Geo log i cal Quar terly, 2008, 52 (3): 225–238

Ver te brate re mains from the Lower Muschelkalk of Raciborowice Górne (North-Sudetic Basin, SW Poland)

Alina CHRZ¥STEK

Chrz¹stek A. (2008) — Verte brate re mains from the Lower Muschelkalk of Raciborowice Górne (North-Sudetic Ba sin, SW Po land). Geol. Quart., 52 (3): 225–238. Warszawa.

Verte brate rem ains, mostly fish teeth and scales, are de scribed from the Lower Muschelkalk of Raciborowice Górne, North-Sudetic Ba- sin, SW Poland. The assem blage oc curs in dark grey organodetrital limestone of unit C. Ver te brate re mains, rep resented mainly by verte - brate bones and copro lite s, are also known from unit B. Five taxa of chondrichthyan teeth — Acrodus lateralis, Acrodus cf. lateralis, Acrodus sp., Palaeobates angustissimus, Palaeobates sp. and, for the first time from this region, two taxa of osteichthyan rem ains — teeth of Birgeria sp., scales from Gyrolepis sp. as well as scales from un clas si fied actinopterygians and enigm atic bones (fishes?) are de- scribed from the Lower Muschelkalk at Raciborowice Górne. Rep tile teeth rep re sent ing the Nothosauridae or Cymatosauridae have been found for the first time at this local ity . They were discov ere d in the Bone Bed of unit C, that had previ ously only yielded fish teeth. The ma te rial col lected has al lowed re con struc tion of the ver te brate as sem blage of the Lower Muschelkalk of the North-Sudetic Basin. It has also helped to constra in re construc tions of the palaeoenvironment, suggest ing that it repre sente d a deepen ing lagoon. The assem blage has been corre late d with age-equiva lents from other regions of Europe, the faunas from the Holy Cross Mts. (Centra l Poland) being the closest analogy. The evi dence in dicate s that, during the de posi tion of units B and C that, contain the ver tebrat e re mains, connec tion with the Tethys Ocean was through the Silesian–Moravian and East Carpathian marine gate ways.

Alina Chrz¹stek, Insti tute of the Earth Sci ences, Wroc³aw Uni ver sity, Maksa Borna 9, PL-50-204 Wroc³aw, Poland; e-mail: [email protected] (receiv ed: Febru ary 18, 2008; accepte d: June 16, 2008).

Key words: North-Sudetic Ba sin, Lower Muschelkalk, ver te brate re mains, fish teeth, Chondrichthyes, Osteichthyes.

INTRODUCTION Nothosaurus cf. mirabilis and Nothosaurus sp. Ver tebrate re - mains from the Roetian and Lower Muschelkalk of the North-Sudetic Basin and Opole region were also descri bed by Chrz¹stek and NiedŸwiedzki (1998). The first re cord of verte brate rem ains from the North-Sudetic Re cent in ves ti ga tions have re sulted in the dis cov ery of an Ba sin (Noetling, 1880) menti oned teeth and bones of fish and as sem blage of chondrichthyan teeth: Acrodus lateralis, labyrinthodont am phib i ans from the Roetian near Raciborowice Acrodus cf. lateralis, Acrodus sp., Palaeobates angustissimus Górne as well as fish scales and rep tile teeth from the Lower and Palaeobates sp. Osteichthyan teeth and scales (Gyrolepis Muschelkalk near Stara Warta. The fos sils com prise Colobodus sp., Birgeria sp. and other actinopterygians) as well as rep tile chorzowiensis Mey, Gyrolepis sp., Pleurolepis silesiacus Eck., teeth (Nothosauroidea or Cymatosauroidea) and bones of un- Placodus sp. and Nothosaurus sp. Holdefleis (1915) reporte d the known taxon om y have been reporte d from the Lower oc cur rence of fish scales and ver tebrate bones in the Lower Muschelkalk at Raciborowice Górne for the first time. Muschelkalk at Raciborowice Górne. Chrz¹stek (1995a, b) discov ered a tooth of the selachian Lissodus sp. in the Roetian near Czapla and reporte d beds of unit A of the Lower Muschelkalk at Jerzmanice Zdrój abound- GEOLOGICAL SETTING ing in fish scales. She also char acter ized an as semblage con sist- ing of skele tal rem ains of fishes, am phibi ans and repti les from The best and most com plete secti on of the Lower units B and C exposed at Raciborowice Górne (Chrz¹stek, Muschelkalk in the North-Sudetic Basin crops out in the 2002). It com prised Acrodus lateralis, Acrodus cf. lateralis, Grodziec Syncline at Raciborowice Górne and its fragm ents Acrodus sp., Palaeobates angustissimus, Palaeobates sp., are exposed in the Leszczyna Syncline at Jerzmanice Zdrój 226 Alina Chrz¹stek

Fig. 1A — geolog i cal map of the North-Sudetic Ba sin after Sawicki and Teisseyre (1978), modi fied by the author; B — schem atic plan of the Raciborowice Górne quarry with places where indi vid ual units (B–E) are ex posed are marked

