Bertrand-Sarfati, 1995, Geology, First Ediacaran Fauna Found in Western Africa and Evidence For An Early Cambrian Glaciation
Bertrand-Sarfati, 1995, Geology, First Ediacaran Fauna Found in Western Africa and Evidence For An Early Cambrian Glaciation
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1
Universidade Federal de Pernambuco/UFPE, Laboratório de Geologia Sedimentar e Ambiental/LAGESE,
Av. Acadêmico Hélio Ramos, s/n, Cidade Universitária, 59740-530 Recife, PE, Brasil
2
Universidade Estadual Vale do Acaraú, Museu Dom José, Laboratório de Paleontologia,
Av. Dom José Tumpinambá, 878, Centro, 62010-290 Sobral, CE, Brasil
Manuscript received on May 24, 2013; accepted for publication on September 9, 2013
ABSTRACT
This study reports the first known occurrence of the Ediacaran fauna in northeastern Brazil (at Pacujá
Municipality, northwestern state of Ceará) and presents preliminary interpretations of its significance.
Regional correlation indicates that the fossils originated in the Jaibaras Basin and that they may represent a
new geological system. The depositional environment can be attributed to a fluviomarine system. Nine Ediacaran
species can be identified, including members of pandemic groups (e.g., Charniodiscus arboreus Glaessner, 1959;
?Charniodiscus concentricus Ford, 1958; Cyclomedusa davidi Sprigg, 1947; Ediacaria flindersi Sprigg, 1947;
and Medusinites asteroides Sprigg, 1949) and endemic groups (e.g., Kimberella quadrata Glaessner & Wade,
1966; Palaeophragmodictya reticulata Gehling & Rigby, 1996; Parvancorina minchami Glaessner, 1958; and
Pectinifrons abyssalis Bamforth, Narbonne, Anderson, 2008). Three ichnogenera are also present: Arenicolites
Salter, 1857; Palaeophycus Hall, 1987; and Planolites Nicholson, 1873. The relative age of the deposits is between
?Ediacaran and Cambrian, and the fauna resembles the White Sea Assemblage. The bioturbation presents typical
unbranched Ediacaran ichnogenera with little depth in the substrate. This previously unknown occurrence of the
Ediacaran fauna reinforces the importance of the state of Ceará to Brazilian and global palaeontology.
Key words: Brazil, state of Ceará, Ediacaran fauna, fluviomarine environment, White Sea Assemblage.
reported occurrences of icnhofauna associated to Basin to the north. It is filled by immature silici
the Ediacaran biota in the states of: Santa Catarina clastic sediments, and its full sedimentary sequence
(Itajaí Basin), Paraná (Camarinha Basin) and Rio (formally known as the Jaibaras Group) comprises,
Grande do Sul (Camaquã Basin). The ichnofossils from bottom to top, the Massapê, Pacujá and
found in the Southern region were: Beltanelliformis Aprazível Formations (Oliveira 2000).
Menner, 1974; Bergaueria hemisphaerica Crimes, The Ubajara Graben (Neoproterozoic) repre
Legg, Marcos & Arboleya, 1997; Cochlichnus sents the basal sequence of the Jaibaras Basin. Its rocks
Hitchcock, 1858; Palaeophycus Hall, 1987; and underwent continuous low-grade metamorphism
Planolites Nicholson, 1873. The associated Edia during the Brasiliano Cycle, generating strongly
caran biota were: Aspidella Billings, 1872; Intrites penetrative mineral foliation (Oliveira and Mohriak
Fendonkin, 1980; and Sekwia Hofmann, 1981. 2003). This Graben was filled under fluviomarine
Numerous publications have addressed the conditions, and an important carbonate facies (the
regional geology of northwestern Ceará, including Frecheirinha Formation) is present in the intermediate
the Jaibaras Basin (Costa et al. 1973, Kegel et al. portion of the sequence (Delgado et al. 2003).
1958, Oliveira 2001, Oliveira and Mohriak 2003, The Jaibaras Graben was filled by cogenetic
Santos and Brito Neves 1984, Schobbenhaus and processes resulting from sedimentary input and
Brito Neves 2003, Torquato 1995, Torquato and magmatic events (Oliveira 2000), mainly during
Nogueira Neto 1996). the Ediacaran period (Table I). The Coreaú Dyke
To date, however, little palaeontological Swarm, of Vendian age, represents the initial stage
information has been available for the study area, of rift opening. Prior to the filling of the graben,
and no Ediacaran fossils have been recorded in the evolution of this distension system during the
northeastern Brazil. Lower Cambrian permitted the accommodation of
the Mucambo Pluton. During the Middle Cambrian,
GEOLOGICAL FRAMEWORK
basin sedimentation was accompanied by a consid
Northwestern state of Ceará is located within the erable volume of magma (the Parapuí Suite).
