Jurassic Dinosaur Footprints from Southern Italy: Footprints as Indicators of Constraints in...

534 RESEARCH REPORT

Copyright Q 2005, SEPM (Society for Sedimentary Geology) 0883-1351/05/0020-0534/$3.00

Jurassic Dinosaur Footprints from Southern Italy:Footprints as Indicators of Constraints in

Paleogeographic Interpretation

MARIA ALESSANDRA CONTIDipartimento di Scienze della Terra, Universita ‘‘La Sapienza’’, P.le A. Moro 5, 00185, Roma, Italy

MICHELE MORSILLIDipartimento di Scienze della Terra, Universita di Ferrara, C.so Ercole I d’Este 32, 44100, Ferrara, Italy

UMBERTO NICOSIA*, EVA SACCHIDipartimento di Scienze della Terra, Universita ‘‘La Sapienza’’, P.le A. Moro 5, 00185, Roma, Italy;

Email: [email protected]

VINCENZO SAVINO, ALEXANDER WAGENSOMMER, LEONARDO DI MAGGIOSpeleo Club Sperone, San Giovanni Rotondo, 71013, Foggia, Italy

PIERO GIANOLLADipartimento di Scienze della Terra, Universita di Ferrara, C.so Ercole I d’Este 32, 44100, Ferrara, Italy

PALAIOS, 2005, V. 20, p. 534–550 DOI 10.2110/palo.2003.p03-99

Three loose blocks, rich in dinosaur footprints, were foundin a small pier at Mattinata (Gargano Promontory, Foggia,Italy), most probably quarried from the Upper JurassicSannicandro Formation. All of the footprints in the blocksare ascribed to medium-sized theropod trackmakers. Re-cent track discoveries from both the Early Cretaceous SanGiovanni Rotondo Limestone and the Late Cretaceous Al-tamura Limestone, as well as this new discovery, reveal theconsistency of terrestrial associations along the southernmargin of the Tethys Ocean in the peri-Mediterranean areaat the end of Jurassic through Cretaceous times. The pres-ence of these dinosaur-track-rich levels within marine sedi-ments of the Apulia Platform underlines the relevance of di-nosaur footprints as a means of constraining paleogeo-graphic reconstructions.

INTRODUCTION

In early 2001, two of the authors (V.S., A.W.) found di-nosaur footprints on the surface of three loose limestoneblocks in a small pier that protects the entry to the smallharbor of Mattinata on the Gargano Promontory (Puglia,southern Italy, Fig. 1). The blocks were removed and arestored in the Mattinata town museum (Museo Civico Sto-rico-Archeologico). This is the third find of dinosaur foot-prints in the Apulia region, where several single dinosaurfootprints and trackways have been found previously. Thefirst site was found in 1999 in the Murge area near thetown of Altamura (Andreassi et al., 1999; Nicosia et al.,2000a, b; Dal Sasso, 2003), and other footprints were dis-covered in the Gargano area (Gianolla et al., 2000a; Dal

*Corresponding Author

Sasso, 2003). The discovery of terrestrial animal tracksimprinted in marine carbonates is considered important,and the Mattinata footprints appear distinct from the ma-terial already known from this same region.

GEOLOGICAL AND STRATIGRAPHICAL SETTING

The Gargano Promontory is part of a large paleogeo-graphical unit known as the Apulia Carbonate Platform,which, during the Mesozoic, was part of the southern mar-gin of the Tethys Ocean. The Apulia Platform is consid-ered one of the so-called peri-Adriatic platforms that arequite similar in facies architecture, size, and shape to themodern Bahamian Banks (Bernoulli, 1972; D’Argenio,1976; Eberli et al., 1993). The Gargano Promontory andother parts of the Apulia Region make up part of the fore-land of the Apennine chain. Structurally, the Garganoarea is folded into a gentle anticline with a WNW axis(Martinis, 1965). This broad structure contains numerousfaults with various trends and kinematics (Funiciello etal., 1992; Bertotti et al., 1999). The most prominent struc-tural feature is the Mattinata fault, a regional E–W shearzone that crosses the entire Gargano area, and continuesoffshore for many tens of kilometers (De Dominicis andMazzoldi, 1989).

The Gargano area, together with Maiella Mountain, isthe only place where the transition between platform andslope-to-basin facies crops out. In other areas, the easternmargin of this platform lies offshore under the AdriaticSea, about 20 to 30 km from the present coastline (DeDominicis and Mazzoldi, 1989).

The stratigraphy of the Gargano area has been reinter-preted frequently in the last two decades. In the late1960s–early 1970s, geological surveys introduced manyformational units (Martinis and Pavan, 1967; Merla et al.,1969; Cremonini et al., 1971), complicating stratigraphicalcorrelations in this area. Other reviews and original stud-

DINOSAUR FOOTPRINTS FROM SOUTHERN ITALY 535

FIGURE 1—Location map of the Gargano Promontory with main roads and towns.

FIGURE 2—Chronostratigraphic chart showing the Upper Jurassic–Lower Cretaceous formations of the Gargano Promontory in an idealizedE–W transect. 1 5 inner-platform facies; 2 5 margin facies; 3 5 slope to base-of-slope facies; 4 5 basin facies; 5 5hiatuses.

ies defined new stratigraphical schemes for this area con-cerning platform and slope-to-basin units (e.g., LupertoSinni and Masse, 1986, 1994; Bosellini et al., 1993, 1999;Claps et al., 1996; Luperto Sinni, 1996; Bosellini and Mor-silli, 1997, 2001), and proposed new models for the evolu-tion of this carbonate platform (Masse and Borgomano,1987; Bosellini et al., 1993, 1999; Morsilli and Bosellini,1997).

The successions cropping out were divided into differentsecond-order stratigraphical sequences, bounded by un-conformities, of different types and origins (see Bosellini etal., 1999, for a review). Herein, the focus mainly is on thelithostratigraphical units that contain shallow-water sed-iments in which footprints of terrestrial reptiles couldhave been impressed.

The main sequences that contain shallow-water sedi-ments are the Monte Sacro Sequence and the MattinataSequence (sensu Bosellini et al., 1999; Fig. 2). During theLate Jurassic–Early Cretaceous, two inner-platform unitswere deposited in the Gargano area: the Sannicandro For-mation and the San Giovanni Rotondo Limestone. Inner-platform facies of Late Cretaceous age also are present insome small outcrops, and have been named the Calcari diCasa Lauriola (Merla et al., 1969); these units correspondto the well-known Altamura Limestone (Laviano and Ma-rino, 1998; Bosellini and Morsilli, 2001).