(Chrz¹stek, 1995b, 2002; Fig. 1). In the North-Sudetic Ba sin, organodetrital limeston es with oc ca sional intraclasts. In ter ca la - apart from the Trias sic rocks, sequence s of Upper Carbon if er - tions of thin-bedded platy limestone and marls are comm on. ous, Permian, Lower Creta ceous and Ceno zoic strata crop out, The depos it s of unit B are about 15 m thick. over laying the Eocambrian–Lower Car bon if er ous base ment The strata of unit C are com posed of thick-bedded (Baranowski et al., 1990). A detai led analy sis and organodetrital limeston e with oc ca sional intraclasts, in ter ca - lithostratigraphic subdi vi sion of the Lower Muschelkalk into lated with thin-bedded wavy or nodu lar limestone s and marls units A–E were given by Chrz¹stek (2002). The same depos it s (Figs. 1 and 3). They com prise the Spiriferina Bed, a very char - were also described by Leœniak (1978) and Szulc (1991). acter istic cor re lation ho ri zon that may be traced in the up per Unit A is exposed only at Jerzmanice Zdrój. In the and lower levels of the quarry (Holdefleis, 1915; Chrz¹stek, Raciborowice Górne quarry the Lower Muschelkalk sec tion 2002). The thickness of unit C reaches about 50 m. The comm ences with unit B, repre sent ed by thick-bedded cri - upperlying 18 m-thick se quence of thick-bedded oncolitic, noid-rich organodetrital limeston e in the low er most part crys tal line and organodetrital lime stones in ter ca lated with (G³uchowski and Salamon, 2005; Figs. 1 and 2). The re main - wavy, nodu lar and platy vari et ies forms unit D. The upper m ost ing part of unit B is built of cellu lar and thick-bedded part of the Lower Muschelkalk exposed in the North-Sudetic Vertebrate remains from the Lower Muschelkalk of Raciborowice Górne (North-Sudetic Basin, SW Poland) 227

Fig. 2. Lithostratigraphic sec tion of unit B Fig. 3. Lithostratigraphic sec tion of unit C

Other ex pla na tions as in Fig ure 2 228 Alina Chrz¹stek

Basin is built of the 3.5 m-thick unit E and is repre sent ed by This organodetrital limestone layer con tains fossiliferous (brachio pods and crinoids) thick-bedded chondrichthyans: Acrodus lateralis, Acrodus cf. lateralis, organodetrital limestone s inter calat ed with platy and nodu lar Acrodus sp., Palaeobates angustissimus and Palaeobates sp., lime stones. teeth and scales of osteichthyans: Birgeria sp., Gyrolepis sp. and other actinopterygians, as well as bones of unknown affin - ity (Ta ble 1). Repti le bones and teeth (nothosaurids or VERTEBRATE REMAINS IN THE LOWER cymatosaurids) were also noted. The limestone is so rich in ver - MUSCHELKALK tebrate re mains that it may be re ferred to as a Bone Bed. Bone fossil s are not encoun ter ed in unit C outside of this layer. I ear- lier identi fied a few bone fragm ents in unit D in Raciborowice Verte brat e rem ains are most abundant in units B and C. Górne (Chrz¹stek, 2002) and fish scales in unit A in Jerzmanice Bones and copro li tes dom inate the assem blage in unit B. Poor Zdrój (Chrz¹stek, 1995b). pres er va tion hin ders their pre cise tax o nomic iden ti fi ca tion though a single verte bra doubt lessly belongs to the repti le Nothosaurus cf. mirabilis, a taxon known from this local ity and SYSTEMATIC PALAEONTOLOGY de scribed by Chrz¹stek and NiedŸwiedzki (1998) and Chrz¹stek (2002). Bone rem ains occur within the thick-bedded organodetrital limestone at the bottom of unit B. The rock Tax on omy of chondrichthyans, osteichthyans (actinoptery - abounds also in cri noids of the genus Dadocrinus and the spe- gians) and rep tiles after Cappetta (1987), Nel son (1994 in cies Holocrinus acutangulus as well as in bivalve s, gas tropods Bürgin, 1999) and Rieppel (2000), respec tively. All the speci - and foraminifera (Hagdorn and G³uchowski, 1993; Chrz¹stek, mens de scribed are depos it ed in the col lec tion of the Geolog i - 2002; Fig. 2). Nu mer ous ver te brate fos sils were en coun tered cal Mu seum, Uni ver sity of Wroc³aw. also in the upper m ost part of unit B. They occur within the thick-bed ded organodetrital limeston e that is rich in the bi valve Myophoria vulgaris (Fig. 2). Cop ro lites are pres ent in all the Super class: Pis ces above-menti oned strata (Fig. 2). Class: Chondrichthyes Huxley, 1880 Fish teeth were discov ered only in unit C. They occur Subcla ss: Elasmobranchii Bonaparte, 1838 within a sin gle layer of pyrite- miner al ized dark grey Or der: Euselachii Hay, 1902 organodetrital limestone with abundant gastro pods, bi valves Superfamily: Hybodontoidea Zangerl, 1981 and less num erous ichnofossils Planolites sp. (Chrz¹stek, Acrodontidae Casier, 1959 2007). The layer lies about 4.5 m below the Spiriferina Bed, Acrodus Agassiz, 1837 and is com posed of thick-bedded organodetrital limeston e with Acrodus lateralis Agassiz, 1837 hardground intraclasts and the trace fos sil Trypanites sp. The (Fig. 4A–B) brachi o pod Punctospirella fragilis appears in the Spiriferina MGUWr 5387s; 5388s Bed for the first and only time in the enti re secti on of the Lower Muschelkalk. It is ac com panied by abundant encrinids, Holocrinus acutangulus, bi valves, gas tro pods and echinoid 1928 Acrodus lateralis, Schmidt, figs. 927, 928. spines (Chrz¹stek, 2002; Salamon et al., 2003). G³uchowski 1973 Acrodus lateralis, Liszkowski, ryc. 2, fig. 10. and Salamon (2005) also descri bed the cri noid Eckicrinus 1981 Acrodus lateralis, Rieppel, fig.11/3. 2001 Acrodus lateralis, Dorka, fig. 1/a. radiatus. 2002 Acrodus lateralis, Chrz¹stek, pl. 23, fig. 1. Material. — Two well-pre served crowns; the roots are not visi ble on the rock surface . Ta ble 1 Description. — The crowns are oval and elongate d. Tax o nomic dis tri bu tion of the Lower Muschelkalk ver te brate They measure 2.5 and 4 mm in length, 1.5–2 mm in width and rem ains from the Raciborowice Górne sec tion 1–1.5 mm in height. The length/width rati o ranges from 1.6 to 2. Crowns are globu lar centra lly, lower and narrow margin all y. No. Taxa Unit B Unit C The crown or na men tation is gen er ally restricted to the cen tral 1 Acrodus lateralis – + part. It consis ts of irreg u lar furrows and deli cat e ridges di verg- ing from the longi tu di nal crest towards the crowns’ edges. The 2 Acrodus cf. lateralis – + crown profil e, orna m enta ti on and size suggest that the teeth be- 3 Acrodus sp. – + long to Acrodus lateralis. They are slightly smaller then the 4 Palaeobates angustissimus – + ones de scribed by Rieppel (1981) and Dorka (2001). Occurrence. — Bone Bed of unit C at Raciborowice. 5 Palaeobates sp. – + The teeth of this specie s are known from the Pecten and 6 Birgeria sp. – + Dadocrinus ho ri zon of the Gogolin Beds in the Opole region 7 Gyrolepis sp. – + (Chrz¹stek and NiedŸwiedzki, 1998). Liszkowski (1981, 1993) 8 Nothosaurus cf. mirabilis + – de scribed sim ilar teeth from the Lower Muschelkalk of the Opole region (Myophoria Beds, Gogolin Beds, Góra¿d¿e Beds 9 Nothosauroidea or Cymatosauroidea + – and Terebratula Beds) as well as from the Middle and Upper Vertebrate remains from the Lower Muschelkalk of Raciborowice Górne (North-Sudetic Basin, SW Poland) 229

