Carvalho and Leonardi Pereiros Ichno Final

Cretaceous Research 144 (2023) 105446
Contents lists available at ScienceDirect
Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes
Dinosaur tracks from the Sítio Pereiros ichnosite, Triunfo Basin (Lower
Cretaceous) and the dinosaur diversity in the Rio do Peixe basins,
Northeastern Brazil
Ismar de Souza Carvalho a, b, *, Giuseppe Leonardi a, c, **
a ria e Ilha do Funda
Universidade Federal do Rio de Janeiro, Departamento de Geologia, CCMN/IGEO 21.949-900 Cidade Universita ~o, Rio de Janeiro, Brazil
b
Universidade de Coimbra, Centro de Geoci^ encias, Rua Sílvio Lima, 3030-790, Coimbra, Portugal
c
Istituto Cavanis, Dorsoduro, 898, 30123, Venezia, Italy
a r t i c l e i n f o a b s t r a c t
Article history: The dinosaur tracks in the Rio do Peixe basins (Lower Cretaceous, Rio da Serra-Aratu stages) occur in at least 39
Received 28 June 2022 individual tracksites through approximately 98 stratigraphic levels in the western part of the State of Paraíba,
Received in revised form Brazil. The Triunfo basin (one of the four Rio do Peixe basins) is a 480 km2 asymmetric graben, located in the
22 November 2022 ~o Jo~
counties of Sa ao do Rio do Peixe, Uiraúna, Poço, Brejo das Freiras, Triunfo, and Santa Helena, controlled by a
Accepted in revised form 28 November
2022
NE transcurrent fault system. To date, only four isolated footprints and two incomplete trackways have been
Available online 6 December 2022 identified in the Antenor Navarro Formation. Among the isolated footprints, three probably belong to the-
ropods. One incomplete trackway consists of just two digitigrade, rounded digits, suggesting they were made
by a small ornithopod. In this study we describe a new ichnosite, located at Sítio Pereiros, Sa ~o Jo~
ao do Rio do
Keywords:
Footprints Peixe county, Paraíba State. The one and a half meter thick succession of fine-grained sandstones, siltstones,
Dinosaur tracks mudstones and shales with ripple marks, climbing ripples and mud cracks of the Sousa Formation reveals a
Triunfo basin bedding plane with three trackways, with a total of 19 tridactyl, mesaxonic footprints. These trackways are
Lower Cretaceous interpreted as produced by theropods, two large and one smaller. In these beds there are also ostracods,
spinicaudatans (conchostracans), and fragments of microvertebrates (fish scales, teeth and bones). The Sítio
Pereiros ichnosite represents a deposition in a floodplain area, with temporary aerial exposure of the su-
perficial sediments in which tracks were impressed. The ichnofauna from this locality increases knowledge of
the theropod fauna from the Triunfo basin and the distribution of the dinosaur tracks throughout the interior
basins of Northeastern Brazil.
Description of these new theropod tracks permits evaluation of the behavior of these three theropods,
including inferences about trackmaker speed and the type of gait of the three animals, and also of their
possible size. This is the 40th ichnosite in the Rio do Peixe basins, extending analysis of the types of
trackmaker associations present at such ichnosites, as well as the dinosaur diversity represented at each
of them. New interpretations are presented about the environments, and the relationship between the
various groups represented in this region during the Early Cretaceous.
© 2022 Elsevier Ltd. All rights reserved.
1. Introduction
The Triunfo basin (also called the Uiraúna-Brejo das Freiras