Sannicandro FormationThe Sannicandro Formation is poorly known when com-

pared to other units in the area because previous works in-

536 CONTI ET AL.

cluded part of this formation in the San Giovanni RotondoLimestone (Mattavelli and Pavan, 1965; Pavan and Pirini,1966; Cremonini et al., 1971). More recent investigations(Luperto Sinni and Masse, 1986; Claps et al., 1996) onlyexamined the Lower Cretaceous succession of inner-plat-form facies. Luperto Sinni and Masse (1994) proposed thatthe Sannicandro Formation should be confined to inner-platform facies of Jurassic age, and other units (Formazio-ne di Monte La Serra, Calcari di Sannicandro, and Calcaridi Rignano; sensu Cremonini et al., 1971) should be includ-ed into the Sannicandro Formation. Based on a Valangi-nian-age-drowning unconformity (Bosellini and Morsilli,1997), Morsilli and Bosellini (1997) suggested that theSannicandro Formation could be extended to include theBerriasian–Early Valanginian interval.

The Sannicandro Formation crops out only in the west-ern and central sectors of Gargano. The base of this unit isunknown in outcrop, and it is overlain by the San Giovan-ni Rotondo Limestone. Its lateral eastern boundary is dif-ficult to map because it passes very gradually into theMonte Spigno Formation. The exposed thickness is esti-mated to be at least 400–500 m (Morsilli, 1998).

The lowermost-known tract of this unit crops out in thearea of Rignano Garganico (description after Morsilli,1998). At the base of the unit, thick beds of whitish, peloi-dal-bioclastic mudstone to wackestone with oncoids alter-nate with peloidal-bioclastic packstone beds (10–30 cmthick). This lower part of the unit is frequently dolomi-tized. At the top of the unit, there are peritidal cycles (1–2m thick) composed of peloidal wackestone, with localizedpackstones containing micritic intraclasts, and cryptomi-crobial laminites (sensu Demicco and Hardie, 1994) show-ing a planar or domal shape. The same lithofacies organi-zation has been recognized in the Monte Calvo area nearSan Giovanni Rotondo, where it is associated with thinbeds of oolitic grainstone that become more frequent to-wards the transition to the adjacent Monte Spigno For-mation. In this area, there also are numerous fenestralstructures and small lenses of flat-pebble breccia (clasts3–4 cm). The upper part of the Sannicandro Formationmainly consists of subtidal lithofacies and rare, thin pla-nar-stromatolite beds. The main textures are peloidalwackestone and bioclastic packstone, locally very rich indasycladalean algae (Campbelliella striata and C. milesi).The microfossil association partially covers the Callovian–Berriasian time interval, but the peritidal facies indicatesa Kimmeridgian–Tithonian age. The facies associations ofthis formation could be referred to an inner-platform set-ting with protected lagoons, tidal-flats, and supratidal de-positional environments, sometimes affected by stormevents (Morsilli and Bosellini, 1997).

Many of the lithofacies in the Sannicandro Formationare the same as the ones observed on the Mattinatablocks. Dinosaur footprints from this formation have notbeen found in place, but more research could fill this gap.Therefore, the potential presence of dinosaur footprints orbones in the Monte Spigno Formation, in particular in thetransition area with the Sannicandro Formation, cannotbe excluded.

San Giovanni Rotondo Limestone

This unit has been studied in detail both in the typearea (Luperto Sinni and Masse, 1986, Claps et al., 1996)

and in the Apricena-Poggio Imperiale area (Luperto Sinniand Masse, 1986). This unit partially corresponds to theBari Limestone (Azzaroli et al., 1968; Ricchetti, 1975) interms of age and sedimentary environments, and repre-sents Lower Cretaceous, inner-platform facies of the Apu-lia Platform.

The San Giovanni Rotondo Limestone is a thick succes-sion (500–600 m) of shallow-water limestone that can besubdivided into three members (see Claps et al., 1996 for amore detailed description). Member 1 (Borgo CelanoMember of Luperto Sinni and Masse, 1986) consists of amonotonous and acyclic subtidal unit with meters-thickmudstone to wackestone that is intensely bioturbated.This member can be referred to a shallow-subtidal, la-goonal setting. Member 2 (Loferitic Member of LupertoSinni and Masse, 1986) is a thick, cyclic unit characterizedby quasi-periodic alternation of loferitic beds and centi-meter-thick layers of green shale. Many lithofacies havebeen documented in this unit: mudstone–wackestone beds(0.5–2 m thick), intensely bioturbated with occasionalblack pebbles at the base, green clayey interlayers,planar-cryptomicrobial or domal-shape laminites (LLH-type sen-su Logan et al., 1964), and thin lenses of ooidal grainstone.Sometimes karst infilling, characterized by reddish shales(terra-rossa-like soils) with floating limestone clasts, dis-conformably cuts the previous lithofacies. These featurescan be referred to a tidal-flat setting with subaerial expo-sure. Member 3 consists of various lithofacies, such asthin-bedded packstone to grainstone with peloids, bio-clasts, and micritic intraclasts that are normally gradedand evenly laminated. Lags of ostreids, requienids, andnerineids also may occur. The peritidal cycles consist ofpeloidal–bioclastic packstone to wackestone at the base,and planar stromatolites (20–100 cm) at the top. There arelens-shaped beds (30–70-cm thick) of clast-supported car-bonate breccia with clasts of various lithofacies and size(reaching 20–25 cm). The age of the San Giovanni RotondoLimestone is between Late Valanginian and Early Aptian(Claps et al., 1996).

Many dinosaur footprints were discovered in this for-mation in a quarry near the Borgo Celano section (Gian-olla et al., 2000). They occur in three distinct layers ofMember 2 of the San Giovanni Rotondo Limestone. Mem-ber 2 belongs to the Campanellula capuensis Zone of LateHauterivian–Early Barremian age (Claps et al., 1996).

