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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 CONTI
Dipartimento di Scienze della Terra, Universita` ‘‘La Sapienza’’, P.le A. Moro 5, 00185, Roma, Italy
MICHELE MORSILLI
Dipartimento di Scienze della Terra, Universita` di Ferrara, C.so Ercole I d’Este 32, 44100, Ferrara, Italy
UMBERTO NICOSIA*, EVA SACCHI
Dipartimento di Scienze della Terra, Universita` ‘‘La Sapienza’’, P.le A. Moro 5, 00185, Roma, Italy;
Email: umberto.nicosia@uniroma1.it
VINCENZO SAVINO, ALEXANDER WAGENSOMMER, LEONARDO DI MAGGIO
Speleo Club Sperone, San Giovanni Rotondo, 71013, Foggia, Italy
PIERO GIANOLLA
Dipartimento 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 found
in a small pier at Mattinata (Gargano Promontory, Foggia,
Italy), most probably quarried from the Upper Jurassic
Sannicandro Formation. All of the footprints in the blocks
are ascribed to medium-sized theropod trackmakers. Re-
cent track discoveries from both the Early Cretaceous San
Giovanni Rotondo Limestone and the Late Cretaceous Al-
tamura Limestone, as well as this new discovery, reveal the
consistency of terrestrial associations along the southern
margin of the Tethys Ocean in the peri-Mediterranean area
at 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 limestone
blocks in a small pier that protects the entry to the small
harbor of Mattinata on the Gargano Promontory (Puglia,
southern Italy, Fig. 1). The blocks were removed and are
stored 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 dinosaur
footprints and trackways have been found previously. The
first site was found in 1999 in the Murge area near the
town 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 tracks
imprinted 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 are
quite similar in facies architecture, size, and shape to the
modern Bahamian Banks (Bernoulli, 1972; D’Argenio,
1976; Eberli et al., 1993). The Gargano Promontory and
other parts of the Apulia Region make up part of the fore-
land of the Apennine chain. Structurally, the Gargano
area is folded into a gentle anticline with a WNW axis
(Martinis, 1965). This broad structure contains numerous
faults with various trends and kinematics (Funiciello et
al., 1992; Bertotti et al., 1999). The most prominent struc-
tural feature is the Mattinata fault, a regional E–W shear
zone that crosses the entire Gargano area, and continues
offshore for many tens of kilometers (De Dominicis and
Mazzoldi, 1989).
The Gargano area, together with Maiella Mountain, is
the only place where the transition between platform and
slope-to-basin facies crops out. In other areas, the eastern
margin of this platform lies offshore under the Adriatic
Sea, about 20 to 30 km from the present coastline (De
Dominicis and Mazzoldi, 1989).
The stratigraphy of the Gargano area has been reinter-
preted frequently in the last two decades. In the late
1960s–early 1970s, geological surveys introduced many
formational units (Martinis and Pavan, 1967; Merla et al.,
1969; Cremonini et al., 1971), complicating stratigraphical
correlations 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 idealized
E–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., Luperto
Sinni 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 different
second-order stratigraphical sequences, bounded by un-
conformities, of different types and origins (see Bosellini et
al., 1999, for a review). Herein, the focus mainly is on the
lithostratigraphical units that contain shallow-water sed-
iments in which footprints of terrestrial reptiles could
have been impressed.
The main sequences that contain shallow-water sedi-
ments are the Monte Sacro Sequence and the Mattinata
Sequence (sensu Bosellini et al., 1999; Fig. 2). During the
Late Jurassic–Early Cretaceous, two inner-platform units
were 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 in
some small outcrops, and have been named the Calcari di
Casa Lauriola (Merla et al., 1969); these units correspond
to the well-known Altamura Limestone (Laviano and Ma-
rino, 1998; Bosellini and Morsilli, 2001).
