, Brzyski & Heflik 1994, Sulej et al. 2012 Radoszyce 3 borehole Bocheński 1957 Ratajów Samsonowicz 1929 Studzianna Karaszewski 1962, Reymanówna 1986, Barbacka et al. 2009, Barbacka et al. in prep. Wieluń Goeppert 1836, Znosko 1955 Woźniki Limestone (Ligota, Roemer 1870, Raciborski 1892, 1893, Różycki 1930, Znosko 1955, Wcisło-Luraniec 1984, Myszków, Nowa Wieś, Poręba, Reymanówna 1986, Szulc et al. 2006, Szulc & Becker 2007 Woźniki)

(Orłowska-Zwolińska 1985), are proposed. with their seed scales Glyptolepis. Pseudo The Lower and Middle Triassic floras are hirmerella, Pachylepis, and a new genus. too poor in taxa for macrofloral assemblage Coniferous foliage incertae sedis Desmiophyl zones to be defined. Only the Upper Triassic lum, is also present. Pterophyllum leaves are strata have a plant macrofossil record rich very rare. This zone comprises the Upper Gip- enough for assemblage zones to be delineated. skeuper strata and corresponds to the Carnian. Three macrofossil assemblage zones are pro- In the palynological zonation of Poland there posed and defined here for the Polish Upper is a hiatus, but it corresponds to the upper Triassic: Camerosporites secatus zone in other parts The Voltzia assemblage zone, in which of Europe (Cirilli 2010, Kürschner & Hern- Voltzia (sensu lato) leaf species predominate, green 2010). 22

The Brachyphyllum assemblage zone is an evolutionary and palaeoecological context. dominated by conifers with Brachyphyllum- The new research on these localities is in its type foliage, probably signalling the emergence infancy (Dzik & Sulej 2007, Staneczko 2007, of modern conifer families. Whole-plant recon- Dzik et al. 2008a, b, Wawrzyniak & Ziaja 2009, structions of those conifers are possible, based Wawrzyniak 2010a, b, Barbacka et al. 2012, on new findings (wood, seed scales with ovules, Pacyna et al. 2013a, b). Intensive research on male cones with pollen in situ, shoots). Other plant remains from drill cores is also in pro- plant taxa are very rare; single fern leaves of gress (Ociepa et al. 2008, Barbacka et al. 2009, Cladophlebis and Clathropteris meniscoides, 2011, Krupnik et al. 2011, Pieńkowski et al. and bennettitalean foliage Pterophyllum and 2012). When good, comprehensive taxonomic Equisetites muensteri shoots have been found. descriptions have been completed, the floras This zone comprises the Lower Rhaetian sensu from these sites may serve as reference floras polonico (Zbąszynek, Jarkowo and Drawno for future evolutionary and palaeoecological beds) and corresponds to the Norian. In the studies of the world’s Triassic plants. palynological zonation of Poland it corresponds to the Corollina meyeriana zone. ACKNOWLEDGMENTS The Lepidopteris ottonis assemblage I thank J. Dzik and T. Sulej (Institute of Paleobiol- (Lepidopteris zone of Harris 1937) is zone ogy, Polish Academy of Sciences) for allowing me to col- characterised by the index species Lepidopte lect plant specimens at their research localities (Krasie- ris ottonis and very abundant Neocalamites jów, Patoka, Lipie Śląskie-Lisowice) and for lending lehmannianus, or locally Equisetum chalubin their collection of fossil plants from Krasiejów for this skii (Tatra Mountains). The genus Pterophyl research; J. Zaja and M. Barbacka (W. Szafer