Fig. 4. Fish teeth from the Bone Bed (unit C) of the Lower Muschelkalk at Raciborowice Górne

A–B — Acrodus lateralis; C — Acrodus cf. lateralis; D–E — Acrodus sp.; F–I — Palaeobates angustissimus; J — Palaeobates sp.

Muschelkalk. They are most abundant in the Gogolin Beds, Acrodus lateralis in the Lower Muschelkalk and Keuper of Wilkowice Beds and Boruszowice Beds. The author re ported Thuringia. Dorka (2001) gave records of Acrodus lateralis the same species from the en tire Muschelkalk sec tion ex cept teeth from the Mid dle Trias sic of Schöningen (Lower Saxony, for the middle and upper Middle Muschelkalk of the Holy Ger many) while Rieppel (1981) reporte d it from the Middle Cross Mts. It is most nu merou s within the Ceratite Beds and Trias sic of Monte San Giorgio (Southern Alps, Switzer land) . £ukowa Beds. Schmidt (1928) noted the oc currence of 230 Alina Chrz¹stek

Acrodus cf. lateralis The crown is wider and overhangs the root. The teeth are cov - (Fig. 4C) ered with irre gu lar granu l ati on composed of small oval ele va - MGUWr 5389s tions and hol lows. Occurrence. — Bone Bed of unit C at Raciborowice. Chrz¹stek and NiedŸwiedzki (1998) and Chrz¹stek (2002) de- Material. — One well-pre served spec im en with scribed sim ilar teeth from this bed. Ac cording to Schmidt (1928) poorly marked or na men ta tion. this specie s occurs in the Upper Roetian and Muschelkalk of Description. — The tooth is oval and rather high. It Jena (Thuringia, Germ any). Liszkowski (1973) observe d sim ilar measures 3–3.5 mm in length, 2–2.5 in width and 2 mm in teeth from the Lower Muschelkalk of Wolica (£ukowa Beds). height. The length/width rati o is 1.4. Faint orna m enta ti on con- He also menti oned (Liszkowski, 1993) Palaeobates sists of folds running from the crown top toward its margins. angustissimus from the Lower (Góra¿d¿e Beds and Terebratula The profil e and size suggest its assign m ent to Acrodus lateralis Beds), Middle and Upper Muschelkalk of the Opole region. The but becaus e of the poorly pre served orna m enta ti on it is de - spe cies is par tic u larly nu mer ous in the Wilkowice Beds and scribed as Acrodus cf. lateralis. Boruszowice Beds. Liszkowski (1993) also reporte d Occurrence. — Bone Bed of unit C at Palaeobates angustissimus from the Roetian and Lower Raciborowice. Muschelkalk of the Holy Cross Mts. (Wolica Beds, Wavy Beds and £ukowa Beds) as well as from the Up per Muschelkalk (Entolium discites Beds and Ceratites Beds). Dorka (2001) dis - Acrodus sp. covered such teeth in Lower Saxony (Germ any) while Rieppel (Fig. 4D–E) (1981) found them in the Middle Trias sic of Swit zerland. MGUWr 5390s; 5391s Schultze and Möller (1986) de scribed sim i lar spec i mens from the Mid dle Muschelkalk of Göttingen (Germ any).