Sannicandro Formation
The 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 Rotondo
Limestone (Mattavelli and Pavan, 1965; Pavan and Pirini,
1966; Cremonini et al., 1971). More recent investigations
(Luperto Sinni and Masse, 1986; Claps et al., 1996) only
examined the Lower Cretaceous succession of inner-plat-
form facies. Luperto Sinni and Masse (1994) proposed that
the 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 Calcari
di 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 the
Sannicandro Formation could be extended to include the
Berriasian–Early Valanginian interval.
The Sannicandro Formation crops out only in the west-
ern and central sectors of Gargano. The base of this unit is
unknown 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 the
Monte 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 the
area 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 cm
thick). This lower part of the unit is frequently dolomi-
tized. At the top of the unit, there are peritidal cycles (1–2
m thick) composed of peloidal wackestone, with localized
packstones 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 near
San Giovanni Rotondo, where it is associated with thin
beds 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 fenestral
structures and small lenses of flat-pebble breccia (clasts
3–4 cm). The upper part of the Sannicandro Formation
mainly consists of subtidal lithofacies and rare, thin pla-
nar-stromatolite beds. The main textures are peloidal
wackestone and bioclastic packstone, locally very rich in
dasycladalean algae (Campbelliella striata and C. milesi).
The microfossil association partially covers the Callovian–
Berriasian time interval, but the peritidal facies indicates
a Kimmeridgian–Tithonian age. The facies associations of
this formation could be referred to an inner-platform set-
ting with protected lagoons, tidal-flats, and supratidal de-
positional environments, sometimes affected by storm
events (Morsilli and Bosellini, 1997).
Many of the lithofacies in the Sannicandro Formation
are the same as the ones observed on the Mattinata
blocks. Dinosaur footprints from this formation have not
been found in place, but more research could fill this gap.
Therefore, the potential presence of dinosaur footprints or
bones in the Monte Spigno Formation, in particular in the
transition area with the Sannicandro Formation, cannot
be excluded.
San Giovanni Rotondo Limestone
This unit has been studied in detail both in the type
area (Luperto Sinni and Masse, 1986, Claps et al., 1996)
and in the Apricena-Poggio Imperiale area (Luperto Sinni
and Masse, 1986). This unit partially corresponds to the
Bari Limestone (Azzaroli et al., 1968; Ricchetti, 1975) in
terms 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 be
subdivided into three members (see Claps et al., 1996 for a
more detailed description). Member 1 (Borgo Celano
Member of Luperto Sinni and Masse, 1986) consists of a
monotonous and acyclic subtidal unit with meters-thick
mudstone to wackestone that is intensely bioturbated.
This member can be referred to a shallow-subtidal, la-
goonal setting. Member 2 (Loferitic Member of Luperto
Sinni and Masse, 1986) is a thick, cyclic unit characterized
by quasi-periodic alternation of loferitic beds and centi-
meter-thick layers of green shale. Many lithofacies have
been documented in this unit: mudstone–wackestone beds
(0.5–2 m thick), intensely bioturbated with occasional
black 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 features
can be referred to a tidal-flat setting with subaerial expo-
sure. Member 3 consists of various lithofacies, such as
thin-bedded packstone to grainstone with peloids, bio-
clasts, and micritic intraclasts that are normally graded
and evenly laminated. Lags of ostreids, requienids, and
nerineids also may occur. The peritidal cycles consist of
peloidal–bioclastic packstone to wackestone at the base,
and planar stromatolites (20–100 cm) at the top. There are
lens-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 Rotondo
Limestone 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 of
Member 2 of the San Giovanni Rotondo Limestone. Mem-
ber 2 belongs to the Campanellula capuensis Zone of Late
Hauterivian–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 as
Calcari di Casa Lauriola, only crops out in two small areas
in the Gargano region south of San Giovanni Rotondo and
near 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 the
better-known Altamura Limestone. The Altamura Lime-
stone represents the return of a transgressive marine suc-
cession after the prolonged mid-Cretaceous emersion of
the Apulia platform (D’Argenio et al., 1987; Mindszenty et
al., 1995), which is indicated by the presence of bauxitic
deposits (Crescenti and Vighi, 1964). In the San Giovanni
area, it consists of mudstone to wackestone with thin lay-
ers of green shale of late Turonian?–Coniacian age (Luper-
DINOSAUR FOOTPRINTS FROM SOUTHERN ITALY 537
FIGURE 3—Map of two of the Mattinata blocks. (A) MPA. (B) MPC.