basin) is one of the four sedimentary sub-basins of the Rio do Peixe
basins. The others are Sousa, Pombal, and Vertentes (Fig. 1), located
* Corresponding author. Universidade Federal do Rio de Janeiro, Departamento
in the western part of the Brazilian State of Paraíba. The Triunfo
de Geologia, CCMN/IGEO 21.949-900 Cidade Universita ria e Ilha do Fund~
ao, Rio de basin occur in the counties of S~ao Joa~o do Rio do Peixe, Uiraúna,
Janeiro, Brazil. Poço, Brejo das Freiras, Triunfo and Santa Helena, with 480 km2
** Corresponding author. Universidade Federal do Rio de Janeiro, Departamento area. The Precambrian basement is composed of igneous (granites,
de Geologia, CCMN/IGEO 21.949-900 Cidade Universita ria e Ilha do Fund~
ao, Rio de
gabbros, and diorites) and metamorphic (migmatites, gneisses,
Janeiro, Brazil.
E-mail addresses: ismar@geologia.ufrj.br (I.S. Carvalho), leonardigiuseppe879@
quartzites, and marbles) rocks. It is an intracratonic asymmetric
gmail.com (G. Leonardi). graben, controlled by a NE transcurrent fault system, developed
https://doi.org/10.1016/j.cretres.2022.105446
0195-6671/© 2022 Elsevier Ltd. All rights reserved.
I.S. Carvalho and G. Leonardi Cretaceous Research 144 (2023) 105446
Fig. 1. Location map of the Triunfo basin in the context of the interior Northeastern Cretaceous basins of Brazil and location (marked with a star) of Sítio Pereiros ichnosite
(Carvalho, 2004).
2
along preexisting structural trends of the basement during the during the opening of the South Atlantic Ocean (Ponte, 1992;
opening of the South Atlantic Ocean (Carvalho et al., 2013a,b). The Mabesoone, 1994; Valença et al., 2003; Araújo et al., 2018; Rapozo
Mesozoic age of the sedimentary deposits in the Rio do Peixe ba- et al., 2021; Oliveira et al., 2022). Deposition occurred along the
sins, based on ostracods (Sousa et al., 2018, 2019) and palynological faulted borders of the basins as alluvial fans (Mabesoone et al.,
material, is characteristic of the Rio da Serra (Berriasian to Hau- 1979), changing to an anastomosing fluvial system more distally
terivian) and Aratu (early Barremian) local stages (Lima and Coelho, and also a meandering fluvial system with a wide floodplain with
1987; Regali, 1990). perennial and temporary lakes (Lima Filho et al., 1999; Carvalho,
An important aspect of the Rio do Peixe basins is the abundant 2000a; Co rdoba et al., 2008).
tetrapod ichnofauna consisting of footprints and trackways, mainly There are a few body fossil occurrences, including Ostracoda,
of theropods, sauropods, and ornithopods. There are also inverte- Spinicaudata, Crocodyliformes and Dinosauria in the central-south
brate ichnofossils, such as traces and burrows produced by ar- region of the basin, and both invertebrate and vertebrate ichno-
thropods and annelids (Fernandes and Carvalho, 2001). The fossils close to the northern margin (Carvalho, 1989; Leonardi,
abundance, distribution in the same stratigraphic levels and similar 1994; Carvalho, 1996, 2004, 2014; Carvalho and Nobre, 2001;
age allows us to interpret the numerous ichnosites from these ba- Carvalho et al., 2002; Carvalho et al., 2017). The skeletal elements
sins as a megatracksite (Viana et al., 1993, sensu Lockley, 1991; assigned to a dinosaur (Sauropoda), named Triunfosaurus leonardii,
Carvalho et al., 1993a,b, 1995a,b; Siqueira et al., 2011; Lockley and were found in the context of gravel bars of channeled flows (Rio
Meyer, 2022). Leonardi and Carvalho (2021) had recorded 37 ich- Piranhas Formation). This is one of the oldest Cretaceous Titano-
nosites; two new ones were discovered by Leonardi (2021); sauriformes of South America, so common in the Late Cretaceous of
another one is registered here, for a total of 40 ichnosites. Although Brazil and Argentina, which opens new perspectives on paleogeo-
the Triunfo basin presents few occurrences of tracksites in the graphical and temporal distribution of titanosaurian sauropods
context of the Rio do Peixe megatracksite (Leonardi and Carvalho, (Carvalho et al., 2014; Carvalho et al., 2017).
2021; Lockley and Meyer, 2022) it is a good instance in which In the Cretaceous sediments of the Triunfo basin there are no
fossil footprints provide the main information about the dinosaur microfossils to dating. By analogy with the sediments dated by
fauna of the region. microfossils and palynology in the Sousa basin (Lima and Coelho,
The analysis of a new tracksite, located at Sítio Pereiros, in- 1987; Regali, 1990; Sousa et al., 2018, 2019), and considering the
creases knowledge concerning the stratigraphic, spatial distribu- similarities among the vertebrate ichnofaunas, the main deposi-
tion and environments in which dinosaur tracks occur in the tional phase in the Triunfo basin probably dates from between the
Triunfo basin, showing the diversity of preservation modes of a Rio da Serra and Aratu stages (Berriasianelower Barremian).
theropod track assemblage.
3. Methodology
2. Geological context
The ichnological terminology and methods used in this study
The Triunfo basin is a depositional area located in the Borbor- mainly follow Leonardi (1979a, 1987, 1994, 1997). The tracks are
ema Tectonic Province, Northeastern Brazil. The succession is distributed on the surface of a stratum with a partial surface
composed of Devonian marine deposits of the Santa Helena Group exposure. Then, to allow better observation of the distribution of
(Roesner et al., 2011; Silva et al., 2014), overlain by Cretaceous the tracks they were partially excavated using chisels and ham-
fluvio-lacustrine and Recent alluvial deposits (Sousa et al., 2019). A mers, removing the overlying strata that covered the surface with
formal lithostratigraphic subdivision of the Cretaceous established the footprints. After the excavation of the rock surface with foot-
the Rio do Peixe Group, and subdivided it into the Antenor Navarro, prints at Sítio Pereiros three trackways attributed to two large and
Sousa, and Rio Piranhas formations (Mabesoone, 1972; Mabesoone one smaller theropod were observed, with a total of nineteen
and Campanha, 1973/1974). The Cretaceous origin of this basin was footprints.
a result of reactivation of basement transcurrent faults. The tilted This surface was gridded using nylon threads stretched at right
blocks created a pronounced rupture in terrain topography, and in angles forming squares with sides 1 m long, allowing the graphic
the southern part of the basin the decrease in gradient favored the representation of the parameters of footprints and trackways. The
establishment of meandering fluvial and lacustrine environments. drawing of each 1 1 m square was then reproduced on a suitable
Roesner et al. (2011) identified Early Devonian rocks through smaller scale on graph paper, following the trackway meter by
palynological analysis from boreholes, showing a pre-Mesozoic meter. The individual photographs of the footprints were made
history related to a Paleozoic deposition in a very distinct context with a digital camera, and when contrast with the surrounding
of the Cretaceous sedimentation. Arai (2006) showed that the matrix was not clear they were delimited using chalk. The main
Mesozoic deposition (Rio do Peixe Group) resulted from crustal procedures established by Falkingham et al. (2018) for document-
extension, at the inflection of NEeSW and E-W faults during Rio da ing fossil ichnological data and for distinguishing tetrapod tracks
Serra time (BerriasianeHauterivian). from other structures (Lallensack et al., 2022) were also followed,
The Antenor Navarro and Rio Piranhas formations are composed despite the use of 3D imagery.
of clastic rocks: breccias, conglomerates, sandstones, siltstones,
shales, and mudstones, near the faulted margins of the basin. The 3.1. Sítio Pereiros tracks
main sedimentary structures are cross-channel and planar strati-
fication. The deposits of the Sousa Formation comprise shales and The dinosaur track occurrences in the Rio do Peixe Group (Rio do
mudstones interbedded with fine sandstones and siltstones. Peixe basins), that crop out over an area of about 1730 km2, are
The main sedimentary structures are ripple marks, climbing regarded as a megatracksite (megatracksite Type 3 sensu Lockley
ripples, mud cracks, convolute lamination, raindrops and features and Meyer, 2022) composed of the 39 tracksites (40 with this
indicating liquefaction (Carvalho, 2000a,b; Carvalho and Leonardi, new one described here). Altogether, the 21 sites of the Sousa
1992; Fernandes and Carvalho, 2001; Nogueira et al., 2015; Formation in the Sousa Basin represent at least 98 stratigraphic
Leonardi and Carvalho, 2021). levels dominated by isolated footprints and trackways of carnivo-
These deposits reflect synrift sedimentation controlled by tec- rous dinosaurs (Leonardi and Carvalho, 2021; Leonardi, 2021). In
tonic activity along preexisting structural trends of the basement total then, in the three formations that correspond to the Rio do
3
Peixe Group, there are 100 levels with ichnofaunas (including the right feet placed directly beneath the body. In the pendulum on
description in this study). The locality of Sítio Pereiros ichnosite is a parasagittal planes that they performed walking or running, if they
new one, located at Sítio Pereiros (6 470 2000 S/38 290 1300 W), Sa ~o made a slight turn laterally with the legs, they did so, in the specific
Jo~
ao do Rio do Peixe county, Paraíba State, in the Triunfo basin. It is case, almost just to avoid crossing them.
an instance of the importance of the footprints to provide infor- In a few instances, in the basins of the Rio do Peixe, as in other
mation about the dinosaur fauna of a region. basins, the mark of the tail is preserved, but not here. The rareness
The site a 1.5-m-thick succession of fine-grained sandstones, car- of tail impressions is usual for dinosaur trackways. Bipeds as well
bonate sandstones, mudstones and shales with ripple marks, climbing as quadrupeds, all of them almost always kept their tails clear of
ripples and mud cracks of the Sousa Formation. In these strata, besides the ground. This theme is well played by Kim and Lockley (2013),
the dinosaur footprints, there are also ostracods, spinicaudatans who describe and comment on 38 records of dinosaur tails in the
(conchostracans), and microvertebrate fragments (fish scales, teeth world. Then one can add the four cases of probable tail tracks of
and bones). The age of this deposit probably dates from Rio da Serra- dinosaurs from the Rio do Peixe basins (Leonardi and Carvalho,
Aratu stages (Berriasianelower Barremian, Lower Cretaceous) by 2021, pp. 376).
analogy with the sediments dated by ostracods and palynology in the
Sousa Basin, and the similarities among the ichnofaunas. 3.4. Bearings
The exposed superficial bedding plane (Fig. 2), of about 60
square meters, shows three trackways, with a total of 19 tridactyl, The two trackways SJSP 1 and SJSP 3 are almost parallel, and
mesaxonic (i. e. roughly symmetrical about the axis of the 3rd digit) only slightly divergent (by 7 ), and are roughly directed towards an
footprints preserved in a mudstone partially covered by a fine approximate bearing of SW, their makers advancing more or less
sandstone surface with ripple marks (Fig. 3A). Two trackways are parallel to the ripple mark crests. In contrast, SJSP 2 crosses these
subparallel (heading ~ SW) and one crosses in a ~E to W direction previous trackways, as well as the ripple mark crests, and its maker
(Fig 3B). The two SW trackways (SJSP 1 and SJSP 3) present tridactyl proceeded in an approximate E-W direction. Consequently, we can
footprints, with pointed digits. The other trackway (SJSP 2) shows 5 say that both rather large theropods that produced the trackways
smaller tridactyl footprints with pointed digits. They are inter- SJSP 1 and SJSP 3 were probably paralleling the water line or water's
preted as produced by two large and one smaller theropods. edge of a most likely alkaline lake. The trackmaker of the SJSP 2
The trackway SJSP 1 (Fig. 4, Appendix A and B) has eight trackway (a smaller theropod), instead, was probably exiting or
consecutive footprints regularly spaced. The footprints are tridactyl, entering the water.
mesaxonic and show pointed digits. In six of them it is possible to It can also be observed that both theropods that produced the
observe distinct impressions of claws. The trackway shows that its almost parallel trackways SJSP 1 and SJSP 3, walked more or less
maker proceeded on a bearing of approximate SW (213 ), had an parallel to the direction of the transcurrent fault NE separating inter
average inner width of 0.10 m and a total length of at least 7.8 m. alia the Triunfo basin from the Sousa basin. As we have observed
The length of steps (oblique pace) varies from 1.00 m to 1.40 m, elsewhere (Leonardi, 1979b, 1991; Leonardi and Carvalho, 2021) the
with an average value of 1.14 m. The stride varies from 2.10 m to dinosaurs were often influenced in their direction of progression by
2.70 m, with an average value of 2.37 m. The step angle varies from features of the landscape, and particularly by the margins of the
150 to 164 , with an average value of 158 400 (Appendix B). bodies of water; and in turn the landscape and bodies of water were
Trackway SJSP 2 (Fig. 5, Appendix A and B) shows five consec- controlled by the regional fault lines, transcurrent or not. It should
utive footprints (Fig. 5A). They are tridactyl, mesaxonic and show also be noted that the margins of the bodies of water conditioned,
acuminate digits, with digit III the longest one. This trackway pre- anyway, apart from their direction, the impression and preserva-
sents, as a whole, almost a WeE bearing (towards an azimuth of tion of footprints of the local fauna. Without any doubt, in a semi-
86 ). This trackway has an average inner width of 0.10 m and a total arid region such as the Triunfo basin, it was only on the edge of the
length of at least 5.50 m. bodies of water e as today e that the ground was sufficiently wet
The trackway SJSP 3 (Fig. 6, Appendix A and B) has six consecutive and plastic to be deformed, and then receive and maintain the
regularly spaced footprints. They are tridactyl, mesaxonic and show footprints of the feet of the “passers-by”. In addition, again, it was
pointed digits. The trackway presents an almost SW (206 ) bearing, only in environments of this type that the soil could gradually leave
an average inner width of 0.18 m and a total length of at least 4.00 m. the tracks to consolidate and dry, so that they could possibly sur-
The length of steps (oblique paces) varies from 1.00 m to 1.20 m, with vive the next rainstorm or elevation of the level of the lakes. The
an average value of 1.13 m. The stride varies from 2.15 m to 2.65 m, presence of bacteria and microscopic algae also helped the pres-
with an average value of 2.36 m. The step angle varies from 162 to ervation of the footprints, not only in general, but also specifically
178 , with an average value of 171150 (Appendix B). in these sediments e the microbialites e of the Rio do Peixe basins
(Carvalho et al., 2013a,b).
3.2. The behavior of the trackmakers
3.5. Manners of gait
As already noted, the footprint/sediment relationship indicates
that the three dinosaurs recorded in the locality Sítio Pereiros based In SJSP 1 trackway, the index SL/FL (stride length/footprint
on three fossil trackways of the Early Cretaceous did not act as an length) is 9.12, which indicates that the animal was walking in a
organized group, and that they were not inferred to have been certain speed, but did not run. As for the 2nd trackway, SJSP 2, the
registered at the same time. The same result is also shown by the fact that the animal was running is also indicated by the very high
behavior of these animals. SL/FL index (stride length/footprint length) ¼ 16.87. We will later
see at what speed this index can correspond. The trackmaker of the
3.3. Posture SJSP 3 trackway walked with a hasty gait, but did not run, as can be
seen from the fact that the index SL/FL (stride length/footprint
Theropods, large and small, were obviously nearly all bipedal length) is 9.64. In this, its gait was very similar to that of the SJSP 1
(Molnar and Farlow, 1990, as for the large theropods), and produced trackway.
very narrow trackways. Theropods, and especially the three from Trackways SJSP 1 and 3 are straight; SJSP 2 is instead slightly
Sítio Pereiros ichnosite, kept the hind legs erect, with the left and curved to the right (see Fig. 5A). Such fossil curved trackways are
4
Fig. 2. Outcrop of Sousa Formation in the Triunfo basin at Sítio Pereiros ichnosite (A) and the footprints distribution in the surface strata (B) of Sousa Formation.
not very common. For example, in the Sousa basin, among 75 dinosaur tracks in Brazilian basins (and in all the others) are usually
reasonably long trackways that can be examined for this purpose, indeed straight, because the trackmakers went from one place to
only two are really curved rather than straight, those of Engenho another in a purposeful way (cf. Leonardi and Carvalho, 2021: fig.
Novo 1 (ANEN 1) and Piau-Caiçara 43 (SOCA 43). In contrast, the 4.66, Passagem das Pedras site).
5
Fig. 3. Surface of the Sousa Formation strata (Triunfo basin) with the theropod trackways of the Sítio Pereiros ichnosite. Footprints preserved in a mudstone with mud cracks and