Insti- tute of Botany, Polish Academy of Sciences) for discus- lum is widespread and species-rich. The ferns sions on Triassic palynology and stratigraphy, for help Cladophlebis roesserti and Clathropteris menis in compiling the literature and for making available coides were also found. There are conifers typi- fossil collections in their care for comparative study; cal of the Triassic (Pseudohirmerella) with spe- B. Kietlińska-Michalik (Geological Museum, Polish cies more evolutionarily advanced than in the Academy of Sciences) for kindly providing the M. Raci- borski collection from Czerwone Żlebki for examina- Voltzia assemblage zone, and species of Lower tion, and also some useful papers; G. Pieńkowski (Pol- Jurassic type (Palissya). The plant assemblage ish Geological Institute) for discussion on the age of is taxonomically rich and somewhat similar Krasiejów flora; D. Zdebska (Jagiellonian University) in composition to Lower Jurassic floras (pers. for discussions and help with many problems during observ. of Lipie Śląskie-Lisowice flora). This the preparation of this article; M. Popa and E. Kustat- scher for their very useful comments and suggestions; zone comprises the Upper Rhaetian sensu M. Jacobs for skilful language correction; and A. Sojka polonico (Wielichowo Beds) and corresponds for preparing Figure 1. I especially thank Editor-in- to the Rhaetian. In the palynological zonation Chief L. Stuchlik for his help and encouragement dur- of Poland it corresponds to the Riccisporites ing the work on this article. The project was financed tuberculatus zone. by funds from the Polish National Science Centre (no. DEC-2012/05/B/NZ8/00990) and from the Jagiellonian The abrupt change in flora at the Carnian- University (no. DS/MND/WBiNoZ/IB/5/2012). Norian boundary could be connected with the mass extinction observed in faunas worldwide (Benton 1994a, b), but this needs to be inves- REFERENCES tigated more carefully, as new data based on the Polish Upper Triassic vertebrates may ANDERSON H.M. & ANDERSON J.M. 1997. Why challenge this conclusion (Dzik et al. 2008a, b). not look for Proangiosperms in the Molteno Forma- Full analyses of the floras from the Trias- tion? In: Herngreen G.F.W. (ed.), Proceedings 4th sic sites in Poland will extend our knowledge European Palaeobotanical and Palynological Con- (still very scant) of Triassic palaeoecosystems ference. Meded. Nederl. Inst. Toegep. Geowetens. TNO, 58: 73–80. in Poland and generally in the Northern Hem- isphere. Recent new finds of taxonomically ANDERSON J.M. & ANDERSON H.M. 2003. Heyday of the gymnosperms: systematics and biodiversity rich, well-preserved floras with numerous of the Late Triassic Molteno fructifications. Strelit- specimens accompanying vertebrate remains zia, 15: 1–398. in Silesia provide researchers studying Pol- ANTEVS E. 1914. Lepidopteris ottonis (Göpp.) Schimp. ish Triassic floras with an opportunity to and Antholithus zeilleri Nath. Kungl. Sven. Vet- upgrade the taxonomy and describe them in ensk. Handl., 51(7): 1–18. 23