to Sinni, 1996). In the Apricena area, it consists of thick
beds (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 radiolitidae
in life position, organized into bouquets or clusters (sensu
Ross and Skelton, 1993). This interval in the Apricena
area is Late Turonian–Early Senonian in age (Laviano
and Marino, 1996; Morsilli, 1998; Morsilli et al., 2002).
DISCOVERY AND STUDY OF THE FOOTPRINTS
This is not the first case of footprints discovered in a
pier. Thus, this study followed the approach previously
chosen by Dalla Vecchia and Venturini (1995). Although
the pier was built some tens of years ago, attempts were
made 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 above
authors, and a definitive result was not reached because
the company documents only mentioned the source area
for the material and not the quarry. Consequently, the
place of origin of the footprint-bearing material is still un-
known. Moreover, the blocks with the footprints are the
only three that differ in lithological characteristics from
the other hundreds of blocks in the pier. How and why
they 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 detailed
study of the sedimentological features of the Mattinata
blocks was conducted in order to compare the various lith-
ofacies and sedimentary structures recognized on the
blocks with ones present in outcrops that showed similar
features.
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 presence
of natural casts. Blocks also were labeled on each side in
order to have reference points for subsequent studies. The
trampled surfaces were carefully mapped (Figs. 3, 4) and
each footprint was numbered and drawn; some specimens
also were cast. Morphologic and sedimentologic features of
blocks MPA and MPC revealed that these two blocks orig-
inally were joined together (side B of the MPA block with
side D of the MPC block). Contiguous trampled surfaces
were found between the two blocks with only a few centi-
meter-sized fragments missing. No correspondence was
found between prints (MPA, MPC) and natural casts
(MPB), so the surface of block MPB represents a different
portion of a track-bearing surface with respect to MPA
and MPC. Nearly 15 m
2
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 was
carried 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—side
C, 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 the
blocks 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 MPA
fits very well to side D of block MPC. An erosional surface
or 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 a
planar shape occur at the base. The texture of this lime-
stone is a peloidal wackestone–packstone, with abundant
rhombohedral dolomite crystals. An interval with a brec-
ciated texture is visible in this lower part. The breccia is
divided into two parts by a small bed showing the same
characteristics as the unit previously described. Above the
breccia interval, the layers are characterized mainly by
very thin laminations that are more-or-less planar to
slightly 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 from
shallow-water settings. These features can be correlated
physically 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 the
disturbance of the entire upper layer are clearly visible.
This structure is very similar to tridactyl imprints (Dalla
Vecchia 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 base
of 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 is
stored upside down in the museum to show the natural
casts, the description here follows the stratigraphic order.
Two slabs were cut to obtain a continuous interval; the
slabs were then polished and described. The first was tak-
en at the edge between sides A and C (upper interval) and
the 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,
DINOSAUR FOOTPRINTS FROM SOUTHERN ITALY 539
FIGURE 6—Polished slab of the block MPA cut on the edge between
side C and D; scale bar 5 20 cm. In this figure, the sedimentary
features 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 chaotic
infill 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 dissolution
and precipitation phenomena (karst features) is visible in
the middle part. The top consists of a massive dolomite
with 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 compiled
from the middles of sides A and D. Because these two logs
are not representative of the entire heterogeneous block, a
draft of the entire geometry of side A has been reconstruct-
ed. The main feature is a small, channeled bed that has its
long axis oriented between side A and C. It pinches or
thins out in the other directions.
