UP PER TRI AS SIC VER TE BRATE TRACKS FROM KRAKÓW CZÊSTOCHOWA UP LAND, SOUTH ERN PO LAND

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1 Annales Societatis Geologorum Poloniae (2013), vol. 83: UP PER TRI AS SIC VER TE BRATE TRACKS FROM KRAKÓW CZÊSTOCHOWA UP LAND, SOUTH ERN PO LAND Grzegorz SADLOK 1 & Zuzanna WAWRZYNIAK Leadside Road AB25 1RX, Aberdeen, Aberdeenshire, Scot land (UK); gregsadlok@gmail.com 2 Fac ulty of Earth Sci ences, Uni ver sity of Silesia, Bêdziñska 60 Street, Pl , Sosnowiec, Po land; zuza.wawrzyniak@gmail.com Sadlok, G. & Wawrzyniak, Z., Up per Tri as sic ver te brate tracks from Kraków Czêstochowa Up land, south ern Po land. Annales Societatis Geologorum Poloniae, 83: Abstract: The first Up per Tri as sic ver te brate fos sil tracks are doc u mented from the Zawiercie lo cal ity, Kraków Czêstochowa Up land, south ern Po land. The most char ac ter is tic com po nents of the as sem blage are tracks, as sign able to archosaurs and dicynodonts. The in ferred com po si tion of the fauna is com pa ra ble to those of much better stud ied, con tem po ra ne ous sites in It aly, Ar gentina and the USA. Key words: ver te brate tracks, Up per Tri as sic, Kraków Czêstochowa Up land, Po land. Manu script re ceived 7 Sep tem ber 2013, ac cepted 25 No vem ber 2013 INTRODUCTION South ern Po land is an area con tin u ous with the Tri as sic Ger manic Ba sin (Szulc, 2007). The Up per Tri as sic strata of this ba sin out crop ping in south ern Po land have yielded new palaeontological sites for the last 20 years (see over view in Sulej et al., 2011). Al though the find ings of Tri as sic ter res - trial body fos sils are not new in this area (see ref er ences in Rauhut and Hungerbühler, 1998), clearly, there is a grow ing in ter est in this field of study. The revitalisation started with the dis cov ery of the fos sil site in Krasiejów in Silesia, southern Po land (Dzik and Sulej, 2007), which has yielded, amongst oth ers, body fos sils of dinosauromorphs show ing some af fin i ties with ornithischians (Dzik, 2003). Sub se - quently, new sites have been added to the list. These sites are Lipie Œ¹skie, WoŸniki and Porêba. The two sites (Lipie Œ¹skie and WoŸniki) have yielded, amongst oth ers, the body fos sils of archosaurs (?di no saurs) and large dicynodonts (Dzik et al., 2008; Sulej et al., 2011; 2012; NiedŸwiedzki et al., 2012). Up to now, ver te brate tracks have been re ported from only two of the three Up per Tri as sic Silesian sites, Lipie Œl¹skie and WoŸniki (Dzik et al., 2008; NiedŸwiedzki and Sulej, 2008; NiedŸwiedzki, 2011; Sulej et al., 2011). Here, by re cord ing the new site and il lus trat ing tracks, the au thors sup ple ment the data on the Up per Tri as sic ver te - brate tracks from south ern Po land. The new fos sil tracks come from Zawiercie, in the Kraków Czêstochowa Up - land, south ern Po land (Fig. 1). The ma te rial de scribed here is the first per tain ing to Up per Tri as sic ver te brate tracks from the Kraków Czêstochowa Up land. GEOLOGICAL BACKGROUND The ma te rial stud ied co mes from the Zawiercie site, which is lo cated in the Kraków Czêstochowa Up land in south ern Po land (Fig. 1). The fos sil site is a se quence of Norian (Up per Tri as sic) strata (see Szulc et al., 2006). These strata were ex ca vated dur ing con struc tion of a city dump in Zawiercie. As a re sult, all spec i mens dis cussed and fig ured in this pa per have been lo cated ex situ (on a pile of sed i - ment). Un for tu nately, cur rently there is no a sin gle eas ily ac ces si ble out crop, where these strata are ex posed and suit - able for study. This emphasises the sig nif i cance of the Zawiercie site as a valu able source of palaeontological in - for ma tion on Late Tri as sic eco sys tems in the area. The Up per Tri as sic strata, oc cur ring in the Zawiercie site (see Kotlicki, 1967), be long to the belt of Up per Tri as - sic strata which stretches be tween Opole in the west and Olkusz in the east (Szulc and Becker, 2007; Fig. 1). These Up per Tri as sic strata from Zawiercie are