Early Jurassic sauropod footprints of the Southern Carpathians, Romania: palaeobiological and palaeogeographical significance

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Geo log i cal Quar terly, 2009, 53 (4): 461–470

Early Ju ras sic sauropod foot prints of the South ern Carpathians, Ro ma nia:

palaeo bio logi cal and palaeo geo graphi cal sig nif i cance

Grzegorz PIEŃKOWSKI, Mihai E. POPA and Artur KĘDZIOR

Pieńkowski G., Popa M. E. and Kędzior A. (2009) — Early Ju ras sic sauropod foot prints of the South ern Carpathians, Ro ma nia: palaeo - bio logi cal and palaeo geo graphi cal sig nif i cance. Geol. Quart., 53 (4): 461–470. Warszawa.

The di no saur foot prints cf. Parabrontopodus isp. Lockley, Farlow et Meyer, 1994, at trib uted to sauro pods, have been found in Hettangian (ear li est Ju ras sic) al lu vial de pos its in Anina (Colonia Cehă Quarry, Reşiţa Ba sin), be long ing to the Getic Nappe in the South - ern Carpathians, West ern Ro ma nia. Heteropodous pes-manus sets and one short, nar row-gauge trackway have been rec og nized on a large sand stone sur face tram pled by sauro pods. A greater load was car ried by the in ner dig its of the pes, par tic u larly digit I, and the heel pad is deeply im printed, which points to a sub-plantigrade pes and gravipodal pos ture, typ i cal for Eusaropoda. A pentadactyl manus im - print sug gests that manus dig its of early sauro pods might have been sep a rate and per haps more func tional when sup port ing walk ing on un sta ble, sticky ground. These tracks, the first find of Ju ras sic di no saur foot prints in Ro ma nia, add an im por tant site to the rel a tively rare re cord of ear li est Ju ras sic sauropod foot prints. These tracks also con firm that Pangaean is lands and pen in su las around the West ern Tethys were in hab ited by early sauro pods. These pen in su las or is lands, in clud ing a hy po thet i cal “Moesian Is land”, must have been at least tem po rarily con nected with the main land. The sizes of the Ro ma nian foot prints are sim i lar to the Hettangian Parabrontopodus isp.

tracks de scribed from Po land (main land Pangaea — Eur asian area) and It aly (Tethyan do main) and do not in di cate in su lar dwarf ism.

Grzegorz Pieńkowski, Pol ish Geo log i cal In sti tute–Na tional Re search In sti tute, Rakowiecka 4, PL-00-975 Warszawa, Po land; e-mail:

grzegorz.pienkowski@pgi.gov.pl; Mihai E. Popa, Uni ver sity of Bu cha rest, Fac ulty of Ge ol ogy and Geo phys ics, Lab o ra tory of Palae on - tol ogy, 1, N. Balcescu Ave., 010041, Bu cha rest, Ro ma nia; e-mail: mihai@mepopa.com; Artur Kędzior, Pol ish Acad emy of Sci ence, In - sti tute of Geo log i cal Sci ences, Kraków Re search Cen tre, Senacka 1, PL-31-002 Kraków, Po land; e-mail: ndkedzio@cyf-kr.edu.pl (re ceived: Oc to ber 27, 2009; ac cepted: De cem ber 12, 2009).

Key words: Parabrontopodus, sauro pods, Ro ma nia, Hettangian, gravipodal pos ture, palaeobiogeographyn.

INTRODUCTION

Rel a tively very few un equiv o cal ex am ples of sauropod tracks are known in Early Ju ras sic strata world wide (Lockley et al., 1994b), even though true sauro pods (Eusauropoda — Upchurch, 1993, 1994) bones are known from this ep och, such as Vulcanodon karibaensis Raath 1972 from Af rica (Raath, 1972; Cruickshank, 1975; Coo per, 1984) and frag men tary pre - served re mains from other con ti nents, in clud ing Sanpasaurus, Zizhongosaurus and Kunmingosaurus from China (Dong et al., 1983; Dong, 1992), Ohmdenosaurus from Ger many (Wild, 1978), and Barapasaurus from In dia (Jain et al., 1979). The most im por tant sites with Early Ju ras sic sauropod foot prints com prise Hettangian sites in North ern It aly (Leonardi and Lanzinger, 1992; Dalla Vecchia, 1994; Avanzini and Petti, 2008; Avanzini et al., 2008) and Cen tral Po land (Gierliński, 1997; Gierliński and Sawicki, 1998; Gierliński and Pień kowski,

1999; Gierliński et al., 2004). Be sides Hettangian strata, sauropod tracks have been also found in Pliensbachian strata of Mo rocco (Ishigaki, 1989; Farlow, 1992). Finds of both body and trace fos sils, scat tered around the globe, sug gest that Early Ju ras - sic sauro pods were geo graph i cally wide spread. How ever, true sauropod (Eusauropoda) foot prints and body fos sils are ab sent from Early Ju ras sic de pos its in North Amer ica (Hunt et al., 1994;