Material. — Two well-pre served spec im ens comprise one com plete and one fragm entar y crown with very weak orna - Palaeobates sp. menta ti on. The latter has one half of the crown miss ing. (Figs. 4J and 5A–C) Description. — Teeth are elongate d, flat and not MGUWr 5396s–5399s very high. The well-pre served orna m enta ti on consis ts of folds, in places bi fur cat ing, run ning down from the lon gi tu di nal crest. The speci m ens are 2.5–5 mm long, 1–2.5 mm wide and about Material. — Two fragm entar y and two com plete 1.5 mm high. The length/width rati o is 1.6–2.5. The profil e and well-pre served teeth. or na men ta tion sug gest that they be long to Acrodus. Description. — The teeth are elon gated, measur ing Occurrence. — Bone Bed of C unit at Racibo - 3–5 mm in length, 1.5–3 mm in width and 1–1.5 mm in height. rowice. Accord i ng to Dorka (2001) they are also pres ent in the The length/width rati o is 2.25–3.3. Fine striations radi ate from Up per Tri assic of Lower Saxony (Germ any). Rieppel (1981) the longi tu di nal ridge, running across the crown (Fig. 5A). The de scribed Acrodus teeth from the Mid dle Tri as sic of Swit zer - gran u la tion is com posed of small ir reg u lar, gen er ally oval el e - land. Acrodus is also num erous in the Tri assic to lower va tions and hol lows an or na men ta tion typ i cal of Palaeobates. Campanian inter val of Europe and Russia (Cappetta, 1987). It is less char acter istic in the spec im ens (Fig. 5B) where it slightly re sembles that of Acrodus. The tooth (Fig. 5C) also shows the or na men ta tion some what dif fer ent from typ i cal one. Polyacrodontidae Glückman, 1964 The crown is charac ter isti cally granu lat ed on one side of the Palaeobates Meyer, 1849 longi tu di nal crest but is covered by folds, in places bifur cat ing, Palaeobates angustissimus Agassiz, 1838 which run in vari ous direc ti ons from the crest on the other side (Fig. 4F–I) (re sem bling that of Acrodus). MGUWr 5392s–5395s Occurrence. — In the North-Sudetic Ba sin the spe - cies occurs in the Bone Bed of unit C. Palaeobates was de - scribed from the Middle to Upper Tri assic in Europe and North 1973 Palaeobates angustissimus, Liszkowski, ryc. 2, fig. 5. Amer ica (Cappetta, 1987). 1981 Palaeobates angustissimus, Rieppel, figs. 8, 9; fig. 13a–c. 1986 Palaeobates angusstissimus, Schultze and Möller, fig. 3c, e. 1987 Palaeobates angustissimus, Cappetta, fig. 40 L–N. Class: Osteichthyes Huxley, 1880 1998 Palaeobates angustissimus, Chrz¹stek and NiedŸwiedzki, pl. II, fig. 6. 2001 Palaeobates angustissimus, Dorka, fig. 1L. Sub class: Actinopterygii Klein, 1885 2002 Palaeobates angustissimus, Chrz¹stek, pl. 23, fig. 5. Or der: Saurichthyiformes Hay, 1902 Birgeriidae Aldinger, 1937 Material. — Four well-pre served teeth com prise one Birgeria Stensiö, 1919 com plete speci m en (crown and root), two com plete crowns and Birgeria sp. a half of a crown. (Fig. 5D) Description. — Teeth are 2–3 mm wide, 5–8 mm MGUWr 5400s long, of rounded margins with a rather low (ca. 1 mm) crown. Vertebrate remains from the Lower Muschelkalk of Raciborowice Górne (North-Sudetic Basin, SW Poland) 231

Fig. 5. Fish teeth, scales and bones from the Bone Bed (unit C) of the Lower Muschelkalk at Raciborowice Górne

A–C — Palaeobates sp.; D — Birgeria sp.; E–F — scales of Gyrolepis sp.; G–H — Actinopterygian scales; I–J — enigm atic bones (fish?)

Material. — One fragm entar y but well-pre served tooth. Occurrence. — In Raciborowice Górne this spe cies Description. — The tooth is small, very slightly occurs in the Bone Bed of unit C. Vickers-Rich et al. (1999) de- curved and broad-based. It measures 1.5 mm in length and scribed sim i lar Birgeria teeth from the Middle Trias sic of Saudi 0.5 mm in width. The surface is smooth without noti ceable or- Arabia. They also noted that the spe cies is known from the na men ta tion. Early Tri as sic to Rhaetian and is of wide geographi c extent (North Amer ica, Green land, Spitsbergen, Mad a gas car). 232 Alina Chrz¹stek

Or der: Palaeonisciformes Hay, 1929 Super class: Terapoda Acrolepididae Aldinger, 1937 Class: Reptilia Gyrolepis Agassiz, 1833 Sub class: Euryapsida Gyrolepis sp. Or der: Eosauropterygia Rieppel, 1994 (Fig. 5E–F) Suborder: Eusauropterygia Tschanz, 1989 MGUWr 5401s; 5402s Infraorder: Nothosauroidea Baur, 1889 Superfamily: Nothosauria Baur, 1889 Nothosauridae Baur, 1889 Material. — Two com plete scales with faint or na men - Nothosaurus Münster, 1834 ta tion. Nothosaurus cf. mirabilis Description. — The scale are rhomboidal, 0.8 and (Fig. 6J) 2.6 mm across respectivelly. Only one of the speci m ens shows del i cate per pen dic u lar striation. Occurrence. — In the North-Sudetic Ba sin these Material. — One very well pre served ver tebra. scales were described from the Bone Bed of unit C. Liszkowski Description. — The roughly square-shaped verte - (1981) dis covered teeth belong ing to vari ous specie s of the ge- bra of the rep tile measures 1.5´2.0 cm. A cen trally located pit nus Gyrolepis from the Silesia–Kraków region and from the broadens towards the edges. The spec im en does not differ from Holy Cross Mts. Accord ing to Schultze and Möller (1986) sim - ver tebrae of the rep tile Nothosaurus mirabilis de scribed by ilar spec im ens oc cur in the Middle Muschelkalk of Göttingen Schmidt (1928, fig. 1112a). (Ger many). Occurrence. — In thick-bedded organodetrital lime- stone with abundant crinoids of unit B. Sim ilar verte brae were descri bed from the same hori zon by Chrz¹stek and Scales of Actinopterygii NiedŸwiedzki (1998) and by Chrz¹stek (2002). Schmidt (1928) (Fig. 5G–H) noted sim i lar spec i mens from the Muschelkalk of Germ any. MGUWr 5403s; 5404s

