Journal of South American Earth Sciences 25 (2008) 196–202

www.elsevier.com/locate/jsames

Taphonomy and paleoecology inferences of vertebrate ichnofossils

from Guará Formation (Upper Jurassic), southern Brazil
Paula C. Dentzien-Dias *,1, Cesar L. Schultz 2, Cristina Bertoni-Machado 1
UFRGS, PPGGeo, Av. Bento Goncßalves, 9500, CEP 91509/900, Porto Alegre, RS, Brazil

Received 11 May 2007; accepted 29 August 2007

Abstract

In southern Brazil, the eolian facies of the Guará Formation (Late Jurassic) reveal footprints and trackways of vertebrates (dino-
saurs), as well as burrows made by small vertebrates. All the footprints and trackways are preserved in dunes and sand sheets. The
footprints made in the sand sheets are not well preserved due to intense trampling and can be distinguished only by the deformation
of the sandstone laminations. In some cases it is possible to see this deformation in plan and in section. Tracks of theropods, orni-
thopods and middle-sized sauropods are present. Two footprints preserved in the foreset of a paleodune permitted recognition of
slide structures and identification of the trackmaker, a theropod. Burrows horizontally across the foresets were found at this same
paleodune. Ribbons of massive sandstone – interpreted as the partial filling of the base of the burrows – covered by little blocks of
stratified sandstone – suggest the collapse of the burrow roof inward. There are no body fossils in the Guará Formation, conse-
quently the preservation of these tracks provides unique evidence of widespread dinosaurs activity in southern Brazil near the
end of the Jurassic.
Ó 2007 Elsevier Ltd. All rights reserved.

Keywords: Taphonomy; Vertebrate ichnofossils; Guará Formation; Eolian deposits; Paleoecology

1. Introduction Knowledge on track taphonomy has increased in recent

years (Cohen et al., 1991; Radley et al., 1998; Loope et al.,
Over the last decade, knowledge about the southwest 1998; Fornós et al., 2002; Carvalho, 2004; Milàn et al.,
region of Rio Grande do Sul State, southern Brazil, 2004; Loope, 2006a; Milàn and Bromley, 2006; Milán
(Fig. 1) has increased considerably, especially about the and Loope, 2007; Lockley et al., 2007), but this field still
Guará Formation of the Paraná Basin. Scherer and Lavina remains a part of paleontology needing further develop-
(2005) portrayed four distinct facies associations: simple to ment (Lockley, 1997, 1991; Gillette and Lockley, 1986;
locally composite crescentic eolian dune sets, eolian sand Lockley et al., 1994).
sheets, distal floodflows, and fluvial channels. Dentzien- The main aim of this work is to present taphonomic
Dias et al. (in press) described the fossil record found in data on the footprints and burrows from the Guará For-
the Guará Formation, which includes vertebrate and inver- mation in order to contribute to our understanding of
tebrate ichnofossils and conchostracans. The vertebrate the Late Jurassic paleoecology of southern Brazil.
ichnofossils are composed of sauropod, ornithopod and
theropod tracks and burrows probably made by small 2. Geological setting
mammals (Dentzien-Dias et al., in press).
The Guará Formation has a wide geographical distribu-
*
tion (see Fig. 1), cropping out on the southwestern portion
Corresponding author. Fax: +55 51 3316 6340.
E-mail address: [email protected] (P.C. Dentzien-Dias).
of the Rio Grande do Sul. Lithologically, it is composed of
1
CNPq graduate student. fine to coarse-grained sandstone, and rare mudstones,
2
CNPq research. deposited by fluvial and eolian depositional systems

0895-9811/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jsames.2007.08.008
 P.C. Dentzien-Dias et al. / Journal of South American Earth Sciences 25 (2008) 196–202 197

Fig. 1. Geological map of the Permian and Mesozoic lithostratigraphic units of the Paraná Basin in the Rio Grande do Sul State, Brazil (After Scherer
and Lavina, 2005).