superimposed ripple marks (A); two trackways (SJSP 1 and SJSP 3) are subparallel (SW) and one (SJSP 2) crosses them proceeding from W to E (Fig. 3B).
3.6. Speeds the hip joint from the ground) using the formula: h ¼ yFL, where y
is an allometric coefficient, specific to each clade or form of dino-
We calculated the three trackmakers speeds (km/h), following saur. The approximate speeds on the basis of the above data and the
the method and formulas of Alexander (1989) as improved by formulas are as follows: SJSP 1: Vc z 8.7 km/h; SJSP 2:
Thulborn (1989, 1990). We calculated the value of h (the height at Vc z 12.9 km/h; SJSP 3: Vc z 9.09 km/h. These data also correspond
6
Fig. 4. Footprints from the SJSP 1 trackway, Sítio Pereiros ichnosite, Sousa Formation (Triunfo basin). Scale bar 5 cm.
7
Fig. 5. Trackway SJSP 2 (A) and detail the its footprints (B, C, D, E and F), at Sítio Pereiros ichnosite, Sousa Formation (Triunfo basin). Scale bar 20 cm (A) and 5 cm (B, C, D, E, F).
8
Fig. 6. Footprints from the SJSP 3 trackway, Sítio Pereiros ichnosite, Sousa Formation (Triunfo basin). Scale bar 5 cm.
9
rather well to the calculation of the relative length of stride, or kind in this basin (Sousa Formation) is certainly related to the role of
of gait (Thulborn, 1989, 1990). biofilms in their consolidation. Petrographic slices show biofilms,
The three trackways on Sítio Pereiros ichnosite come to be in the disrupted carbonate microbial laminites, interbedded with very
middle band of the speeds of the tracks already calculated else- thin terrigenous microclastic beds, associated with footprints of
where for the basins of the Rio do Peixe, and also in that of the Sousa Basin (Carvalho et al., 2013a,b).
world ichnosites calculated elsewhere. There are few previously discovered tracks from the Triunfo
For comparison, the mean calculated speed (Vc) for the medium basin. To date, only four isolated footprints and two incomplete
and large theropods of the Sousa Basin, (n ¼ 62 individuals, trackways have been identified there in the Antenor Navarro For-
including SOCA 493, 991, 150; 82.67% of all the trackways of this mation (Carvalho, 1989, 1996; Leonardi, 1994; Leonardi and
sample) is 5.57 km/h (Leonardi and Carvalho, 2021, p. 364). Again, Carvalho, 2021). The lithologies of these units are coarse, imma-
the average calculated speed for all dinosaur tracks from the Sousa ture, poorly sorted, detrital sediments interbedded with siltstones,
Formation alone (n ¼ 55; 73.33% of all the trackways) is 5.23 km/h that were certainly a restrictive factor for fossil track preservation.
(Leonardi and Carvalho, 2021). The interpretation of these deposits is fan delta, alluvial fan, and
braided/anastomosing fluvial environments. The footprints were
3.7. Social or individual behavior probably produced during periodic interruption of sedimentation
owing to discharge fluctuations toward the base of an alluvial fan
We have already observed above that the three animals passed deposit (Carvalho, 2000a,b). Despite the low preservation potential
through the affected area at different times (though rather close), of alluvial fans and braided fluvial systems, dinosaur tracks and
and that therefore there was no direct interaction between them, trackways are found in these contexts in the Sousa basin in fine
except that they lived, hunted, they fought for life in the same area sediments that were accumulated as subaerial sand bars (Carvalho,
and at the same time. All of them are interpreted as theropod 1989, 1996; Carvalho and Leonardi, 1992). Among the previously
trackways of large size (SJSP 1 and SJSP 3) and respectively of described dinosaur tracks from Triunfo basin there are isolated
smaller size (SJSP 2). Thus, in this case, we cannot speak of social footprints, three of them probably belonging to theropods. The
behavior among these theropods, but rather of individual behavior, others' poor preservation does not allow the identification of their
as indeed is quite common for theropods. They seem to have been makers. One incomplete trackway consists of just 2 digitigrade,
lone travelers, not only in these basins of the Rio do Peixe, but also tridactyl, and mesaxonic footprints. There is a protuberance at the
in many other ichnosites in the world, unlike herbivores, especially proximal outline of the footprints that would correspond to digit I
sauropods, which in the Sousa Basin e and in many other places or to a more basal pad of digit IV. The rounded extremities of the
and times in the world e often traveled in large herds. digits, without clear claw marks, suggest they were made by a small
Another surprising fact, here and elsewhere, is that there is no ornithopod (Carvalho and Leonardi, 2021).
trace of any interest of a track-maker with regard to trackways The new tracks of Triunfo basin, the ones described herein,
imprinted by individuals that passed before. No deviation, no occur in a distinct geological context (Carvalho and Leonardi, 2022).
deceleration, no sign of interest. The tracks continue (here on Sítio They are found in mudstones intercalated with fine sandstones and
Pereiros ichnosite and elsewhere) without showing that their siltstones of the Sousa Formation, in a surface with primary and
makers have moved away to take an interest in other dinosaur secondary mud cracks. The sediments that formed the substrate
tracks. over which the dinosaurs traveled allowed a better footprint
preservation when compared with the coarse sediments of an al-
4. Discussion luvial fan. As the Sítio Pereiros ichnosite shows footprints preserved
in fine clastic sediments (mudstones) they present a more detailed
The way a footprint is preserved relates to its geological context. morphology.
Its preservation for posterity requires specific substrate cohesive- The morphology of the tracks results from a set of factors such as
ness, plasticity, grain size, texture, water content and microbial the anatomy and behavior of the producer, syn-depositional and
processes (Leonardi, 1979a,b, 1984a,b, 2011; Lockley et al., 1989; post-depositional processes (Peabody, 1955; Falkingham et al.,
Lockley and Meyer, 2000; Avanzini et al., 2000; Leonardi and 2010; Avanzini et al., 2012; Belvedere and Farlow, 2016;
Mietto, 2000; Dalla Vecchia, 2008; Marty et al., 2009; Li et al., Marchetti, 2018; Marchetti et al., 2019, 2020). An important aspect
2011; Lockley and Xing, 2015; Getty et al., 2017; Pe rez-Lorente, concerning the morphology is the consistency of the sediments
2015, 2017; Citton et al., 2015; Castanera et al., 2016; Falkingham (Laporte and Behrensmeyer, 1980; Lockley, 1991; Nadon, 2001;
et al., 2016; Melchor et al., 2019; Noffke et al., 2001, 2019; Gatesy, 2003; Mila n, 2003; Dalman and Weems, 2013; Falkingham
Abrahams et al., 2020; Romano and Citton, 2020; Belvedere, 2008; and Gatesy, 2014; Belvedere et al., 2017; Gatesy and Falkingham,
Belvedere et al., 2022; Carvalho et al., 2022; Figueiredo et al., 2022). 2017), and this close relationship with the nature of the substrate
The presence of microbial mats in the sediments where the foot- allows the paleoenvironmental interpretations (Lockley, 1986;
prints are produced provide an early lithification favoring their Avanzini, 1998; Gatesy et al., 1999; Mil n and
an et al., 2004; Mila
preservation (Lockley, 1991; Avanzini et al., 1997; Paik et al., 2001; Bromley, 2006; Falkingham et al., 2010, 2011; Platt et al., 2012;
Marty, 2005; Conti et al., 2005; Phillips et al., 2007; Marty et al., Díaz-Martínez et al., 2018; Menezes et al., 2019). There are some
2009; Noffke, 2010; Carvalho et al., 2013a,b; Cariou et al., 2014; requirements concerning the substrate cohesiveness, plasticity,
Noffke et al., 2019) as they can stabilize those surfaces by precipi- grain size, texture and water content to allow the footprint to
tation of calcium carbonate (Chafetz and Buczynski, 1992; Dupraz register (Lockley et al., 1989; Avanzini et al., 2000; Leonardi and
et al., 2004; Dupraz and Visscher, 2005; Noffke, 2010) and/or Mietto, 2000; Dalla Vecchia, 2008; Getty et al., 2017; Melchor
covering the tracks and protecting them from erosion with an et al., 2019). Therefore, their preservation is controlled by the
organic felt (Avanzini, 1998; Conti et al., 2005; Marty, 2005). They sedimentation events, which are minimal during long-lasting pe-
also enhance the preservation potential of primary structures like riods of exposure, but favored by rapid and episodic sedimentation
ripple marks and mud cracks (Dai et al., 2015), as those observed in (Razzolini et al., 2014).
the Sítio Pereiros ichnosite. The occurrence of microbial structures The footprints of Sítio Pereiros ichnosite are all preserved as
in the Sousa basin was described by Silva Filho (2009). The large concave epireliefs, sometimes with claws and digital pads (Fig. 4C).
number of dinosaur tracks and associated sedimentary structures They are bordered by mud cracks and some of them show a
10
displacement rim (Fig. 4D) surrounding the rear outline and the be observed in the footprints from trackway SJSP 1 (footprints SJSP
digits (SJSP 1 trackway). The mud cracks have their origin related to 1e5, SJSP 1e6, SJSP 1e7 and SJSP 1e8).
the outline of the footprint (Fig. 4E) or as an extension of the digits. The ostracods, spinicaudatans (conchostracans), skeletal re-
In this way, it is seen that the general shape and anatomical details mains and scales of fishes in the successive layers where the foot-
of the foot, especially the digits, after being impressed in the mud, prints occurr are strong evidence of ponds in this environmental
somehow, in the phase of mud dehydration and desiccation, con- setting. The presence of spinicaudatans indicates that the ponds
trol the opening and the direction of the mud cracks. were mostly small and temporary, hot and shallow, and their water
This was not a subaqueous environment, as in this context there chemistry had an alkaline character (pH between 7 and 9) as
is a decrease in the morphological clarity of the footprints, with observed in the Sousa basin (Carvalho and Carvalho, 1990;
poorly defined claws or pad marks (Lockley and Conrad, 1991; Carvalho, 1993). The sedimentary succession of fine sandstones,
Prince and Lockley, 1991; Paik et al., 2001; Belvedere et al., 2010; siltstones, mudstones and shales with mud cracks and ripple
~ uela Sua
Pin rez, 2015; Sciscio et al., 2016; Carvalho et al., 2021, marks, in a cyclic deposition, interpreted as a deposition in a
2022). The footprints of SJSP 1 trackway, with well-defined mor- floodplain, with temporary aerial exposure of the superficial sedi-
phologies are interpreted to have been produced in sediments with ments, in which footprints could be impressed.
high plasticity and low water content, probably in a subaerial Floodplains of meandering fluvial rivers presents the finest
setting of floodplains and lake margins. This setting is easily grain size of all alluvial sediments where generally perennial and
recognized in the Sousa basin where there are the association of temporary lakes may develop in this area. Mud cracks, bio-
footprints with raindrop impressions and mud cracks, the latter of turbation, and other surface features are widespread because of
which sometimes originated at the margins of the footprints or as repeated exposure (Reineck and Singh, 1986). This is a favorable
distal extensions from the digits (Leonardi and Carvalho, 2021). The environment for footprint preservation as floodplains cover a wide
footprints of the other two trackways (SJSP 2 and SJSP 3) are distribution area, with repetitive track-bearing strata. Thus, fluvial
shallow concave epireliefs associated with mud cracks that and lacustrine environments are favorable for recording and pre-
bordered them and frequently cut across the footprints (Fig. 5B). In serving dinosaur footprints, especially due to cyclic deposition of
this case they should be related to a later time of the substrate microclastic sediments in episodic flooding events and shoreline
deformation, when the muddy sediment was drier. variations (Reineck and Singh, 1986; Lockley and Conrad, 1991) as
The relationship of mud cracks and footprints shows distinct also interpreted for the Sousa Formation in the Sousa basin
patterns of cracking related to the moment when the processes of (Leonardi, 1979b, 1991, 1994; Carvalho, 1996, 2000a,b, 2004;
dehydration of the sediments occurred. The impression of a foot in Carvalho and Leonardi, 1992; Santos et al., 2016; Calvo and Rivera,
a sediment prior to the dehydration processes induces a preferen- 2018).
tial cracking in the surrounding margins of the footprint and from The identification of the theropods responsible for the foot-
the extremities of the digits (Carvalho and Leonardi, 2021). In SJSP 1 prints from Sítio Pereiros ichnosite is difficult as there are few data
footprints, there is the indication that the track originated when on South American theropod skeletons, and especially skeletons of
there was a high water content in the substrate. Other examples of their feet, from Early Cretaceous. Abelisauroids, carchar-
this pattern are found in Bemaraha Formation (Middle Jurassic, odontosaurids and spinosaurids may have been theropod track-
Madagascar), but these rarely affect the footprint itself makers of the Rio do Peixe basins. Some of the large theropod
(Wagensommer et al., 2012, 2022). Therefore, the footprints of SJSP tracks are possibly attributed to large and very large (8e17 m long;
2 and SJSP 3 trackways show a distinct pattern of mudcracks that Hendrickx et al., 2015) Spinosauridae, although they are only
frequently cross the footprints. We therefore conclude that the recognized from the fossil record in the Aptian deposits of the
footprints of Sítio Pereiros ichnosite probably were produced in a Araripe basin (Martill et al., 1996; Kellner and Campos, 2000; Sues
subaerial setting in two distinct events. The footprints with well- et al., 2002) and Cenomanian of Sa ~o Luís basin (Medeiros, 2006).
defined morphologies with impressions of claws, and sole pads The Carcharodontosauridae, medium-size to very large allosauroid
(trackway SJSP 1) and surrounded by displacement rims are theropods (6e14 m long; Hendrickx et al., 2015) are known from
considered to have been produced in sediments with high plasticity South America after the Valanginian (Novas et al., 2005; Coria et al.,
and low water content, probably in a subaerial setting of flood- 2020). We remember here also the great Meraxes, which however is
plains. The mud cracks mainly have their origin related to the much later (Late Cretaceous; Canale et al., 2022). Nevertheless, it is
contour of the track or as an extension of the digits. This pattern is more plausible that the large theropod tracks from the Rio do Peixe
also observed in other footprints from the Sousa basin (Carvalho, basins were produced by abelisauroids, as they are already present
2000a,b; Carvalho and Leonardi, 2021). When the footprints are in the BerriasianeValanginian of the Bajada Colorada Formation,
produced in a more cohesive substrate, as with the ichnocoenosis Argentina (Canale et al., 2016). The smaller footprints (SJSP 2
from the Late Jurassic Iouaride ne Formation, Morocco (Belvedere, trackway), with digit III substantially longer than digits II and IV
2008; Belvedere et al., 2010; Boutakiout et al., 2019), in which should also be related to abelisauroids such as the Noasauridae
there are microbial mat laminations, there is a high potential to from the BarremianeAptian of La Amarga Formation, Argentina
preserve footprints produced in a firm ground. (Bonaparte, 1996). Other possible small trackmakers could be
The brecciated bottom of footprints and the separation of mud similar to Santanaraptor Kellner, 1999 and Mirischia Naish, Martill
polygons are common aspects within the footprints of SJSP 2 and and Frey, 2004 (Aptian, Araripe basin), a velocisaurid and a basal
SJSP 3 trackways from Triunfo basin, which indicate that the di- coelurosaur respectively (Bonaparte, 2007).
nosaurs stepped on a harder, drier substrate than in the case of It would be interesting to add some predictions or projections
trackway SJSP 1. The absence of displacement rim around these about the size of the theropod makers of these tracks from Pereiros
footprints are also indicative of a compact and firm substrate. site. It should be noted, first, that the estimation of the size of a
The displacement rims of footprints are related to the substrate dinosaur, even starting from the finding of bones, is usually much
cohesiveness. The mud bulges developed when an animal discussed and questionable, when the complete or almost complete
impressed its feet into a surface of very plastic and/or waterlogged skeleton is not found. Even more has been discussed the possibility
mud. In theropods, there is also compressed sediment between of making projections relating to the weight of these animals.
toes, in the shape of a wedge, especially in footprints made by Even more cautious one has to be when you want to attempt
running theropods (Carvalho and Leonardi, 2021). This aspect can projections of this type starting from footprints and trackways. For
11
I.S. Carvalho and G. Leonardi
12
Cretaceous Research 144 (2023) 105446

