A di no saur track as sem blage from the Up per Hettangian (Lower Ju ras sic) mar ginal-ma rine de pos its of Zapniów, Holy Cross Moun tains, Po land
Grzegorz NIEDZWIEDZKI1, * and Grzegorz PIEÑKOWSKI2
1 Uppsala Uni ver sity, De part ment of Organismal Bi ol ogy, Evo lu tion ary Bi ol ogy Cen tre, Norbyvägen 18A, 752 36 Uppsala, Swe den
2 Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, Rakowiecka 4, 00-975 Warszawa, Po land
NiedŸwiedzki, G., Pieñkowski, G., 2016. A di no saur track as sem blage from the Up per Hettangian (Lower Ju ras sic) mar - ginal-ma rine de pos its of Zapniów, Holy Cross Moun tains, Po land. Geo log i cal Quar terly, 60 (4): 840–856, doi:
10.7306/gq.1310
Up per Hettangian (Lower Ju ras sic) mar ginal-ma rine Przysucha Ore-bear ing For ma tion ex posed at Zapniów mine and clay-pit (northwest ern part of the Holy Cross Moun tains area, cen tral Po land) re vealed an in ter est ing as sem blage of di no - saur tracks. Al though mostly yield ing poorly pre served and iso lated tracks prob a bly left in shal low wa ter con di tions, the site nev er the less in di cates a di ver sity of early saurischian (theropods and sauropodomorphs) and ornithischian (thyreophorans) di no saur trackmakers. This new as sem blage is partly con sis tent with the Lower-Mid dle Hettangian ichnorecord from the same re gion. The tracks are pre served in sed i ments of a bar rier-fore shore or i gin within a bar rier/la goonal depositional sys - tem, be long ing to the highstand sys tems tract, lo cated be low a marked ero sional sur face (se quence bound ary), which is as - so ci ated with a sub stan tial fall in sea level at the end of Hettangian times. Here we de scribed all pre vi ously and newly col lected or ob served in the field di no saur tracks from both sur face and un der ground ex po sures at Zapniów. Four ichnospecies: Grallator isp., Anchisauripus isp., cf. Tetrasauropus isp., and cf. Anomoepus isp. were iden ti fied. The theropod and ornithischian tracks show dis tinct sim i lar i ties to those de scribed from the rich est in this re gion and most fa mous Gliniany Las di no saur track as sem blage, in age ap prox i mately co eval to Zapniów. Two sizes of theropod tracks (small and me dium) in di cate the pres ence of two dif fer ent size classes or spe cies of pred a tors in this area. The de scribed cf.
Tetrasauropus isp. from Zapniów is the first un ques tioned ev i dence of basal sauropodomorphs in the Up per Hettangian of the Holy Cross Moun tains and first re cord of this ichnotaxa in the Lower Ju ras sic of Po land. Ad di tion ally, two theropod trackways (Anchisauripus isp.) show ev i dence for trot ting. The new finds sug gest sim i lar i ties be tween mar ginal-ma rine en vi - ron ments (delta-plain and fore shore-bar rier/la goon lithofacies) as so ci a tion of di no saurs con tain ing low-brows ing thyreophorans ac com pa nied by small or ju ve nile sauropodomorphs and small to me dium sized theropods. Pres ence of the ornithischian foot prints sug gests their prom i nent role as a ma jor com po nent in Mid dle-Up per Hettangian di no saur fau nas in mar ginal-ma rine en vi ron ments dom i nat ing in the re gion.
Key words: di no saur tracks, Przysucha Ore-bear ing For ma tion, Up per Hettangian, Lower Ju ras sic, Holy Cross Moun tains, Po land.
INTRODUCTION
Trace fos sils pro vide un par al leled in for ma tion about the be hav iour of ex tinct an i mals in the en vi ron ments in which they lived. One of the aims of ver te brate palaeoichnological stud ies must be the re con struc tion of the an i mal com mu ni ties rep re - sented by the trace fos sils. Re search of the di no saur ichno - fauna of the Lower Ju ras sic of the Holy Cross Moun tains in Po land il lus trates some of these pos si bil i ties (Gierliñski and Pieñkowski, 1999; Gierliñski et al., 2004). The in for ma tion that can be de duced from Holy Cross Moun tains ichnofaunas is
par tic u larly vi tal for our un der stand ing of the Early Ju ras sic tetrapod com mu nity ecol ogy, be cause of the scar city of skel e - tal re mains of this age (e.g., Shubin and Sues, 1991; Sues et al., 1994; Irmis, 2005; Smith et al., 2007). Glob ally, Early Ju - ras sic tetrapod as sem blages are re mark ably ho mo ge neous, rather dom i nated by di no saur taxa and it is well-vis i ble in body and trace fos sil re cord (e.g., Olsen and Galton, 1977, 1984;
Attridge et al., 1985; Sues and Reisz, 1995; Olsen et al., 1998;
Irmis, 2004).
The Lower Ju ras sic tetrapod track re cord is wide spread through out large area of the north ern Me so zoic mar gin of the Holy Cross Moun tains (see NiedŸwiedzki et al., 2009). First di - no saur foot prints in the Holy Cross Moun tains were dis cov ered in the Up per Hettangian de pos its of Gliniany Las (Karaszewski, 1969, 1975). Sub se quently, they were iden ti fied in nu mer ous ho ri zons of ter res trial and mar ginal-ma rine Lower Ju ras sic for - ma tions in lo cal i ties such as: So³tyków, Zapniów, Jakubów, Œmi³ów, Idzikowice, Gromadzice A and B (lower and up per), Podole, Œmi³ów, Kontrewers, Paszkowice, Starachowice,
* Corresponding author, e-mail: grzegorz.niedzwiedzki@ebc.uu.se Received: February 21, 2016; accepted: September 6, 2016; first published online: September 30, 2016
Szwa r szowice, Chyby, ¯arnów, Bielowice, Szyd³ówek (Gier - liñski and Potemska, 1987; Pieñkowski and Gierliñski, 1987;
Gierliñski and Sabath, 1998; Gierliñski and Sawicki, 1998; Gier - liñski, 1990, 1991, 1994, 1995a, b, 1996, 1997, 1999, 2004;
Gierliñski and Pieñkowski, 1999; NiedŸ wiedzki, 2003, 2006, 2011; Gierliñski et al., 2001a, b, 2004, 2005, 2009; NiedŸwie - dzki and NiedŸ wiedzki, 2001, 2004; Gierliñski and NiedŸwie - dzki, 2002, 2005; NiedŸwiedzki and Pieñkowski, 2004, 2014;
NiedŸwiedzki et al., 2005, 2006, 2008a, b, 2009; NiedŸwiedzki and Remin, 2008). This ichnofauna is im por tant for sev eral rea - sons. Firstly, this is one of the rich est re cord of post-ex tinc tion, end-Tri as sic ex tinc tion, ear li est Ju ras sic tetra pod ichnofauna in Eu rope and one of the few re cords of very large Early Hettangian theropod tracks world wide (Gierliñski et al., 2001b;
Pieñkowski et al., 2014). Sec ondly, ver te brate ichno fauna from the Holy Cross Moun tains area shows a uni que early orni - thischian trace fos sil di ver sity (Gierliñski, 1999; Gierliñski and Pieñkowski, 1999).