(Scherer et al., 2000). Although highly variable, it has an tion), reflects changes in the substrate wetness associated
average thickness of 200 m and rests unconformably over with either modifications of the rate of water table fluctua-
the fluvial deposits of the Early Triassic Sanga do Cabral tion or dry sand availability (Chakraborty and Chaudhuri,
Formation. Above, the Guará Formation is unconform- 1993). The Guará Formation accumulation was controlled
ably overlain by the eolian deposits of the Early Cretaceous by oscillations between arid and semi-arid conditions
Botucatu Formation (Scherer et al., 2000). The Guará For- (Scherer and Lavina, 2006).
mation is poorly lithified compared with the other Meso- The Guará Formation extends from southwest Rio
zoic units of Rio Grande do Sul and is easily eroded. Grande do Sul to Uruguay, where it corresponds, litho-
The Guará Formation displays a well-defined facies stratigraphically, to the basal member of the Tacuarembó
shift along its outcrop belt. On its northern portion it is Formation (Lavina et al., 1985), which reveals a rich and
characterized by coarse-grained to conglomeratic sand- diversified fossil record, including crocodile, semionotiform
stones with trough and planar cross-bedding, as well as fishes, gastropods, and conchostracans (Mones & Figueira,
low-angle lamination, which are interpreted to represent 1980; Ferrando et al., 1987). Nevertheless, no taxon com-
braided river deposits. Southwards these fluvial facies thin mon to both the Tacuarembó and Guará Formations has
out and interfinger with fine- to medium-grained sand- been previously described.
stones with large-scale cross-stratification and horizontal
lamination, interpreted as eolian dune and sand sheets
deposits, respectively (Scherer and Lavina, 2005). These 3. Data gathering
eolian sediments reveal large cross-beddings with grain-
flow, grain-fall and wind-ripple lamination (Hunter, For each fossiliferous outcrop, a stratigraphic section
1977), interpreted as large eolian dune deposits, and/or was made, in which the layers with ichnofossils were delin-
horizontal wind-ripples strata, composed of fine to eated. Sedimentary facies were described following the
coarse-grained sandstones interpreted as eolian sand sheet model of Reading (1986).
deposits (Scherer and Lavina, 2006). We collected two separated footprints and a pair of
In the eolian sand sheet deposits a vertical transition footprints from the same trackway. They were registered
between wind ripples and adhesion strata (crinkled lamina- in the Laboratory of Paleovertebrates of the Universidade
198 P.C. Dentzien-Dias et al. / Journal of South American Earth Sciences 25 (2008) 196–202

The first one consists of rounded footprints, with med-

ium diameter about 50 cm, without traces of digits
(Fig. 3). Based on the pace angulation pattern and foot-
print shape (Faria dos Santos et al., 1992), they are better
attributed to the pes of a sauropod. No half-moon mark
was associated to them, which would characterize the
manus tracks. Probably this pattern is due to the poor pres-
ervation of the footprints and/or to the overlap of the pes
overstepping the manus footprints, a common phenome-
non in sauropod trackways (Moreno and Benton, 2005).
The morphology and the size of the footprints suggest
Fig. 2. Distribution map of the sauropod footprints from Santana do the presence of a sauropod with a body size similar to an
Livramento (Cerro Paloma) outcrop. The quadrangles have 1 m2.
extant African elephant. These sauropod footprints could
be observed in section, and the undertracks reach around
Federal do Rio Grande do Sul (UFRGS PV 0003 J/K , 45 cm in depth.
UFRGS PV 0004 J/K and UFRGS PV 0005 J/K). Inside some of these footprints, several small (8–10 mm
All the ichnological material was photographed and in diameter), vertical burrows can be observed. One of
measured, using the parameters of Leonardi (1987) for them was excavated to allow its observation in section.
footprints, as well as data regarding trackways. These small burrows begin as vertical tubes which become
To visualize some footprints both in surface and in sec- horizontally enlarged at their bases, forming little cham-
tion, we cut through the sandstone horizontally and/or bers and were attributed as burrows made by insects
vertically, allowing for better observation of some struc- (Dentzien-Dias et al., in press).
tures. Surface mapping of one of the outcrops was under- The second type of footprint found is tridactyl, with
taken to record the distribution and orientation of the marks of sharp claws. These measure between 17–35 cm
footprints (Fig. 2). To assist in the construction of the long and 15–26 cm wide (Fig. 4). This morphology
map, the surface was divided into quadrangles of 1 m2. indicates that these footprints were made by a
To better visualize the tracks, they were marked with theropod.
chalk. Another track is 25 cm long and 23 cm wide. The heel
We measured the length, wide and thickness of the bur- has a ‘‘U” shape and no claw marks are visible. These were
rows. After that, one of them was cut through to observe attributed to a bipedal ornithopod. Other isolated foot-
possible internal structures. prints of the same shape occur, but they are very poorly
preserved, or almost totally eroded, to permit a better
4. Description and interpretation of tracks and burrows classification.
Associated with theropod footprints in a paleodune,
Trackways and isolated footprints were found in five several ribbons of massive sandstone can be observed
different outcrops, all in the eolian facies, and three differ- crossing the sets of the dune. In some portions, these rib-
ent kinds of footprints were found. bons are covered by little blocks of stratified sandstones.
The ribbons of massive sandstones are here interpreted
as partially filled burrows, and the stratified blocks as
the collapsed parts of the burrow roofs. The burrows
typically are straight to slightly curved in plan view
and horizontal to gently inclined in lateral view. These
structures tend to be rectilinear, but some of them
describe curves and bifurcate (Fig. 5). Burrow diameters
range from 15 cm to 20 cm and are elliptical in cross sec-
tion, with shorter axe about 8 cm and lengths from
0.40 m to 2.80 m. The external surface has been weath-
ered, therefore no well-defined burrow linings, or scratch
marks, were observed.
The size and shape of these burrows would be
compatible with excavations done by small reptiles or
mammals (Miller et al., 2001; Loope, 2006b) and can
be illustrated by the extant Ctenomys sp. (the ‘‘tuco–
tuco”), that build extensive tunnels in the coastal eolian
Fig. 3. Sauropod footprint with diameter of 50 cm, in detail the dunes at the South of Brazil (Dentzien-Dias et al., in
deformational features in the stratification that suggest that we are seeing press). However, no tracks are directly associated with
actually the undertracks. the burrows.
 P.C. Dentzien-Dias et al. / Journal of South American Earth Sciences 25 (2008) 196–202 199