known in for mally as the WoŸniki Lime stone. Lime stones, in clud ing bi valve coquinas (Racki, 2010, fig. 2), car bon ate con cre tions and con glom er ates al ter nat ing with red beds, form the WoŸniki Lime stone, a unit de pos ited un der an arid and semiarid cli - mate (see Pieñkowski, 1988; Szulc et al., 2006). The conglomerates contain the remains of freshwater bivalves and ver te brate bones (Szulc et al., 2006; Budziszewska-Karwowska et al., 2010; Racki, 2010; Skawina and Dzik, 2011). Ver te brate tracks are pre served on soils of these lay ers. Car bon ate con cre tions of ten con tain rich plant ma te ri als, ver te brate fos sils, as well as miospores (Corollina meyeriana, Ovalipolis ovalis, Brachysaccus neomun-

2 106 G. SADLOK & Z. WAWRZYNIAK Fig. 1. Lo ca tion of all Up per Tri as sic fos sil sites in Po land at the University of Silesia (collection abbreviation is WNoZ). Some of the larg est spec i mens of ver te brate tracks that have been fig ured and dis cussed in this pa per were left in the field. In such cases, the spec i mens were cov ered with a pro tec tive ve neer of plas ter of Paris. The mea sure ments of track pa ram e ters were made in ImageJ and graphics were pre pared with Inkscape and Gimp. All three ap pli ca tions are readily ac ces si ble in the Ubuntu (Lu cid Lynx) Soft ware Cen tre. Ow ing to the lim ited num ber of spec i mens col lected and in par tic u lar their weak state of pres er va tion, the au - thors have de cided not to give pre cise ichnotaxonomic la - bels to this ma te rial. They note only the sim i lar ity of the tracks con sid ered to known ichnotaxa. danus, Enzonalasporites sp.), which are well-known morphotypes of the Pol ish Up per Tri as sic (cf. Or³owska-Zwoliñska, 1985). Stromatolite-like struc tures also have been found at the site (com pare Szulc et al., 2006; Racki, 2010, fig. 3), and these struc tures are in ter est ing palaeoenvironmental indicators awaiting further study. The age of the Zawiercie site Ac cord ing to Szulc et al. (2006), Lipie Œl¹skie is con - tem po rary with the site at Zawiercie. Ac cord ing to this au - thor, both these sites are prob a bly of Norian age (Up per Tri - as sic; see palynological zone IVb in Szulc et al., 2006). How ever, this view has been chal lenged by Dzik et al. (2008) and sub se quently by NiedŸwiedzki and Sulej (2008), who pos tu lated a Rhaetian age for the Silesian site (Lipie Œl¹skie). This sig nif i cant in con sis tency un der mines the ap - pli ca tion of fos sil finds from the area for com par i son with other sites in the world and re stricts their po ten tial util ity in any dis cus sion of fau nal turn over (ex tinc tion) at the Ju ras - sic Tri as sic bound ary (e.g., Benton, 2005). The Norian age of the Zawiercie site might be con firmed by the pres ence of the miospore Corollina meyeriana (characteristic for sedi - ments of that age) found by Wawrzyniak dur ing palynological anal y sis of sed i ments in which the tracks were found. This would sup port the re sults of Szulc et al. (2006) about the Norian age. A sep a rate sci en tific pro ject has been ini ti - ated to help in solv ing this strati graphic is sue (Racki, 2010). Un til this prob lem is solved, the Zawiercie site should be as - signed only on a gen eral ba sis to the Up per Tri as sic. MATERIAL AND METHODS The field work (stu dent sum mer camps) at the Zawiercie site was con ducted ev ery sum mer since The sum - mer stu dent camps are or ga nized by the Mu seum of Fac ulty of Earth Sci ences at the Uni ver sity of Silesia (Sosnowiec, Po land) with lo gis tic sup port from the city of Zawiercie. The ma te rial col lected is di verse and in cludes: fos sil bones of dicynodonts (some bear ing bite marks; Budziszewska-Karwowska et al., 2010), archosaurs, fishes, bi - valves, re mains of plants and ver te brate tracks. This ma te - rial is held by the Mu seum of the Fac ulty of Earth Sci ences VERTEBRATE TRACKS All of the ver te brate tracks de scribed are pre served as hypichnia (see ter mi nol ogy in Martinsson, 1970), which