A distinctive crouching theropod trace from the Lower Jurassic of Poland

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1 Geo log i cal Quar terly, 2009, 53 (4): SHORT COMMUNICATION A distinctive crouching theropod trace from the Lower Jurassic of Poland Gerard D. GIERLIŃSKI, Mar tin G. LOCKLEY and Grzegorz NIED WIEDZKI Gierliński G. D., Lockley M. G. and Niedźwiedzki G. (2009) A dis tinc tive crouch ing theropod trace from the Lower Ju ras sic of Po - land. Geol. Quart., 53 (4): Warszawa. A well-pre served, es sen tially com plete and dis tinc tive trace fos sil of a crouch ing, me dium-sized theropod di no saur is re ported from the north ern slope of the Holy Cross Moun tains, Po land. Crouch ing theropod traces are rare, and cur rently known from only six spec i mens from the Ju ras sic of North Amer ica and Asia. Thus, this new spec i men adds a sev enth spec i men to the di no saur track re cord. The spec i - men was found in the Early Ju ras sic (Late Pliensbachian) sand stones mined in the Szydłówek quarry, in the vi cin ity of Szydłowiec. Gerard D. Gierliński, Pol ish Geo log i cal In sti tute Na tional Re search In sti tute, Rakowiecka 4, PL Warszawa, Po land; JuraPark, ul. Sandomierska 4, PL Ostrowiec Świętokrzyski, Po land; hoo.com; Mar tin G. Lockley, De part ment of Ge ol ogy, Uni ver sity of Col o rado at Den ver, P.O. Box , Den ver, CO , USA; Grzegorz Niedźwiedzki, De part ment of Zo ol ogy, Fac ulty of Bi ol ogy, Uni ver sity of War saw, S. Banacha 2, PL Warszawa, Po land (re ceived: June 09, 2009; ac cepted: Sep tem ber 09, 2009). Key words: Po land, Holy Cross Moun tains, Lower Ju ras sic, di no saur crouch ing trace. INTRODUCTION A new dinosaur-sitting trace was found by two of us (G. D. G. and M. L.) at the Szydłówek site, in the Holy Cross Mountains of Poland, in August The first dinosaur tracks at this locality were discovered a year earlier by Joanna Roszkowska, Urszula Meissner and Zbigniew Remin. Those previous finds were reported by Niedźwiedzki and Remin (2008), and Niedźwiedzki et al. (2009). Tracks are found in the so-called Szydłowiec Sandstones mined in a small quarry located at GPS coordinates N and E Szydłowiec Sandstones (informal regional lithostratigraphic unit) belongs to the Drzewica Formation of late Pliensbachian age and represents the sixth sequence of the epicontinental Lower Jurassic deposits in Poland. This part of Drzewica Formation represents a succession com - prised of (from the oldest to youngest) nearshore-fore - shore-backshore/eolian facies (Pieńkowski, 2004). A few previous finds of sand stone slabs with dinosaur footprints were destroyed during regular quarrying activity (e.g., Fig. 1A). How ever, most of the discovered specimens, including the one re ported here with the catalogue number J484, were bought by Stowarzyszenie Delta (Delta Association) and protected in the JuraPark theme park com plex in Bałtów (e.g., Fig. 1B F). Specimens were re cently stored in a spe cially de - signed open-air ex hibit named Sabathówka, in memorial of late Polish palaeontontologist Karol Sabath. Only five previously known specimens of crouch ing theropod tracks have been re ported in the literature. These specimens are from the Lower Jurassic of Massachusetts (USA), St. George (Utah, USA) and the Lower or Mid dle Ju - rassic of China (Gierliński, 1994; Lockley et al., 2003; Mi lan et al., 2008; Milner et al., 2009). A sixth, previously undocumented specimen, was discovered in 2004 at a site in the Lower Jurassic Glen Can yon Group of the Glen Can yon (Lake Powell) National Recreation area, Utah. This specimen is illustrated here for the first time (Fig. 2), but is not de scribed in de tail; a mold and rep lica are housed at the University of Colorado at Den ver, in the Dinosaur Tracks Mu seum (CU ). Fi nally, the new Szydłówek find represents the sev - enth example, and most importantly is among the most com - plete and best pre served. TRACK DESCRIPTION The dinosaur-sitting trace fos sil is pre served as a natural cast on the fine-grained sand stone slab and associated with numerous bivalve resting traces of Lockeia. The dinosaur