Two stratigraphic logs were measured on various sides
of the blocks. Log A–A’ shows an irregular base with very
thin beds of limestone, sometimes with stylolitic boundar-
ies (Fig. 10A). The textures are mainly wackestone with
some dolomitic crystals. Some laminae may have been re-
lated to a weak traction current. Above this thin interval
of 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 the
bottom 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 out
in the upper part of this layer. A poorly laminated or bio-
turbated interval in the middle part of the succession is
followed by a very well defined laminated interval with a
wavy to planar shape. In the upper part of the block, there
is an interval without lamination containing small depres-
sions or deformations that can be related to pressure on a
weak 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 different
organization, mainly in the lower part (Fig. 10B). The
breccia interval here is very thin and some flat clasts are
still in place (mud cracks). Other deformations are very
clear 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 part
of this block is characterized by the presence of thin lamination. The
middle part shows karst features, while the upper part is comprised
of dolomitic limestone withoutsedimentary structures, probablyrelated
to bioturbation of a subtidal interval.
←
FIGURE 8—Polished slab of the block MPB cut on the edge between
side A and C; scale bar 5 20 cm. The main feature to note is the
interval with karst structures (halfway up the slab) that obliterate the
original rock texture.
when viewing all of side A (Fig. 11). The most impressive
feature is the channeled breccia body that abruptly cuts
the lower part of the layers and has a more-or-less flat up-
per boundary. This body is elongated in the direction of
sides A and C, and gives some information about the
shape of this bed and its formation (Fig. 12).
Paleoenvironmental Interpretation of the Track-Bearing
Blocks
Some lithological features are useful for reconstructing
the depositional environment of the dinosaur footprints.
The various lithofacies found in the Mattinata blocks
could be interpreted as indicators of a subtidal (shallow-la-
goon) setting, and intertidal- to supratidal-flat settings. A
small tidal channel is indicated by the lens-shaped, flat-
pebble breccia bed along side A of Block MPC. Desiccation
cracks and the abundance of flat clasts testify to the com-
mon occurrence of supratidal conditions. Laminated inter-
DINOSAUR FOOTPRINTS FROM SOUTHERN ITALY 541
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 bed
with irregular geometric features (see Fig. 11), and the lower part is
less dolomitized. The other lithofacies are an alternation of massive
and laminated dolostone.
vals, interpreted as cryptomicrobial laminites, indicate
the presence of algal or microbial mats. Most of the dino-
saur footprints, therefore, must be related to an intertidal
to supratidal setting.
DINOSAUR FOOTPRINTS
The Mattinata pier material is composed of 29 natural
molds and 8 natural casts; 14 molds and 3 casts are pre-
served well enough to allow good descriptions. All of the
footprints lack convex rims. This seems to confirm the
presence 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 impressed
areas. 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 the
footprints, or sliding traces in elongated footprints were
recognized. Consequently, most of footprints correspond
well 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 trackway
from incomplete and ambiguous evidence. Three speci-
mens (MPB4, MPB5, MPC2) have a very pronounced me-
tapodium impression, as long (or longer) than digit III, so
that the length of these footprints is nearly double that of
the other Type 1 specimens. The proportions suggest these
impressions are from a large part of the metatarsals made
during plantigrade gait or squatting. There are no exam-
ples of paired parallel prints, therefore suggesting a sit-
ting posture. This elongation and the hallux impression
are evident in four footprints (MPC3, MPC2, MPC12,
MPC1) associated in an irregular sequence, evidently
made 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 hallux
touching the ground. Wide-gauge theropod tracks recently
were described by Day et al. (2002, 2004) and were associ-
ated with a slow-walking phase of locomotion. A partial
trackway of two consecutive footprints (MPC19, MPC13)
was reconstructed graphically (Fig. 14), and both track-
ways suggest normal trackmaker progression, not resting
phases. This type of gait may be unusual: thus, trackway
parameters should be considered with caution.
Footprints MPA1 (Fig. 15), MPC3, and MPC11, pre-
served as natural molds, clearly show phalangeal pads on
digits I, II, and III, and less clearly on digit IV, with the
digit II metatarsophalangeal pad being diagnostic. MPA 1
seems to show a web-like structure between digits III and
IV, most probably due to folding of thin microbial laminae
around 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 this
figure, which is a channelized body with an erosional base. The irregularities in the upper part of this block are related to dinoturbation, with
normal print (compare with Fig. 7), and underprint deformation.