Di no saur trace fos sils from Hettangian (Zagaje, Sk³oby and Przysucha Ore-bear ing) for ma tions in clude ma te rial from a va - ri ety of theropod, sauropodomorphs and basal ornithischians.
How ever, still there were some gaps of knowl edge con cern ing di no saur ichnofaunas from the Up per Hettangian of the Holy Cross Moun tains due to sparse out crops. The only site con tain - ing nu mer ous and well-pre served di no saur tracks have been known from the Up per Hettangian strata, up per part of the Przysucha Ore-bear ing For ma tion, ex posed at Gliniany Las.
Few di no saur tracks have been re ported also from the same for ma tion ex posed at Zapniów (Pieñkowski and Gierliñski, 1987; Gierliñski, 1995b; Gierliñski and NiedŸwiedzki, 2005), but this tracksite has never been well-stud ied in terms of tracks, lack ing de tailed de scrip tions and fig ures. How ever, this site is sig nif i cant also for its palaeoenvironmental and palaeo eco logi - cal con text. Dur ing the Early-Mid dle Hettangian brack ish sea in - vaded the Holy Cross Moun tains area (Pieñkowski, 2004) and this gen eral change of Hettangian en vi ron ment had a pro found im pact on di no saur fau nal com po si tion (Gierliñski and Pieñ - kowski, 1999).
Be tween 2003 and 2005, new di no saur tracks were found in the Zapniów mine and clay-pit near Przysucha (51°18’53"N, 20°34’27"E), lo cated in the north ern mar gin of the Holy Cross Moun tains area (Fig. 1). Hith erto, only a poorly pre served iso - lated ornithischian (orig i nally de scribed as Moyeni sauropus isp.) track and nu mer ous theropod tracks (Anchi sauripus isp.) were re ported from this lo cal ity (Pieñ kowski and Gierliñski, 1987; Gierliñski, 1995a, b; Gierliñski and NiedŸwiedzki, 2005).
Newly dis cov ered ma te rial con sists of small and me dium sized theropod tracks, me dium sauro podomorph manus and pes tracks, and also two iso lated tracks of me dium-sized early ornithischians (pos si bly thyreo phorans). This new ma te rial from the Zapniów mine and clay-pit had never been ad e quately ex - am ined be fore.
The Zapniów un der ground mine and clay-pit (both now dis - used and closed) are sit u ated on the edge of the Mazowieckie voivodship (prov ince), near the bor der with the Œwiêtokrzyskie voivodship. The mine was es tab lished in 1975 and ex ploited high-qual ity ce ramic claystone (kaolinite rich in ter vals of the Przysucha Ore-bear ing For ma tion), un til it was closed in 2010.
The un der ground ex ca va tions reached a depth of 55 m and con sisted of sev eral kilo metres long sys tem of tun nels. Cur - rently (2011–2015), the clay-pit and un der ground mine are flooded and there is no ac cess to track-bear ing de pos its in all de scribed tracksites (A–D).
The first di no saur track was found at Zapniów (tracksite A;
51°18’80"N, 20°34’30"E) in 1985 (Pieñkowski and Gierliñski, 1987; Muz. PIG 1572.II.2, spec i men of ornithischian track pre -
vi ously de scribed as Moyenisauropus isp. – and herein as cf.
Anomoepus isp.). In 1993, was dis cov ered iso lated and poorly pre served me dium-sized theropod tracks [Anchi sauripus isp.;
Muz. PIG 1560.II.35; orig i nally de scribed as Gralla tor (Grallator) tenuis] in area lo cated close to the tracksite B (51°18’83"N, 20°34’32"E; see Gierliñski, 1995b).
Af ter many years of re search on the di no saur tracks from Hettangian and col lect ing new im por tant ma te rial from the Przysucha Ore-bear ing For ma tion ex posed at Zapniów, sys - tem ati za tion, up date and sum mary of hith erto pub lished re sults are needed. The new ma te rial pre sented here was dis cov ered in four places named as tracksite A–D, dur ing oc ca sional vis its in the Zapniów mine and clay-pit. Some of these spec i mens (tracksite C) were briefly de scribed by Gierliñski and NiedŸ - wiedzki (2005), but other tracks (tracksite A, B, D) have not been pre vi ously pub lished. The sur face from the tracksite C had been reburied in 2006 with rock de bris to pre vent the many tracks that were there from be ing dam aged by clay-pit ve hi cles.
We have iden ti fied the tracks to the ichnogeneric level and em - pha sized iden ti fy ing in di vid ual.
This pa per con trib utes to the knowl edge on Hettangian di - no saur ichnofaunas of the Holy Cross Moun tains and their re la - tions with en vi ron ment, as well as to the knowl edge on an Early Ju ras sic evo lu tion of dinosaurs.
GEOLOGICAL AND PALAEOENVIRONMENTAL BACKGROUND
The Lower Ju ras sic di no saur tracks of the north ern mar gin of the Holy Cross Moun tains oc cur within the six Lower Ju ras sic lithoformations (Fig. 2; Pieñkowski, 2004; NiedŸwiedzki et al., 2009). The Early Ju ras sic ep och rep re sents more than 25 Myr long pe riod of al ter nat ing non-ma rine and mar ginal-ma rine de - po si tion in the north ern mar gin of the Holy Cross Moun tains.
Dur ing this time the area was oc cu pied by sev eral flo ras (Barbacka et al., 2010, 2014; Pacyna, 2013) and fau nas, com - po si tion of which was strictly con trolled by ma rine trans gres - sions and cli ma tic changes (Gierliñski et al., 2004; Hesselbo and Pieñkowski, 2011; Pieñkowski et al., 2012).
The most di ver si fied di no saur track as sem blages from this re gion are mostly Hettangian and Late Pliensbachian in age, be cause out crops of con ti nen tal or mar ginal-ma rine de pos its of Sinemurian or Toarcian age are very rare (Gierliñski, 1995b;
NiedŸwiedzki et al., 2009). Herein, we de scribe di no saur foot - prints com ing from the Late Hettangian (ca. 199.5 Ma – Co hen et al., 2015) up per part of the Przysucha Ore-bear ing For ma tion (Fig. 2). Lithofacies and sed i men tary cyclicity de vel op ment, which was ob served in the Zapniów ex po sure is also known from fully-cored bore holes (Fig. 1). The near est, ref er ence one is Gródek OP-2 (Fig. 1), in which the in ter val 135–150 metres cor re sponds to Zapniów sec tion (see Pieñkowski, 2004;
http://www.pgi.gov.pl/docman-tree/publikacje-2/spe cial -pa pers/
/12/1546 -sp12-figury/file.html). The data from bore holes and out - crops al lowed high-res o lu tion lithological, biostratigra phical and se quence stra tig ra phy cor re la tion of the Hettangian strata (Pieñkowski, 2004).