Altamura Limestone

This unit comprises the uppermost track-bearing hori-zon in the Apulia region. The unit, previously known asCalcari di Casa Lauriola, only crops out in two small areasin the Gargano region south of San Giovanni Rotondo andnear Apricena (Merla et al., 1969; Luperto Sinni and Mas-se, 1986). Laviano and Marino (1996) pointed out the cor-respondence in terms of facies and age of this unit with thebetter-known Altamura Limestone. The Altamura Lime-stone represents the return of a transgressive marine suc-cession after the prolonged mid-Cretaceous emersion ofthe Apulia platform (D’Argenio et al., 1987; Mindszenty etal., 1995), which is indicated by the presence of bauxiticdeposits (Crescenti and Vighi, 1964). In the San Giovanniarea, it consists of mudstone to wackestone with thin lay-ers of green shale of late Turonian?–Coniacian age (Luper-


FIGURE 3—Map of two of the Mattinata blocks. (A) MPA. (B) MPC.

to Sinni, 1996). In the Apricena area, it consists of thickbeds (1–2.5 m) of peloidal to bioclastic mudstone–wacke-stone alternating with planar or domal (also LLH) stro-matolite beds (Morsilli, 1998). In the upper part of the ex-posed succession, there are many beds rich in radiolitidaein life position, organized into bouquets or clusters (sensuRoss and Skelton, 1993). This interval in the Apricenaarea is Late Turonian–Early Senonian in age (Lavianoand Marino, 1996; Morsilli, 1998; Morsilli et al., 2002).

DISCOVERY AND STUDY OF THE FOOTPRINTS

This is not the first case of footprints discovered in apier. Thus, this study followed the approach previouslychosen by Dalla Vecchia and Venturini (1995). Althoughthe pier was built some tens of years ago, attempts weremade to trace the source of the material back to the quar-ries that furnished the blocks by examination of docu-ments from the companies that built the pier. Unfortu-nately, this was not as successful as it was for the aboveauthors, and a definitive result was not reached becausethe company documents only mentioned the source areafor the material and not the quarry. Consequently, theplace of origin of the footprint-bearing material is still un-known. Moreover, the blocks with the footprints are theonly three that differ in lithological characteristics fromthe other hundreds of blocks in the pier. How and whythey reached their place in the pier is still a mystery.

Subsequently, micropaleontological analyses were at-tempted, but samples examined in thin section were prac-tically barren and lacked diagnostic paleontological evi-dence for age determination. Consequently, a detailedstudy of the sedimentological features of the Mattinatablocks was conducted in order to compare the various lith-ofacies and sedimentary structures recognized on theblocks with ones present in outcrops that showed similarfeatures.

The blocks were designated as MPA, MPB, and MPC,respectively. Blocks MPA and MPC include footprints pre-served as natural molds, while MPB reveals the presenceof natural casts. Blocks also were labeled on each side inorder to have reference points for subsequent studies. Thetrampled surfaces were carefully mapped (Figs. 3, 4) andeach footprint was numbered and drawn; some specimensalso were cast. Morphologic and sedimentologic features ofblocks MPA and MPC revealed that these two blocks orig-inally were joined together (side B of the MPA block withside D of the MPC block). Contiguous trampled surfaceswere found between the two blocks with only a few centi-meter-sized fragments missing. No correspondence wasfound between prints (MPA, MPC) and natural casts(MPB), so the surface of block MPB represents a differentportion of a track-bearing surface with respect to MPAand MPC. Nearly 15 m2 of trampled surface were exam-ined. The amount of trampling is quite low (;20%), al-though it reveals an area where considerable activity oc-curred.

Sedimentological Features of the Blocks

A detailed study of the sedimentological features wascarried out on the Mattinata blocks. Sedimentological fea-tures were observed on polished slabs taken from edges

538 CONTI ET AL.

FIGURE 4—Map of the Mattinata block MPB.

FIGURE 5—Side views of the blocks. (A) MPA—side C, block thick-ness 70 cm; (B) MPB—side B, block thickness 65 cm; (C) MPC—sideC, block thickness 75 cm.

and sides of the blocks to obtain measurements and de-scriptions for stratigraphical logs, as well as to make de-tailed observations of fresh surfaces of the lithofacies.Thin sections also were made and used for descriptions.Bed-by-bed measurements of the various sides of theblocks revealed great heterogeneity at a centimeter scale,particularly in blocks MPA and MPC.

Block MPA: MPA is ;2.5 m long and 1.8 m wide. The av-erage thickness is ;70 cm (Fig. 5A). Side B of block MPAfits very well to side D of block MPC. An erosional surfaceor other irregularities are present in some layers. A con-tinuous polished slab was obtained by cutting the edge be-tween side A and B (Fig. 6). Well-laminated beds with aplanar shape occur at the base. The texture of this lime-stone is a peloidal wackestone–packstone, with abundantrhombohedral dolomite crystals. An interval with a brec-ciated texture is visible in this lower part. The breccia isdivided into two parts by a small bed showing the samecharacteristics as the unit previously described. Above thebreccia interval, the layers are characterized mainly byvery thin laminations that are more-or-less planar toslightly wavy. In the upper part, some layers are not lam-inated, and other parts are only a little disturbed.

The top layer has features typical of cyclic facies fromshallow-water settings. These features can be correlatedphysically to those in block MPC. A very peculiar struc-ture occurs in the upper part of a parallel polished slab(Figs. 6, 7), where the section of the bottom layer and thedisturbance of the entire upper layer are clearly visible.This structure is very similar to tridactyl imprints (DallaVecchia and Venturini, 1996; Avanzini, 1998; Dini et al.,

1998). In general, this could be referred to as typical dino-turbated structure.

Block MPB: MPB has a trapezoidal shape, with the baseof the triangle ;1.3 m long and a height of ;2.5 m. The av-erage thickness is ;65 cm (Fig. 5B). Although this block isstored upside down in the museum to show the naturalcasts, the description here follows the stratigraphic order.Two slabs were cut to obtain a continuous interval; theslabs were then polished and described. The first was tak-en at the edge between sides A and C (upper interval) andthe second at the edge between B and C (lower interval;Fig. 8). A schematic log also was measured on side B (Fig.9). From the base, corresponding to the tracked layer,


FIGURE 6—Polished slab of the block MPA cut on the edge betweenside C and D; scale bar 5 20 cm. In this figure, the sedimentaryfeatures and the main layers of this block are clearly visible.