FIGURE 12—Detail of the breccia bed of Block MPC (maximumwidth
25 cm, see Fig. 11 for location).
FIGURE 13—Four consecutive tracks referred to the same dynamic
action of an undecided or sliding trackmaker (two tracks of the left pes
precede the impression of two tracks of the right one).
DINOSAUR FOOTPRINTS FROM SOUTHERN ITALY 543
FIGURE 14—Footprints (MPC19, MPC13) geometrically completed
by 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). Note
the central area masked by displaced carbonate mud.
also is present, as sometimes happens in squatting dino-
saurs. In these cases, the interdigit divergences are larger
and 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 partial
impression 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 and
Rainforth, 2003), but they also are known in theropod foot-
prints (e.g., Bu¨ ckeburgichnus sensu Lockley 2000; Eutyni-
chnium;‘‘Gigandipus’’; ‘‘Hyphepus’’; Jalingpus; Kayenta-
pus soltykovensis; Picunichnus; Saurexallopus; Theroplan-
tigrada; Tyrannosauripus; and an unnamed theropod
from Middle Jurassic of Morocco; Nouri et al., 2000). The
hallux often is undervalued as a character (see Harris et
al., 1996 for a review) because it is considered a kind of ex-
tramorphological feature in typical tridactyl forms, and
only is visible in the deepest impressions.
A superficial resemblance between Apulian specimens
and Anomoepus (related to ornithischian trackmakers)
concerns partial metatarsal impressions. However, the
Apulian footprints are not the traces of resting animals
with 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 Jurassic
of Portugal, in which ‘‘depth. . . [10–15 cm]. . . accounts for
the 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 a
theropod (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 appears
to be a rare case within this sample. A partial resemblance
in the position of digit I and the narrowing at the base of
the 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 Lower
Jurassic 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 not
available.
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 footprint
interpreted 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 wet
substrate’’ was described by Kvale (2001, p. 250).
Elongate theropod footprints are quite widespread (e.g.,
Kuban, 1989; Gierlinski, 1994; Pe´rez-Lorente, 1994) and
have been described either as elongated, or as footprints of
plantigrade dinosaurs. As noted above, most of the foot-
prints, including Kayentapus (sensu Gierlinski 1996), Jal-
ingpus, and Megalosauripus, sometimes show digit I and
elongate metapodium impressions. Deep impressions, or a
peculiar semidigitigrade and plantigrade gait may explain
the origin of both features.
The closest fit between the new material and known
footprints is with unnamed footprints described and fig-
ured by Aguirrezabala and Viera (1980, fig. 24) from Late
Jurassic (Kimmeridgian) sediments of Bretun (Soria,
Spain). In these footprints, digit I seems to lie in a slightly
different plane and is less deeply impressed. The similari-
ty is strong enough to consider ascribing this material to
the same trackmaker group.
The same material, subsequently figured by Pe´rez-Lor-
ente as trackways C and G (Pe´rez-Lorente, 1994, figs. 1,
2), allows favorable comparison of trackway characters
with the Mattinata material. The Spanish trackways
slightly differ in having a higher pace angulation (narrow-
er interpedes distance).
Theroplantigrada, an ichnotaxon based on Aptian
tracks from La Rioja (northern Spain) ascribed to a thero-
pod (Casanovas Cladellas et al. 1994), is similar to the
Mattinata material and to trackways described by Aguir-
rezabala and Viera (1980). This monotypic ichnogenus is
based on a single trackway in which, except in size, the
footprints seem quite similar to the Mattinata material in
the presence and position of the hallux and the metapo-
dium impression. All of the Spanish material shows very
elongated footprints. The Apulian material resembles
trackway A and some footprints of trackway D (Casanovas
Cladellas 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 feature
of 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 not
permit confirmation of whether web traces truly are pre-
sent. Apparent web traces are lacking in all the Mattinata
footprints described herein, except in MPA1, just in front
of the hypex III-IV (Fig. 17). However, this structure was
generated by the inflexion of microbial laminae some cen-
timeters away from the real boundaries of the footprint.