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MAREK S. & PAJCHLOWA M. (eds) 1997. Epikonty- NIEDŹWIEDZKI G. 2008. Dinosaur tracks in the nentalny perm i mezozoik w Polsce (summary: The Late Triassic of the Tatra Mountains: 62–63. In: Epicontinental Permian and Mesozoic in Poland). Pisera A., Bitner M.A. & Halamski A.T. (eds.), Pr. Inst. Geol., 153: 1–452. Abstracts, 9th Paleontological Conference, War- szawa, 10–11 October, 2008. Polish Academy of MARYNOWSKI L. & SIMONEIT B.R.T. 2009. Wide- Sciences, Institute of Paleobiology, Warszawa. spread Late Triassic to Early Jurassic wildfire records from Poland: evidence from charcoal and NIEDŹWIEDZKI G. 2011. A Late Triassic dinosaur- pyrolytic polycyclic aromatic hydrocarbons. Palaios, dominated ichnofauna from the Tomanová Forma- 24: 785–798. tion of the Tatra Mountains, Central Europe. Acta MAZUREK D. & SŁOWIAK J. 2009. Silezaur dino- Palaeont. Pol., 56(2): 291–300. zaurem? (Was Silesaurus a dinosaur?). Prz. Geol., NIEDŹWIEDZKI G. & PTASZYŃSKI T. 2007. Large 57(7): 569–571. (in Polish). Chirotheriidae tracks in the Early Triassic of MERCKLIN C.E. 1852. Prospectus der Paläontologis- Wióry, Holy Cross Mountains, Poland. Acta Geol. chen Pflanzenüberresten in Russland, so wie ihrer Pol., 57(3): 325–342. Erforschung. Bull. Cl. Phys.-Mathém. Acad. Imper. NIEDŹWIEDZKI G. & SULEJ T. 2007. Tropy Sci. Saint-Petersbourg, 10(24): 373–384. kręgowców w górnym triasie Polski: 93–94. In: MERCKLIN C.E. 1855 (1856). Palaeodendrologikon Żylińska A. (ed.), Granice Paleontologii, XX Kon- Rossicum, vergleichende anatomisch-mikroskopis- ferencja Naukowa Paleobiologów i Biostratygrafów che Untersuchungen fossile Holzer. Buchdruckerei PTG, Św. Katarzyna pod Łysicą, 10–13 września der Kaiserlichen Akademie der Wissenschaften, St. 2007. Polskie Towarzystwo Geologiczne, Kielce- Petresburg. Warszawa. MEYEN S.V. 1987. Fundamentals of Palaeobotany. NIEDŹWIEDZKI G. & SULEJ T. 2008. Lipie Śląskie Chapman and Hall, London – New York. koło Lisowic – okno na późnotriasowy ekosystem MICHAEL R. 1893. Ueber eine neue Lepidosteiden- lądowy (Lipie Śląskie off Lisowice – a window Gattung aus dem oberen Keuper Oberschlesiens. on the Late Triassic continental ecosystem). Prz. Zeit. Dtsch. Geol. Ges., 45: 710–729. Geol., 56 (9): 821–822. (in Polish). MICHAEL R. 1894. Ueber eine neue Lepidosteiden- NIEDŹWIEDZKI G., BRUSATTE S.L. & BUTLER Gattung aus dem oberen Keuper Oberschlesiens. R.J. 2013. Prorotodactylus and Rotodactylus Jahr. Schles. Ges. Vaterl. Cultur, 71: 71–74. tracks: an ichnological record of dinosauromorphs from the Early-Middle Triassic of Poland. In: Nes- MICHAEL R. 1895a. Ueber fossile Fischer aus dem bitt S.J., Desojo J.B. & Irmis R.B. (eds), Anatomy, oberen Keuper Oberschlesiens. Jahr. Schles. Ges. phylogeny and palaeobiology of early archosaurs Vaterl. Cultur, 72: 20. and their kin. Geol. Soc., London, Spec. Publ., 379: MICHAEL R. 1895b. Ueber zwei neue Pflanzeresten 319–351. aus dem oberschlesischen Muschelkalk. Naturwis- NIEDŹWIEDZKI G., GORZELAK P. & SULEJ T. sen. Wochensch., 10(41): 491–492. 