Suborder: Eusaropterygia Material. — Two scales with faint or na menta tion. (Fig. 6B–D) Description. — Oval and el lip soi dal scales measur - MGUWr 5407s–5409s ing 1.1 mm and 2 mm across. One of them lacks any noti ceable or na ment (Fig. 5G) while the other shows faint ridges paral lel to the edges (Fig. 5H). They do not differ signif i cant ly from the Material. — Three well-pre served teeth — one com - scales of Colobodus descri bed by Schmidt (1928, fig. plete and two with broken tips. 1022a,f–g) but poorly preserve d orna m enta ti on hinders a more Description. — The teeth are tri an gu lar, slightly def i nite tax o nomic as sign ment. curved with lon gi tu di nal striation. The spec i mens mea sure Occurrence. — Bone Bed of unit C at Raciborowice 0.75, 4 and 6.5 mm in length and 0.3, 1 and 1.5 mm in width. Górne. Their profil e and orna m enta ti on suggest that they belong to the fam ily Nothosauridae or Cymatosauridae though pre cise taxo - nomic attri buti on is not possi ble based on isolat ed teeth only. Enig matic bones Occurrence. — For the first time noted at (Figs. 5I–J and 6A) Raciborowice Górne from the Bone Bed of unit C. Reif (1980) MGUWr 5406s; 5405s-1; 5405s-2 found simi lar teeth at Baden–Würtemberg (SW Ger many).

Material. — Three very well pre served spec im ens. Ver te brate cop ro lites Description. — One bone has the shape of an in - (Fig. 6E–J) verted let ter T and a smooth sur face (Fig. 6A). The base is 4–4.5 mm across down to 2 mm at the thinnest sec tion. The spec im en measures 5 mm in length. It re sem bles the skull bone Material. — Seven well pre served spec im ens. of an actinopterygian fish (Hagdorn, pers. comm.). Description. — The spec im ens are elon gated, well Other bones are elongate d and measure 0.75 mm and 1 cm rounded. They measure 1.5–2.5 cm in length and 0.5–1.0 cm in in length and about 0.75 mm in width (Fig. 5I–J). Their surface width. is gen er ally smooth though lon gi tu di nal ridges and del i cate Occurrence. — In two hori zons of organodetrital folds may be observe d (Fig. 5I). They resem ble fish bones limestone , one with abundant Dadocrinus, the other with nu- (Hagdorn, pers. comm.). mer ous Myophoria vulgaris, in the lower and upper parts of Occurrence. — The Bone Bed of unit C at unit B of the Raciborowice Górne Lower Muschelkalk sec tion, Raciborowice Górne. re spec tively. Vertebrate remains from the Lower Muschelkalk of Raciborowice Górne (North-Sudetic Basin, SW Poland) 233

Fig. 6. Ver tebrate re mains from the Bone Bed of unit C (A–D) and from unit B (E–J) of the Lower Muschelkalk at Raciborowice Górne A — enigm atic bone (actinopterigian?); B–D — rep tiles teeth (nothosaurids or cymatosaurids); E–G — cop ro lites; H — c — ver te brate coprolite, d — Dadocrinus sp.; I — c — ver te brate coprolite, b — bi valves (Myophoria vulgaris); J — r — ver te bra of Nothosaurus cf. mirabilis, c — ver te brate coprolite