Hettangian sec tion in north ern slopes of the Holy Cross Moun tains is ex panded, ex ceed ing in the depocentre 300 m. It is com posed of three for mal lithoformations: Zagaje For ma tion, it’s the low er most part of which com prise also some de pos its of the Rhaetian age, Sk³oby For ma tion and Przysucha Ore-bear ing For ma tion, ap prox i mately cor re spond ing to the Hettangian depositional se quence – num bered as the se quence I (Pieñ - kowski, 2004). Con ti nen tal Zagaje For ma tion and deltaic/bra ck - A dinosaur track assemblage from the Upper Hettangian (Lower Jurassic)... 841
ish-ma rine Sk³oby For ma tion are of Lower-Mid dle Hettangian age and rep re sent the transgressive sys tems tract (TST). The max i mum transgressive sur face oc curs in the Mid dle Hettan gian Sk³oby For ma tion and its up per most part, along with the over ly - ing Przysucha Ore-Bear ing For ma tion, rep re sent the progra - dational, highstand sys tems tract (HST). The end of progra - dational sed i men ta tion at the top of Przysucha Ore -Bear ing For -
ma tion is marked by the re gional ero sion sur face (Fig. 2). In most places, ero sion re moved sed i ments of the un der ly ing parasequences. Based on com par i son to the more com plete suc ces sion re vealed at the nearby Gródek OP-2 bore hole, at least 4 m or more metres of sed i ments of the up per most parasequence were re moved at the ero sional se quence bound - ary at Zapniów. This ero sional sur face is as so ci ated with a sub - Fig. 1A – lo ca tion of sites with the Lower Ju ras sic di no saur tracks in the Holy Cross Moun tains, Po land; B – a sketch map show ing lo ca tion of the Zapniów mine and clay-pit near Przysucha; C – de tailed map of the Zapniów mine and clay-pit area
with po si tions of di no saur track-bear ing sur faces
stan tial fall in sea level and it forms a typ i cal se quence bound ary, end ing progradational HST and com menc ing the TST be long ing to the next se quence II of a Lower Sinemurian age. Char ac ter of clay min er al ogy, palynomorphs, rare plant macroremains and also some in ver te brate body and trace fos sils point to warm and rather hu mid cli mate in the area stud ied, in the Late Hettangian times (Pieñkowski, 1985, 2004; Pacyna, 2013).
The Late Hettangian Przysucha Ore-bear ing For ma tion and cor re spond ing HST is com posed of in creas ingly pro grading
parasequences, point ing to progradational, re gres sive trend (Pieñkowski, 1983, 1991, 2004). Bar rier-la goon depositional cy - cles dom i nate (like in the Zapniów ex po sure, Figs. 2 and 3A), but also flu vial-dom i nated deltaic cy cles are com mon. Within the Pol - ish Ba sin, this for ma tion oc curs only in the Holy Cross Moun tains re gion, thus point ing to a spe cific char ac ter of this part of ba sin in the Late Hettangian times. Par tic u larly char ac ter is tic are la - goonal de pos its, rep re sented by si der it ic clays and mudstones form ing char ac ter is tic “ore-bear ing” ho ri zons. La goons were A dinosaur track assemblage from the Upper Hettangian (Lower Jurassic)... 843
Fig. 2A – over all sec tion of the Lower Ju ras sic of the Holy Cross Moun tains with po si tion of Zapniów sec tion; for more de tailed sec tion (135–150 m) in the ref er ence Gródek OP-2 bore hole see Pieñkowski (2004); http://www.pgi.gov.pl/docman - -tree/publikacje-2/spe cial-pa pers/12/1546-sp12-figury/file.html; B – lithostratigraphical-log of the Zapniów mine and clay-pit sec tion, show ing oc cur rences of some fos sils, sed i men tary struc tures and ho ri zons from where di no saur tracks have been in situ re corded (pro files af ter Pieñkowski, 2004, mod i fied)
broad but shal low and are of ten as so ci ated with marsh de pos its with palaeosol ho ri zons. Vast marsh/swampy ar eas de vel oped around la goon coasts pro vided high quan ti ties of iron. In the Holy Cross Moun tains subbasin, there are usu ally three ore-bear ing ho ri zons, cor re spond ing to three ep i sodes of wide spread de vel - op ment of la goon depositional sub sys tem. To wards the ba sin cen tre, which was lo cated in the NE Po land, con nec tions with the open ba sin were wider which led to for ma tion of more open embayments and mi grat ing bar ri ers, such as in Zapniów (Fig. 4).
The land rimmed by a delta sys tem, was lo cated fur ther to the N–NE, be yond the Nowe Miasto-I³¿a Fault Zone (Fig. 4), which formed the north ern tectonical frame of the Early Ju ras sic ba sin in the Holy Cross Moun tains area (Pieñ kowski, 2004). Embay - ment lo cated at Zapniów was open ing to the NW, to wards the more open brack ish ba sin, ex tend ing to the Cen tral Po land and fur ther to Pomerania, north Ger many and Den mark, where it mer ged with the open bo real sea (Pieñ kowski, 2004). Bar rier de - pos its at Zapniów are built of well- sorted quartz sand stones (are - nites) and form two con spic u ous lay ers, sep a rated by la goonal de pos its (Fig. 3A). The bar ri ers were flat, of a low-re lief char ac - ter. The lack of ae olian de pos its and traces of veg e ta tion, abun - dance of wave rip ple marks (Fig. 3B) and in ver te brate trace fos - sils, in clud ing Sko lithos, Mono craterion, Calycraterion, Lockeia, Cochlichnus (Fig. 3C), point to their partly sub merged/tem po - rarily emerged char ac ter. Mea sure ments of az i muths of wave rip ple crests at the top of the lower bar rier sand stone (rang ing be tween 100 and 125 de grees), al lowed for ap prox i mate es ti ma - tion of the bar rier shore line ori en ta tion, which was roughly par al - lel to the WWN–EES di rec tion.
Tem po rarily emerged bar ri ers pro vided good en vi ron ment both for di no saur pen e tra tion and pres er va tion of their tracks.
The Anomoepus–Moyenisauropus–Anchisauripus–Grallator track as so ci a tion was de fined as a typ i cal di no saur track as so - ci a tion for this kind of en vi ron ment (Gierliñski and Pieñ kowski,
1999). Di no saur foot prints in Zapniów come en tirely from the up per sur faces of two bar rier sys tems where they were left.
Most of the tracks were found on the top of the lower bar rier level (Fig. 2). Tracks were quickly filled with fine-grained la - goonal de pos its, which pro tected them from ero sion.
MATERIAL AND METHODS
Di no saur tracks found at Zapniów are pre served as true tracks (con cave epireliefs), undertracks and also as nat u ral casts on the bot tom sur faces of fine-grained, clayish or sid er - ite-rich sand stone. They show dif fer ent states of pres er va tion.
Some tracks con tain an a tom i cally con trolled struc tures (e.g., claw or pad im pres sions), other are with out such characters.
In June 2002 and May 2003, a track-bear ing sur face (tracksite C; 51°18’92"N, 20°34’42"E) was ex posed. Once the tracksite C, mea sur ing 28 m2, was fully ex posed (Fig. 5C), the foot prints were iden ti fied and the sur face was mapped onto a graph pa per. Seven spec i mens of Anchisauripus isp. foot prints were pre served on ex posed in situ sand stone sur face in the mine drive way lead ing to the front face of the Zapniów clay-pit (Gierliñski and NiedŸwiedzki, 2005). In 2004, the au thors found the sur face with two iso lated theropod di no saur tracks (Anchi - sauripus isp.) in the un der ground part of the Zapniów mine (tracksite D; level min ing 2, –12 m, tun nel B, about 23 m from the in ter sec tion with main tun nel). Both foot prints were left in the place of the dis cov ery (in the floor of min ing tun nel), but their pic tures and clay casts were taken.