Olsen et al., 2002), sug gest ing the ex is tence of a bar rier sit u ated along the Cen tral At lan tic Rift, which pre vented mi gra tion of di - no saurs to North Amer ica from the rest of Pangaea (Fig. 1). An al ter na tive ex pla na tion would in volve cli ma tic dif fer ences, com - bined with hab i tat re quire ments of sauro pods (drier con di tions in North Amer ica), caus ing bioprovincialism of di no saurs. The Hettangian time in ter val was an im por tant stage in the evo lu tion of sauro pods, also in terms of prin ci pal pos ture changes in the fore- and hindlimbs, which in flu enced the shape of their foot - prints (Wil son and Sereno, 1998). There fore, any find of sauropod tracks of that age is sig nif i cant in an swer ing both

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Fig. 1. A — Lo ca tions of the Reşiţa Ba sin and Colonia Ceh² Quarry on a geo log i cal map of the Anina Anticline; B — Early Ju ras sic palaeo ge ogra phy with ma jor sites of Early Ju ras sic sauro pods — bone re mains (Zim ba bwe) and foot prints;

C — out lined area of “B” show ing palaeo ge ogra phy of the West ern Tethys with in ferred sauropod mi gra tion routes Palaeo ge ogra phy af ter Golonka, 2004 (sim pli fied, mod i fied)

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palaeo bio logi cal and palaeobiogeographical ques tions of the early evo lu tion of these an i mals.

Thus we are pleased to an nounce the dis cov ery of the di no - saur foot prints cf. Parabrontopodus isp. Lockley, Farlow et Meyer, 1994, at trib uted to sauro pods in Hettangian (ear li est Ju - ras sic) al lu vial de pos its at Anina (Colonia Cehă Quarry and Reşiţa Ba sin), in the South ern Carpathians of West ern Ro ma - nia. To date, no Ju ras sic di no saur foot prints have been found in Ro ma nia. Re ports of Cre ta ceous di no saur foot prints were pub - lished by Vremir and Codrea (2002) from Maastrichtian con ti - nen tal beds of the Sebeş area (Transylvanian De pres sion), while ear lier Cre ta ceous foot prints of un known af fin ity were de scribed by Koch (1900) from Neocomian beds of Lăpuş (Maramureş area). Di no saur bones (Benton et al., 1997) were de scribed from Early Cre ta ceous (Berriasian) beds of the Brusturi-Cor net area, north west ern Ro ma nia, while bones and eggs (Grigorescu, 2005; Grigorescu and Csiki, 2008) were de - scribed from Up per Cre ta ceous (Maastrichtian) beds of the Haţeg Ba sin (Transylvania). The small size of these di no saur re mains from the Haţeg Ba sin points to an in su lar dwarf ism of these an i mals (Grigorescu, 2005; Grigorescu and Csiki, 2008), as well as for those of the Brusturi-Cor net area (Benton et al., 2006). Our new ev i dence, how ever, shows nor mal-sized sauro - pods and hence no signs of dwarf ism.

GEOLOGICAL SETTING

Anina (South ern Carpathians, West ern Ro ma nia), for merly known as Steierdorf, was an im por tant coal min ing lo cal ity in the South Carpathians, oc cur ring in the cen tral area of the Reşiţa Ba sin, Getic Nappe, and 35 km south of Reşiţa (Fig. 1).

This area has been in ten sively mined since 1792 for Lower Ju - ras sic coals, and later for re frac tory clays and bi tu mi nous shales, all of these ac tiv i ties be ing al most to tally halted to day.

The min ing was un der taken through ver ti cal shafts, ex ten sive and in tri cate un der ground works, and large open-cast mines, such as the Ponor, Colonia Cehă and Hildegard quar ries (Popa, 2009). The Colonia Cehă Quarry is an im por tant open-cast mine in the area, next to the Steierdorf old neigh bor hood of Anina, and was opened for the ex trac tion of bi tu mi nous shales in the early 1980s. The di no saur tracks cf. Parabrontopodus isp. Lockley, Farlow et Meyer, 1994 de scribed in this pa per were re corded from its east ern and north ern flanks, along steeply in clined bed ding sur faces, ex posed in the up per most lev els of the quarry and cut by trans verse faults (Figs. 2 and 3).

The Getic Nappe (Murgoci, 1905) rep re sents one of the main struc tural units of the South Carpathians in Ro ma nia, a nappe yield ing both a crys tal line base ment and a sed i men tary cover.