PALAEOENVIRONMENT Holocrinus acutangulus (Hagdorn and G³uchowski, 1993; Chrz¹stek, 2002). However a hori zon of cellu lar limestone (Fig. 2) in di cates that pe ri odic sea level changes and ep isodic The beds of unit B, rich in the rem ains of verte brat es, emer gences took place. Such a type of limestone is typ ically mainly repti les, were depos it ed in a shal low envi ron m ent. Wa - formed in the sabkha en vi ron ment (sensu Wilson, 1975). ter sa lin ity was gen er ally nor mal marine as shown by a mass Lithological and palaeontological data suggest a la goon with occur rence of the cri noids Dadocrinus and (less frequent) re stricted wa ter cir cu la tion as a dom i nant en vi ron ment dur ing 234 Alina Chrz¹stek the de posi ti on of unit B (Chrz¹stek, 2002). Szulc (1991) pro- CORRELATION posed sim i lar sed i men tary con di tions for unit B and in ter preted the cel lu lar limestone as a postevaporitic fabric . The dom ina - Palaeontological data on verte brat es from the Lower tion of bones and verte brae of repti les (Nothosaurus cf. Muschelkalk showed that fish re mains are com mon in the Ger- mirabilis) as well as a lack of scales and teeth of fish in the skel- manic Ba sin (Cen tral Eu rope), in clud ing Po land. Ac cord ing to etal assem blage seem to sup port that con cept. The rep tiles Liszkowski (1981) the ver tebrate re mains are rep re sented proba bly lived in a very shal low lagoon. mainly by iso lated frag ments of ex ter nal (scales, fin rays) and The as semblage of macro- and ichnofossils pres ent in inter nal (verte brae, teeth) skele ton. This is supporte d by the as - unit C in di cates a some what deeper sed i men tary en vi ron ment sem blage of verte brat e fossil s from the Lower Muschelkalk at for these beds (Chrz¹stek, 2002). Limestone s of the unit are Raciborowice Górne where num erous isolat ed teeth, sporadi c dark, py rite-bearing with the ichnofossil Planolites isp. and scales, bones of repti les and proba bly of fish were discov ered. scarce ver tebrate re mains that sug gest a lower ox ygen con tent The col lected ver tebrate re mains from the North-Sudetic Ba sin in the seawa ter . Low oxy gen a ti on might have been caused by shows a great simi lar ity to the age-equiv alent as semblage from the decomposition of fish rem ains. Abundant teeth may indi - Wolica near Chêciny in the Holy Cross Mts. (Liszkowski, cate that sharks were common in the Germanic Basin. 1973, 1981, 1993). It occurs in the £ukowa Beds, in the Lower Based on the col lected teeth as semblage it may be con - Muschelkalk sec tion of the Holy Cross Mts., an equiva lent of cluded that during the sedi m enta ti on of unit C the shark popu - unit C of the North-Sudetic Ba sin (G³uchowski and Salamon, la tion was dom i nated by euryhaline bot tom-dwell ing se- 2005). Although the fauna from the Holy Cross Mts. is richer in lachians that fed on benthos (Acrodus sp. and Palaeobates sp.). taxa, sim i lar chondrichthyans (Acrodus lateralis, Acrodus sp., This is indi cat ed by the presence of crushing teeth and frequent Palaeobates angustissimus and Palaeobates sp.) and lay ers con tain ing gas tro pods and bi valves — or gan isms that osteichthyans (Birgeria sp. and Gyrolepis sp.) occur in both con sti tute a com mon sclerophages’ diet (Fig. 3). Reif (1982) regions. con sid ered “gen er al ized sharks” as con tem po rary equiv a lents The skel etal rem ains from the Lower Muschelkalk of of the Trias sic and Cre taceous sharks. They are rather Raciborowice Górne are also simi lar to the age-equiv alent as - slow-swimming bot tom-dwell ers that feed on small fish, sem blages from the Gogolin Beds of the Up per Silesia. The lat- thin-shelled bi valves, worms and other benthi c organ ism s. ter are dom inated by Hybodus sp., Acrodus lateralis, Boss (1982) dis tinguishe d 3 types of chondrichthyan teeth and Palaeobates sp. and Lissodus sp. with less com mon classi fied the Trias sic as semblage as of type I, i.e. teeth adapted Polyacrodus sp. (Liszkowski, 1981, 1993). Osteichthyans — for crushing shells. Accord ing to him a Tri assic Saurichthys sp., Colobodus sp. and Gyrolepis sp. — also oc cur chondrichthyan had an elongate d body and large mouth to prey (Liszkowski, 1981). Thus the Sudetic as sem blage is com pa ra - on larger organ ism s. Scales and teeth of osteichthyans are less ble to both of these age-equiva lent faunas though it most num erous, which may suggest that the Sudetic basin was too closely re sembles the as semblage from the Holy Cross Mts. As shal low for them. in the North-Sudetic Basin the maxi m um abundance s of such Bürgin (1999) reporte d verte brat e rem ains from taxa as Acrodus lateralis and Palaeobates angustissimus were lithologically simi lar dark-grey limeston e in the Lower observe d in the £ukowa Beds, an equiva lent of unit C at Muschelkalk of southern Switzer land (Monte San Giorgio). Raciborowice Górne. Accord ing to him the limestone was formed in the anoxic envi - In Lower Silesia, Up per Silesia and in the Holy Cross Mts. ronm ent of a shallow lagoon. chondrichthyan teeth domi nate (mostly Acrodus sp. and Thus the sedi m entar y envi ron m ent of unit C was shal low Palaeobates sp.) while teeth and scales of Actinopterygii are but somewhat deeper than the one of unit B (Szulc 1991; rare. On the other hand Hybodus sp., Lissodus sp., Polyacrodus Chrz¹stek, 2002). The anal ysis of ichnoassociations sp. and Saurichthys sp., present in the Holy Cross Mts. and in (Chrz¹stek, 2007) indi cat es that the part of unit C contai ning the Silesia-Kraków region, were not observe d in the the Bone Bed shows a regres sive trend. This is shown by the North-Sudetic Ba sin. pres ence of hardground intraclasts in the Spiriferina Bed that The tax o nomic com po si tion of the skel e tal re mains from the occur a lit tle higher in the secti on. The break in sedi m enta ti on, Lower Muschelkalk of Raciborowice Górne shows also many as shown by hardground for mation, may be re lated to sim ilar iti es to the equiva lent faunas from other regions of Eu - transgressive-re gres sive events (Chrz¹stek, 2004). The maxi - rope. It does not differ from the Germ an as sem blage that in- mum marine transgres sion in the studied region, caused proba - cludes such fish taxa as: Acrodus sp., Hybodus sp., Palaeobates bly by the opening of the Silesian-Moravian marine gate way, sp., Gyrolepis sp., Saurichthys sp. (Reif, 1980; Kriwet and started im me di ately with the de po si tion of the Spiriferina Bed Schultz, 1998; Bürgin, 1999). Sim ilar Chondrichthyes and (Kêdzierski and Szulc, 1996; Chrz¹stek, 2002). It may be con- Osteichthyes are known from the Middle Trias sic of the cluded that the sedi m entar y envi ron m ent of the Lower Netherland: Acrodus sp., Palaeobates angustissimuss, Gyrolepis Muschelkalk deepened gradu all y from the be ginning of unit B sp., Colobodus sp., Saurichthys sp. and Birgeria sp., France: depo si ti on. The ba sin reached its maxi m um depth dur ing the Gyrolepis sp., Saurichthys sp., Birgeria sp. and Colobodus sp. as sedi m enta ti on of the upper part of unit C and unit E. This is well as north ern It aly: Acrodus sp., Hybodus sp., Saurichthys sp. confirm ed by the assem blage of macro- and ichnofossils and Colobodus sp. (Bürgin, 1999). In the Tri assic of north-east - (Fig. 3; Chrz¹stek, 2002, 2007). ern Spain chondrichthyans do not occur but osteichthyan re- mains such as Saurichthys sp. and Colobodus sp. are comm on Vertebrate remains from the Lower Muschelkalk of Raciborowice Górne (North-Sudetic Basin, SW Poland) 235