FIGURE 7—Dinoturbated structure. This part corresponds to the top part of the previous figure. The deformation of the layers and the chaoticinfill of the depression caused by dinosaur imprint (maximum width 5 40 cm) are clearly visible.

there is an interval with wavy laminae, sometimes dis-turbed or poorly visible. A thick interval in which the orig-inal texture has been obliterated completely by dissolutionand precipitation phenomena (karst features) is visible inthe middle part. The top consists of a massive dolomitewith evidence of bioturbation.

Block MPC: MPC is ;3 m long and 1.5 m wide. The av-erage thickness is ;75 cm (Fig. 5C). Based on certain ir-regular layers, a detailed reconstruction of some bed ge-ometries has been carried out. Two logs were compiledfrom the middles of sides A and D. Because these two logsare not representative of the entire heterogeneous block, adraft of the entire geometry of side A has been reconstruct-ed. The main feature is a small, channeled bed that has itslong axis oriented between side A and C. It pinches orthins out in the other directions.

Two stratigraphic logs were measured on various sidesof the blocks. Log A–A’ shows an irregular base with verythin beds of limestone, sometimes with stylolitic boundar-ies (Fig. 10A). The textures are mainly wackestone withsome dolomitic crystals. Some laminae may have been re-lated to a weak traction current. Above this thin intervalof limestone (the only part of the block that is not dolomi-tized), there are dolomitic, millimeter-thick, slightly un-dulating laminae. The laminae are truncated both later-ally and in the upper part by an irregular, channelized,dark-gray, flat-pebble breccia with centimeter-sized clasts(maximum size of 5 cm). The breccia is concentrated at thebottom and top of the bed and is separated by a matrix-rich interval. On top of the breccia bed, very thin, undulat-ing laminae have a hemispheroidal shape and flatten outin the upper part of this layer. A poorly laminated or bio-turbated interval in the middle part of the succession isfollowed by a very well defined laminated interval with awavy to planar shape. In the upper part of the block, thereis an interval without lamination containing small depres-sions or deformations that can be related to pressure on aweak substrate. On the top of block MPC, where the foot-prints occur, laminations frequently are disturbed by din-oturbation. Log B–B’ on the same block shows a differentorganization, mainly in the lower part (Fig. 10B). Thebreccia interval here is very thin and some flat clasts arestill in place (mud cracks). Other deformations are veryclear and are considered good examples of dinoturbation.

The remarkable heterogeneity of this block is apparent

540 CONTI ET AL.

FIGURE 9—Log of MPB measured along the side B. The lower partof this block is characterized by the presence of thin lamination. Themiddle part shows karst features, while the upper part is comprisedof dolomitic limestone without sedimentary structures, probably relatedto bioturbation of a subtidal interval.

←

FIGURE 8—Polished slab of the block MPB cut on the edge betweenside A and C; scale bar 5 20 cm. The main feature to note is theinterval with karst structures (halfway up the slab) that obliterate theoriginal rock texture.

when viewing all of side A (Fig. 11). The most impressivefeature is the channeled breccia body that abruptly cutsthe lower part of the layers and has a more-or-less flat up-per boundary. This body is elongated in the direction ofsides A and C, and gives some information about theshape of this bed and its formation (Fig. 12).

Paleoenvironmental Interpretation of the Track-BearingBlocks

Some lithological features are useful for reconstructingthe depositional environment of the dinosaur footprints.The various lithofacies found in the Mattinata blockscould be interpreted as indicators of a subtidal (shallow-la-goon) setting, and intertidal- to supratidal-flat settings. Asmall tidal channel is indicated by the lens-shaped, flat-pebble breccia bed along side A of Block MPC. Desiccationcracks and the abundance of flat clasts testify to the com-mon occurrence of supratidal conditions. Laminated inter-


FIGURE 10—Logs of block MPC measured along (A) side A and (B)side D. This block is characterized by the presence of a breccia bedwith irregular geometric features (see Fig. 11), and the lower part isless dolomitized. The other lithofacies are an alternation of massiveand laminated dolostone.

vals, interpreted as cryptomicrobial laminites, indicatethe presence of algal or microbial mats. Most of the dino-saur footprints, therefore, must be related to an intertidalto supratidal setting.

DINOSAUR FOOTPRINTS

The Mattinata pier material is composed of 29 naturalmolds and 8 natural casts; 14 molds and 3 casts are pre-served well enough to allow good descriptions. All of thefootprints lack convex rims. This seems to confirm thepresence of a microbial mat on the track-bearing surface.On blocks MPA and MPC, footprints are obscured in plac-es by laminae that partially fill the more deeply impressedareas. This suggests the footprints were made on the ex-posed surfaces and are not undertracks. Moreover, over-printing is very rare, and no extramorphological features,such as retro-scratching, reflux of sediment inside thefootprints, or sliding traces in elongated footprints wererecognized. Consequently, most of footprints correspondwell to the underside morphology of the trackmakers’ feet.The better-preserved footprints are discussed below, fol-lowing Thulborn (1990), Leonardi (1987), and Padian(1992), and can be separated easily into three types.

Type 1

Material referred to this type consists of 12 footprints,nearly 30 cm long, which can be ascribed to a medium-sized bipedal dinosaur. The material includes both natu-ral molds (MPB4, MPB5) and natural casts (MPA1,MPB2, MPC1, MPC2, MPC3, MPC5, MPC11, MPC12,MPC13, MPC19), which allow reconstructing a trackwayfrom incomplete and ambiguous evidence. Three speci-mens (MPB4, MPB5, MPC2) have a very pronounced me-tapodium impression, as long (or longer) than digit III, sothat the length of these footprints is nearly double that ofthe other Type 1 specimens. The proportions suggest theseimpressions are from a large part of the metatarsals madeduring plantigrade gait or squatting. There are no exam-ples of paired parallel prints, therefore suggesting a sit-ting posture. This elongation and the hallux impressionare evident in four footprints (MPC3, MPC2, MPC12,MPC1) associated in an irregular sequence, evidentlymade during a very slow progression (Fig. 13). The result-ing trackway morphology is quite wide, with short steps,and with a large part of the metatarsals and the halluxtouching the ground. Wide-gauge theropod tracks recentlywere described by Day et al. (2002, 2004) and were associ-ated with a slow-walking phase of locomotion. A partialtrackway of two consecutive footprints (MPC19, MPC13)was reconstructed graphically (Fig. 14), and both track-ways suggest normal trackmaker progression, not restingphases. This type of gait may be unusual: thus, trackwayparameters should be considered with caution.