are nat u ral casts pre served on the lower sur faces of beds rep re - sent ing car bon ate con glom er ates. The pres ence of the fos sil tracks on the soles of con glom er atic beds may in di cate that these trace fos sils rep re sent the casts of par tially scoured deep tracks and/or scoured and reburied undertracks. This is be cause some ero sion is ex pected to be as so ci ated with en - ergy pulses, re sult ing in the de po si tion of gravel-grade ma - terial. Tracks of archosaurs Fig. 2 Ma te rial. Two spec i mens: WNoZ/SS/2/32 and WNoZ/SS/2/38. De scrip tion. These tracks have traces of three dig its (II IV). In the case of the track il lus trated in Fig ure 2, the traces of the an te - rior phalangeal pads are miss ing. How ever, this is a track with well pre served mor phol ogy. There are phalangeal pads de mar cated by nar rower inter-pad creases (Fig. 2). The trace of digit II has pads 1 and 2 pre served (start ing the count from the prox i mal pad) and is 10 cm long. The trace of digit III has pads 1 (com plete) and 2 (frag men tary) pre served. It is 7 cm long. The trace of digit IV has pads 1 3 and a frag ment of pad 4 pre served. The trace of digit IV is 12 cm long and pad 1 ex tends pos te ri orly, giv ing the asym met ric ap pear ance to the pos te rior part of the track (com pare Farlow et al., 2000). The length of the track (mea sured in axis of digit III) is 14 cm, and this is the lon gest, al though still in com plete spec i men. The inter-dig i tal an gle is low (~39 ). Re marks on po ten tial track-mak ers. These tracks are pre served well enough to al low some ba sic con clu sions. These tracks dis play fea tures, such as low inter-dig i tal an gle, asym met ric heel and in some cases (Fig. 2) vis i ble pads. These fea tures oc cur in tracks of theropods (see Farlow et al., 2000). In gen eral, the mor phol ogy of these tracks re sem bles that of grallatorid tracks (Grallator, Anchisauripus; e.g., Olsen et al., 1998; Melchor and Valais, 2006). Grallator Anchisauripus Eubrontes are three typ i cal theropod-made ichnogenera re ported from ter res trial de pos its of the Up per Tri as sic and Lower Ju ras sic strata of North Amer ica, South Amer ica and Eu rope (e.g., Hunt et al., 1998; Olsen et al., 1998; Melchor and Valais, 2006; NiedŸwiedzki, 2006; Lockley, 2009). These three ichnogenera form a spec trum of size with Grallator as the small est and Eubrontes the larg est (Lockley, 1991, 2009; NiedŸwiedzki, 2006). The morphological differences between these ichnogenera may mir ror the tax o nomic dif fer ences of their re spec -

3 UP PER TRI AS SIC VER TE BRATE TRACKS 107 Fig. 2. The fos sil track with three traces of dig its (Ro man nu - mer als: I, II, III in di cate traces of dig its), with low inter-dig i tal an - gle and asym met ric heel ; ar rows show nar rower zones, which are bound aries be tween digit pads (WnoZ/SS/2/32) Fig. 3. The iso met ric fos sil track with large meta tar sal pad (493 cm 2 ) and with short traces of dig its I V (field pho tog ra phy, spec i - men left in the field, Ro man nu mer als: I V in di cate traces of dig - its) tive track-mak ers or re cord the on tog eny of one trace maker (Olsen et al., 1998). How ever, sim i lar track morphologies may oc cur also in sauropodomorph di no saur tracks (Evazoum, see Lucas et al., 2006; Porchetti et al., 2008) and in in com plete pre served tracks, be long - ing to the Chirotheriidae ichnofamily (see ichnofamily over view in King et al., 2005). There fore, ow ing to the lim ited ma te rial, the as sign ment to an archosaur trace-maker ap pears to be the most cau tious. Tracks of?dicynodonts Fig. 3 Ma te rial. Two large tracks, both spec i mens were left in the field, be cause of the large size of the blocks. De scrip tion. These tracks are large and their max i mum length is com pa ra ble to their width. For ex am ple, for the track shown in Fig ure 3, the length/width ra tio is = 0.96 (max i mum length: 26 cm, max i mum width is 27 cm). These tracks clearly be long to a plantigrade trace-maker. This is be