Fig. 7. Reconstruction of the environmental scenary with the theropod tracks of Sítio Pereiros, Sousa Formation, Triunfo basin (art by Deverson Silva). The three theropods recorded, based on the three trackways, are represented here
together, although in reality they passed into the area at different times, with no evidence of social behavior.
an accurate study of such awkward problems see Thulborn (1990, especially where sediments are most likely to be deposited, i. e. in
pp. 234e256) and Xing et al. (2022). low lands. If the dry season in the Early Cretaceous in the basins of
For simplicity, given the small amount of ichnological material the Rio do Peixe (especially where the sediments of the Sousa
on this site, we will make a brief reference to this subject. The most Formation were deposited) had been characterized by rare pools of
likely hypothesis is the two SJSP 1 and 3 trackways are abelisauroid, water, these would have been places where animals tended to
as mentioned above, and that they can be attributed to two spec- gather. This would have been especially true for the banks of
imens quite similar to each other in size (but not necessarily to the temporary lakes during the lowering of the water level. Here the
same form or kind). In this case, they may have been 4e4.5 m in meat eater theropods would have probably waited for the big
length, and be high about 1.5e1.8 m on the back, just above the hip herbivores, or they would have frequented one after the other the
joint. The front part of the body and especially the head were rare points of water, in the necessity of hunting and feeding. In
normally held higher, but in different ways, according to the gaits short, meat eaters were more active than herbivores, which almost
and according to the occasions, as it happens to all bipedal animals. stopped, grazing the vegetation. Carnivores walked much more, in
However, these last two measurements cannot be calculated from a search of prey, so they may have made many more footprints than
bipedal trackway. The measures of length mentioned here are large sauropods and ornithopods.
meant as approximate total length, from snout to the tip of the tail; With 40 ichnosites in the basins of the Rio do Peixe, the situation
although for the truth the dinosaur skeletons so completely pre- is as follows. Against what one would expect in theory, with respect
served, for a comparison, are very scarce. The more, they are non- to values related to different groups of dinosaur tracks, ichnosites in
existent, as for abelisauroids. This suggested total length was then which meat eaters outnumber plant eaters (32 sites out of 40; that
based on comparisons made on reconstructions, with the sole is 80% of all sites) are more abundant than those in which the
purpose of having an approximate idea of the linear size of the opposite occurs, (5 sites, 12.5%). There are also two sites with
trackmakers. insufficient data or with tracks of dinosaurs that cannot be classi-
It must be said, in favor of the total lengths suggested above, fied with safety (5%); and one site where the parity between the-
although they have not been calculated following the method ropods and herbivores is reached (2.5%). Besides, ichnosites in
proposed by Ellemberger (1972, p. 60), that they correspond rather which there are only tracks of theropods are rather numerous in
well to the result that would give that method, that is 4.68 m for the basins of the Rio do Peixe. They are 19, out of 40 ichnosites, and
SJSP 1 and, respectively, 4.41 m for SJSP 3. The correspondence may correspond today to the 47.5% of all 40 ichnosites. One among them
be causal, but it is interesting. If one wanted to give a rough guess is the case of Sítio Pereiros ichnosite that we present herein. This
about the body mass of the two trackmakers mentioned, you could high number of sites with apparent exclusive presence of theropods
suggest a weight between 500 and 1000 kg, more likely for the first is well explained by Pe rez-Lorente (2015, p. 325): “Because
value. Following instead the formula (Hip height 2.63) suggested theropod footprints are the most abundant, so are outcrops with
for theropods by Xing et al. (2021), total body lengths of the SJSP 1 theropod footprints”.
and 3 trackmakers, from snout to the tip of the tail, would be 342 In addition to these 19 ichnosites where only theropods are
cm and 329 cm respectively. The poor quality of the footprints of represented with tracks, there are: two ichnosites with tracks of
SJSP 2 trackway and the fact that the animal was running in an theropods, sauropods, ornithopods and ankylosaurs (four clades,
irregular way, makes it more unlikely the possibility of giving data 5%); five ichnosites with tracks of theropods, sauropods, ornitho-
on its size and body mass. Without a doubt, however, it was a pods (three clades, 12.5%); four ichnosites with tracks of theropods
smaller and lighter animal than the other two. and sauropods (two clades, 10%); six ichnosites with tracks of
theropods and ornithopods (two clades, 15%); one site with only
4.1. Why are theropods the only Sítio Pereiros ichnosite sauropod tracks (2.5%); 1 site with only large unclassifiable herbi-
trackmakers? vore tracks (2.5%); two ichnosites with uncertain or unclassifiable
dinosaur tracks (no clades, 5%). There are also a few sites where rare
As also found in the Sousa basin, the dominance of theropod tracks of animals (reptiles) of the mesofauna are also recorded, they
footprints in the floodplain areas is attributed to an ecological are three (7.5% of all the 40 sites). These localities are Caiçara-Piau,
zonation of the dinosaurian biota and/or a taphonomic artifact Serrote do Pimenta and Tapera. In addition to tetrapod tracks and
(Carvalho, 2000a,b; Leonardi and Carvalho, 2021). The more com- traces, fish-trails (Leonardi and Carvalho, 2021) also occur.
mon habitat of the theropods is the lowland areas, where the main
environments are the low floodplain areas, with more humid and 5. Conclusions
fine-grained sediments, that enhances the chances of their foot-
print preservation (Fig. 7). The ichnofauna from Sítio Pereiros enhances the knowledge of
It is not uncommon that theropod tracks are more abundant the dinosaur fauna from Triunfo basin indicating the presence of
than those of herbivores. Without considering the three most large and small theropods, probably related to abelisauroids. This
recently discovered localities, including the one presented here, the ichnosite registers a new geological context for the occurrence of
percentage rate of theropod tracks in the basins of the Rio do Peixe dinosaur tracks in the Triunfo basin related to a temporary aerial
in general is 79%. In particular, in the beds of the Sousa Formation, exposition of the sediments in a floodplain area. There are mud
the percentage of theropods is even higher: up to 89.16%. These cracks and ripple marks associated with the footprints. Some
data differ from the theoretical model of a large prevalence of plant footprints show a displacement mud rim related to the substrate
eaters and a small number of carnivores. The abundance of cohesiveness when the theropod dinosaurs impressed their feet
theropod footprints against herbivorous traces could be caused by a into a surface of very plastic and humid sediment. The relationship
different level of activity of the respective trackmaker and/or the of mud cracks and footprints show distinct patterns of cracking
grouping of some taxa in particular environments. Furthermore, interpreted as related to the moment when the processes of
the prevalence of theropod tracks is typical of many ichnosites dehydration of the muddy sediments occurred. Two patterns were
worldwide, and especially in South America (Leonardi, 1991, 1994; here recognized. One pattern where the impression of a foot in a
Leonardi and Carvalho, 2021). muddy humid sediment previously to the dehydration processes
Indeed, a disproportion of plant-eaters and meat-eaters can induces a preferential cracking in the surrounding contours of the
sometimes develop in special environments (and facies), and footprints and starting from the extremities of the toes; the other
13
pattern generally crosses and brecciated the bottom of footprints Avanzini, M., Frisia, S., Rinaldo, M., 2000. I Lavini di Marco nel Giurassico Inferiore:
la ricostruzione di un antico ambiente di vita. In: Leonardi, G., Mietto, P. (Eds.),
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tracks permits inferences about trackmaker speed and the type of resti fossili italiani. Accademia Editoriale, Pisa-Roma, pp. 247e272.
gait of the three animals, and of their possible size. Avanzini, M., Pin ~ uela, L., García-Ramos, J.C., 2012. Late Jurassic footprints reveal
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prevalence of theropods throughout the territory of the basins of in the Iouaride ne area (Demnat, Central High-Atlas, Morocco). PhD. thesis.
Dipartimento di Geoscienze, Universita degli Studi di Padova, p. 121.
the Rio do Peixe. This predominance has been confirmed. The result
Belvedere, M., Farlow, J.O., 2016. A numerical scale for quantifying the quality of
was also a review of the 40 ichnosites discovered so far in these preservation of vertebrate tracks. In: Falkingham, P.L., Marty, D., Richter, A.
basins, and an overview of the number and type of clades present in (Eds.), Dinosaur Tracks: The Next Steps. Indiana University Press, Bloomington,
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from the siliciclastic Iouaride ne Formation (Central High Atlas, Morocco).
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This study was supported by Fundaça ~o Carlos Chagas Filho de top of Mt. Pelmo: new data for Early Jurassic palaeogeography of the Dolomites
(NE Italy). Bollettino della Societa Paleontologica Italiana 56 (2). https://doi.org/
Amparo a Pesquisa do Estado do Rio de Janeiro (Proc. E-26/200.828/
10.4435/BSPI.2017.10, 2017, i-viii.
2021, Brazil) and by Conselho Nacional de Desenvolvimento Cien- Belvedere, M., Budka, M., Wiseman, A.L.A., Bennett, M.R., 2022. When is enough,
tífico e Tecnologico (CNPq 303596/2016e3, Brazil), as well as many enough? Questions of sampling in vertebrate ichnology. Palaeontology 64 (5),
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Ismar de Souza Carvalho, conducted the fieldwork, conceived Giant theropod footprints in the Upper Jurassic of Morocco. Aït Mazigh site
(Central Atlas). Geogaceta 66, 83e86.
and designed the research; performed the analysis; interpreted
Calvo, J.O., Rivera, C., 2018. Huellas de dinosaurios en la costa oeste del embalse
obtained data; wrote the manuscript. Ezequiel Ramos Mexía y alrededores (Cret acico Superior, Provincia de Neuque n,
Giuseppe Leonardi, conceived and designed the research; per- República Argentina). Boletín de la Sociedad Geolo gica Mexicana 70 (2). https://
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Canale, J.I., Apesteguía, S., Gallina, P.A., Gianechini, F.A., Haluza, A., 2016. The oldest
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Acknowledgments j.cretres.2016.11.010.
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We thank the Brazilian National Mining Agency (Age ^ncia Shinya, A., Haluza, A., Gianechini, F.A., Makovicky, P.J., 2022. New giant
carnivorous dinosaur reveals convergent evolutionary trends in theropod arm
Nacional de Mineraça ~o), through the geologist T. Yamamoto, for reduction. Current Biology 32 (14), 3195e3202.e5. https://doi.org/10.1016/
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Janeiro, Brazil (unpublished).
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16
Roesner, E.H., Lana, C.C., He risse