The Getic Nappe oc curs be tween the Supragetic Nappe (west - wards) and the Severin and Danubian units (eastwards), as parts of a nappe pile overthrust from west to wards east in the South Carpathians. The Reşiţa Ba sin, also known as the Reşiţa–Moldova Nouă Sed i men tary Zone, rep re sents the larg est sed i men tary ba sin of the Getic Nappe, oc cur ring in the South Carpathians, be tween Reşiţa and the Dan ube River (Fig. 1). This ba sin in cludes de pos its be long ing to Palaeozoic (Up per Car bon - if er ous–Lower Perm ian) and Me so zoic (Lower Ju ras sic–Lower

Cre ta ceous) strata, un con form ably over ly ing the crys tal line base ment of the Getic Nappe. Dur ing Early Ju ras sic times, the Reşiţa Ba sin with the de pos its of the whole Getic Nappe be - longed to a large Moesian–Rhodope Plat form, and ac cord ing to palaeo geo graphi cal maps (Scotese, 2002; Golonka, 2004;

Golonka et al., 2005; Blackley, 2009), de vel oped on a shelf area with is lands and pen in su las, in clud ing an emer gent “Moesian Is - land”, at lat i tudes of about 30 de grees N (Fig. 1).

The Me so zoic sed i men tary suc ces sion be gins with the con - ti nen tal Steierdorf For ma tion (Bucur, 1991, 1997; Popa and Kędzior, 2008; Fig. 2), un con form ably over ly ing the crys tal - line base ment or the Up per Palaeozoic strata, and be ing con - form ably over lain by the brack ish-ma rine Uteriş For ma tion (Pliensbachian–mid dle Toarcian). The whole Steierdorf For - ma tion is 250 m thick in the north, and it de creases to 50–60 m south wards, reach ing about 150–160 m in thick ness in the area of the Colonia Cehă Quarry. The age of the Steierdorf For ma - tion is Hettangian–Sinemurian (Bucur, 1991, 1997; Popa, 2000a, b, 2009; Popa and Kędzior, 2008). How ever, be cause of a lack of di ag nos tic fos sils in its lower part, the low er most part of the Steierdorf For ma tion (Dealul Budinic Mem ber, at least in its lower part) may be older (?Up per Tri as sic/Rhaetian). At the top of the Dealul Budinic Mem ber, a pyroclastic layer oc - curs, in di cat ing a vol ca nic event. Above this layer, con ven tion - ally ac cepted as the bound ary marker be tween the Dealul Budinic Mem ber and suc ceed ing Valea Tereziei Mem ber, the col our of sed i ment changes into grey. Fur ther more, some 10 m above (Fig. 2), the first coal seams ap pear, point ing to a sig nif i - cant cli ma tic change.

The sauropod tracks oc cur just above the pyroclastic level mark ing the bot tom of the Valea Tereziei Mem ber, at the top of a fine-grained sand stone (Figs. 2 and 3). The Valea Tereziei Mem ber is late Hettangian–Sinemurian. In the Colonia Cehă Quarry, the low er most part of the Valea Tereziei Mem ber (in - clud ing the foot print-bear ing ho ri zon) may be mid dle Hettangian, be cause this part still lacks age-di ag nos tic fos sils;

the in dex flo ral re mains ap pear some 10 metres above, in the coal-bear ing strata. The Valea Tereziei Mem ber (ex cept for its low er most 10 m) is the main coal-bear ing unit of the Steierdorf For ma tion, con tain ing black, car bo na ceous sand stones, mud - stones, and clays of var i ous types, with abun dant fos sil plants in di cat ing a up per Hettangian (a range zone with Thaumato - pteris brauniana) to Sinemurian age (an acmezone with Nilssonia cf. orientalis) — Popa (1997, 2000a, b, c, 2005, 2009); Popa and Van Konijnenbrug-Van Cittert (2006), Popa and Kędzior (2006). The ex cep tional fos sil con tent in di cates Anina as a fos sil Lagerstatte lo cal ity, both for di ver sity and for de gree of pres er va tion (Andrae, 1855). The clos est com pa ra ble flo ras to those of the Ro ma nian Early Ju ras sic flo ras be long ing to the Reşiţa, Sirinia, or other bas ins in the South Carpathians are the co eval as sem blages of Iran and Af ghan i stan, which are also the clos est flo ras palaeogeographically (Givulescu, 1998;

Popa, 1998, 2000a). The Valea Tereziei Mem ber con tains 8 main bi tu mi nous coal seams and a char ac ter is tic re frac tory clay bed and shows a con stant strati graphic po si tion and a wide lat - eral de vel op ment in the cen tral part of the Reşiţa Ba sin, thus serv ing as a valu able strati graphic marker for the Hettangian-Sinemurian bound ary (Fig. 2). The re frac tory clay bears a rich pol len and spore as sem blage (Antonescu, 1973),

Early Jurassic sauropod footprints of the Southern Carpathians, Romania: palaeobiological and palaeogeographical significance 463

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Fig. 2. Geo log i cal/sedimentological pro files of the flu vial Steierdorf For ma tion with po si tion of sauropod foot prints shown

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and marks a tem po rary but wide spread lac us trine sed i men ta - tion ep i sode within a gen er ally flu vial/al lu vial plain-swamp en - vi ron ment. Be sides rich plant fos sils with pre served or gans, in - ver te brate trace fos sils as well as ver te brate tracks (Batrachopus cf. deweyi) have been found in the Valea Tereziei Mem ber (Popa, 2000a, c, 2009).