The most nu mer ous is me dium-sized theropod track Anchisauripus isp., oc cur ring mainly in the form of true tracks or in part undertracks in two ho ri zons of the sec tion in Zapniów (Figs. 2 and 5–7). Four spec i mens have been found ex situ in
Fig. 3A – de pos its of the bar rier-la goon-embayment depositional sys tem from the Zapniów ex po sure near Przysucha (Late Hettangian, Przysucha Ore-bear ing For ma tion); B – sur faces with wave rip ple marks; C – Cochlichnus isp.,
si nu soi dal in ver te brate traces
Red ar rows – foot print-bear ing ho ri zons; f.s. – B – fore shore-bar rier lithofacies with Anchisauripus isp. tracks;
o.s. – L – embayment-la goon mudstone lithofacies; s.f. – B – shoreface heterolith-sand stone lithofacies with re cord of tem po ral emersions marked with di no saur tracks; scale bar – 3 cm
loose sand stone blocks with re cord of ripplemarks (Figs. 6B, D, E), nu mer ous in ver te brate bioturbation (Fig. 8A, C) or in the sid - er ite-rich sand stone sur face (Fig. 6C). Other spec i mens have been found in situ (Figs. 6A, 7 and 8B, D). Pho to graphs were taken un der nat u ral light in the field and mu seum (Geo log i cal Mu seum, Pol ish Geo log i cal In sti tute – Na tional Re search In sti - tute, War saw). Mea sure ments were taken on the orig i nal ma te - rial or casts (Ta ble 1) fol low ing the meth ods of Leonardi (1987) with mea sure ments of tridactyl foot prints digit ra tios [L/W, L/III, III/II, III/IV, te/fw, (fl-te)/fw] af ter Weems (1992) and Olsen et al.
(1998). The ichnological ter mi nol ogy used in this pa per for the de scrip tion of these di no saur tracks mainly fol lows the works of Thulborn (1990).
Two cal cu la tions were made from the mea sure ments. First, we cal cu lated the hip height of the Anchisauripus trackamker us ing the morphometric ra tios sug gested by Thulborn (1990) as fol lows: h » 4.5 x foot print length (for mula for foot prints smaller than 25 cm). Sec ond, a hip height was used to es ti mate speeds us ing an equa tion de vel oped by Al ex an der (1976). The equa - tion for speed is:
v = 0.25 g0.5 l 1.67 h –1.17
where: v – rep re sents ve loc ity, g – is the ac cel er a tion due to free fall, l – rep re sents stride length, h – rep re sents height of the an i mal at the hip).
A dinosaur track assemblage from the Upper Hettangian (Lower Jurassic)... 845
Fig. 4. Lo ca tion of the Zapniów mine and clay-pit in the north ern mar gin of the Holy Cross Moun tains area on the palaeo geo graphi cal back ground dur ing Late Hettangian; in set – gen eral palaeo geo graphi cal ex tent
of the Early Ju ras sic epicontinental bas ins in Eu rope (af ter Pieñkowski, 2004, mod i fied)
Orig i nal spec i men or plas ter casts of some spec i mens from Zapniów are housed in the Geo log i cal Mu seum of the Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute in War saw, Po land (ab bre vi a tion: Muz. PIG) and the Mu seum of Na ture and Tech nol ogy (for mer Mu seum of His tory of Ma te rial Cul ture) in Starachowice (abbreviation: MPT).
DESCRIPTION OF DINOSAUR TRACKS
THEROPOD TRACKS
Ichnofamily: Grallatoroidae Lull, 1904 Ichnogenus: Grallator Hitch cock, 1858
Grallator isp. (Fig. 6D, E) Fig. 5. Dis cov er ies of di no saur tracks in Zapniów mine and clay-pit, Przysucha Ore-bear ing For ma tion
(Up per Hettangian), Holy Cross Moun tains, Po land
A – tracksite C; two tracks of Anchisauripus isp. left on the top of lower bar rier de pos its (photo. G. Gierliñski); B – tracksite C; ex - ca va tion of the sur face with nu mer ous Anchisauripus isp. tracks (Zapniów, 2003; photo. G. Gierliñski); C – tracksite C; map ping of the sur face with nu mer ous Anchisauripus isp. tracks (Zapniów, 2003; photo. G. Gierliñski); D – tracksite D; ex a m i na tion of two Anchisauripus isp. tracks pre served on the top of sand stone (ar row) ex posed in un der ground mine (Zapniów, 2004); E – tracksite D; sur face with Anchisauripus isp. tracks (ar rows) pre served as true tracks (Zapniów, 2004)
M a t e r i a l. – Two un col lected slab with iso lated pes im - prints pre served on the sur face with ripplemarks and ob served in the tracksite A (orig i nal spec i mens left in field; 51°18’82"N, 20°34’28"E).
D e s c r i p t i o n. – Small sized (up to 120 mm long), digitigrade and func tion ally tridactyl pedal im prints pre served as nat u ral casts. The pedal digit ra tios are: III/II = 1.52–1.73 and III/IV = 1.05–1.12; L/III = 1.37–1.53; te/fw = 0.56–0.67;
(fl-te)/fw = 0.89–1.01.The to tal length to to tal width ra tio is 1.75–1.87. The digit II is shorter than IV, but both are shorter than III. Im pres sions of two phalangeal pads are vis i ble in digit II, three in digit III (Fig. 6E) and digit IV con tains more than three phalan geal pads. Divarication an gle of outer dig its is nar row: 30 and 35°.
D i s c u s s i o n. – Here we use the term Grallator in the tra di tional, re stricted, ichnotaxonomic sense, only to de scribe small, tridactyl foot prints that show strong sim i lar ity to var i ous New ark Supergroup ma te rial (Olsen et al., 1998). The Grallator ichnogenus is widely dis trib uted in the Lower to Up per Hettan - gian of the Holy Cross Moun tains (Gierliñski and Pieñkowski, 1999; NiedŸwiedzki, 2011). Very sim i lar small theropod tracks have been de scribed from the Przysucha Ore-bear ing For ma - tion ex posed at Gliniany Las (Gierliñski, 1996). The nar row divarication an gle is typ i cal of Grallator de scribed by Olsen et al. (1998). Grallator is also a long-rang ing di no saur ichnotaxon and there fore of lit tle biostratigraphic util ity. Tracks in the Zapniów ichnoassemblage have a mean foot length and width of 98 and 51 mm re spec tively. Most au thors agree that Grallator rep re sents a theropod di no saur, pos si bly a small theropod sim i - lar to Coelophysis and Megapnosaurus (Colbert, 1989; Irmis,
2004). The mean sizes of all Grallator foot prints col lected from the Przysucha Ore-bear ing For ma tion (Gliniany Las = 2, Pasz - kowice = 1, Zapniów = 2) sam ple are around 15–20% larger than those re corded from some clas si cal Up per Tri as sic site of North Amer ica (e.g., Lockley and Eisenberg, 2006; Lockley and Gierliñski, 2009). This sug gests that there are some dif fer ences be tween the Late Tri as sic and Early Ju ras sic rep re sen ta tives of the Grallator ichnogenus.