Footprints MPA1 (Fig. 15), MPC3, and MPC11, pre-served as natural molds, clearly show phalangeal pads ondigits I, II, and III, and less clearly on digit IV, with thedigit II metatarsophalangeal pad being diagnostic. MPA 1seems to show a web-like structure between digits III andIV, most probably due to folding of thin microbial laminaearound the footprint. In three cases (MPB4, MPB5,MPC2) the impression of a long and narrow metapodium

542 CONTI ET AL.

FIGURE 11—Drawing of side A of the block MPC. The main feature to note is the shape of the flat-pebble breccia bed, shaded gray in thisfigure, which is a channelized body with an erosional base. The irregularities in the upper part of this block are related to dinoturbation, withnormal print (compare with Fig. 7), and underprint deformation.

FIGURE 12—Detail of the breccia bed of Block MPC (maximum width25 cm, see Fig. 11 for location).

FIGURE 13—Four consecutive tracks referred to the same dynamicaction of an undecided or sliding trackmaker (two tracks of the left pesprecede the impression of two tracks of the right one).


FIGURE 14—Footprints (MPC19, MPC13) geometrically completedby a third virtual footprint (MPv) to complete a stride.

FIGURE 15—Photograph and outline drawing of Type 1 footprint(MPA1).

FIGURE 16—Natural cast of a very elongated footprint (MPB4). Notethe central area masked by displaced carbonate mud.

also is present, as sometimes happens in squatting dino-saurs. In these cases, the interdigit divergences are largerand much mud was displaced, hiding the metatarsal-pha-langeal region (Fig. 16). MPC5, although apparently tri-dactyl and slightly smaller, could be ascribed to a partialimpression of this type.

Footprints of functionally tridactyl peds with hallux im-pressions often are present among ornithischian tracks(e.g., Anomoepus and/or Moyenisauropus; see Olsen andRainforth, 2003), but they also are known in theropod foot-prints (e.g., Buckeburgichnus sensu Lockley 2000; Eutyni-chnium; ‘‘Gigandipus’’; ‘‘Hyphepus’’; Jalingpus; Kayenta-pus soltykovensis; Picunichnus; Saurexallopus; Theroplan-tigrada; Tyrannosauripus; and an unnamed theropodfrom Middle Jurassic of Morocco; Nouri et al., 2000). Thehallux often is undervalued as a character (see Harris etal., 1996 for a review) because it is considered a kind of ex-tramorphological feature in typical tridactyl forms, andonly is visible in the deepest impressions.

A superficial resemblance between Apulian specimensand Anomoepus (related to ornithischian trackmakers)concerns partial metatarsal impressions. However, theApulian footprints are not the traces of resting animalswith both hind limb impressions parallel to each other(Lull, 1904; Gierlinski, 1994, 1996; Avanzini et al., 2001;Lockley et al., 2003).

The impression of digit I is a common feature in old fig-ures of Eutynichnium lusitanicum, from the Late Jurassicof Portugal, in which ‘‘depth. . . [10–15 cm]. . . accounts forthe preservation of hallux traces’’ (Lockley et al., 2000a, p.323; but see dos Santos, 2002). A small footprint figured by

544 CONTI ET AL.

FIGURE 17—Specimen MPA1. A thin microbial lamina, in front of hip-ex III–IV, mimics a web-like structure.

Kuban (1989, p. 68, fig. 7.17J) as ‘‘Jalinapus’’ [sic], origi-nally attributed to an ornithischian, but more recently to atheropod (Gierlinski, 1994), also is similar. The type spec-imen (in Zhen et al., 1983) is a single footprint with a hal-lux impression in a well-impressed series of 38; it appearsto be a rare case within this sample. A partial resemblancein the position of digit I and the narrowing at the base ofthe digit II also can be recognized in Picunichnus benedet-toi from Cenomanian sediments of Argentina (Calvo,1991). The central posterior elongation and the hallux im-pression resemble Kayentapus soltykovensis from LowerJurassic of Hungary (Gierlinski, 1996), but specimens dif-fer in relative proportions and in digit III-IV divergence.Megalosauripus footprints (sensu Lockley et al., 2000a)sometimes show the hallux impression when deeply im-pressed, although detailed published information is notavailable.

The metapodium, or posterior elongation of the foot-prints, appears in Mattinata material consistently. Pitt-man (1989, figs. 15.8, 15.9) illustrated a similar footprintinterpreted as a very deep impression, and rejected the hy-pothesis of an animal walking on its metatarsals. An ex-ample of elongate footprints from a walking dinosaur in-terpreted as the tracks of ‘‘an animal slipping on a wetsubstrate’’ was described by Kvale (2001, p. 250).

Elongate theropod footprints are quite widespread (e.g.,Kuban, 1989; Gierlinski, 1994; Perez-Lorente, 1994) andhave been described either as elongated, or as footprints ofplantigrade dinosaurs. As noted above, most of the foot-prints, including Kayentapus (sensu Gierlinski 1996), Jal-ingpus, and Megalosauripus, sometimes show digit I andelongate metapodium impressions. Deep impressions, or apeculiar semidigitigrade and plantigrade gait may explainthe origin of both features.

The closest fit between the new material and knownfootprints is with unnamed footprints described and fig-ured by Aguirrezabala and Viera (1980, fig. 24) from LateJurassic (Kimmeridgian) sediments of Bretun (Soria,Spain). In these footprints, digit I seems to lie in a slightlydifferent plane and is less deeply impressed. The similari-ty is strong enough to consider ascribing this material tothe same trackmaker group.

The same material, subsequently figured by Perez-Lor-ente as trackways C and G (Perez-Lorente, 1994, figs. 1,2), allows favorable comparison of trackway characterswith the Mattinata material. The Spanish trackwaysslightly differ in having a higher pace angulation (narrow-er interpedes distance).

Theroplantigrada, an ichnotaxon based on Aptiantracks from La Rioja (northern Spain) ascribed to a thero-pod (Casanovas Cladellas et al. 1994), is similar to theMattinata material and to trackways described by Aguir-rezabala and Viera (1980). This monotypic ichnogenus isbased on a single trackway in which, except in size, thefootprints seem quite similar to the Mattinata material inthe presence and position of the hallux and the metapo-dium impression. All of the Spanish material shows veryelongated footprints. The Apulian material resemblestrackway A and some footprints of trackway D (CasanovasCladellas et al., 1994, figs. 3 and 15, respectively).