SEDIMENTOLOGICAL BACKGROUND OF THE STEIERDORF FORMATION

(HETTANGIAN–SINEMURIAN)

The lower part of the Dealul Budinic Mem ber rests on an ero sional sur face, over ly ing ei ther ?Perm ian redbeds or crys tal - line base ment. It is mainly com posed of poorly sorted, ma - trix-sup ported con glom er ates and coarse-grained sand stones (Fig. 2). Peb bles (as much as 5 cm in di am e ter) are an gu lar to sub-an gu lar and are scat tered through out sand stone lay ers.

Coarse-grained units are usu ally structureless, and in some places show large-scale cross strat i fi ca tion. Oc ca sion ally, up - per sur faces of sand stone bod ies have po lyg o nal struc tures with sec ond ary fer ric ox ide crusts, in dic a tive of des ic ca tion.

Fine-grained de pos its are sub or di nate, and are rep re sented by hor i zon tally lam i nated, red dish mudstones. Ad di tion ally, a fire clay ho ri zon within this se quence has been found. These fea tures in di cate high vis cos ity flow, typ i cal of al lu vial fans.

The age of the basal se quence of the Dealul Budinic Mem ber can not be pre cisely iden ti fied, due to the lack of fos sils, so it may be ei ther of Late Tri as sic (Rhaetian) or ear li est Ju ras sic (Hettangian) age. The up per part of the mem ber shows a tran si - tion to a more sta ble, lower-en ergy sed i men tary re gime. The de pos its be come finer, the first drifted flo ral re mains oc cur, and palaeosol ho ri zons ap pear. As the en ergy of de po si tion de - creased, al lu vial fans changed into an al lu vial plain en vi ron - ment. In the up per most, still red dish part of this mem ber, ver te -

brate bur rows have been found (Popa and Kędzior, 2006). Just above these bur rows the pyroclastic level oc curs, mark ing the bound ary be tween the two mem bers.

The Valea Tereziei Mem ber is com posed mainly of sand - stones, with a few con glom er atic lay ers and sub or di nate mudstones, claystones and coal seams. Flo ral re mains are very rich and well-pre served, ex cept in the low er most part of the mem ber. Sand stone bod ies are char ac ter ized by large-scale trough cross-bed ding and part ing lineations, point ing to a high en ergy of river flow. Rip ple lami na tions oc cur at the top of some sand stone lay ers. Sand stone lay ers oc cur ring within fine-grained de pos its (at trib uted to cre vasse-splay de pos its) show hor i zon tal or rip ple-drift lami na tions and are strongly bioturbated by roots. Drifted logs are abun dant and ori ented par al lel to the bed ding. Tree roots and trunks in ver ti cal, life po - si tion have also been found. Sand stone bod ies show channelized ge om e tries; they of ten pinch out lat er ally af ter short dis tances (~15 m). Traces of for est fires (fre quent char - coal frag ments) are also pres ent, and the so called “burnt coal-seam” forms a lo cal cor re la tive ho ri zon. These fea tures point to a hu mid al lu vial plain en vi ron ment, with al ter nat ing flu vial and lac us trine/swamp sed i men ta tion. Hu mid cli mate con di tions dur ing the de po si tion of the Valea Tereziei Mem ber are also in di cated by the flo ral re mains (Popa, 1997, 2000a, b, c, 2005, 2009; Popa and Van Konijnenbrug-Van Cittert, 2006;

Popa and Kędzior, 2006). It should be noted, how ever, that the Colonia Ceh² Quarry is af fected by a dense net work of faults, thus dif fer ent parts of the pro file, in volv ing both the up per most Dealul Budinic and the low er most Valea Tereziei mem bers, oc cur in lat eral con tact with each other (Fig. 3).

The sauropod foot prints, which oc cur at the top of the first sand stone layer of the Valea Tereziei Mem ber, are cov ered by a grey mudstone con tain ing plant roots in its top most part. The sauropod tracks were prob a bly left on the top of a wide spread cre vasse-splay/cre vasse-delta layer.