Fig. 4. Theropod footprint with 17 cm long and 15 cm wide.

Fig. 5. (a) Bifurcated burrow with 1.50 m of length and 20 cm of width, the arrow shows a stratified block interpreted as the collapsed parts of the burrow
roofs. (b) The longest burrow found in the Guará Formation, with 2.80 m of length and 20 cm of width.

5. Taphonomy by the superimposition of footprints and tracks, resulting

in chaotic perturbation (dinoturbation). The footprints
The footprint preservation occurred in two contexts of were produced and, then eroded and, covered by sand, so
an eolian environment: eolian sand sheets and paleodunes. that no lithological discontinuities (which could have pre-
In the first case the tracks are preserved in fine-grained served them in relief and with more details) are present.
sandstones with both horizontal and wavy stratified depos- So, they are completely flattened on the surface, and their
its. In the paleodunes, footprints occur cutting the foresets. presence is evidenced just by deformational features in
the stratification (see Fig. 3) suggesting that we are seeing
5.1. Tracks in the eolian sand sheets actually the undertracks.
In section, it is possible, in some cases, to see the foot-
All types of footprints above cited (made by sauropods, prints and respective undertracks; the deformation of the
ornithopods and theropods) occur in the eolian sand sheets stratification inside the footprints is clearly asymmetric
facies. They occur at different levels inside the same pack- (Fig. 6). In these cases, a deeper portion is always present
age and it is common for strata to be strongly trampled in each footprint, which resulted from the pressure per-
200 P.C. Dentzien-Dias et al. / Journal of South American Earth Sciences 25 (2008) 196–202

and the pin stripes were down-folded as the animal feet

penetrated the sand, showing that the substrate was rela-
tively firmer (Fig. 7). On the bedding surface, it is possible
to identify the trackmaker and see, again, slide structures
that show that the theropod was climbing the dune obli-
quely. Some details of the footprint were preserved making
it possible to identify which kind of animal made the foot-
print. This was possible because the sand should be
moisten, otherwise it would not be preserved, since they
are easily destroyed by erosion and do not allow preserving
details of the tracks (Fornós et al., 2002). The quality of the
details preserved in the Guará Formation paleodunes
makes us believe that sand was moist, in spite of Loope
(2006a), who says that moisture is not needed to preserve
Fig. 6. Sauropod footprint in section, the undertracks reach around 45 cm tracks.
in depth; the deformation of the stratification inside the footprints is The good preservation of these footprints indicates that
clearly asymmetric. they escaped erosion or further trampling, by rapid burial
(Radley et al., 1998). This rapid burial must be the result
formed by the anterior portion of the foot during the step. of low energy process that, in dunes, is characterized by
This spatial orientation indicates to which direction the the grain-fall. However only in small dunes grain-fall strata
animal was moving. reach the base of the slip face (Hunter, 1977). Grain-flows
To preserve a footprint, even a tiny amount of water erode only the upper parts of tracks (Loope, 2006a; Loope
should be present in the substrate (<1%) in order to pro- et al., 1998), but they are erosive near their point of origin,
mote a significant increase of sediment cohesion, hence becoming depositional down-dip (Hunter, 1977). There-
allowing footprint preservation with well-developed verti- fore, as Guará Formation dunes were large, and only the
cal features (Loope, 1986, 2006a). Therefore, preservation lower part of those dunes was preserved, these tracks were
of this facies occurs because of a certain degree of wetness buried by a grain-flow.
in the substrate, evidenced by the adhesion strata. According to Loope (2006a) the deepest-penetrating
animal tracks have the highest preservation potential and
5.2. Tracks and burrows in the paleodunes on an active dune, the softest, most easily deformed sub-
strate underlies the slip face. Because the slip face lies
The only kind of tracks found in the cross-strata of the within the zone of flow separation, tracks made on grain-
paleodunes was made by a theropod. In section, slide struc- flows are better protected from wind erosion than those
tures formed during this climbing can be clearly visualized made on any other dune surface.