, A.L., Melo, J.H.G., 2011. Bacia do Rio do Peixe (PB): possible to distinctly observe the impression of claws. The interdigital angles IIeIII
novos resultados biocronoestratigra ficos e paleoambientais. In: Carvalho, I.S., range from 23 to 35 and angles IIIeIV from 27 to 40 . Footprints SJSP 1e1, SJSP
Srivastava, N.K., Strohschoen Jr., O., Lana, C.C. (Eds.), Paleontologia: Cena rios de 1e2, and SJSP 1e3 (Figs. 4A, 4B and 4C) present an angular rear outline, while SJSP
Vida. Editora Intercie ^ncia, Rio de Janeiro, pp. 135e141. 1e4, SJSP 1e5, SJSP 1e6 and SJSP 1e7 (Figs. 4D, 4E, 4F, 4G) show a rounded rear
Romano, M., Citton, P., 2020. Tetrapod ichnology in Italy: the state of the art. Guest outline. Claw impressions are found in SJSP 1e1 (digits II and III), SJSP 1e2 (digit IV),
editorial. Journal of Mediterranean Earth Sciences 12 (2020), 1e12. https:// SJSP 1e3 (digits II, III and IV), SJSP 1e4 (digits II, III and IV), SJSP 1e5 (digits II, III and
doi.org/10.3304/jmes.2020.17068. IV), SJSP 1e6 (digits II, III and IV), SJSP 1e7 (digits II, III and IV), SJSP 1e8 (digits III and
Santos, W.F.S., Carvalho, I.S., Brilha, J.B., Leonardi, G., 2016. Inventory and Assess- IV, Fig. 4H). There usually are not digital or “plantar” pads, although there are
ment of Palaeontological Sites in the Sousa Basin (Paraíba, Brazil): Preliminary probably “plantar” pads in SJSP 1e3 and SJSP 1e4. The third toe consists practically
Study to Evaluate the Potential of the Area to Become a Geopark. Geoheritage 8 of only one large rounded digital pad, and of a narrow peduncle that unites it to the
(4), 315e332. https://doi.org/10.1007/s12371-015-0165-9. other parts of the foot. In some cases, for example, in SJSP 1e3 and 6, this peduncle
Sciscio, L., Bordy, E.M., Reid, M., Abrahams, M., 2016. Sedimentology and ichnology can be considered an additional digital pad. The index SL/FL (Stride length/footprint
of the Mafube dinosaur tracksite (Lower Jurassic, eastern Free State, South Af- length) is 9.12.
rica): a report on footprint preservation and palaeoenvironment. PeerJ 4, e2285. The first two footprints (SJSP 1e1 and SJSP 1e2) are also preserved as concave
https://doi.org/10.7717/peerj.2285. epirelief, but they are filled in with the similar sediment of the overlying contiguous
rdoba, V.C., Caldas, L.H.O., 2014. Proposta de novas unidades litoes-
Silva, J.G.F., Co layer, so they seem to be convex in epirelief; but this is just a result of the infilling (it
tratigra ficas para o Devoniano da Bacia do Rio do Peixe, Nordeste do Brasil. is a case of pseudoreverse relief). The others are concave epirelief with higher rims of
Brazilian Journal of Geology 44 (4), 561e578. https://doi.org/10.5327/ displaced sediments.
Z23174889201400040004. These footprints, unlike those of the other two trackways, have a distinctive
Silva Filho, R.P., 2009. Ana lise de facies da Formaça ~o Sousa (Cretaceo Inferior), bacia appearance that Li et al. (2011) call “tulip-shaped”, or even “lily-shaped”. We accept
de Sousa (PB), e seu contexto em um sistema petrolífero. In: Monografia de these truly expressive terms, but we could also characterize them with the name of
Conclus~ ao de Curso. Universidade Federal do Rio de Janeiro. Departamento de “flame-shaped” tracks, because they recall the shape of a little campfire with three
Geologia (Unpublished). flames, of which the central (III) is much larger than the other two. This 3rd toe
Siqueira, L.M.P., Polck, M.A.R., Hauch, A.C.G., Silva, C.A., et al., 2011. Sítios Paleon- impression, in fact, is rather narrows to a pointed end, with the clear impression of
tolo gicos das Bacias do Rio do Peixe: Georreferenciamento, Diagno stico de the claws in some cases. The footprints that present this type of “flame” bulge, or lily
Vulnerabilidade e Medidas de Proteç~ ao. Anua rio do Instituto de Geocie^ncias 34 petal, are the SJSP 1, and the following numbers 3, 5, 6 and 8. The other two toes, on
(1), 9e21. e-ISSN 1982-3908. the other hand, are much shorter than III, and very similar in length, and less
Sousa, A.J., Carvalho, I.S., Ferreira, E.P., 2018. Western Gondwana non-marine os- expanded in the center. They arrive with the distal end where the bulge of toe III
tracods from Early Cretaceous lowlatitude ephemeral lake, Northeastern Brazil. begins. Sometimes these outer toes are more or less straight; sometimes instead
Journal of South American Earth Sciences 86 (2018), 23e37. https://doi.org/ they bend, again in the form of lily petals or flames, respectively externally or
10.1016/j.jsames.2018.06.001. medially. This happens especially in the footprints of serial number 4, 6 and 7.
Sousa, A.J., Carvalho, I.S., Ferreira, E.P., 2019. Non-marine ostracod biostratigraphy of The hypices seem to be generally rounded and not angular, probably because toe
cretaceous rift lake deposits (Sousa Basin, Brazil): Paleogeographical implica- III, in its proximal free part, just above the junction line of the two corresponding
tions and correlation with Gondwanic basins. Journal of South American Earth hypices, it is very narrow. This situation probably depends on the fact that this toe
Sciences 2019, 102345. https://doi.org/10.1016/j.jsames.2019.102345. had to be curved on the sagittal plane of the foot, and therefore almost entirely
Sues, H.D., Frey, E., Martill, D.M., Scott, D.M., 2002. Irritator challengeri, a Spinosaurid raised from the ground.
(Dinosauria: Theropoda) from the Lower Cretaceous of Brazil. Journal of The impressions of the claws, when clearly visible in the III toe, as they are
Vertebrate Paleontology 22 (3), 535e547. https://doi.org/10.1671/0272- sometimes in the footprints (SJSP 1, 3e8), seem to be usually directed outwards
4634(2002)022[0535:ICASDT]2.0.CO;2. (laterally). Especially noticeable, and separated from the toe, is the impression of the
Thulborn, R.A., 1989. The Gaits of dinosaurs. In: Gillette, D.D., Lockley, M.G. (Eds.), claw of toe III of the left footprint 8.
Dinosaur Tracks and Traces. Cambridge University Press, New York, pp. 39e50. This type of footprint with a swollen toe III and the rounded hypices is not
Thulborn, R.A., 1990. Dinosaurs Tracks. Chapman & Hall, London, p. 410pp. uncommon in the Basin of Sousa, the most generous of tracks. See for example the
Valença, L.M.M., Neumann, V.H., Mabesoone, J.M., 2003. An overview on Callovian- followings: in the Sousa Formation, the footprints or trackways Engenho Novo
Cenomanian intracratonic basins of Northeast Brazil: onshore stratigraphic (ANEN) 16, Matadouro (SOMA) 2, Piau-Caiçara (SOCA) 5bis, 54bis, 75, 91, 92, 97,
record of the opening of the Southern Atlantic. Geolo gica Acta 1 (3), 261e275. 13498, 13499, 13693, 150, 151, 152, Poço do Motor (SOPM) 8, Sítio Saguim (SOSA) 4;
https://doi.org/10.1244/105.000001614. in the Rio Piranhas Formation, Curral Velho (SOCV) 1 and 4. There are no examples of
Viana, M.S.S., Lima Filho, M.F., Carvalho, I.S., 1993. Borborema Megatracksite: uma this style of footprints in the Antenor Navarro Formation (Leonardi and Carvalho,
base para correlaç~ ao dos “arenitos inferiores” das bacias intracontinentais do 2021). It is not easy to decide whether this morphotype with swollen digit III re-
Nordeste do Brasil. In: Simpo sio de Geologia do Nordeste, Sociedade Brasileira flects the anatomy of their makers, or whether there is also an extra-morphological
de Geologia/Núcleo Nordeste, Boletim, vol. 13, pp. 23e25. aspect, deformation of the footprint depending on the type of sediment and/or the
Wagensommer, A., Latiano, M., Leroux, G., Cassano, G., Porchetti, S.d'O., 2012. New behavior of the animal. This could reflect, for example, movements of the foot in the
Dinosaur Tracksites from the Middle Jurassic of Madagascar: Ichnotaxonomical, stages of initially placing the foot on the ground, any movement with the foot during
Behavioural and Palaeoenvironmental Implications. Palaeontology 55 (1), the stance phase, or liftoff of the foot. One would have to evaluate this on the type of
109e126. https://doi.org/10.1111/j.1475-4983.2011.01121.x. sediment and/or the behavior of the animal. This could be, for example, the way of
Wagensommer, A., Dolch, R., Ratolojanahary, T., Donato, S., Porchetti, S.d'O., 2022. moving the foot in the stages of placing the foot on the ground, any intermediate
Defining the Bemaraha megatracksite: an update on dinosaur ichnology in movement with the foot on the ground, and movement of lifting the foot. One have
Madagascar. In: Co nsole-Gonella, C., de Valais, S., Díaz-Martínez, I., Citton, P., to see it case by case. In the case of the trackway SJSP 1, it seems that the shape of the
Verde, M., McIlroy, D. (Eds.), Ichnology in Shallow-marine and Transitional footprint depends mainly on the anatomy of the foot and particularly of the III digit,
Environments, Geological Society, London, Special Publications, vol. 522. given the constancy of the shape in the repetition of this pattern.
https://doi.org/10.1144/SP522-2021-86.
Xing, L.-D., Lockley, M.G., Klein, H., Zhang, L.-J., Romilio, A., Persons IV, W.S.,
Peng, G.-Z., Ye, Y., Wang, M.-Y., 2021. The new ichnotaxon Eubrontes nobitai SJSP 2 (Fig. 5, Appendix B)
ichnosp. nov. and other saurischian tracks from the Lower Cretaceous of
Sichuan Province and a review of Chinese Eubrontes-type tracks. Journal of The length of steps (oblique paces) varies from 1.10 m to 1.50 m, with an average
Palaeogeography 10 (17). https://doi.org/10.1186/s42501-021-00096-y. value of 1.30 m. The stride varies from 2.64 m to 3.25 m, with an average value of
Xing, L.-D., Lockley, M.G., Klein, H., Wang, T., 2022. Sauropod tracks from the Middle 2.91 m. The step angle varies from 162 to 179 , with an average value of 170 200 .
Jurassic Chuanjie Formation of Yunnan Province and the pre-Cretaceous sau- The trackmaker had to be a rather light and lean animal, and/or the soil had to be
ropodomorph trackways from China. Palaeoworld 30 (2021), 495e502. https:// rather dry and not very plastic; with the result that the footprints are shallow and
doi.org/10.1016/j.palwor.2020.07.007. not very detailed.
The footprints of the SJSP 2 trail range from 16.5 to 18.0 cm in length with an
average value of about 17.25 cm; and 12.0e13.0 cm in width, with an average value
Appendix A. Description of the footprints and trackways of about 12.2 cm. They are mesaxonic, tridactyl, with possible claw impression.
There are no evident digital or plantar pads. Interdigital angles IIeIII range from 30
The description of the footprints and tracks follows the methodology of Leonardi to 36 and angles IIIeIV from 30 to 40 . The footprints SJSP 2e1 (Fig. 5B), SJSP 2e4
(1979a,b, 1987) concerning the definitions and measurement of the footprints. The (Fig. 5E) and SJSP 2e5 (Fig. 5F) show a rounded rear outline, while SJSP 2e2 (Fig. 5C)
code SJSP is a reference to the county where the ichnosite was found (S~ ~o do Rio
ao Joa and SJSP 2e3 (Fig. 5D) present an angular rear outline. Claw impressions are found
do Peixe e SJ) and to the locality of Sítio Pereiros (SP), in the Paraíba State, Brazil. All in SJSP 2e1 (in digits III and IV), SJSP 2e3 (in digits III and IV), SJSP 2e5 (in digits III
the footprints occur on a same bedding plane. and IV) and SJSP 2e4 (in digits II and III). No digital or plantar pads are seen. All
footprints are preserved as concave epireliefs. As for the manner of measurement of
the “free-digit”, the III toe is without doubt the longest; and between the outer toes,
SJSP 1 (Fig. 4, Appendix B) the IV is longer than the II. The index SL/FL (Stride length/footprint length) is
approximately 16.87.
The footprints of SJSP 1 range from 24.0 to 28.0 cm in length and 16.6 to 28.5 cm The five footprints of this trackway are absolutely distinct and well visible when
in width. They are mesaxonic, tridactyl, with pointed digits. In some of them it is the sunlight comes grazing, very low, in the early morning and at sunset. They are
17
very shallow, and then almost invisible in the hours when the sun is high. The claw marks are not evident, the relative narrowness of the interdigital angles and
displacement rim is also very low and not very distinct. For this reason, among other above all the extreme narrowness of the trackway do not suggest an ornithopodian
things, they were marked with chalk so that they could be better seen, both in their identification. The trackway is straight and regular over the first three footprints,
sequence and in order to better recognize the outline of each individual footprint. after which the animal made a small deviation towards the right, but then resumed
The shape of the footprints of this trackway, although very difficult to interpret, movement in the initial direction. There is some irregularity in the lengths of the
is rather characteristic and uncommon, due to the length of the third toe compared strides and in the step angle. This is always high, particularly in the last measurable
to that of the two outer digits, and also because of the triangular shape. Digit III is step-angle, which is close to 180 (178 ). The type of trackway, with very high step
much longer than the peripheral toes, and does not have the sinuous shape that is angles and poor footprints, makes it difficult to determine which tracks are right and
characteristic of the footprints of small theropods, of the group that are attributable which are left; this in turn makes it even more difficult to interpret the shape and
to the plexus Grallator/Eubrontes. footprint, instead, is more like some footprints of style of the tracks.
the Sousa Formation of the Sousa basin: for example, the footprints and trackways The footprints of SJSP 3 trackway are 23.0e26.0 cm in length with an average
Piau-Caiçara (SOCA) 153, 154, of the level 15 of this locality; and Passagem das value of about 24.5 cm and 15.5e26.0 cm in width, with an average value of
Pedras (SOPP) 8 (Leonardi and Carvalho, 2021). about 21 cm. There are no digital or plantar pads. The interdigital angles IIeIII
range from 30 to 38 and angles IIIeIV from 28 to 34 . The footprints SJSP
3e1 (Fig. 6A), SJSP 3e3 (Fig. 6C), and SJSP 3e5 (Fig. 6E) and SJSP 3e6 (Fig. 6F)
SJSP 3 (Fig. 6, Appendix B) present a rounded rear outline, while SJSP 3e2 (Fig. 6B) and SJSP 3e4 (Fig. 6D)
show an irregular rear outline with a probable projection as a “heel”. However,
All in all, this is a bipedal and tridactyl trackway, clearly visible and recognizable this expansion of the back of the footprint may be, instead, the impression of toe
as such for the periodicity of the sequence of its six footprints, and also because 1st. The digits of all these footprints do not have claw impressions. There are no
overall it is quite straight. However, the footprints are almost all shallow and of poor digital or plantar pads. All footprints are preserved as concave epireliefs. The
or modest quality; only the second and third are clearly tridactylous; perhaps even index SL/FL (stride length/footprint length) is 9.64; it is an index quite similar to
the first was. The fourth footprint looks incomplete, it shows something as a heel on that of the SJSP 1 trackway.
the rear, but the three toes are indistinct. The last two (5 and 6), then, are only
rounded cavities, and perhaps, after the impression of the foot in the mud, have been
washed by a small wave, effacing the anatomical details. With such poor quality, it is
not possible to compare these prints with other tracks of the basins of the Rio do Appendix B. Table with measurements of the trackways and
Peixe, except for the broad identification of this as a theropod trackway. Although
footprints
Trackway-Footprints Length of the trackway (m) External width (m) Inner width (m) Stride (m) Step Angle ( ) Velocity
SJSP 1 7.8 1.00 0.10 e e SL/h ¼ 1.97