Casanovas Cladellas et al. (1994) considered the pres-ence of an interdigital web as the main diagnostic featureof Theroplantigrada. This ichnotaxon differs from the ma-

terial of Aguirrezabala and Viera (1980) in age, but this in-terdigital web cannot be considered a defining character.First-hand observation of the Spanish material does notpermit confirmation of whether web traces truly are pre-sent. Apparent web traces are lacking in all the Mattinatafootprints described herein, except in MPA1, just in frontof the hypex III-IV (Fig. 17). However, this structure wasgenerated by the inflexion of microbial laminae some cen-timeters away from the real boundaries of the footprint.

If webbing really is present in Theroplantigrada, itwould be a rare case among dinosaurs. More likely, it is anextramorphological feature. Only three purported caseswere cited by Thulborn (1990)—Otouphephus magnificus,Swinnertonichnus mapperleyensis, and Talmontopus ter-si—and in all three cases, the web traces subsequentlywere interpreted as extramorphological features. ForOtouphephus magnificus, ‘‘the web-like trace . . . was latershown to be an artefact’’(Thulborn, 1990, p. 80). Swinner-tonichnus mapperleyensis, based on a single tridactyl coe-lurosaur footprint with web-like traces (Sarjeant, 1967),subsequently lacks web traces in the figures of Haubold(1971, pl. 42, fig. 14), and was reinterpreted as a crocodil-ian imprint, because ‘‘webbing. . . (is). . . a feature un-known in dinosaur footprints. . . ’’(Sarjeant, 1996, p. 14).The same specimen, re-examined by King and Benton(1996), was ascribed more convincingly to Chirotherium,an ichnotaxon that lacks a web. Regarding Otouphephusand Talmontopus, King and Benton (1996, p. 221) reported


FIGURE 18—Photograph and outline drawing of the best-preservedType 2 footprint (MPC10).

FIGURE 19—Outline drawings of other Mattinata footprints. (A) Type3 footprint (MPC17). (B) Undetermined specimen (MPA3).

that ‘‘. . . Lockley (pers. comm., 1994) cannot confirm thepresence of webbing in either taxon.’’

In the opinion of some authors, ichnogenera should cor-respond to footprints revealing diagnostic differences indefinite skeletal structures of the trackmakers (Carranoand Wilson, 2001). Consequently, if the supposed webbingin Theroplantigrada is extramorphological, the main di-agnostic features of Theroplantigrada would be the pres-ence of hallux and metatarsal impressions that corre-spond to skeletal structure; the presence of web traces con-ceivably could be considered a character at the ichnospe-cies level. If the above rationale is accepted, the Mattinatamaterial, trackways C and G described by Aguirrezabalaand Viera (1980), trackway D of Casanovas Cladellas et al.(1994), and the trackway ascribed to Theroplantigrada en-cisensis all could be included in the same ichnogenus. Con-sequently, the Mattinata material described here is clas-sified as cf. Theroplantigrada isp.

Comparisons are possible with similar tracks. Similartrackways and tracks are known from the Upper Jurassicof Soria, Spain (Aguirrezabala and Viera, 1980); ‘‘Eutyni-chnium’’ lusitanicum in the Uppermost Jurassic of Portu-gal; an unnamed trackway from Middle Jurassic depositsof Morocco (Nouri et al., 2000); Jalingpus yuechiensis fromthe Upper Jurassic of Sichuan, China; Kayentapus solty-kovensis from the Lower Jurassic of Poland, Sweden, andHungary; and Theroplantigrada encisensis from EarlyCretaceous beds (probably Aptian in age) of northernSpain. All represent medium-sized dinosaurs that left hal-lux and metapodium traces. This kind of footprint cannotbe ascribed easily to a zoological group below the level oftheropod trackmaker or, perhaps, to a cursorial, light or-nithopod (Thulborn, 1990; Viera and Torres, 1992; Farlowand Lockley, 1993). Accepting the more probable hypoth-esis of a theropod trackmaker, it is inferred that thesefootprints represent a medium-sized, non-derived thero-pod (a ceratosaur) or a basal tetanuran where digit I is notreduced.

Type 2

Only three stout, tridactyl footprints preserved as nat-ural casts (MPC4, MPC10, MPB7) are included in thistype (Fig. 18). MPC4 and MPC10, although deeply im-pressed, lack many details of pads; MPC10 and MPB7

show a characteristic, folded proximal margin and well-developed claws.

Often, similar material has been described under aplethora of names or, more frequently, has remained un-named, perhaps due to the generalized shape of the prints(e.g., Olsen and Galton, 1984). However, recent papers byLockley et al. (1996; 2000a) and Olsen et al. (1998) haveshown that detailed analyses can allow separation of atleast some theropod footprints and trackways. Accordingto older literature, material similar to the Mattinataprints could be referred to Eubrontes or to Megalosauripus(for a review on the debate around this last name and asuggested solution, see Lockley et al., 2000a; Lockley andMeyer, 2000). However, studied specimens lack the so-called heel, or proximal area, made by the impression ofmetatarsal-phalangeal pad of digit IV. The Mattinataspecimens differ in this character from most other thero-pod footprints, which are characterized by the impressionof the metatarsophalangeal pad of digit IV. Two excep-tions are Carmelopodus untermannorum, a small footprintfrom Middle Jurassic deposits of northeastern Utah (Lock-ley et al., 1988), and Skartopus australis from the Creta-ceous of Australia (Thulborn and Wade, 1984). In the di-agnosis of Carmelopodus, the ‘‘lack of any impression of afourth proximal pad on digit IV is stressed’’ (Lockley et al.,1998, p. 260). The Mattinata footprints and the Utah andAustralian prints differ in other characters, but it is inter-esting that this is another example of the lack of such animportant feature in mid-Jurassic material. Consequent-ly, the Mattinata specimens may share this functionalcharacter with Carmelopodus. Due to the lack of relatedichnotaxa, no stratigraphical inferences are possible. It isonly possible to note that Carmelopodus comes from Mid-dle Jurassic deposits, Skartopus from the Mid-Cretaceous,and there are similar subdigitigrade theropod tracks inthe Lower Jurassic of Poland (Gierlinski and Pienkowski,1999).