Fig. 7. Theropod footprint in section (a) and on surface (b). Slide structures formed during this climbing can be clearly visualized and the pin stripes were
down-folded as the animal feet penetrated the sand, showing that the substrate was relatively firmer (Scales: Brush 15 cm; Ruler 5 cm).
 P.C. Dentzien-Dias et al. / Journal of South American Earth Sciences 25 (2008) 196–202 201

There is no lithologic difference between the infilling of moist, otherwise it would not be preserved, since they
the burrows and the surrounding matrix, the absence of are easily destroyed by erosion and do not allow pre-
stratification. Burrows filled with the same sediment as that serving details of the tracks (Fornós et al., 2002).
in which they were excavated, easily are overlooked (Smith, – The good preservation of some footprints indicates that
1987). However, this stratification difference gave the bur- they escaped erosion or further trampling, by rapid
rows some relief, allowing us to identify them easily. burial.
Unfortunately no footprints that could be attributed to – The strongly trampled nature of some eolian sand sheets
the constructer of the burrows have been found to this could indicate that large dinosaurs were passing by
moment. Probably these footprints did not have the same southern Brazil, searching for water, considering that
luck of those made by theropods, maybe because they were there is no evidence that the environment of the Guará
too shallow and even low energy processes could have Formation could support a large population, as corrob-
destroyed them. orated by the dry eolian deposits.
– The burrows are the only evidence of vertebrate burrows
6. Paleoecology inferences from the Mesozoic of Rio Grande do Sul and prove that
some small animals could live in the semi-arid environ-
Using the ichnological and taphonomic data some ment of the Guará Formation.
paleoecology inferences can be made, such as the speed, – Body fossils are unknown to the Guará Formation, con-
weight, and behavior of the organism (Lockley, 1997), cor- sequently the chance preservation of these tracks pro-
relating the fossil record with the sedimentological data. vides unique evidence of widespread dinosaurs activity
Here, we speculate about the animals and the behavior that in southern Brazil during the end of Jurassic.
lead them to make some of the ichnofossils described
above.
The burrows were found in the same outcrop that the Acknowledgements
theropod footprints occur, in a paleodune. It is possible
that theropods were in that dune to hunt little animals P.C.D.-D. and C.B.-M. thank CNPq for graduate schol-
(mammals?) that used to live in that kind of burrows. arship. We thank Juan Cisneros for revision of an earlier
It is known that in arid and semi-arid environments, version of this manuscript. We are also grateful for the con-
trampled layers occur near lakes and other sources of water structive reviews by Dr. David Loope and an anonymous
(Sung Paik et al., 2001) and the rate of trampling is depen- referee.
dent on the number of large vertebrates in the sedimentary
basin and their behavior (Laporte and Behrensmeyer, References
1980). Therefore, the strongly trampling that occurs in
some eolian sand sheets, could indicate that the dinosaurs, Carvalho, I.S., 2004. Dinosaur footprints from northeastern brazil:
specially the sauropods, in southern Brazil, were passing taphonomy and environmental setting. Ichnos 11, 311–321.
Chakraborty, T., Chaudhuri, A.K., 1993. Fluvial–Eolian interactions in a
by, perhaps searching for water, considering that there is Proterozoic alluvial plain: example from the Mancheral Quartzite,
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