8.7 km/h
1e3 e e e 2.40 150 e
3e5 e e e 2.50 162 e
5e7 e e e 2.25 160 e
2e4 e e e 2.25 162 e
4e6 e e e 2.70 154 e
6e8 e e e 2.10 164 e
SJSP 2 5.50 0.45 0.10 e e SL/h ¼ 2.5
12.9 km/h
1e3 e e e 2.85 170 e
3e5 e e e 2.64 162 e
2e4 e e e 3.25 179 e
SJSP 3 4.00 0.82 0.18 e e SL/h ¼ 1.89
9.09 km/h
1e3 e e e 2.65 172 e
3e5 e e e 2.15 173 e
2e4 e e e 2.50 162 e
4e6 e e e 2.15 178 e
Trackway- Morphology Claw impression Digital or Plantar Digital angles IIeIII Preservation -
Footprints pads and IIIeIV
SJSP 1 Mesaxonic, tridactyl, pointed digits, swollen toe III, rounded hypices. e - Mean IIeIII e 28.28 Concave epirelief -
Minimum IIeIII e
23
Maximum IIeIII e
35
Mean IIIeIV e
31.62
Minimum IIIeIV e
23
Maximum IIIeIV e
40
SJSP 1 e 1 Mesaxonic, tridactyl, pointed digits, angular rear outline Digits II and III No digital or 35 and 40 Concave epirelief. e
plantar pads Infilled.
SJSP 1 e 2 Mesaxonic, tridactyl, pointed digits, angled rear outline Digit IV No digital or 30 and 27 Concave epirelief. e
plantar pads Infilled.
SJSP 1 e 3 Mesaxonic, tridactyl, pointed digits, rounded rear outline Digits II, III and IV Probably plantar 23 and 29 Concave epirelief e
pads