Grallator foot prints are rel a tively rare in the Lower Ju ras sic of Eu rope (Lockley and Meyer, 2000). What seems to be in ter - est ing is a com mon and rather wide spread ichnogenus in the Mid dle-Up per Tri as sic of Ger many and France (Haubold, 1984;
Klein and Lucas, 2010). This is con sis tent with the high di ver sity Hettangian ichnofaunas from Po land with rare Grallator and sug gests that typ i cal Grallator-dom i nated as sem blages dis ap - peared in this part of the world across the Tri as sic-Ju ras sic bound ary.
Ichnofamily: Grallatoroidae Lull, 1904 Ichnogenus: Anchisauripus Lull, 1904 Anchisauripus isp. (Figs. 6A–C and 7)
M a t e r i a l. – One col lected pes spec i men (Fig. 6C; Muz.
PGI 1560.II.35) from tracksite A; three plas ter cast (Fig. 7B, Muz. PGI 1688.II.1; Fig. 7D, MPT.P/6 and un reg is tered cast de - pos ited in the Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, War saw) of the pes spec i mens from the tracksite C (spec i mens in situ); clay cast (un reg is tered, de pos ited in the Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, War - saw) of the pes spec i men from un der ground mine (tracksite D, A dinosaur track assemblage from the Upper Hettangian (Lower Jurassic)... 847
Fig. 6. Theropod tracks from Zapniów mine and clay-pit near Przysucha, Przysucha Ore-bear ing For ma tion (Up per Hettangian), north ern mar gin of the Holy Cross Moun tains, Po land (pho to graphs and sche matic draw ings;
IV – po si tion of the digit num ber four)
A–C – Anchisauripus isp. (A – clay cast, un reg is tered, de pos ited in the Geo log i cal Mu seum, Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, War saw, spec i men from tracksite D; B – spec i men found ex situ and left in the field on the loose sand stone; C – Muz. PGI 1560.II.35, spec i men from tracksite A); D, E – Grallator isp., both spec i mens were found ex situ and left in the field on loose sand stone slabs; scale bar – 5 cm
Fig. 7. Theropod tracks from Zapniów mine and clay-pit near Przysucha, Przysucha Ore-bear ing For ma tion (Up per Hettangian), north ern mar gin of the Holy Cross Moun tains, Po land
(pho to graphs and sche matic draw ings; IV – po si tion of the digit num ber four)
A – sur face with nu mer ous Anchisauripus isp. tracks and map of the sur face with marked trackways (R – right, L – left im print); note ap prox i mately W–E di rec tion of the trackways, which is ap prox i mately par al lel to the shore line di rec tion, in ferred from az i muths of wave rip ples; B–D – plas ter casts of se lected Anchisauripus isp. tracks from the sur face ex posed at tracksite C (B – Muz. PGI 1688.II.1, plas ter cast; C – plas ter cast, un reg is tered; D – MPT.
P/6, plas ter cast); scale bar – 5 cm
spec i men pre served in situ) (Fig. 6A); iso lated pes (not col - lected) pre served in situ in tracksite D (not il lus trated); slab with iso lated pes pre served on sur face with ripplemarks and ob - served in the tracksite A (Fig. 6B; orig i nal spec i men left in field;
51°18’86"N, 20°34’40"E); four pes spec i mens (not col lected) pre served in situ on sur face in the tracksite C (Fig. 7A).
D e s c r i p t i o n. – Me dium sized (up to 235 mm long), digitigrade and func tion ally tridactyl pedal im prints pre served as true tracks or undertracks (Figs. 6A, B and 7) and nat u ral cast (Fig. 6C). The pedal digit ra tios are: III/II = 1.54–1.65 and III/IV = 1.10–1.16; L/III = 1.67–1.83; te/fw = 0.52–0.62; (fl-te)/fw = 1.13–1.32. The digit II is shorter than IV, but both are shorter than III. The to tal length to to tal width ra tio is 1.70–1.82.
Divarication an gle of outer dig its is nar row: 35–55°. The sur face with Anchisauripus isp. from tracksite C (Fig. 7A and Ta ble 1) shows seven foot prints (trackway I with two steps is 96 cm long;
trackway II with three steps is 193 cm long; two iso lated foot - prints), rep re sent ing prob a bly four an i mals (see Ta ble 1). Both trackways (I and II; Fig. 7A) trend ap prox i mately to wards the E–
EESast; in ter est ingly, this di rec tion is roughly par al lel to the WWN–EES di rec tion of palaeoshoreline, which was in ferred from az i muths of wave rip ples. It should be noted that the dif fer - ence in depth and mor phol ogy of these trackways sug gests that they were made at dif fer ent times. Two shal lower foot prints (first from trackway I, and one iso lated foot print) have in di ca - tions of pad and claw im pres sions (Fig. 7B, C). Other foot prints show lit tle more than the out line of the foot.
D i s c u s s i o n. – These me dium-sized and rel a tively nar row forms can be as signed to Grallator Hitch cock, 1858, fol - low ing grallatorid des ig na tion con cept of Lockley and Hunt (1995), or to the ichnogenus Anchisauripus sensu Olsen et al.
(1998). The pes mea sure ment ra tios of these spec i mens cor re - spond also to the ra tios of Grallator tuberosus sensu Weems, 1992. Olsen et al. (1998) di ag nosed this type of foot print (me - dium sized and nar row pes about length be tween 15 and 25 cm) as ichnogenus Anchisauripus. In Anchisauripus the digit III pro jects rel a tively fur ther an te ri orly than in small spec i mens of Eubrontes and Kayentapus, but not as far as in Grallator. We be lieve that the clas si fi ca tion of Olsen et al. (1998) better re - flects di ver sity of the Lower Ju ras sic theropod ichnotaxa. As has been pre vi ously re ported (NiedŸwiedzki and Pieñkowski, 2004) in com par i son to fre quen cies of other theropod ichnotaxa a me dium-sized theropod foot prints (Anchisauripus) are gen er - ally rare in the Lower Ju ras sic di no saur track as sem blages of Po land (Gierliñski, 1995a, b; NiedŸwiedzki, 2011; NiedŸwiedzki and NiedŸwiedzki, 2001, 2004; Gierliñski and NiedŸwiedzki, 2005). Newly de scribed Anchisauripus spec i mens from the Zapniów slightly in crease a gen er ally low fre quency of these ichnotaxa oc cur rences in the Lower Ju ras sic of the Holy Cross Moun tains area. Ichnogenus Anchisauripus rep re sents a theropod di no saur, pos si bly a me dium sized Coelophysis-like theropod.