2011. Bite traces on dicynodont bones and the early MICHALÍK J., PLENDEROVÁ E. & SÝKORA M. evolution of large terrestrial predators. Lethaia, 1976. To the stratigraphic and paleogeographic 44: 87–92. position of the Tomanová Formation in the Upper- NIEDŹWIEDZKI G., SULEJ T. & DZIK J. 2012. most Triassic of the West Carpathians. Geol. Zb., A large predatory archosaur from the Late Trias- Geol. Carpath., 27(2): 299–318. sic of Poland. Acta Palaeont. Pol., 57(2): 267–276. MICHALÍK J., KÁTLOVSKÝ V. & HLUŠTÍK A. 1988. Plant remains in the Tomanová Formation (Upper- NIEDŹWIEDZKI G., KIN A., REMIN Z. & MAŁKIE most Triassic, West Carpathians): their origin, WICZ M. 2007. Środkowotriasowa ichnofauna composition and diagenetic alteration. Geol. Zb., kręgowców z „warstw z Krynek” w Górach Geol. Carpath., 39(5): 523–537. Świętokrzyskich – wstępny przegląd (summary: Middle Triassic vertebrate ichnofauna from the MILLER C.N. 1977. Mesozoic conifers. Bot. Rev., “Krynki Beds” in the Holy Cross Mountains – pre- 43(2): 217–280. liminary review). Prz. Geol., 55(10): 870–879. NATHORST A.G. 1910. Lés depôts mésozoiques OCIEPA A.M., STANECZKO K., FELDMAN-OLSZEW- précrétacés de la Scanie. Geol. Förenin. Stockholm SKA A. & BARBACKA M. 2008. Nowe stanowiska Förhandl., 32(3): 487–532. Lepidopteris ottonis (Goeppert) Schimper w Polsce. NAWROCKI J. & SZULC J. 2000. The Middle Triassic In: Krobicki M. (ed.), Utwory przełomu jury i kredy magnetostratigraphy from the Peri-Tethys basin in w zachodnich Karpatach fliszowych polsko-cze- Poland. Earth Planet. Sci. Lett., 182: 77–92. skiego pogranicza, Jurassica VII, 27–29.09.2008, Żywiec/Štramberk, Abstrakty. Kwartalnik AGH NIEDŹWIEDZKI G. 2005. Nowe znalezisko śladów Geologia, 34(3/1): 199–200. dinozaurów w górnym triasie Tatr (summary: A new find of dinosaur footprints in the Upper Tri- OLEMPSKA E. 2004. Late Triassic spinicaudatan assic of the Tatra Mountains, southern Poland). crustaceans from southwestern Poland. Acta Pal- Prz. Geol., 53(5): 410–413. aeont. Pol., 49: 429–442. 30

OLEMPSKA E. 2011. Fresh-water ostracods from the triasie i dolnej jurze południowej Polski: 44–45. In: Late Triassic of Poland. Joann. Geol. Paläontol., Kędzierski M. & Kołodziej B. (eds), „Aktualizm 11: 154–155. i antyaktualizm w paleontologii” 22 Konferencja Naukowa Sekcji Paleontologicznej Polskiego Towa- ORŁOWSKA-ZWOLIŃSKA T. 1967. Mikroflorystyczne rzystwa Geologicznego, 27–30.09.2013, Tyniec. kryteria oceny wieku warstw z pogranicza triasu Materiały konferencyjne. Polskie Towarzystwo i jury na terenie Polski Pozakarpackiej (summary: Geologiczne, Kraków. Microfloristic criteria for age dermination of the beds occuring At the Triassic-Jurassic boundary in PAWŁOWSKA K. 1979. Utwory triasowe w południowo- the Extra-Carpathian areas of Poland). Biul. Inst. wschodniej części Gór Świętokrzyskich (summary: Geol., 203: 47–57. Triassic rocks of south-eastern Góry Świętokrzyskie Mts.). Kwart. Geol., 23(2): 337–362. ORŁOWSKA-ZWOLIŃSKA T. 1971. On several strati- graphically important species of sporomorphs PAUTSCH M. 1958. Keuper sporomorphs from Świer- occurring in the Keuper of Poland. Acta Soc. Botan. czyna, Poland. Micropalaeontology, 4(3): 321–325. Pol., 40(4): 633–651. PAUTSCH M. 1971. Sporomorphs of the Upper Trias- ORŁOWSKA-ZWOLIŃSKA T. 1976. Palynological sic from the borehole at Trzciana near Mielec (S. chracteristics of the reed sandstone in Polish Low- Poland). Acta Palaeobot., 12(1): 1–59. land area. Acta Geol. Pol., 26(4): 557–567. PAUTSCH M. 1973. Upper Triassic spores and pollen ORŁOWSKA-ZWOLIŃSKA T. 1977. Palynological cor- from the Polish Carphatian Foreland. Micropalae- relation of the Bunter and Muschelkalk in selected ontology, 