If webbing really is present in Theroplantigrada,it
would be a rare case among dinosaurs. More likely, it is an
extramorphological feature. Only three purported cases
were cited by Thulborn (1990)—Otouphephus magnificus,
Swinnertonichnus mapperleyensis, and Talmontopus ter-
si—and in all three cases, the web traces subsequently
were interpreted as extramorphological features. For
Otouphephus magnificus, ‘‘the web-like trace . . . was later
shown 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 Otouphephus
and Talmontopus, King and Benton (1996, p. 221) reported
DINOSAUR FOOTPRINTS FROM SOUTHERN ITALY 545
FIGURE 18—Photograph and outline drawing of the best-preserved
Type 2 footprint (MPC10).
FIGURE 19—Outline drawings of other Mattinata footprints. (A) Type
3 footprint (MPC17). (B) Undetermined specimen (MPA3).
that ‘‘. . . Lockley (pers. comm., 1994) cannot confirm the
presence of webbing in either taxon.’’
In the opinion of some authors, ichnogenera should cor-
respond to footprints revealing diagnostic differences in
definite skeletal structures of the trackmakers (Carrano
and Wilson, 2001). Consequently, if the supposed webbing
in 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 Mattinata
material, trackways C and G described by Aguirrezabala
and 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. Similar
trackways and tracks are known from the Upper Jurassic
of Soria, Spain (Aguirrezabala and Viera, 1980); ‘‘Eutyni-
chnium’’ lusitanicum in the Uppermost Jurassic of Portu-
gal; an unnamed trackway from Middle Jurassic deposits
of Morocco (Nouri et al., 2000); Jalingpus yuechiensis from
the Upper Jurassic of Sichuan, China; Kayentapus solty-
kovensis from the Lower Jurassic of Poland, Sweden, and
Hungary; and Theroplantigrada encisensis from Early
Cretaceous beds (probably Aptian in age) of northern
Spain. All represent medium-sized dinosaurs that left hal-
lux and metapodium traces. This kind of footprint cannot
be ascribed easily to a zoological group below the level of
theropod trackmaker or, perhaps, to a cursorial, light or-
nithopod (Thulborn, 1990; Viera and Torres, 1992; Farlow
and Lockley, 1993). Accepting the more probable hypoth-
esis of a theropod trackmaker, it is inferred that these
footprints represent a medium-sized, non-derived thero-
pod (a ceratosaur) or a basal tetanuran where digit I is not
reduced.
Type 2
Only three stout, tridactyl footprints preserved as nat-
ural casts (MPC4, MPC10, MPB7) are included in this
type (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 a
plethora 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 by
Lockley et al. (1996; 2000a) and Olsen et al. (1998) have
shown that detailed analyses can allow separation of at
least some theropod footprints and trackways. According
to older literature, material similar to the Mattinata
prints could be referred to Eubrontes or to Megalosauripus
(for a review on the debate around this last name and a
suggested solution, see Lockley et al., 2000a; Lockley and
Meyer, 2000). However, studied specimens lack the so-
called heel, or proximal area, made by the impression of
metatarsal-phalangeal pad of digit IV. The Mattinata
specimens differ in this character from most other thero-
pod footprints, which are characterized by the impression
of the metatarsophalangeal pad of digit IV. Two excep-
tions are Carmelopodus untermannorum, a small footprint
from 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 a
fourth proximal pad on digit IV is stressed’’ (Lockley et al.,
1998, p. 260). The Mattinata footprints and the Utah and
Australian prints differ in other characters, but it is inter-
esting that this is another example of the lack of such an
important feature in mid-Jurassic material. Consequent-
ly, the Mattinata specimens may share this functional
character with Carmelopodus. Due to the lack of related
ichnotaxa, no stratigraphical inferences are possible. It is
only possible to note that Carmelopodus comes from Mid-
dle Jurassic deposits, Skartopus from the Mid-Cretaceous,
and there are similar subdigitigrade theropod tracks in
the 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-defined
pads 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 that
they 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 of
Therangospodus oncalensis, an ichnospecies first consid-
ered Early Cretaceous in age, still seems questionable (see
Lockley et al., 2000b).