Two Anchisauripus isp. trackways from the tracksite C, in di - cate trot ting be hav iour. Both trackways, have stride to hip A dinosaur track assemblage from the Upper Hettangian (Lower Jurassic)... 849
Fig. 8. Sauropodomorph and ornithischian tracks from Zapniów near Przysucha, Przysucha Ore-bear ing For ma tion (Up per Hettangian), north ern mar gin of the Holy Cross Moun tains, Po land (pho to graphs and sche matic draw ings;
IV – po si tion of the digit num ber four; ci – claw im pres sion)
A – cf. Tetrasauropus isp., ju ve nile or small sauropodomorph tracks, manus (m) and pes (p) tracks (spec i men left in the field); B–D – cf. Anomoepus isp., an early ornithischian tracks (B, C – spec i mens left in the field;
D – Muz. PGI 1572.II.2, plas ter cast); scale bar – 5 cm
height ra tios close to the trot (data in cluded in Ta ble 1) and this in di cates that both trackmakers were run ning. The ev i dence from the tracksite C at Zapniów is in agree ment with ob ser va - tions made by other work ers (e.g., Thulborn, 1989, 1990; Irby, 1996; Getty, 2005) in that di no saurs us ing a trot ting or run ning gait were small- to me dium-sized an i mals.
SAUROPODOMORPH TRACKS
Ichnofamily: Navhopodidae Olsen and Galton, 1984 Ichnogenus: Tetrasauropus Ellenberger, 1972
cf. Tetrasauropus isp. (Fig. 8A)
M a t e r i a l. – Un col lected slab with manus and pes im - prints pre served on the sur face with nu mer ous in ver te brate trace fos sils and ob served in the tracksite A (orig i nal spec i men left in field; 51°18’85"N, 20°34’39"E).
D e s c r i p t i o n. – Nat u ral cast of the pes-manus set of qua dru pe dal di no saur, plantigrade, with a tetradactyl pes. The pes is asym met ri cal. The pes is tetradactyl and elon gated, 317 mm long, 249 mm wide an te ri orly and 115 mm wide pos te - ri orly. The pos te rior part of pes shows heel-like area. Diva - rication be tween pes dig its is low. Digit I lies pos te ri orly with re - spect to dig its II–IV. Digit III ex tended for ward more than the oth ers. Manus smaller then pes, semi cir cu lar, 204 mm wide and 134 mm long. The manus is sit u ated anteromedially to the pes. The ra tios of the pes length to manus length equal 3.01;
pes width to manus width equals 2.37; pes length to pes wide (an te ri orly) equals 1.36.
D i s c u s s i o n. – The ichnogenus Tetrasauropus Ellen - berger, 1972 was de scribed for the first time from the Late Tri - as sic-Early Ju ras sic ichnofauna of south ern Af rica (Le sotho).
The ichnogenus was re vised by D’Orazi Porchetti and Nicosia (2007) and con sid ered as monotypic with T. unguiferus as the only valid type ichnospecies. The pres ence of Tetrasauropus out side Le sotho has of ten been in ferred, es pe cially in North Amer ica (e.g., Lockley and Hunt, 1993, 1995; Lockley et al., 2001; Gaston et al., 2003) but foot prints re sem bling Tetra - sauropus were de scribed from from the Up per Tri as sic (Norian–Rhaetian) of Swit zer land (Furrer, 1993), Up per Tri as - sic (Norian–Rhaetian) of Green land (Jenkins et al., 1994; Loc - kley and Meyer, 2000; Sulej et al., 2014), Up per Tri as sic of of South Wales (Lockley et al., 1996), Up per Tri as sic (Carnian) Portezuelo For ma tion of Ar gen tina (Marsicano and Barredo, 2004), Up per Tri as sic (Norian) of Po land (Gierliñski, 2007). The only re port about oc cur rence of Tetrasauropus in Early Ju ras sic strata (Corniola For ma tion) from Cen tral It aly (Arduini, 1996) has al ready been ques tioned (Manni et al., 1999). How ever, in the Lower Ju ras sic strata of north ern It aly there were sauro - podomorph tracks re ported (named as Lavinipes cheminii Avanzini, Leonardi et Mietto 2003), whose char ac ter is tics are com pa ra ble with those of Tetrasauropus (D’Orazi Porchetti and Nicosia, 2007).
Cf. Tetrasauropus isp. from Zapniów shows sev eral dis tinc - tive fea tures (Fig. 9). Manus was rest ing on the ground, in an an te rior and ex ter nal po si tion – with re spect to the pes. Asym - met ri cal pes and manus ex hibit a def i nite heteropody. Manus digit I mainly shows an im print of a large, bent claw. Ichnogenus Tetrasauropus is a re cord of a large quad ru ped and the cal cu -
T a b l e 1 Mea sure ments (in cm) of di no saur tracks from Zapniów mine and clay-pit
Spec i men Ichnotaxa TR TL TW TD Pa l h v
Left in the field Grallator isp. A 11.9 5.9 30 – – 53.5 –
Left in the field Grallator isp. A 9.5 5.6 35 – – 42.7 –
Un reg is tered, clay cast Anchisauripus isp. D 21.1 12.5 42 – – 94.9 –
Left in the field Anchisauripus isp. D 19.2 13.1 35 – – 86.4 –
Muz. PGI 1688.II.1, plas ter cast Anchisauripus isp. C (TR I) 22.8 12.5 39 77 154* 102.6 3.5
Left in the field Anchisauripus isp. C (TR I) 21.5 11.8 55 96.7
Left in the field Anchisauripus isp. C (TR II) 22.5 13.6 47 96 174 101.2 3.9
Un reg is tered, plas ter cast Anchisauripus isp. C (TR II) 22.3 14.8 36 100.3
MPT.P/6, plas ter cast Anchisauripus isp. C (TR II) 23.1 12.5 39 78 103.9
Left in the field Anchisauripus isp. C 23.5 14.2 42 – – 105.7 –
Left in the field Anchisauripus isp. C 23.2 13.5 39 – – 104.4 –
Muz. PGI 1560.II.35 Anchisauripus isp. A 17.2 10.2 46 – – 77.4 –
Left in the field Anchisauripus isp. A 20.5 12.8 42 – – 92.2 –
Left in the field cf. Tetrasauropus isp.
manus (pes) A 13.4
(31.7) 20.4
(24.9) – – – 155.3 –
Left in the field cf. Anomoepus isp. A 14.1 17.5 74 – – 63.4 –
Left in the field cf. Anomoepus isp. B 13.3 12.7 68 – – 59.8 –
TR – trackway, TL – to tal length, TW – to tal width, TD – to tal divarication, Pa – pace, l – stride, h – height at hip, * – stride that was de ter mined by mul ti ply ing a sin gle pace by two, v – ve loc ity in m/s
lated height at the hip of this trackmaker is about 2 metres, a to - tal body length is about 10 metres (D’Orazi Porchetti and Nicosia, 2007). Since first re ported, the ichnogenus has been re ferred both to dino saur ian taxa and to non-dino saur ian groups (see D’Orazi Porchetti and Nicosia, 2007).
Ellenberger et al. (1969, 1970) and Ellenberger (1970) sug - gested a large melanorosarid sauropodomorph as a sup posed trackmaker. Baird (1980), Olsen and Galton (1984), Haubold (1984), Thulborn (1990), An der son et al. (1998) and Lockley and Meyer (2000) re ferred Tetrasauropus to a prosauropod (=
sauropodomorph). Lockley et al. (2001) pro posed a sauro - podan af fin ity for the North Amer i can Tetrasauropus which, as we note, are ap par ently some what dif fer ent from the South Af ri - can type ma te rial. Rainforth (2003) con sid ers T. unguiferus from Le sotho as a poorly pre served Brachychirotherium, and im plic itly as a non-dino saur ian track. Re cently, D’Orazi Por - chetti and Nicosia (2007), based on di rect com par i son of foot - prints and manus and pes osteology, sug gested Yunnano - saurus sauropodomorph from the Early Ju ras sic of China as the best can di date to be the Tetrasauropus trackmaker.