19(2): 129–149. profiles from Western Poland. Acta Geol. Pol., 27: PIECHOWSKI R. & DZIK J. 2010. The axial skeleton 417–430. of Silesaurus opolensis. J. Verteb. Paleontol., 30: ORŁOWSKA-ZWOLIŃSKA T. 1983. Palinostratygra- 1127–1141. fia epikontynentalnych osadów wyższego triasu PIEŃKOWSKI G. 1988. Analiza facjalna najwyższego w Polsce (summary: Palynostratigraphy of the triasu i liasu Wyżyny Krakowsko-Wieluńskiej oraz upper part of Triassic epicontinental sediments in perspektywy występowania surowców ilastych Poland). Pr. Inst. Geol., 104: 1–89. (summary: Facial analysis of the Uppermost Tri- ORŁOWSKA-ZWOLIŃSKA T. 1984. Palynostratigra- assic and the Liassic of the Cracow-Wieluń Upland phy of the Buntsandstein in sections of western and prospectes for occurrence of clay deposits). Prz. Poland. Acta Palaeont. Pol., 29(3–4): 161–194. Geol., 36(8): 449–456. ORŁOWSKA-ZWOLIŃSKA, T. 1985. Palynological PIEŃKOWSKI G. 2004. The epicontinental Lower zones of the Polish epicontinental Triassic. Bull. Jurassic of Poland. Pol. Geol. Inst. Special Papers, Pol. Acad. Sci., Earth Sci., 33: 107–117. 12: 1–122. ORŁOWSKA-ZWOLIŃSKA T. 1986. Flora. Micro- PIEŃKOWSKI G., NIEDŹWIEDZKI G. & WAKS- flora. Miospores: 126–155. In: Malinowska L. (ed.), MUNDZKA M. 2012. Sedimentological, palynologi- Geology of Poland, vol. III. Atlas of guide and cal and geochemical studies of the terrestrial Tri- characteristic fossils, part 2a. Mesozoik. Triassic. assic–Jurassic boundary in northwestern Poland. Wydawnictwa Geologiczne, Warszawa. Geol. Mag., 149(2): 308–332. ORŁOWSKA-ZWOLIŃSKA T. 1988. Palinostraty- PIWOCKI M. 1970. Lepidopteris ottonis z retyku grafia utworów triasu w okolicach Brzegu –SE południowej części monokliny przedsudeckiej (sum- część monokliny przedsudeckiej (summary: mary: Lepidopteris ottonis from the Rhaetic depos- Palynostratigraphy of Triassic deposits in the its of the southern part of the Fore-Sudetic Mono- vicinity of Brzeg – SE part of the Fore-Sudetic cline). Kwart. Geol., 14(1): 101–106. Monocline. Geol. Quart., 32(2): 349–366. POTONIÉ H. 1897. Lehrbuch der Pfanzenpalaeontolo- ORŁOWSKA-ZWOLIŃSKA T. & SENKOWICZOWA H. gie. Ferd. Dümmlers Verlagsbuchhandlung, Berlin. 1970. Flora triasu: 17–19. In: Budowa Geologiczna PRETO N., KUSTATSCHER E. & WIGNALL P.B. Polski. Tom II. Katalog Skamieniałości. Część 2. 2010. Triassic climates – state of the art and per- Mezozoik. Wydawnictwa Geologiczne, Warszawa. spectives. Palaeogeogr., Palaeoclimat., Palaeoecol., OOSTENDORP C. 1987. The Bryophytes of the Palae- 290: 1–10. ozoic and the Mesozoic. Bryophytorum Bibliotheca, PTASZYŃSKI T. 1996. Ślady gadów w najniższym 34: 1–112. pstrym piaskowcu okolic Ostrowca Świętokrzyskiego PACYNA G. 2013. Critical review of research on the (summary: Reptile tracks in the lowermost Bunt- Lower Jurassic flora of Poland. Acta Palaeobot., sandstein near Ostrowiec Świętokrzyski – Central 53(2): 141–163. Poland). Prz. Geol., 44(10): 1042–1043. PACYNA G., ZDEBSKA D., BARBACKA M. & ZIAJA J. PTASZYŃSKI T. 2000a. Lower Triassic vertebrate 2013a. Co jadły polskie dinozaury? Wszechświat, footprints from Wióry, Holy Cross Mountains, 114(7): 250–252. Poland. Acta Palaeont. Pol., 45: 151–194. PACYNA G., ZDEBSKA D., BARBACKA M. & ZIAJA J. PTASZYŃSKI T. 2000b. Tropy kręgowców z piaskowca 2013b. Ewolucja i sukcesja flor towarzyszących tumlińskiego Góry Grodowej – Góry Świętokrzyskie szczątkom kostnym i tropom dinozaurów w górnym (summary: Vertebrate tracks from the Tumlin 31