2 472 Gerard D. Gierliński, Martin G. Lockley and Grzegorz Niedźwiedzki Fig. 1. Late Pliensbachian track as sem blage from Szydłówek, Po land A Parabrontopodus trackway; B Anchisauripus sp.; C Kayentapus sp.; D Eubrontes sp.; E Anomoepus trackway; F Moyenisauropus sp. ichnite is com prised of an ischial trace (Fig. 3C), left and right pes (Fig. 3B), and left and right manus (Fig. 3A). The manus tracks are much smaller than the pes and slightly wider (5.0 cm) than long (4.5 cm). The manus is tridactyl and dig its increase in length in the se quence: I, II, III. The ischial trace is tear-shaped, lon ger (12 cm) than wide (6.5 cm). The ischiadic callosity trace is lo cated al most mid way be tween the meta tar - sal proximal ends. The pedal tracks are mesaxonic, tridactyl and plantigrade. The phalangeal portion of the footprint is 23 cm long, whereas the entire footprint (including the metapodium) is 37 cm long. Angles between the digit axes are: II III = 13 18, III IV = The third pedal digit is the longest one. The ra - tio of the third pedal digit projection beyond the lateral toes to the length of the pes phalangeal portion equals The ratios of the digits (entire lengths) are: III/II = 1.17 and III/IV = However, these ratios measured according to the method of Olsen et al. (1998) are: III/II = 1.37 and III/IV = 0.89, which possibly reflect the ratios of the combined lengths of trackmaker phalanges III2 + III3 + III4 to II2 + II3 and IV1 + IV2 + IV3 + IV4 + IV5.

3 A distinctive crouching theropod trace from the Lower Jurassic of Poland 473 Fig. 2. Theropod crouch ing trace (spec i men CU ) from the Lower Ju ras sic, Glen Can yon Group, Utah A pes; B hallux; C other foot print of a large theropod (Eubrontes sp.) DISCUSSION As shown as shown in Figure 2, the Glen Can yon spec i men is quite sim i lar in size and mor phol ogy to the Szydłówek find, al - though part of the right pes is miss ing. The pedal tracks are sim i lar in length (both spec i mens are ~40 cm, when in clud ing meta tar sal traces), and com prise ap prox i mately 50% of the to tal length, and are width about ~12 cm. The most no ta ble dif fer ences are that the Glen Can yon tracks lack manus traces, but have clearer hallux traces. In ad di tion, the pes traces are more widely spaced and the right pes trace is lo cated an te rior to the left pes by the equiv a lent of about one-half of a track length (~20 cm). The ra tios of pedal digit lengths in the new Pol ish spec i men (JuraPark J484) are sim i lar to those given by Olsen et al. (1998) for the ichnospecies Eubrontes giganteus and Anchisauripus sillimani, as well as for the foot mor phol ogy of the basal theropod gen era Coelophysis and Dilophosaurus. Thus, this is

4 474 Gerard D. Gierliński, Martin G. Lockley and Grzegorz Niedźwiedzki Fig. 3. Theropod crouch ing trace (spec i men JuraPark J484) from the late Pliensbachian of Szydłówek, Po land A manus; B pes; C ischial cal los ity trace a unique trace fos sil of a crouch ing ceratosaurian theropod with well-de fined man ual digit prints left by the hands, and with its palms fac ing ven trally, not me di ally like in the St. George spec - imen SGDS.18.T1 (see, Milner et al., 2009). The pres ence of a well-pre served man ual im print in the pre - vi ously de scribed squat ting theropod im pres sion, AC 1/1 from the Lower Jurassic of Massachusetts (see, Gierliński, 1994), was later ques tioned by Lockley et al. (2003). Ac cord ing to the original interpretation, the Massachusetts specimen shows man ual toe traces, which in crease in length in the se quence: I, II, III. The same fea ture is ob served in the pres ent spec i men from Po land. As noted by Gierliński (1994), this fea ture seems important in understanding the origin of the Atreipus trackmaker (Olsen and Baird, 1986). Atreipodids are dis tinc tive Late Tri as sic ichnites, which com bine a small theropod-like (grallatorid) pes with the prim i - tive archosaurian (chirotheriid) re duced, tridactyl or tetradactyl manus. Fur ther more, their man ual dig its in crease in length in the clearly dif fer ent se quence: I, IV, II, III. Olsen and Baird (1986) have ar gued that such a com bi na tion might have been made by a basal ornithischian trackmaker, How ever, later au - thors, such as Thulborn (1990, 1993) and Weems (1992), sug - gested a theropod or i gin for those ichnites.