cause the track mor phol ogy is dom i - nated by a large meta tar sal pad (with an area of 493 cm 2 ; Fig. 3). On the other hand, the traces of dig its (I V) are short and of nearly equal length (2 3.5 cm). Traces of ex ter nal dig its (I and V) are slightly shorter than traces of in ter nal dig its (II, III and IV). The digit traces have blunt tips, and there fore there is no ev i dence for sharp claws on the foot of the track-maker. Re marks on po ten tial track-mak ers. The iso met ric shape of the track with short digit traces and blunt end ings re sem bles the tracks of dicynodontids (Hunt et al., 1993; Lucas et al., 2003; Surkov et al., 2007). An iso met ric track was also fig ured from WoŸniki and at trib uted to a dicynodont (Sulej et al., 2011). Un as signed track of?archosauromorphs Figs 4, 5 Ma te rial. Two spec i mens: WNoZ/SS/2/34 and WNoZ/SS/2/37. De scrip tion. These poorly-pre served tracks have three traces of dig its, all of them sim i lar in length. The lengths of dig its for the track from Fig. 4 are as fol lows: digit II: 5.4 cm, digit III: 5.8 cm, digit IV: 5.4 cm and for track from Fig. 5 are as fol lows: digit II: 4.9 cm, digit III: 5.8 cm, digit IV: 4.7 cm. These tracks also have a sym met ric pos te rior ( heel ) re gion. These tracks have a wide Fig. 4. The fos sil track with traces of three dig its (Ro man nu - mer als: I, II, III in di cate traces of dig its), be ing of equal length and with sym met ric heel and wide inter-dig i tal an gle (136, WnoZ/ SS/2/37) inter-dig i tal an gle be tween dig its II and IV, which is 136 in the track, shown in Fig. 4, and 76 for the track from Fig. 5. Re marks on po ten tial track-mak ers. The sym met ri cal ap pear - ance of the pos te rior part of the tracks, the wide inter-dig i tal an gle and digit traces of nearly equal lengths all sug gest that the tracks are sim i lar to those of ornithischians (e.g. Anomoepus; Lockley, 2003; Olsen and Rainforth, 2003; NiedŸwiedzki, 2011). On the other hand, these tracks also might rep re sent theropod tracks, if part of the heel is miss ing (which may be the case, Fig. 4). The track in ques tion also could fit the Coelurosaurichnus (Gand and Demathieu, 2005) or even prosauropod tracks (i.e. Evazoum (Pseudotetrasauropus;, see Lucas et al., 2006), if the digit I is miss ing. In such a case, a sauropodomorph foot print could readily be mis taken for a theropod or ornithischian foot print (see Farlow and Lockley, 1993). How ever, the poor state of pres er va tion pre cludes the pos - si bil ity of an ac cu rate de ter mi na tion of a track-maker in this case.

4 108 G. SADLOK & Z. WAWRZYNIAK Fig. 5. The fos sil track with traces of three dig its (Ro man nu - mer als: I, II, III in di cate traces of dig its), be ing of equal length and with sym met ric heel and wide inter-dig i tal an gle (76, WnoZ/ SS/2/34) Un as signed track of a plantigrade an i mal Figs 6, 7 Ma te rial. Sin gle spec i men: WNoZ/SS/2/33. De scrip tion. The track shown in Fig ure 6 is a struc ture 10 cm deep. The length of the track along the axis of the lon gest digit IV is 29 cm. This track be longs to a plantigrade an i mal with a mor - phol ogy dom i nated by a sin gle meta tar sal pad with an area of 217 cm 2. This pad is wider (max i mal width: 20.5 cm) than long (max i - mal length: 12 cm). There are four digit traces in the front of the pad. The lengths of digit traces in crease from digit trace I to IV (digit I: 5.5 cm, digit II: 7 cm, digit III: 8 cm, digit IV: 9.3 cm). A wide de pres sion sep a rates the meta tar sal pad and digit group II IV. The width of the de pres sion in creases to ward the trace of digit IV. An additional longitudinal depression separates digit IV from digit group I III. Re marks on po ten tial track-mak ers. The large meta tar sal pad, the pat tern of digit lengths (I<II<III<IV) and con fig u ra tion of de - pres sions are all fea tures which, in this com bi na tion, can be found in prosauropod tracks (pes). Good ex am ples are Navahopus coyoteensis, an ichnotaxon from the Lower Ju ras sic strata of USA (com pare Mil n et al., 2008) and Up per Tri as sic Eosauropus (Tetrasauropus, see Lockley et al., 2001; Lucas et al., 2006) whose (pes) shows some mor pho log i cal sim i lar ity with the track con sid ered (Fig. 6). Tak ing into con sid er ation the poor state of pres er va tion of the ma te rial stud ied, the au thors only are able to de scribe this track as at trib ut able to a plantigrade an i mal. How - ever, it is worth men tion ing that some work ers (see Hunt and Lucas, 2006) re turned to the synapsid con cept and sug gested that Navahopus rep re sents an extramorphological vari ant of a large tritylodont trackway. There fore, the pos si bil ity of a synapsid or i - gin of the fos sil track also should be taken into ac count. DISCUSSION OF FAUNAL COMPOSITION Fig. 6. The fos sil track with over all mor phol ogy, com parable to pes of prosauropods (WNoZ/SS/2/33) Fig. 7. Draw ing of fos sil track, shown in Fig. 6, not drawn to scale (Ro man nu mer als: I IV in di cate traces of dig its) The sam ple stud ied, al though re stricted in quan tity, in - di cates the pres ence of a di ver si fied ver te brate fauna and is a sig nif i cant con tri bu tion to the knowl edge of Late Tri as sic eco sys tems of south ern Po land. Tracks as sign able to the archosaurs show some sim i lar ity to tracks of the grallatorid ichnogenera (Grallator, Anchisauripus). Ac cord ingly, the au thors pro pose the pos si bil ity of link ing the or i gin of these tracks to the theropods, prob a bly to mem bers of the Coelophysoidea (Lockley, 1991), as all of the de ter mi na ble theropod re mains from the Up per Tri as sic strata of Eu rope be - long to the Coelophysoidea (see Rauhut and Hungerbühler, 1998). Tracks, in ter preted as formed by mem bers of the Coelophysoidea, have been re ported from Lipie Œl¹skie and WoŸniki (Dzik et al., 2008; NiedŸwiedzki and Sulej, 2008; NiedŸwiedzki, 2011; Sulej et al., 2011). The ichnological ev i dence for the pres ence of Coelophysoidea at the site in Zawiercie would be of spe cial in ter est. Pre vi ously, from the same site, Budziszewska-Karwowska et al. (2010) described dicynodontid tibia with tooth marks and at trib uted those marks to the life ac tiv ity of mem bers of the Coelophysoidea. How ever, though as signed to the archosaurs, the tracks also may rep re sent in com plete sauropodomorph tracks (e.g. Evazoum, Lucas et al., 2006) or weakly pre served tracks of the Chirotheriidae ichnofamily (King et al., 2005). The large, iso met ric plantigrade tracks are at trib uted here to dicynodonts. Al though not proven, this in ter pre ta - tion is quite likely, be cause the body fos sils of dicynodonts

5 UP PER TRI AS SIC VER TE BRATE TRACKS 109 are al ready known from the Pol ish Up per Tri as sic, in clud - ing that of the Zawiercie site (Dzik et al., 2008; Budziszewska-Karwowska et al., 2010; Sulej et al., 2011). Other sup port for this in ter pre ta tion co mes from a com par i son of the out line of one of the stud ied tracks with the pro por tions of the osteology of the dicynodontid foot, as ex em pli fied by the foot of Mid dle Tri as sic Tetragonias njalilus (Fröbisch, 2006; Fig. 8). The most prob lem atic ma te rial is rep re sented by the sin gle track, il lus trated in Fig. 6. This is be cause its mor - phol ogy re sem bles tracks, re fer able to the prosauropods and the dicynodonts. How ever, the re mains (body and trace fos - sils) of prosauropods are ab sent from other Pol ish sites (Krasiejów, Lipie Œl¹skie, WoŸniki; Dzik and Sulej, 2007; Dzik et al., 2008; Sulej et al., 2011). In the con tem po ra ne - ous strata of Ger many, prosauropods are quite com mon oc - cur rences (Weishampel and Chap man, 1990; Sander, 1992). At pres ent, the au thors only can note a sim i lar ity of the stud ied ma te rial to the pes mor phol ogy of some Up per Tri as sic and Lower Ju ras sic ichnogenera tracks, in ter preted as pro duced by sauropodomorphs (Mil n et al., 2008; Porchetti et al., 2008). How ever, the pres ent au thors also note that the al ter na tive in ter pre ta tion as