(Beltan, 1972 in Bürgin, 1999). Birkenmajer and Jerzmañska Gyrolepis sp. and Birgeria sp., reached the Ger manic Ba sin (1979) de scribed sim i lar taxa: Hybodus sp., Acrodus sp., though they dom inate clearly in the Alpine Basin (Aus tria, Polyacrodus sp. and Saurichthys sp. from the Lower Trias sic of Swit zerland) and in other epicontinental bas ins lo cated close to Spitsbergen. Hybodus sp., Gyrolepis sp. and Lissodus sp. were the Tethys Ocean (Spain, France). on the other hand noted from the Upper Tri assic of Luxem bourg Ac cord ing to Minikh (1998) a connec ti on betwe en the Rus- (Godefroit et al., 1998). sian Basin and the Boreal and Tethys oceans exist ed in the Descri bing the teeth of Chondrichthyes and Actinopterygii Anisian. The Middle Trias sic transgres sion spread out towards from the Middle Trias sic of southern Switzer land, Bürgin the Urals result ing in the dom ina ti on of Osteichthyes over (1999) reporte d Gyrolepis sp. and Acrodus sp., which occur Chondrichthyes taxa in the Russia n Trias sic depos it s, as in the also at Raciborowice Górne. The dif fer ence be tween the Al pine se quences. Sudetic as semblage and its Swiss age equiv alent lies in a Rieppel and Hadgorn (1997), based on the analy sis of rep- greater num ber of actinopterygian taxa in the lat ter. Rieppel tiles dis tri bu tion, pro posed that sauropterygians migrate d from (1981) also menti oned Hybodus sp., Acrodus lateralis and the Asiati c Province into the Germ anic Basin through the Palaeobates angustissmus from the Middle Trias sic of south- Silesian–Moravian marine gate way. They sug gested this direc - ern Swit zer land. tion of transgres sion be cause Nothosaurus sp. did not appear in Ac cord ing to Minikh (1998) many Trias sic verte brat es oc - the Al pine Tri as sic un til the Anisian/Ladinian, when the west- cur also in the age-equiva lent depos it s of Rus sia (Hybodus sp., ern marine gate way opened. Acrodus sp., Lissodus sp., Cera to dus sp., Saurichthys sp., As sem blages of skel e tal re mains in the North-Sudetic Ba - Colobodus sp.). sin and in the rem aining part of the Germ anic Basin are sim ilar . The taxo nom ic com posi ti on of the assem blage of fish re- Thus they do not offer a suffi cie nt basis to conclude whether mains from the Lower Muschelkalk of the North-Sudetic Ba sin the East Carpathian or the Silesian–Moravian marine gate way does not differ signif i cant ly from the faunas of the Germ anic was the dom inant connec ti on with the Tethys Ocean. In the late facie s from other regions of Poland (Holy Cross Mts., Olenekian and early Anisian (sensu Nawrocki and Szulc, 2000) Silesia–Kraków region) and the rest of Europe: Germ any, the faunal migra ti on took place most likely through both gateway s Nether lands and Luxem bourg. It is also sim ilar to the Al pine as sug gested al ready by Szulc (2000) and NiedŸwiedzki and fa cies as semblages of the same age, though osteichthyan taxa Salamon (2002). The Silesian–Moravian marine gate way be - are consid er ably less num erous in the Lower Muschelkalk of came dom inant from the de posi ti on of the Spiriferina Bed, the North-Sudetic Basin com pared to the assem blages known when the brachio pod Punctospirella fragilis and num erous cri - from Swit zerland, Austri a and northern It aly (Bürgin, 1999). noids and ammonites appeare d in the North-Sudetic Ba sin These sharks preferred proba bly deeper envi ron m ents and the (Chrz¹stek, 2002). At that time the Sudetic Ba sin started to Sudetic Basin was too shallow. Conse quentl y Osteichthyes, es - deepen con sid er ably, which may be re lated to the ex istence of a pe cially Actinopterygii, dom i nated and tax o nom i cally more di- conve nient connec ti on with the Tethys Ocean proba bly verse in the Tethys Ocean. The as semblages of ver tebrate re - through the Silesian–Moravian marine gate way (Kêdzierski mains from the epicontinental basins of Spain and France, lo - and Szulc, 1996; Chrz¹stek, 2002). The data of G³uchowski cated closer to the Tethys Ocean, also dif fer from those from and Salamon (2005) suggest that this event could take place at the Ger manic Ba sin. They are richer in Osteichthyes but poorer the be ginning of the Pelsonian. The posi ti on of the Bone Bed in, or lack com pletely, Chondrichthyes taxa. Sim ilar conclu - about 4.5 m below the Spiriferina Bed in unit C of the sec tion sion may be drawn from the analy sis of taxo nom ic di versit y of may in di cate its early Anisian (Bithynian) age. ver tebrate re mains in the Lower Muschelkalk of Russia . Apart from chondrichthyans, typi cal of the Trias sic , the num ber of osteichthyan gen era and spe cies is sig nif icant. SUMMARY AND CONCLUSIONS