THYREOPHORAN TRACKS
Ichnofamily: Anomoepodidae Lull, 1904 Ichnogenus: Anomoepus Hitch cock, 1848
cf. Anomoepus isp. (Fig. 8B–D)
M a t e r i a l. – Un col lected slab with pes im print pre served on the sur face with in ver te brate trace fos sils (Fig. 8C) and ob - served in the tracksite A (orig i nal spec i men left in field;
51°18’83"N, 20°34’37"E); un col lected pes im print pre served on the sur face with ripplemarks (Fig. 8B) in the tracksite B (orig i nal spec i men left in field; 51°18’83"N, 20°34’32"E); plas ter cast (Fig. 8D; Muz. PGI 1572.II.2) of the pes spec i mens dis cov ered in 1985 in the tracksite A (spec i mens was found in situ).
D e s c r i p t i o n. – Three iso lated small to me dium sized (130–160 mm long and 105–125 mm wide) tridactyl pes im - prints, pre served as nat u ral moulds (Fig. 8B, D) or nat u ral cast (Fig. 8C). All pes spec i mens are slightly lon ger than wide by 5–10%. The pro jec tion of the third digit be yond the lat eral dig its var ies from 12–25% of the whole pes length. All spec i mens show wide digit divarication: II–III = 35–40°, III–IV = 30–34°, II–IV = 65–74°. Dig its III and IV are subequal in length with small claw im prints (Fig. 8C), but in poorly pre served spec i men
(Muz. PGI 1572.II.2) the fourth digit is the lon gest. Digit II is clearly shorter than III and IV. Digit III is slightly curved in the end in two ob served spec i mens (Fig. 8B, C). All ichnites show two poorly pre served phalangeal pads on digit III.
D i s c u s s i o n. – The two foot prints (Fig. 8B, C) show an anomoepodid pat tern and typ i cal func tion ally tridactyl pes with rel a tively short dig its and also typ i cal Anomoepus sizes (Gier - liñski, 1991; Thulborn, 1994; Lockley and Gierliñski, 2006).
How ever, the spec i men Muz. PGI 1572.II.2 shows also some spe cific mor pho log i cal as pects (blunted dig its, short digit II, long digit IV), char ac ter is tic for ichnogenus Moyenisauropus Ellenberger, 1974. Two de scribed spec i mens (Fig. 8B, C) show fea tures iden ti fied in the Early Ju ras sic ichnospecies Anomo - epus pienkovskii Gierliñski, 1991, such as its size, pro por tions of digit length and rel a tively nar row dig its. Some au thors con sid - ered Ellenberger’s Moyenisauropus as a ju nior syn onym of Anomoepus (Olsen and Galton, 1984; Thulborn, 1994). Re - cently, Lockley and Meyer (2000), Olsen and Rainforth (2003) con tended that all ichnospecies of Moyenisauropus (ex cept Moyenisauropus karaszevskii Gierliñski, 1991) should be trans - ferred to Anomoepus – how ever, fur ther de bate on these tax o - nom i cal ques tions would be be yond scope of this pa per.
Pieñkowski and Gierliñski (1987) pro posed a hy po thet i cal
“wad ing and swim ming sce nario” to ex plain the or i gin of the first anomoepodid spec i men dis cov ered in the Zapniów (Fig. 8D;
Moyenisauropus isp. here in redescribed as cf. Anomoepus isp.). Lockley (1991) pro posed his “undertracks sce nario” in - stead, rea son ing that there were no con tin u ous trackways show ing tran si tion from com plete to in com plete tracks, left by an an i mal pro gress ing into deeper wa ter. Ac cord ing to Gierliñski and Pieñkowski (1999) the “undertracks sce nario” is un likely, be cause the tracks are pre served on two cor re spond - ing slabs: as a mould (slab Muz. PGI 1560.II.9A), and as a cast (slab Muz. PGI 1560.IL9B). On both track-bear ing sides one can ob serve the same mor pho log i cal pat tern of the same tracks and there are nei ther tracks nor undertracks on the op po site sides of these rel a tively thin slabs.
DISCUSSION
The Up per Hettangian mar ginal ma rine sed i ments in Zapniów mine and clay-pit are not rich in fos sils but nev er the - less yield quite di ver si fied di no saur track as sem blage (Fig. 10).
The di no saur ichnofauna de scribed from Zapniów com prises A dinosaur track assemblage from the Upper Hettangian (Lower Jurassic)... 851
Fig. 9. Com par a tive sil hou ettes of sauropodomorph pes tracks (I – first digit)
A – cf. Tetrasauropus isp. from the Up per Hettangian of Po land; B – Tetrasauropus Ellenberger, 1972 from the Late Tri as - sic–Early Ju ras sic of south ern Af rica (Le sotho); C – Lavinipes Avanzini, Leonardi et Mietto, 2003 from the Lower Ju ras sic of north ern It aly; scale bar – 10 cm (mod i fied from Ellenberger, 1972; Avanzini et al., 2003)
tracks of small and me dium theropods (Grallator isp., Anchi - sauripus isp.), large basal sauropodomorph (cf. Tetrasauropus isp.) and an ornithischian di no saur (cf. Anomoepus isp.), which prob a bly rep re sent early thyreophorans. No typ i cal swim ming traces (scratches) from ei ther di no saurs, croc o diles, tur tles or fishes have so far been found in the Zapniów ex po sure.
Dur ing Late Hettangian times, the area rep re sented mar - ginal ma rine set tings, lo cated be tween the emerged land and the shal low epicontinental sea (Fig. 4). It was area with shal low la goons, marshes, bar ri ers and other eco sys tems of mar - ginal-ma rine coastal plain and at tracted the di no saur biota. In the Zapniów sec tion di no saur tracks oc cur in fore shore – bar rier lithofacies, which prob a bly rep re sented a en vi ron ment fre - quented by these an i mals for com mu ni ca tion (likely long-shore wan der ing) and feed ing op por tu ni ties. How ever, as has been dem on strated above, the more de tailed study of these tracks pres ents some dif fi cul ties linked to the pres er va tion of in di vid - ual tracks and lon ger trackways.