Type 3

Only two middle-sized footprints (MPC9, MPC17; natu-ral casts) pertaining to the same partial trackway (Fig.19A) are ascribed to Type 3 prints. These footprints resem-ble Therangospodus, a recently formalized ichnogenus at-tributed to a theropod track-maker that lacks well-definedpads on the digit impression (Lockley et al., 2000b), and

546 CONTI ET AL.

includes only two ichnospecies: T. pandemicus and T. on-calensis. Gierlinski et al. (2001, p. 445) reported a speci-men with more distinct phalangeal pads, but stated thatthey were ‘‘not sure if Therangospodus should be distin-guished from Megalosauripus.’’

Specimens that can be compared to the Mattinata ma-terial are ascribed to deposits of Late Jurassic age. Ther-angospodus pandemicus has been described from the Up-per Jurassic of the United States (Utah) and Turkmenis-tan (Lockley and Meyer, 2000), and from the Upper Juras-sic of Portugal (Lockley et al., 2000b). The correct age ofTherangospodus oncalensis, an ichnospecies first consid-ered Early Cretaceous in age, still seems questionable (seeLockley et al., 2000b).

Other Footprints

The remaining footprints, although easily recognizableas dinosaur traces, are preserved too poorly to be ascribedto a particular group. MPA5 (Fig. 19B) resembles an or-nithopod footprint in the II-IV digit divergence (928), rela-tive shortness of digits, and lack of pads. However, it alsoshows sharp claw traces and thus could represent a the-ropod trackmaker. MPC6 and MPA4 could be ascribed toType 1 traces, and MPC20 and MPC21 likely are two the-ropod traces in a lightly impressed trackway. This mate-rial provides data to evaluate the number of individualsthat crossed the area.

Notes on the Ichnocoenosis

Although the Mattinata pier material still presentsmany unsolved problems, it allows inferences to be madeabout paleobiogeography (discussed below), and throwslight on the presence of a theropod-dominated track as-semblage with inferred correspondence to the assemblagedescribed by Lockley et al. (2000a) and by Lockley andMeyer (2000), which includes Megalosauripus and Ther-angospodus. Such ichnocoenoses have been recorded fromUzbekistan, Spain, and North America, and are consid-ered characteristic of the early Late Jurassic. In the pre-sent case, the dominance of different tracks related toTheroplantigrada is recorded. This difference could reflectage or facies controls affecting the Apulian Platform ichn-ocoenosis.

CONCLUSIONS

Age of the Blocks

Direct biostratigraphic or chronological calibration ofthe age of the material is lacking, but three lines of evi-dence allow inference of the approximate geologic age ofthe Mattinata blocks: (1) provenance of the blocks, (2) thelithofacies recognized in the blocks, and (3) the footprinttypes from the blocks.

(1) Documents about the pier building are related to theextraction of blocks from quarries in the San Giovanni Ro-tondo area and Apricena. The former were opened mainlyin the San Giovanni Rotondo Limestone, and this unit isnever dolomitized. In the Apricena area, the only intervalwhere block extraction continues is in the San GiovanniRotondo Limestone, where no dolomitic layers are known.

(2) Some beds in small, abandoned quarries near MonteCalvo (Fig. 1) in the Sannicandro Formation reveal simi-lar sedimentological features to the footprint-bearingblocks, and are partially dolomitized. The age of this partof the Sannicandro Formation is referred to the Kimmer-idgian–Tithonian interval (Morsilli, 1998). The same lith-ological and sedimentary characteristics also can be foundnear the San Nazario area (see Fig. 1) in the northwestpart of the Gargano. Here, some old quarries, in some cas-es dismantled by the construction of a road, reveal verysimilar lithological and sedimentary features. The age ofthis succession is the same as the one in the Monte Calvoarea (Luperto Sinni and Masse, 1994).

(3) Although cautiously avoiding attribution of the foot-prints to named ichnospecies, the overall character of theassemblage suggests a Late Jurassic age, although theseforms were not exclusive to that time interval.

In conclusion, these lines of evidence suggest that thestudied material comes from the Late Jurassic Sannican-dro Formation.

Paleogeographic Inferences and Problems

During Middle–Late Jurassic and Cretaceous times, theApulian Platform has been interpreted as a small, isolatedcarbonate platform, surrounded by the Tethys Ocean, sep-arated from other peri-Adriatic carbonate shelves (e.g.,the Laziale—Abruzzese, Campana, Sazani, and Kruja car-bonate platforms) by deep-sea areas. It is inside the Apulo-Dinaric structural unit, and is isolated from both thesouthern and northern continents by deeper basins(D’Argenio, 1976; Zappaterra, 1990; Eberli et al., 1993;Masse et al., 1993).

This model obviously was constructed before repeateddiscoveries of dinosaur footprints on the Apulian Plat-form. Recent finds of dinosaur trackways from differentplaces and stratigraphical intervals are very strong paleo-geographic constraints that necessitate reconsideration ofprevious interpretations. At first, this new evidence maybe considered weaker than geophysical or structural data,but dinosaur footprints may serve as powerful new tools tochange the best-built paleogeographic models. Otherworkers have pointed out that paleogeographic recon-structions must account for dinosaur footprints. For ex-ample, Meyer and Lockley (1997, p. 425) stated that ‘‘re-current emergent areas must have been present that con-nected the southern part of the London-Brabant masswith the northeastern part of the Massif Central’’; Kvaleet al. (2001, p. 233) claimed that ‘‘a major change in the pa-leogeographic reconstructions for Wyoming’’ is needed;and a connection of ‘‘emerged area with larger land mas-ses’’ in the Holy Cross Mountains (Poland) was suggestedfor the Late Jurassic by Gierlinski and Niedzwiedzki(2002, p. 58A).