Borborema Province (the northeastern portion of Subsequently, the Meruoca Pluton was formed by
the South American Platform). Thermal, tectonic, passive intrusion (Oliveira 2001).
magmatic and other geological activities occurred in In addition to the data shown in Table I, De
this area during the Late Proterozoic, lasting through Araújo et al. (2012) dated Neoproterozoic zircons
the Cambrian-Ordovician (F.F.M. Almeida et al., in the Pacujá sandstone, indicating that the two
unpublished data). This region, known as the Médio main groups extended from 550 to 598 Ma and
Coreaú Domain, was a source area of substantial from 600 to 630 Ma.
rupture mobility throughout the late Precambrian, The sedimentary environment of the Jaibaras
resulting in a general trend of subsidence processes Group begins with a systematic repetition of con
(Santos and Brito Neves 1984). The structure of glomeratic fans, reflecting the cyclical nature
the area comprises horsts and grabens separated by of continuous basin subsidence. This phase is
ancient deep shear zones (Oliveira 2001). represented by the Massapê Formation. With
This study focused on the Jaibaras Basin (Figure continued basin subsidence, the material supply
2), which forms part of the Médio Coreaú System. was lower than the subsidence rate, permitting
The Jaibaras Basin is bounded by the Palaeozoic the development of lakes and rivers with deltaic
Parnaíba Basin to the west, the Transbrasiliano systems. This phase is represented by the Pacujá
Lineament to the south and east and the Ubajara Formation. Terminating the sedimentary deposits
TABLE I
Geochronological data for the major sedimentary and magmatic events
of the Jaibaras Basin, indicating the minimum and the maximum value of each age.
Rb/Sr (isochronous)
Pacujá Formation 535 + 27 F.R.G. Novais et al. (unpublished data)
metamorphism
in the basin, large alluvial fans brought an input The fossiliferous deposits differ from the Pacujá
of paraconglomerates with fragments that were Formation, presenting coarse, fossiliferous, strongly
compositionally similar to the underlying formations silicified and immature sandstone with cream,
as well as fragments from the plutons and from the yellowish, beige and sometimes grey colours. These
Parapuí Suite, whose assembly is recorded by the deposits exhibit, at the base, horizontal lamination
Aprazível Formation (Oliveira and Mohriak 2003). with planar cross bedding and low diversity of
With the attenuation of the Brasiliano tectonic Ediacaran fauna, followed to the top, trough cross-
cycle, the Borborema Province underwent a tran bedding and symmetrical ripple marks with higher
sition phase with post-orogenetic conditions. At diversity that helped characterise a meandering
the end of this phase, Saharan glaciations were fluvial system near a river mouth. This sandstone,
manifested in the region by the Serra Grande Group, herein informally called the Contra Fogo Sandstone,
which is located in proximity to the area in Africa occurs as a thick lobe oriented from south to north in
where this glaciation was (Mabessone 2002). the Jaibaras Graben and unconformingly rests on the
breccias of the Pacujá Formation.
RESULTS
The Contra Fogo Sandstone is partially eroded
FIELDWORK
and is approximately 100 m thick, with lateral
The research area exhibits the typical lithological variations. The Serra Grande Group laterally
characteristics of the Pacujá Formation, including underlies it to the east (Figure 3). On current maps,
fine/silty, reddish, micaceous sandstone, plane- the sandstones containing Ediacaran fossils are
parallel lamination and ripples. The formation included within the Parnaíba Basin (Cavalcante et
is underlain by unconforming vol canoclastic al. 2003), suggesting the need for a stratigraphic
breccias containing fragments of sandstones and revision of the area based on these newly discovered
basic volcanic rocks surrounded by a strongly fossiliferous deposits.
recrystallised silt matrix and overlain by fine At the Contra Fogo Farm, the deposits
sandstone with plane-parallel lamination. containing the Ediacaran fauna are found within
a 4,800-m2 rectangular area. The fossils, which Along Lameirão Creek, where the action of
occur on eroded surfaces, are mostly basal fluvial waters has accelerated the erosion process
discs and fronds but also include ichnofossils and exposure of the Contra Fogo Sandstone, the
and associated Ediacaran bilaterians. The local fossils extend over a distance of approximately 4
preserved species diversity is approximately 5 km. The diversity found at this locality is lower
species/m2, with an abundance of approximately than that found at Contra Fogo Farm due to the
8 individuals/m2. absence of Ediacaran bilaterians.
THE EDIACARAN FAUNA OF JAIBARAS BASIN represented mainly by discs forms. However,
Some of the Ediacaran fossils described below are most of the fossils were described in the field
stored in the Dom José Museum (MDJ) collection, due to the impossibility of collection. Including
located in the city of Sobral, and managed by the impressions and ichnofossils, the total number
Laboratório de Paleontologia (LABOPALEO) of of species is estimated to exceed 30, but only 12
the Universidade Estadual Vale do Acaraú-UVA. have been identified to date. However, the total
The collection of Ediacaran fauna is composed by number of specimens exceeds thousands in all
80 specimens (MDJ Ed-01 to MDJ Ed-80), being outcrops visited.