Five taxa of chondrichthyan and, for the first time, two taxa PALAEOGEOGRAPHY of osteichthyan fishes (Ta ble 1) were found in the Lower Muschelkalk of the North-Sudetic Ba sin. Ver te brate bones Ac cord ing to Kriwet and Schultz (1998) the migra ti on of (fishes?, repti les) and, not hithert o records from Raciborowice Chondrichthyes and Osteichthyes: Acrodus sp., Hybodus sp., Górne, teeth of the rep tiles Nothosauroidea or Palaeobates sp. and Saurichthys sp. from the Tethys Ocean Cymatosauroidea were also recov ered. through the East Carpathian marine gate way into the Ger manic Rep tile re mains are pres ent mostly in unit B while fish teeth Basin took place in the late Scythian. In the early Anisian oc cur in a char ac ter is tic dark grey py rite-bear ing lime stone (Roetian/Muschelkalk) Chondrichthyes: Acrodus sp., Hybodus layer (Bone Bed) about 4.5 m below the Spiriferina Bed of sp., Palaeobates sp., Polyacrodus sp. and Lissodus sp. becam e unit C. Apart from fish teeth that had been already descri bed, wide spread across the en tire shal low Ger manic Ba sin. This is the Bone Bed contai ns also enigm atic bones (fish?) and teeth linked to a transgres sion from the Tethys Ocean through the from nothosaurids or cymatosaurids. Silesian–Moravian and East Carpathian marine gate ways to - The Sudetic assem blage is com posed mainly of teeth be- wards northern Germ any into the southern part of the North long ing to Chondrichthyes whereas Osteichthyes are rare. The Sea as well as south-west wards to Burgundy. Some of the as semblage is typ ical of the Lower Muschelkalk, in which osteichthyan taxa, such as: Saurichthys sp., Colobodus sp., euryhaline bottom -dwelling se lachians that fed on benthos pre- 236 Alina Chrz¹stek vailed. Layers contai ning gas tropods and bi valves are com mon Spain and France. It may be suggest ed that these fishes pre- in the pro file and in di cate that prey was eas ily available. Ac - ferred deeper en vi ron ments whereas the Sudetic and maybe cord ing to Reif (1982) the so-called “gen er alized sharks” may also the Ger manic Bas ins were too shal low. be con tem po rary equiv a lents of the Tri as sic and Cre ta ceous se - From the late Olenekian to the early Anisian, during the lachians that liv ing in a sim ilar hab itat. sedi m enta ti on of the verte brat e-bearing units B and C, a con- During the sedi m enta ti on of the Bone Bed, the envi ron m ent nec tion with the Tethyan Ocean ran through the East was oxy gen-de fi cie nt as shown by dark col our of the lime- Carpathian and Silesian–Moravian marine gate ways as sug - stone, the presence of pyrit e miner al iza tion and the ichnofossil gested al ready by Szulc (2000) and NiedŸwiedzki and Salamon Planolites isp. The seawa ter proba bly had a low oxy gen con- (2002). The Silesian–Moravian marine gate way be came more tent becaus e of large quanti ty of decom posing chondrichthyans im por tant dur ing the sed i men ta tion of the Spiriferina Bed. Sig - that consti tuted a source of numerous teeth. nifi cant deepen ing of the basin suggest s a good connec ti on The transgres sion de veloped from the be ginning of the with the Tethyan Ocean at that time. This is supporte d by the Lower Muschelkalk. Unit B formed in the shallowe r envi ron - palaeontological record of this part of unit C (Chrz¹stek, 2002; ment of a la goon with re stricted wa ter cir cu la tion (Chrz¹stek, 2007). 2002). Unit C, on the other hand, was depos it ed in a deeper la - G³uchowski and Salamon (2005) suggest ed a Pelsonian age goon with less re stricted wa ter cir cu la tion (Chrz¹stek, 2002). for the Spiriferina Bed. This in di cates an early Anisian This is also indi cat ed by the assem blage of repti le bones (Bithynian) age for the Bone Bed becaus e it occurs about 4.5 m (unit B) as well as of chondrichthyan and osteichthyan teeth be low the Spiriferina Bed in the Lower Muschelkalk sec tion. (unit C). The Sudetic as semblag e shows the clos est simi lar ity to that Ac knowl edge ments. The author would like to thank H. of the age-equiva lent strata from the Holy Cross Mts. As an as- Hagdorn his crit ical review of the taxo nom ic assign m ent of semblag e typ i cal of the Ger manic Ba sin it does not dif fer sig - some teeth and scale taxa and R. NiedŸwiedzki for fruitful dis - nifi cant ly from the faunas of the same age from other regions of cussion and for some of the speci m ens. The comments of the Po land (Up per Silesia), Ger many, France, the Neth er lands, re view ers, J. Szulc and T. Sulej, were of great value in pre par - Spain, northern It aly, Austri a, Switzer land and Russia . It ing the fi nal version of the manuscri pt. M. Ratajczyk, super - should be noted, however, that the Alpine facie s faunas (Aus - vised by the author, gathered a part of the fossil collec tion dur- tria, Swit zer land) con sist mostly of osteichthyan re mains. A ing her MSc field work. The inves ti gati ons were supporte d fi- sim i lar sit u a tion is ob served in the as sem blages from nancia lly by the grants 2022/W/ING/-07 and 1017/S/ING/-4. epicontinental seas lo cated close to the Tethyan Ocean, i.e. in

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