SJSP 1e 4 Mesaxonic, tridactyl, pointed digits, rounded rear outline Digits II, III and IV Probably plantar 23 and 27 Concave epirelief e
pads
18
(continued )
Trackway- Morphology Claw impression Digital or Plantar Digital angles IIeIII Preservation -
Footprints pads and IIIeIV
SJSP 1e 5 Mesaxonic, tridactyl, pointed digits, rounded rear outline Digits II, III and IV No digital or 25 and 27 Concave epirelief e
plantar pads
SJSP 1 e 6 Mesaxonic, tridactyl, pointed digits, rounded rear outline Digits II, III and IV No digital or 30 and 40 Concave epirelief e
plantar pads

SJSP 1e 7 Mesaxonic, tridactyl (two preserved digits), pointed digits, rounded Digits II, III and IV No digital or 32 and 40 Concave epirelief e
rear outline plantar pads
SJSP 1e 8 Mesaxonic, tridactyl, pointed digits, irregular rear outline Digits III and IV No digital or - and 23 Concave epirelief e
plantar pads
SJSP 2 e e e Mean IIeIII e 32.75 e e
Minimum IIeIII e
30
Maximum IIeIII e
36
Mean IIIeIV e
33.40
Minimum IIIeIV e
27
Maximum IIIeIV e
40
SJSP 2 e1 Mesaxonic, tridactyl, pointed digits, rounded rear outline Digits III and IV No digital or 36 and 40 Concave epirelief e
plantar pads
SJSP 2 e 2 Mesaxonic, tridactyl, pointed digits, angled rear outline Without claw No digital or - and 27 Concave epirelief e
impressions plantar pads
SJSP 2 e 3 Mesaxonic, tridactyl, pointed digits, angled rear outline Digits III and IV No digital or 32 and 35 Concave epirelief e
plantar pads