Other Footprints
The remaining footprints, although easily recognizable
as dinosaur traces, are preserved too poorly to be ascribed
to 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 also
shows sharp claw traces and thus could represent a the-
ropod trackmaker. MPC6 and MPA4 could be ascribed to
Type 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 individuals
that crossed the area.
Notes on the Ichnocoenosis
Although the Mattinata pier material still presents
many unsolved problems, it allows inferences to be made
about paleobiogeography (discussed below), and throws
light on the presence of a theropod-dominated track as-
semblage with inferred correspondence to the assemblage
described by Lockley et al. (2000a) and by Lockley and
Meyer (2000), which includes Megalosauripus and Ther-
angospodus. Such ichnocoenoses have been recorded from
Uzbekistan, 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 to
Theroplantigrada is recorded. This difference could reflect
age or facies controls affecting the Apulian Platform ichn-
ocoenosis.
CONCLUSIONS
Age of the Blocks
Direct biostratigraphic or chronological calibration of
the age of the material is lacking, but three lines of evi-
dence allow inference of the approximate geologic age of
the Mattinata blocks: (1) provenance of the blocks, (2) the
lithofacies recognized in the blocks, and (3) the footprint
types from the blocks.
(1) Documents about the pier building are related to the
extraction of blocks from quarries in the San Giovanni Ro-
tondo area and Apricena. The former were opened mainly
in the San Giovanni Rotondo Limestone, and this unit is
never dolomitized. In the Apricena area, the only interval
where block extraction continues is in the San Giovanni
Rotondo Limestone, where no dolomitic layers are known.
(2) Some beds in small, abandoned quarries near Monte
Calvo (Fig. 1) in the Sannicandro Formation reveal simi-
lar sedimentological features to the footprint-bearing
blocks, and are partially dolomitized. The age of this part
of the Sannicandro Formation is referred to the Kimmer-
idgian–Tithonian interval (Morsilli, 1998). The same lith-
ological and sedimentary characteristics also can be found
near the San Nazario area (see Fig. 1) in the northwest
part of the Gargano. Here, some old quarries, in some cas-
es dismantled by the construction of a road, reveal very
similar lithological and sedimentary features. The age of
this succession is the same as the one in the Monte Calvo
area (Luperto Sinni and Masse, 1994).
(3) Although cautiously avoiding attribution of the foot-
prints to named ichnospecies, the overall character of the
assemblage suggests a Late Jurassic age, although these
forms were not exclusive to that time interval.
In conclusion, these lines of evidence suggest that the
studied material comes from the Late Jurassic Sannican-
dro Formation.
Paleogeographic Inferences and Problems
During Middle–Late Jurassic and Cretaceous times, the
Apulian Platform has been interpreted as a small, isolated
carbonate 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 the
southern 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 repeated
discoveries of dinosaur footprints on the Apulian Plat-
form. Recent finds of dinosaur trackways from different
places and stratigraphical intervals are very strong paleo-
geographic constraints that necessitate reconsideration of
previous interpretations. At first, this new evidence may
be considered weaker than geophysical or structural data,
but dinosaur footprints may serve as powerful new tools to
change the best-built paleogeographic models. Other
workers 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 mass
with the northeastern part of the Massif Central’’; Kvale
et 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 suggested
for the Late Jurassic by Gierlinski and Niedzwiedzki
(2002, p. 58A).