a synapsid track should also be con sid ered. Long-term col lect ing is needed to better com pare Pol - ish Late Triassic vertebrate communities with German commu ni ties and the fau nal com po si tion at var i ous Pol ish sites. Pre lim i nary study of the fos sil flora has re vealed slight dif - fer ences be tween the Lipie Œl¹skie and Zawiercie sites. The car bon ate con cre tions found at the Zawiercie site con tain Equisetales stalks and leaves that have not been re corded at Lipie Œl¹skie (Wawrzyniak, 2010). This may in di cate some lo cal cli ma tic dif fer ences (e.g. higher mois ture at the Zawiercie site). Fur ther study is needed to ver ify whether pos - si ble dif fer ences in food com po si tion and/or food avail abil - ity might be par al leled by dif fer ences in the as sem blage of first-or der con sum ers at var i ous Pol ish sites. How ever, at pres ent in view of the poor state of knowl edge of the flo ral as sem blages at both sites, any con clu sions would be highly speculative. In gen eral, the tracks stud ied can be, with some con fi - dence, at trib uted to archosaurs and dicynodonts, the re - mains of which are typ i cal body fos sils that one can find at the Silesian sites. Such a com po si tion would be more or less con sis tent with the better un der stood ichnofaunas of the Up per Tri as sic in Eu rope (Nicosia and Loi, 2003, It aly), South Amer ica (Melchor and Valais, 2006, Ar gen tina) and North Amer ica (Gaston et al., 2003, USA). CONCLUSIONS The Up per Tri as sic site at Zawiercie has yielded the fos sil tracks of ver te brates. This re cord is the first from the Up per Tri as sic of the Kraków Czêstochowa Up land in south ern Po land. The fos sil tracks in di cate the pres ence of archosaurs and dicynodonts. This com po si tion is con sis tent with other ev i dence, al ready re corded from Up per Tri as sic sites in south ern Po land (Krasiejów, Lipie Œl¹skie and WoŸniki) as trace and body fos sils. Fig. 8. Dicynodont track and foot com pared. Su per po si tion of draw ing of dicynodont track based on track shown in Fig. 3 and draw ing of osteological pro por tions of dicynodontid foot based on Fröbisch (2006). The solid black lines in di cate rel a tive lengths of meta tar sals and dig its, whereas the oval shows the tarsalia area. Track and foot draw ings are not to scale, but this su per po si tion is intended to show the general similarities in proportions Acknowledgements We thank Grzegorz Pieñkowski from the Pol ish Geo log i cal Institute-National Research Institute (Warsaw, Poland) and Profes sor Mar tin Lockley from the Uni ver sity of Col o rado (Den ver, USA) for their crit i cal and con struc tive re marks. We thank Spencer G. Lucas from the New Mex ico Mu seum of Nat u ral His - tory and Sci ence (Al bu quer que, USA) for dis cus sion on the ichnotax on omy of the trace fos sils. We thank Grzegorz Racki from the Fac ulty of Earth Sci ences of Uni ver sity of Silesia in Sosnowiec (Sosnowiec, Po land) for some crit i cal re marks and lin guis tic cor - rec tions at an early stage of manu script prep a ra tion. We thank Pawe³ Filipiak (Sosnowiec, Po land) for help with palynological anal y sis. We are grate ful to the of fi cials of Zawiercie for their lo - gis tic and fi nan cial sup port dur ing the field camps and for their never-end ing en thu si asm for our re search. We thank all of the stu - dents and am a teur dino-hunt ers, who par tic i pated in the camps. We hope they will join us at the next camp. The work on this pa per has been par tially fi nanced from Re search Grant N N to Grzegorz Racki. We thank re view ers of this manu script Joachim Szulc from the Jagiellonian Uni ver sity (Kraków, Po land) and Spencer G. Lucas from New Mex ico Mu seum of Nat u ral His - tory and Sci ence (Al bu quer que, USA). We would like to thank Alfred Uchman (Kraków, Po land) and Frank Simpson (Wind sor, Can ada) for help and lin guis tic cor rects. REFERENCES Benton, M. J., Vertebrate Palaeontology, 3rd Edition. 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