SJSP 2 e 4 Mesaxonic, tridactyl, pointed digits, rounded rear outline Digits II and III No digital or 33 and 35 Concave epirelief e
plantar pads
SJSP 2 e 5 Mesaxonic, tridactyl, pointed digits, rounded rear outline Digits III and IV No digital or 30 and 30 Concave epirelief e
plantar pads
SJSP 3 e e e Mean IIeIII e 34 e e
Minimum IIeIII e
30
Maximum IIeIII e
38
Mean IIIeIV e
30.66
Minimum IIIeIV e
28
Maximum IIIeIV e
30
SJSP 3 e 1 Mesaxonic, tridactyl, rounded rear outline Without claw No digital or Unclear digits Concave epirelief e
impression plantar pads
SJSP 3 e 2 Mesaxonic, tridactyl, pointed digits, angled rear outline with a Without claw No digital or 30 and 28 Concave epirelief e
probable heel (digit I) impression plantar pads
SJSP 3 e 3 Mesaxonic, tridactyl, pointed digits, rounded rear outline Without claw No digital or 38 and 34 Concave epirelief e

SJSP 3 e 4 Mesaxonic, probably tridactyl, pointed digits, irregular rear outline, Without claw No digital or e and 30 Concave epirelief e
with a possible heel (digit I) impression plantar pads
SJSP 3 e 5 Mesaxonic, tridactyl, pointed digits II and IV, rounded rear outline Without claw No digital or e Concave epirelief e
SJSP 3 e 6 Mesaxonic, unclear digits, rounded rear outline Without claw No digital or e Concave epirelief e
19

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Cretaceous Research 144 (2023) 105446

Contents lists available at ScienceDirect

The Triunfo basin (also called the Uiraúna-Brejo das Freiras

Cretaceous Research 144 (2023) 105446

Carvalho, I.S., Nobre, P.H., 2001. Um Crocodylomorpha (?Notosuchia) da Bacia de

Roesner, E.H., Lana, C.C., He risse

SJSP 1 7.8 1.00 0.10 e e SL/h ¼ 1.97

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