Fig. 3. Front and lat eral view of the track-bear ing layer at the Colonia Ceh² Quarry

Note trans verse faults ex pos ing dif fer ent strati graphic lev els; v.b. — ver te brate bur rows of the up per most Dealul Budinic Mem ber (Popa and Kędzior, 2006); s.t. — sauropod tracks shown on Fig ure 4; s.tw. — sauropod trackway shown on Fig ure 5 — the sur face is densely tram pled by sauro pods

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SYSTEMATIC DESCRIPTION OF THE DINOSAUR FOOTPRINTS

cf. Parabrontopodus isp. Lockley, Farlow et Meyer, 1994 (Figs. 4, 5)

(type ichnosp. Parabrontopodus mcintoshi Lockley, Farlow et Meyer, 1994)

M a t e r i a l. — Two plas ter casts LPB III ICH 76 (pes-manus set) and LCB III ICH 77 (two-steps trackway) of orig i nal sur faces in the Colonia Cehă Quarry (Fig. 3), reposited at the Uni ver sity of Bu cha rest, Fac ulty of Ge ol ogy and Geo - phys ics, Lab o ra tory of Palae on tol ogy, 1, N. Balcescu Ave., 010041, Bu cha rest, Ro ma nia.

D e s c r i p t i o n. — LPB III ICH 76 — pes-manus sets (Fig. 4) and LCB III ICH 77 (Fig. 5) — a short, nar row-gauge sauropod trackway. Pes and manus foot print of small size, pes ca. 40 cm long, 25 cm wide, manus ca. 15 cm wide and 10 cm long. Pes is oval, strongly nar rowed in its pos te rior (heel) part.

Manus oval (Fig. 4) or star-like, pentadactyl (Fig. 5). Pro - nounced heteropody, manus/pes area ra tio 1:5 or less. Pes im - prints show out wardly ro tated claw (or ungual) im pres sions cor re spond ing to digit I, II, III and IV. Digit I is ro bust, greater than the other dig its. Lead ing mar gin of the pes print (the me - dial side of the pes) is deeper that the trail ing (lat eral) mar gin of the pes. Heel pad is dis tinctly im pressed. All the tracks are sur - rounded by dis place ment rims.

The trackway (Fig. 5) is nar row-gauged, and shows marked heteropody. Both pes and manus are ro tated out wardly, the sec - ond pes-manus spac ing is shorter. Foot prints are deeply im - pressed, the first pes is sur rounded by a high dis place ment rim, the sec ond one was filled by upwelling, sat u rated sed i ment.

The first pes is 32 cm long, the sec ond one lacks a clear heel pad be cause of oblit er a tion caused by sed i ment upwelling. The first manus is pre served as an undertrack (up per part of the sand stone layer is miss ing); sec ond manus well pre served and more or less pentadactyl in shape. The in ferred midline of the trackway is curved, per haps due to pro gres sion on an un sta ble sur face. The pes stride length for the step is ca. 80 cm, mea - sured be tween the mid dle points of two pedal im prints and the pes di ver gence an gle, rel a tive to the in ferred midline, is ir reg u - lar, from 0° (pos te rior pes) to 38° (an te rior pes).

D i s c u s s i o n. — The track-bear ing sand stone sur face at the base of the Valea Tereziei Mem ber is densely tram pled by sauro pods, which makes indentification of in di vid ual tracks and trackways dif fi cult. How ever, in two places (Fig. 3) the last gen er a tion of tracks al low one to rec og nize two prob a ble pes-manus sets in ferred to rep re sent a nar row-gauge trackway (Figs. 4 and 5). Par tic u larly, one pes foot print with digit im - prints is di ag nos tic (Fig. 4, where pes-manus set no. 2 is in the mid dle). The pro nounced heteropody may also be di ag nos tic of Parabrontopodus isp. Lockley, Farlow et Meyer, 1994 — here ten ta tively la belled cf. Parabrontopodus. The best pre served pes no. 2 (Fig. 4) shows other char ac ter is tic fea tures. A ro bust digit I is typ i cal of the Eusauropoda, which can also point to a large first ungual (Wil son and Sereno, 1998). More over, Wil - son and Sereno (1998) men tioned that the assymetrical ro bust -

ness of pedal dig its must be re lated to the un equal weight dis tri - bu tion across the sub-plantigigrade pes in Eusauropoda. In - deed, in the LPB III ICH 76 spec i men (Fig. 4), digit I is the dom i nant digit of the pes and the me dial (in ner) side of the pes is deeper, while the heel pad is sub stan tial. These fea tures are also ob served in trackways pro duced by eusauropods (Pittman and Gilette, 1989) and may re flect how a greater load was car - ried by the in ner dig its of the pes of these sauro pods. The sig - nif i cance of these fea tures is that the col lapse from digitigrade to a sub-plantigigrade (or gravipodal) pos ture was an im por tant turn over in the evo lu tion of sauro pods as the Eusauropoda evolved from basal Sauropoda in the Early Ju ras sic times (Wil - son and Sereno, 1998). Early and mid dle Hettangian sauropod foot prints from Po land (Gierliński, 1997; Gierliński and Sawicki, 1998; Gierliński and Pieńkowski, 1999; Gierliński et al., 2004; Pieńkowski, 2004) show sub stan tial heel pad and con spic u ous traces of pes digit I. There is also a dif fer ence in depth be tween the deeper me dial and shal lower lat eral parts of pedal im prints in sev eral tracks. Such a dif fer ence is not clear in the pho to graphs of sauropod tracks from It aly, where pes prints sug gest rather sym met ri cal dis tri bu tions of pres sure (Leonardi and Lanzinger, 1992; Leonardi and Avanzini, 1994; Dalla Vecchia, 1994; Avanzini and Petti, 2008; Avanzini et al., 2008). The Ro ma nian foot prints are sim i lar in age to the Ital ian ones (mid dle–late Hettangian) and are some what youn ger than most of those from Po land. Ro ma nian finds sug gest that a sub-plantigrade (gravipodal) pos ture of sauro pods de vel oped prob a bly by mid dle Hettangian times.