5 A distinctive crouching theropod trace from the Lower Jurassic of Poland 475 Fig. 4. Atreipus sp. from the Up per Tri as sic of Woźniki near Częstochowa, Po land, the spec i men found and owned by G. N. Recent discoveries support the hypothesis of Olsen and Baird (1986), Gierlinski (1994), and Milner et al. (2009) on the non-theropod af fin ity of the Atreipus trackmaker. It is also noteworthy that a potential atreipodid trackmaker candidate has al ready been found: an ornithischian-like dinosauromorph, such as the re cently-dis cov ered Silesaurus from the Up per Tri - as sic of Po land (Dzik, 2003). Silesaurus is a small, non-theropod, quad ru ped with a theropod-like pes. More over, Atreipus footprins were re cently found in the Up per Tri as sic strata of Po land (Fig. 4). REFERENCES DZIK J. ( 2003) A beaked her biv o rous archosaur with di no saur af fin i - ties from the early late Tri as sic of Po land. J. Vertebr. Paleont., 23 (3): GIERLIŃSKI G. (1994) Early Ju ras sic theropod tracks with the meta - tar sal im pres sions. Prz. Geol., 42 (4): LOCKLEY M., MATSUKAWA M. and LI J. (2003) Crouch ing theropods in tax o nomic jun gles: ichnological and ichnotaxonomic in - ves ti ga tions of foot prints with meta tar sal and ischial im pres sions. Ichnos, 10 (1): MIL N J., LOOPE D. B. and BROMLEY R. G. (2008) Crouch ing theropod and Navahopus sauropodomorph tracks from the Early Ju ras - sic Na vajo Sand stone of USA. Acta Palaeont. Pol., 53 (2): MILNER A. R. C., HARRIS J. D., LOCKLEY M. G., KIRKLAND J. I. and MATTHEWS N. A. (2009) Bird-like anat omy, pos ture, and be hav - ior re vealed by an Early Ju ras sic theropod di no saur rest ing trace. PLoS ONE, 4 (3): e4591. NIED WIEDZKI G. and REMIN Z. (2008) Gi gan tic theropod di no saur foot prints from the up per Pliensbachian of the Holy Cross Moun tains, Po land (in Pol ish with Eng lish sum mary). Prz. Geol., 56 (9): NIED WIEDZKI G., REMIN Z., ROSZKOWSKA J. and MEISSNER U. (2009) New finds of di no saur tracks in the Li assic strata of the Holy Cross Moun tains, Po land (in Pol ish with Eng lish sum mary). Prz. Geol., 57 (3): OLSEN P. E. and BAIRD D. (1986) The ichnogenus Atreipus and its sig nif i cance for Tri as sic biostratigraphy. In: The Be gin ning of the Age of Di no saurs: Fau nal Change Across the Tri as sic-ju ras sic Bound ary (ed. K. Padian): Cam bridge Uni ver sity Press. Cam bridge. OLSEN P. E., SMITH J. B. and Mc DON ALD N. G. (1998) The ma te rial of the spe cies of the clas sic theropod foot print gen era Eubrontes, Anchisauripus and Grallator (Early Ju ras sic, Hart ford and Deerfield bas ins, Con nect i cut and Mas sa chu setts, U.S.A.). J. Vertebr. Paleont., 18 (3): PIEŃKOWSKI G. (2004) The epicontinental Lower Ju ras sic of Po land. Pol. Geol. Inst. Spec. Pap., 12. THULBORN R. A. (1990) Di no saur tracks. Chap man and Hall, Lon don. THULBORN R.A. (1993) A tale of three fin gers: ichnological ev i dence re veal ing the homologies of man ual dig its in theropod di no saurs. In: The Nonmarine Tri as sic (eds. S. G. Lucas and M. Mo rales). New Mex - ico Mus. Nat. Hist. Sc. Bull., 3: WEEMS R. E. (1992) A re-eval u a tion of the tax on omy of New ark Supergroup saurischian di no saur tracks, us ing ex ten sive sta tis ti cal data from a re cently ex posed tracksite near Culpeper, Vir ginia. In: Proc. 26th Fo rum on the Ge ol ogy of In dus trial Min er als (ed. P. C. Sweet). Vir ginia Di vi sion of Miner. Res. Publ., 119: Char - lottes ville.

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