At present, three different track assemblages in the
Apulian region have been found. The first, described here-
in, is a theropod-dominated ichnoassociation, probably
Late Jurassic in age. The second, described from the Early
Cretaceous (Late Hauterivian–Early Barremian) of Borgo
Celano (Gianolla et al., 2000), contains footprints ascribed
to theropods, ornithopods, and perhaps sauropods; and
the third, described from Late Cretaceous (Santonian)
from Altamura (Andreassi et al., 1999; Nicosia et al.,
DINOSAUR FOOTPRINTS FROM SOUTHERN ITALY 547
2000a, b), is ornithopod dominated. These are the most
striking evidence of widespread, terrestrial vertebrates in
the area. Other data, such as the so-called Ruvo varanoid
(Varola, 1999) and vertebrate remains recorded from the
Melissano Limestone (Medizza and Sorbini, 1980), also
are evidence of terrestrial vertebrates in this area. Finds
of 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 events
recognized elsewhere. Moreover, this data set probably is
biased by taphonomic events and by reduced availability
of Jurassic and Cretaceous rock outcrops with track-bear-
ing potential. This kind of research needs extensive, un-
disturbed, well-exposed bedding surfaces, a situation not
found 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 the
trackways only are exceptional occurrences, they could be
related to short-term, repeated connections to larger land
areas. 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 a
paleontological point of view, these hypotheses correspond
either to repeated immigration pulses, or to a single, early
colonization 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 areas
were not separated by deep basins, and sea-level drops
would have allowed emergence of the platform followed by
land-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 need
complex evolutionary hypotheses.
The second hypothesis must, in turn, be subdivided to
two sub-hypotheses: either the platform was as small as
current models suggest, with different portions of the in-
ner platform emerging from time to time, or the emergent
area was much larger than thought, although no traces of
it are preserved. In the first scenario, lacking any evidence
of a single, persistent island area large enough to sustain
a balanced community of huge animals, land vertebrates
must have been wandering continuously from one zone to
another within the same platform area—an idea that does
not consider the need of dinosaurs for vegetation, fresh
water, and nesting sites. Simple emergent episodes, even
if 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 (including
fresh-water sources and nesting sites), in which the persis-
tence of a balanced fauna was possible for ;130 Ma, with
all traces of such a large area removed by subsequent geo-
logical events. In both cases, balanced ecosystems imply
the presence and the coevolution of plants, herds of plant-
eating animals, and meat-eating dinosaurs. It also must
be 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 environment
raises 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 presence
of fossils. In fact, a lack of river systems could have re-
duced the chance for fossilization by reducing deposits
available to preserve dinosaur bones and/or footprints.
Both cases included in the second hypothesis present
another important concern. The recorded Apulian dino-
saur ichnocoenoses apparently differ from one another,
but may correspond in both the faunal composition and
the 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, and
in North America. It is difficult to accept a hypothesis of
parallel evolution on the mainlands and isolated carbon-
ate-platform areas.
In any case, the presence of a large number of dinosaurs
in an area such as the Apulian Platform is problematic. It
underlines the inadequacy of paleogeographic models that
simply suggest a tectonic disjunction of the platforms. On
the contrary, the footprint and other data strongly support
a 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. If
recognized, they might be considered partial explanations
or just the record of the final phase of endemic evolution
after immigration.
Bosellini (2002) used these new data for strong paleo-
geographic constraints. Reviewing various geological and
geophysical data associated with the presence of dino-
saurs around the Ionian Sea and surrounding areas, Bo-
sellini (2002) reached the conclusion that the Apulian
Platform 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 Mesozoic
carbonates in Crete. In Bosellini’s (2002) reconstruction,
the Apulian Platform, and probably other peri-Adriatic
platforms, were not isolated Bahamian banks, but rather
were more like the modern Florida peninsula. This model
conflicts strongly with other paleogeographic models (e.g.,
Catalano et al., 2001). In fact, using these data, an eastern
connection 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 all
data sources, such as geophysical (seismic refraction and
reflection profiles of the Apulian Platform margin and sur-
rounding basins), structural and kinematic (relationships
between thrust-belt chains and foreland areas), strati-
graphical, sedimentological, and paleontological data.
ACKNOWLEDGEMENTS
Marco Avanzini is warmly thanked for his suggestion
about 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 kindly
helped our work.
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ACCEPTED DECEMBER 28, 2004