sandstone of Grodowa Hill, Holy Cross Mts. – Cen- Instytut Nauk Geologicznych UJ, Komisja Paleon- tral Poland). Prz. Geol., 48(5): 418–421. tologii przy Komitecie Nauk Geologicznych PAN. PTASZYŃSKI T. & NIEDŹWIEDZKI G. 2002. Nowe Wydawnictwo AGH, Kraków. znaleziska tropów kręgowców z pstrego piaskowca REYMANÓWNA M. 1986. Macroflora. Classes Pterido- Gór Świętokrzyskich (summary: New finds of ver- phyta and Spermatophyta: 184–186. In: Malinow- tebrate tacks from the Buntsandstein of the Holy ska L. (ed.), Geology of Poland. Volume III. Atlas of Cross Mountains – Central Poland). Prz. Geol., guide and characteristic fossils. Part 2a. Mesozoic. 50(5): 441–446. Triassic. Wydawnictwa Geologiczne, Warszawa. PTASZYŃSKI T. & NIEDŹWIEDZKI G. 2004. Late REYMANÓWNA M. & BARBACKA M. 1981. Lepidop Permian vertebrate tracks from the Tumlin Sand- teris ottonis (Goeppert) Schimper z Wyżyny Ślą- stone, Holy Cross Mountains, Poland. Acta Palae- sko-Krakowskiej i obrzeżenia Gór Świętokrzyskich ont. 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w południowo-zachodniej części arkusza „Żarki” SUES H.-D. 2000. Evolution of herbivory in terrestrial mapy 1:100.000 (Compte-rendu des recherches vertebrates. Perspectives from the fossil record. géologiques effectuées en 1930). Posiedzenia Cambridge University Press, Cambridge. naukowe Państwowego Instytutu Geologicz- SUES H.-D. & FRASER N.C. 2010. Triassic life on nego (Comptes-Rendus des Séances du Service land. The great transition. Columbia University Géologique de Pologne), 28: 24–27. (in Polish). Press, New York. SADLOK G. & WAWRZYNIAK Z. 2013. Upper Tri- SULEJ T. 2002. Species discrimination in the Late assic vertebrate tracks from Kraków-Częstochowa Triassic labyrinthodont Metoposaurus. Acta Palae- Upland, southern Poland. Ann. Soc. Geol. Pol., ont. Pol., 47: 535–546. 83(2): 105–111. SULEJ T. 2003. Krasiejów. Sensacyjne odkrycia tria- SAMPSON S.D. 2012. Dinosaurs of the Lost Conti- sowych pra-dinozaurów. Przygoda Studio, Opole. nent. Scientific American, 306(3): 40–47. SULEJ T. 2004. Krasiejów – okno na świat późnego SAMSONOWICZ J. 1924. Sprawozdanie z badań triasu. Otoczak, 31: 2–22. geologicznych między Wierzbnikiem a Ostrowcem nad Kamienną (Compte rendu des rechereches SULEJ T. 2005. A new rauisuchian reptile (Diapsida: géologiques entre Wierzbnik et Ostrowiec sur la Archosauria) from the Late Triassic of Poland. J. Kamienna). Posiedzenia naukowe Państwowego Verteb. Paleontol., 25: 78–86. Instytutu Geologicznego (Comptes-Rendus des SULEJ T. 2007. Osteology, variability, and evolution Séances du Service Géologique de Pologne), 8: of Metoposaurus, a temnospondyl from the Late 24–27. (in Polish). Triassic of Poland. Palaeont. Pol., 64: 29–139. SAMSONOWICZ J. 1929. Cechsztyn, trias i lias na SULEJ T. 2010. The skull of an early Late Triassic aeto- północnym zboczu Łysogór (summary: Les Zech- saur and the evolution of the stagonolepidid archo- stein, le Trias et le Liasique sur le versant nord saurian reptiles. Zool. J. Linn. 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