The nar row-gauge trackway (LCB III ICH 77, Fig. 5) is short, in ferred to be com posed of only two manus-pes sets, and was left in soft, sat u rated sed i ment, as shown by deeply-im - pressed tracks, high sed i ment-dis place ment rims, and in one case in fill ing of the track with sat u rated sed i ment. Be cause of the ir - reg u lar step, one might in fer a chang ing pes di ver gence an gle, and the bent midline of the short trackway. How ever, pre cise mea sure ments of trackway pa ram e ters could not be taken. Ir reg - u lar ity of the trackway may per haps point to an ir reg u lar pro - gres sion of the trace pro ducer, caused by a soft, un sta ble sub - strate. The track-bear ing layer could be fur ther dis torted by early diagenesis. The first or prox i mal pes is very deeply im pressed, and the cor re spond ing manus is rep re sented by an undertrack as the up per part of the sand stone layer is miss ing in this part of the trackway. The next pes is in com plete (due to sed i ment upwelling), whereas the sec ond manus is well pre served and has re tained a pentadactyl shape with im pres sion of dig its. Digit im - prints in sauropod manus tracks are rarely pre served (Lockley et al., 1994a), al though noted from sev eral lo cal i ties (Lockley et al., 1992; Santos et al., 1994; Leonardi and Avanzini, 1994;

Gierliński and Sawicki, 1998; Avanzini and Petti, 2008;

Avanzini et al., 2008; Santos et al., 2009). Gierliński and Sawicki (1998) and Gierliński and Pieńkowski (1999) il lus trated early Hettangian sauropod trackways of both adult and ju ve nile sauro pods, and in the adult sauropod trackways, some of the manus im prints were clearly pentadactyl. The manus de scribed herein bears re sem blance also to the Late Tri as sic Eosauropus isp. (Lockley et al., 2006), pro duced by mem bers of the Sauropodomorpha (Hunt and Lucas, 2007). Dalla Vecchia (1999) and Dalla Vecchia and Tarlao (2000) sug gested three sauropod manus morphotypes based on the con fig u ra tion of

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digit I — the manus de scribed herein is most sim i lar to their Morphotype A, which com prises manus prints with a well-de vel oped im pres sion of digit I (see also Lockley et al., 1992, their fig. 2:

print 3). As sum ing a soft and sat u rated sub strate, such pres er va tion might sug - gest that man ual dig its of early sauro - pods might oc ca sion ally sep a rate or be splayed, per haps pro vid ing better sup - port on un sta ble ter rain. Sev eral au thors be lieve the sauropod manus prob a bly func tioned as a sin gle, rigid, block-like struc ture with no intermetacarpal move - ments (McIntosh, 1990; Upchurch, 1994; Bonnan, 2003), but the non-tu bu - lar, prim i tive manus of ear li est sauro - pods might have func tioned in a dif fer - ent way.

The ma te rial is con sis tent in terms of shape and size with known Hettan - gian Parabrontopodus isp. trackways re ported from North ern It aly (Leonardi and Lanzinger, 1992; Dalla Vecchia, 1994; Avanzini and Petti, 2008;

Avanzini et al., 2008) and Cen tral Po - land (Gierliński, 1997; Gierliński and Sawicki, 1998; Gierliński and Pień - kowski, 1999). Re cently, Santos et al.

(2009) dis cussed Sauropodomorpha ichno- morphotypes — most of the fea - tures of our ma te rial (nar row-gauge trackway, pes shape and its out wardly di rected claw marks, high heteropody) fits the Parabrontopodus- like morpho - type. Fur ther more, the sin gle penta - dactyl manus im print would rather fit their new Polyonyx-like morphotype (Santos et al., 2009).