At present, three different track assemblages in theApulian region have been found. The first, described here-in, is a theropod-dominated ichnoassociation, probablyLate Jurassic in age. The second, described from the EarlyCretaceous (Late Hauterivian–Early Barremian) of BorgoCelano (Gianolla et al., 2000), contains footprints ascribedto theropods, ornithopods, and perhaps sauropods; andthe third, described from Late Cretaceous (Santonian)from Altamura (Andreassi et al., 1999; Nicosia et al.,


2000a, b), is ornithopod dominated. These are the moststriking evidence of widespread, terrestrial vertebrates inthe area. Other data, such as the so-called Ruvo varanoid(Varola, 1999) and vertebrate remains recorded from theMelissano Limestone (Medizza and Sorbini, 1980), alsoare evidence of terrestrial vertebrates in this area. Findsof bauxite levels (D’Argenio et al., 1987) and land-plant re-mains (De Cosmo and Morsilli, 2002; Morsilli et al., 2002)also are evidence of soil development and emergent land.

These data span different time intervals, and show dif-ferent evolutionary levels consistent with biological eventsrecognized elsewhere. Moreover, this data set probably isbiased by taphonomic events and by reduced availabilityof Jurassic and Cretaceous rock outcrops with track-bear-ing potential. This kind of research needs extensive, un-disturbed, well-exposed bedding surfaces, a situation notfound in Mesozoic deposits in this part of southern Italy.

Available data record three land-vertebrate assemblag-es in the area, but they must be analyzed cautiously. If thetrackways only are exceptional occurrences, they could berelated to short-term, repeated connections to larger landareas. On the other hand, if they are considered a rare re-cord of normal events, then continuous, long-lasting colo-nization by land dwellers must be hypothesized. From apaleontological point of view, these hypotheses correspondeither to repeated immigration pulses, or to a single, earlycolonization followed by endemic evolution. The first hy-pothesis implies a continuous connection and ongoing im-migration between the Apulian Platform and large conti-nental areas. Thus, the carbonate-shelf shallow-sea areaswere not separated by deep basins, and sea-level dropswould have allowed emergence of the platform followed byland-vertebrate colonization. In the source area, evolu-tionary processes could proceed normally, with the immi-grants reflecting worldwide evolutionary trends. This ex-planation may be easier to accept because it does not needcomplex evolutionary hypotheses.

The second hypothesis must, in turn, be subdivided totwo sub-hypotheses: either the platform was as small ascurrent models suggest, with different portions of the in-ner platform emerging from time to time, or the emergentarea was much larger than thought, although no traces ofit are preserved. In the first scenario, lacking any evidenceof a single, persistent island area large enough to sustaina balanced community of huge animals, land vertebratesmust have been wandering continuously from one zone toanother within the same platform area—an idea that doesnot consider the need of dinosaurs for vegetation, freshwater, and nesting sites. Simple emergent episodes, evenif repeated and prolonged, thus are not sufficient to ex-plain the data. In the second case, the scenario must in-clude a large, isolated, complex environment (includingfresh-water sources and nesting sites), in which the persis-tence of a balanced fauna was possible for ;130 Ma, withall traces of such a large area removed by subsequent geo-logical events. In both cases, balanced ecosystems implythe presence and the coevolution of plants, herds of plant-eating animals, and meat-eating dinosaurs. It also mustbe assumed that a complete suite of scavengers and inver-tebrate organisms, as well as the proper physical, chemi-cal, and biological conditions, were present to allow evolu-tionary processes. The carbonate-platform environmentraises particular difficulties in this regard because of lim-

ited vegetation productivity, underdeveloped river sys-tems, and soils that need a long time to form from the re-siduals of karstification. On the other hand, the lack of riv-er environments also could explain the scattered presenceof fossils. In fact, a lack of river systems could have re-duced the chance for fossilization by reducing depositsavailable to preserve dinosaur bones and/or footprints.

Both cases included in the second hypothesis presentanother important concern. The recorded Apulian dino-saur ichnocoenoses apparently differ from one another,but may correspond in both the faunal composition andthe evolutionary level to age-equivalent dinosaur commu-nities known from Europe, central Asia, and North Amer-ica. The recognized evolutionary events appear to corre-spond to events recognized in Europe, in central Asia, andin North America. It is difficult to accept a hypothesis ofparallel evolution on the mainlands and isolated carbon-ate-platform areas.

In any case, the presence of a large number of dinosaursin an area such as the Apulian Platform is problematic. Itunderlines the inadequacy of paleogeographic models thatsimply suggest a tectonic disjunction of the platforms. Onthe contrary, the footprint and other data strongly supporta scenario that discards the hypothesis of many small,peri-Adriatic carbonate shelves and interposed basins,and suggests structural continuity and frequent (or con-tinuous) connections of the platform to the mainlands.Consequently, previous hypotheses of dwarf dinosaur fau-nas that originated by endemic evolutionary phenomena(Dalla Vecchia and Tarlao, 2000; Dalla Vecchia et al.,2002) seem insufficient to explain the evidence fully. Ifrecognized, they might be considered partial explanationsor just the record of the final phase of endemic evolutionafter immigration.

Bosellini (2002) used these new data for strong paleo-geographic constraints. Reviewing various geological andgeophysical data associated with the presence of dino-saurs around the Ionian Sea and surrounding areas, Bo-sellini (2002) reached the conclusion that the ApulianPlatform was connected during the Jurassic and Creta-ceous to Africa through the Peloponnesus, Crete, the Cy-rene Seamount, and the Medina Ridge. Gierlinski (pers.comm., 2002) also has found similar tracks in the Mesozoiccarbonates in Crete. In Bosellini’s (2002) reconstruction,the Apulian Platform, and probably other peri-Adriaticplatforms, were not isolated Bahamian banks, but ratherwere more like the modern Florida peninsula. This modelconflicts strongly with other paleogeographic models (e.g.,Catalano et al., 2001). In fact, using these data, an easternconnection to the mainland cannot be dismissed. Similar-ly, more than one immigration route could be hypothe-sized for these dinosaurs at different stratigraphical lev-els. Future reconstructions of the area must integrate alldata sources, such as geophysical (seismic refraction andreflection profiles of the Apulian Platform margin and sur-rounding basins), structural and kinematic (relationshipsbetween thrust-belt chains and foreland areas), strati-graphical, sedimentological, and paleontological data.

ACKNOWLEDGEMENTS

Marco Avanzini is warmly thanked for his suggestionabout dinoturbation. G. Gierlinski, M. Lockley, and P. Up-

548 CONTI ET AL.

church revised the manuscript and gave important sug-gestions. Prof. G. Andreassi of the Soprintendenza Ar-cheologica per le Puglie and the Mayor of Mattinata kindlyhelped our work.

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