Rel a tively short, less than 2 Myr (Co hen et al., 2015) Hettangian age is im por tant in terms of di no saur evo lu tion as the be gin ning of this age was as so ci ated with a rapid ra di a tion of this group. Nu mer ous work ers (e.g., Benton, 1986; Hunt, 1991;
Olsen et al., 2002; Brusatte et al., 2008, 2010) have drawn at ten - tion to a rel a tively sud den in crease in di no saur abun dance, di - ver sity and body size dur ing the lat est Tri as sic-ear li est Ju ras sic and this event is geo graph i cally wide spread and not lithofacies cor re lated, so prob a bly it is a re cord of evo lu tion ary events con - nected with ter mi nal Tri as sic ex tinc tion of therapsid - -pseudosuchian fau nas on the lands. In deed, the Hettangian track as sem blage from the Holy Cross Moun tains (Fig. 11) re - flects no ta ble di no saur di ver sity (Gierliñski and Pieñkowski, 1999; NiedŸwiedzki, 2011; Pieñkowski et al., 2014), both in con ti - nen tal and mar ginal-ma rine set tings. More over, the rich di no saur track as sem blages reg is ter an ear lier ap pear ance of cer tain di -
no saur groups and more rapid di no saur evo lu tion than it was be - lieved, based on North Amer i can re ports (i.e., Lucas, 1996;
Lockley et al., 1998; Padian, 1989 – see dis cus sion in Gierliñski and Pieñkowski, 1999). Mark edly, al lu vial plain and bar rier-la - goon, deltaic en vi ron ments reg is tered dif fer ent di no saur track as sem blages (Fig. 11). Both large early sauro pods and gi ant theropods are miss ing in the lat ter (Gierliñski and Pieñkowski, 1999), al though bit smaller basal sauropodomorphs (trace - makers of cf. Tetrasauropus isp.) and theropods (not big ger than Dilophosaurus) as well as ju ve nile sauro pods (Gierliñski, 1997;
NiedŸwiedzki and Pieñ kowski, 2004) could oc ca sion ally in vade these mar ginal-ma rine plains. Dis cern ing fac tor was sub strate sta bil ity, dis cour ag ing big ger and heavier an i mals, and low-rise veg e ta tion, fa vour ing spe cial ized, low brows ing ornithischians (early thyreo phorans), which were trace-mak ers of Anomoepus and Moye ni sauropus tracks. Small to me dium-sized theropods would fol low these her biv o rous di no saurs, leav ing Anchi sauripus and Grallator tracks. In ter est ingly, the ear li est Hettan gian tetrapod as sem blages of the Holy Cross Moun tains con tain, be - sides the most com mon di no saur tracks, ichnites of basal crocodylomorphs, early mam mals and other small rep tiles (prob - a bly lepidosauromorphs), while mar ginal-ma rine plains with del - tas, bar rier-la goon en vi ron ments are dom i nated by di no saur tracks. This show ing wide en vi ron men tal ad ap ta tion of di no saurs in their early evo lu tion ary his tory. No tracks from large theropod di no saurs have so far been found at the Zapniów lo cal ity, but tracks from rel a tively large theropods (Kayentapus isp.) are known from the nearby Gliniany Las tracksite of a sim i lar geo log i - cal age (Gierliñski and NiedŸ wiedzki, 2005), and it is likely that larger theropod tracks will turn up dur ing fu ture re search and ex - ca va tions in the Przy sucha Ore-bear ing For ma tion sites lo cated near Zapniów.
This di ver sity of early di no saur tracks in the Hettangian for - ma tions of the Holy Cross Moun tains re flects a di verse fauna,
Fig. 10. Di no saur ichnotaxa of the Przysucha Ore-bear ing For ma tion (Up per Hettangian) of Zapniów, Holy Cross Moun tains, Po land
Trackmakers: A – early ornithischians (pos si bly thyreophorans), B – sauropodomorph, C – small to me dium-sized theropods; TL – to tal length of tracks
A dinosaur track assemblage from the Upper Hettangian (Lower Jurassic)... 853
)ci ssa ruJ rewoL( naignatteH re ppU ot rewoL eht fo eli forp lacigolohtil dez il ar eneg eht ni skcart rua so nid fo sei cneu qerf dna segnar ,snoi ti sop cihpar gitartS .11 .giF no desab( dna loP ,snia tnuoM ssorC yloH eht fo ikswokñeiP ;4002 ,ikzdeiwŸdeiN ;9002 ,.la te ikzdeiwŸdeiN1102 ,) 1 .fc skcartdoporeht ci tna gig –supiruasolageMpsi2 ;. .fc skcartdoporeht egral –setnorbuEpsi .fc/.supatneyaKpsi3 ;.skcartdoporeht egral –supatneyaKpsi4 ;.dezis-mui dem – skcartdoporehtsupiruasihcnApsiskcartdoporeht llams – 5 ;.rotallarGpsi6 ;. .fc –xynonetSpsi7 ;.skcartekil-drib llams –sinroiselPpsi8 ;.egral –hpromodoporuas9 ;skcartllams – hpromodoporuas01 ;skcart–hpromodoporuas .fc skcartmuozotOpsi– 11 ;.hpromodoporuas .fc skcartsuporusarteTpsi .fc skcartnaihcsihtinro llams – 21 ;.sunhcirrotaleDpsi– 31 ;. skcart rua so nid naihcsihtinro dezis-mui demsupeomonApsi/.suporuasineyoMpsi41 ;.skcart rua so nid naihcsihtinro dezis-egral –suporuasineyoMpsi.
which is still poorly known based on osteological dis cov er ies (see NiedŸwiedzki, 2011). This makes the ichnoassemblage from the Przysucha Ore-bear ing For ma tion an im por tant win - dow into the Hettangian di no saur-dom i nated eco sys tem.
CONCLUSIONS
The com bined di no saur ichnofauna of Zapniów con sists of tracks of at least two theropods, a sauropodomorph and early ornithischian, pos si bly thyreophoran. This site re veals a coastal di no saur track as sem blage and con firms the ex is tence of the char ac ter is tic Lower Ju ras sic di no saur ichnofacies (Gierliñski and Pieñkowski, 1999; NiedŸwiedzki and Pieñkowski, 2004), dom i nated by small-me dium sized theropods and ornithi - schians. Palaeoecologically the oc cur rence of cf. Anomoepus isp. in mar ginal ma rine strata sup ports for mer hy poth e ses (Gierliñski and Pieñkowski, 1999; Gierliñski et al., 2009) that ornithischian di no saurs pre ferred coastal en vi ron ments and low-rise veg e ta tion. In ad di tion, Anchisauripus isp. trackways from tracksite C in di cat ing trot ting gaits of me dium-sized thero -
pods. The Zapniów ichnoassemblage en riches our knowl edge about di no saur com mu ni ties in the Late Hettangian, add ing me - dium-sized sauropodomorph as a fur ther com po nent. These finds con firm a high phylogenetical di ver sity of di no saurs, reached as early as in the Hettangian age. The Late Hettangian di no saur com mu ni ties were well es tab lished and de ter mined by en vi ron men tal fac tors.
Ac knowl edge ments. We are deeply in debted to M. Œwirski (Zapniów mine and clay-pit) for his help, ad vice and kind ness dur ing the field works (2002–2005) in the site. The au thors thank Dr. G. Gierliñski of the Pol ish Geo log i cal In sti tute – Na tional Re - search In sti tute for their valu able re marks which im proved the pa per. The re search and field works in 2003-2005 were sup - ported by the Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute. The study was sup ported by grant fi nanced from re - sources of the Pol ish Na tional Sci ence Cen tre, granted on the ba sis of de ci sion no. DEC-2012/06/M/ST10/00478. G.N. is cur - rently funded by grants awarded to Prof. Per E. Ahlberg (Uppsala Uni ver sity).
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