PALAEOGEOGRAPHIC IMPLICATIONS

The palaeoclimatic and palaeogeographic in ter pre ta tion is based largely on the rich Early Ju ras sic (Hettan - gian–Sinemurian) flora found in the coal-bear ing Valea Tereziei Mem ber. Givulescu (1998) re ferred the flo ral as sem - blage to var i ous co eval en vi ron ments of Iran and Af ghan i stan (north ern Tethyan rim). This au thor also sug gested that the palaeovegetation in Anina oc curred along sea shores and beaches, al though gen er at ing coals in pure limnic, fresh wa ter con di tions. Semaka (1962) and Givulescu (1998) em pha sized the paralic fea tures of the Steierdorf For ma tions, fol low ing pre - vi ous geo log i cal works (Răileanu et al., 1957) de scrib ing the coal mea sures of the Steierdorf For ma tion as Gresten-type de - pos its. How ever, any fea tures sup port ing sea shore prox im ity were not dem on strated by the au thors. The first un equiv o cal ma rine in flu ences in the Me so zoic of the Reşiţa Ba sin oc cur in the Uteriş For ma tion (Pliensbachian–Mid dle Toarcian in ter val

— Bucur, 1991, 1997; Popa, 2000a, 2009). There fore, an in ter - pre ta tion of a con ti nen tal intramontane (limnic) ba sin was pro - posed by Popa (2000a) and Popa and Kędzior (2008). Ac cord - ing to these au thors, dur ing Hettangian times, the bas ins of the Getic Nappe (Reşiţa) and of the Danubian Units (Sirina, Presacina and Cerna-Jiu) were con ti nen tal, closely re lated to each other at a short dis tance.

Popa (1997, 1998), Popa and Van Konijnenburg-Van Cittert (2006) and Givulescu (1998) in di cated that the flora of Anina be longs to the Eurosinian Re gion, Eu ro pean Prov ince sensu Vakhrameev (1991). Popa and Van Konijnenburg-Van Cittert (2006) in ter preted the palaeolatitude of the Reşiţa Ba sin as be - tween 20° and 30° north. Mateescu (1958) re corded sec ond ary xy lem tis sues with pseudo-an nual growth rings in fusains from Rudăria, in the Sirinia Ba sin, Danubian Unit, a neigh bor ba sin of the Reşiţa Ba sin in the Hettangian, cit ing sim i lar fea tures in wood tis sues from Anina, Doman (Reşiţa Ba sin) and from the Vul can-Codlea area (Holbav Ba sin). These rings in di cate mon - soonal con di tions at this time, an ap pro pri ate model for the north ern frame of the Tethys realm. Such pseudo -an nual growth

Fig. 4. Iso lated sauropod tracks (last track gen er a tion on the heavily tram pled sur face), deeper parts of the foot prints shad owed: note the pes-manus set (p2–m2), pes with four dig its im - printed, and prom i nent digit I; me dial (in ner) part of the pes is sig nif i cantly deeper, as in all pes tracks on the sur face, and were pro duced by dif fer ent sauropod in di vid u als

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rings were also re corded in permineralized co ni fers from Holbav, Holbav Ba sin, part of the Getic Nappe, in di cat ing a sim - i lar cli mate. Popa (1998, 2000a) con cluded that the Early Ju ras - sic flora of the Reşiţa Ba sin oc curred on the north ern frame of the Tethys realm, within a typ i cal limnic intramontane de pres - sion, with out paralic in flu ences, whereas paralic in flu ences were stron ger in the bas ins of the Danubian Units.

The sauropod tracks come from the low er most 10 m of the Valea Tereziei Mem ber. This part of the sec tion does not con - tain coal seams, thus, the cli ma tic con di tions of Ro ma nian sauro pods could have been some what drier than the con di tions pre vail ing dur ing de po si tion of the main coal mea sures in the Reşiţa Ba sin. Likely, it could have been a warm and hu mid sub - trop i cal (mon soonal) cli mate, per haps with a drier sea son.

Al though rel a tively few un equiv o cal nar row-gauge sauro - pods trackways are re corded from Early Ju ras sic strata world - wide (Hettangian: Leonardi and Lanzinger, 1992; Dalla Vecchia, 1994; Gierliński, 1997; Gierliński and Sawicki, 1998;

Gierliński and Pieńkowski, 1999; Avanzini and Petti, 2008;

Avanzini et al., 2008, and Pliensbachian strata of Marocco:

Ishigaki, 1989; Farlow, 1992; see also Lockley et al., 1994b), it is clear that sauro pods roamed this re gion, in both the Pangaean (Af ri can, Eur asian) main land and sup posed is land/pen in sula ar - eas within the west ern/north ern Tethys do main (Fig. 1). The

palaeogeographic re con struc tions of Blackley (2009), Golonka (2004) and Golonka et al. (2005) show that the area which is now the Ro ma nian Carpathians in cluded large is lands, em brac - ing both Hun gar ian and Ro ma nian coal-bear ing for ma tions (Fig. 1). The palaeogeographic map of Scotese (2002) shows a rather pen in su lar area, but this map is much more gen er al ized.

As men tioned above, in the Early Ju ras sic, the Getic Unit would be long, to gether with the Danubian Units, to the larger Moesian Plat form, in clud ing the “Moesian Is land” of Golonka (2004).

Such an is land could pro vide a hab i tat for lush sub trop i cal plants and an i mals, but this hy po thet i cal is land must have been at least tem po rarily con nected to main land Pangaea to al low mi gra tion of sauro pods. Mi gra tion routes of sauro pods would cross the west ern and north ern frames of West ern Tethys, in clud ing Af - rica and Eur asia (Fig. 1). More over, the size of the sauropod cf.

Parabrontopodus isp. foot prints from Ro ma nia is typ i cal of Early Ju ras sic sauro pods and does not dif fer from those from Po - land (main land Pangaea — Eur asia) and It aly (Tethyan car bon - ate plat form/sea shore). Thus, the size of the Ro ma nian foot - prints would speak against in su lar dwarf ism, with the lat ter con - di tion ex pected in the case of long-last ing is land con di tions. This is con trary to the finds of Maastrichtian di no saurs of the “Haţeg Is land” in Ro ma nia, where di no saur bone re mains clearly in di - cate in su lar dwarf ism (Bojar et al., 2005; Csiki and Grigorescu,

Fig. 5. Short, one-step (two manus-pes sets) trackway with in ferred midline, dis torted in the soft, sat u rated sed i ment Note bent midline, chang ing stride and di ver gence an gle (prob a bly re sult ing from pro gres sion on an un sta ble sur face) and deeply im pressed foot prints with high sed i ment dis place ment rims; pentadactyl manus of the sec ond set (m2) shown in the in sert photo

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2008). Pol ish and Ro ma nian Early Ju ras sic sauropod foot prints come from a con ti nen tal al lu vial en vi ron ment, whereas Ital ian and Mo roc can ones come from car bon ate tidal flat fa cies. This sug gests a wide spec trum of en vi ron ments roamed by early sauro pods and a great mo bil ity of these an i mals. As sum ing this mo bil ity and the var ied en vi ron ments, one can sup pose that the ab sence of sauro pods from North Amer ica in the ear li est Ju ras sic times was rather caused by a per ma nent ob sta cle (rift) be tween this part and other parts (Af rica, Eur asia) of the frag ment ing Pangaean land mass.

CONCLUSIONS

Early Ju ras sic cf. Parabrontopodus isp. tracks and an in - ferred nar row-gauge trackway de scribed herein rep re sent the first Ju ras sic di no saur foot prints de scribed in Ro ma nia. Ro ma - nian foot prints are of sim i lar age to Ital ian sauropod tracks (mid dle to up per Hettangian) and they are some what youn ger than the early to mid dle Hettangian tracks from Po land. Asy - metrically im pressed pes prints with sub stan tial heel pads in di - cate that a greater load was car ried by the in ner dig its of the pes of sauro pods dur ing walk ing. This points to a sub-plantigrade (gravipodal) pos ture of the trace maker, and sug gests that such a pos ture of sauro pods (Eusauropoda) de vel oped from digiti - grade to a sub-plantigrade by the mid Hettangian. The penta - dactyl manus may in di cate that the manus dig its were oc ca sion - ally func tional in Eusauropoda, per haps pro vid ing a better sup -

port when walk ing on un sta ble, sticky ground. In the ear li est Ju ras sic, the “Moesian Is land” was at least tem po rarily at tached to the Pangaean (Eur asian) main land. More over, the nor mal sizes (for Hettangian times) of the Ro ma nian sauropod foot - prints speak against in su lar dwarf ism of these an i mals. Based on our find and on those of other sauropod tracks in the re gion, sauro pods roamed both the north ern and the south ern rim of West ern Tethys. This wide palaeogeographic and palaeo - environmental dis tri bu tion of sauropod foot prints in di cates that the ab sence of sauropod foot prints ob served in Early Ju ras sic strata of North Amer ica was per haps caused by an geo graph ical ob sta cle (rift), rather than by cli ma tic fac tors.

Lastly, we ex pect that this find of di no saur foot prints can con trib ute to the de vel op ment of geotourism and to the lo cal econ omy in this part of Ro ma nia. Per spec tives of geotourism de - pend on fu ture sci en tific re search on di no saur tracks and on other fos sils such as plant and in ver te brate re mains in the re gion.

Ac knowl edge ments. The au thors wish to thank Prof. M.

Sandulescu, Prof. T. Neagu and Dr. Z. Csiki, Uni ver sity of Bu - cha rest, for use ful dis cus sions and com ments. This pa per was sup ported by the CNCSIS (NURC) grant no. 436/1.10.2007 (978) to one of the au thors (MEP). We ex press our thanks to A.

Mar tin and M. Lockley for their most help ful re views and valu - able com ments. This is a con tri bu tion to the IGCP pro ject 506

“Ma rine and Non-ma rine Ju ras sic: Global Cor re la tion and Ma - jor Geo log i cal Events”.

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