Description: A frond-shaped impression preserved Description: Only the basal disc of this species
in epirelief, the petaloid sometimes in negative and was observed. The disc is a bipartite circular im
sometimes in positive epirelief. Branches approxi pression in positive epirelief, 6.9 cm in width and
mately uniform, parallel, with the primary branches 13 cm in total diameter. Outer edge 3.3 cm wide,
diverging from the central shaft at slightly oblique separated from the inner disc by a depression. Inner
angles and the distal extremities connected to an disc 6.0 cm in diameter. Edge of the outer disc
exterior edge, preserved on only one side. Central perpendicularly cut by a long, narrow depression.
shaft 33 cm long and 2.5 cm wide at the base. Largest Material: MDJ Ed-007, found at the Contra
primary branch 9.0 cm long. Additional preserved Fogo Farm.
discs lacking fronds may represent this species. Observation: The depression perpendicular
Material: A single specimen described and to the outer disc represents the contact of the shaft
recorded in the field at the Contra Fogo Farm. on the disc during the fall of the frond onto the
Observation: The specimen was probably substrate. The exclusive preservation of the disc
not buried alive because the detachment of some was probably facilitated due to it being already
primary branches from the central shaft indicates buried in the sediment.
a state of decomposition. Moreover, a new disc is Discussion: This species was initially des
present at the base of the central shaft, indicating that cribed by Ford (1958 apud Ford 1999) from a
the substrate was quickly recolonised after burial. volcanic-sedimentary sequence in Charnwood
Discussion: The main criterion used to Forest, Leicestershire, England. Subsequently,
identify this specimen as C. arboreus is the oval Martin Glaessner found similar individuals in the
shape of the extremity (Laflame and Narbonne Ediacara Member in Australia (McCall 2006).
2008a). According to Laflame and Narbonne A fully preserved frond species, 25 cm in length,
(2008b), the appearance of the branches may found in Charnwood Forest (Ford 1999) facilitated
vary markedly in response to their preservation the identification of the Charniodiscus concentricus
quality. Charniodiscus sp. shafts are usually over specimen from Pacujá. Notably, this specimen
30 cm long. Charniodiscus has been interpreted appears to be associated with Cyclomedusa and
by Seilacher (1992) as Vendobionta, by Narbonne some typically Vendian discrete ichnofossils.
(2005) as a cnidarian and by Peterson et al. (2003)
as a fungus. The latter authors stated that the shafts Genus Cyclomedusa Sprigg, 1947
of the Charniodiscus specimens had apparently Species Cyclomedusa davidi Sprigg, 1947
been filled with sand in their internal spaces, (Figure 4C)
indicating that they were hollow in life and that
a large channel extended over the entire length Description: A bipartite circular impression in
of the shaft. Charniodiscus is a pandemic genus, positive epirelief, 12 cm in diameter. Inner disc
found at most Ediacaran localities. It is also the first surrounded by an outer disc with a slightly raised
Ediacaran group to appear in the fossil record, with edge, 1.5 cm in width. Inner edge raised above
some representatives extending into the Cambrian the outer edge, 7.0 cm in diameter, with a central
(Waggoner 2003). depression 1.5 cm in diameter.
Material: Described and photographed in the Genus Kimberella Glaessner and Wade, 1966
field at the Contra Fogo Farm. Species Kimberella quadrata Glaessner and Wade, 1966
Discussion: The central depression has been (Figure 4E)
interpreted as a Cyclomedusa subtype Ford (1999),
and it has frequently been called Cyclomedusa cliffi. Description: An oval-shaped impression with the distal
These impressions are by far the most common portions showing a relatively inflated outer edge and a
type in Charnwood Forest, with diameters varying distorted, raised internal zone. Anterior extremity with a
from less than 1 cm to 22 cm. The Charnwood neck-shaped structure. Transverse U-shaped partitions
Forest specimens lack striations, but the Australian not present on the inflated edges, possibly due to the
Cyclomedusa species do present these structures sediment particle size. Anteroposterior length 5 cm,
(Ford 1999). Jenkins (1992, p. 157, apud Crimes edge width 1 cm, width of the inner depression 0.5 cm.
and McIlroy 1999) considered C. plana, C. radiata, Material: One specimen recorded and
Springginia annulata Sprigg, 1949, Medusinites photographed in the field at the Lameirão creek.
asteroides Sprigg, 1949 and possibly Ediacaria Discussion: Kimberella quadrata from north
flindersi Sprigg, 1947 to be synonymous because eastern Brazil resembles the “C” subtype of Ivantsov
of their recorded associations. The phylogenetic (2009). Notably, the latter group of specimens is endemic
placement of Cyclomedusa sp. has been attributed to Russia and Australia, where it is included in the White
to cnidarians (McCall 2006). Sea Assemblage and found in association with medusoid
fossils and ichnofossils. Fedonkin and Waggoner (1997)
Genus Ediacaria Sprigg, 1947 reconstructed Kimberella quadrata as a symmetrical
Species Ediacaria flindersi Sprigg, 1947 bilaterian, a benthic animal with a univalvate and non-
(Figure 4D) mineralised shell, similar to a mollusc. This specimen
described here is of average length and size.
Description: A bipartite circular impression in
positive epirelief (though fragmented), 17 cm in Genus Medusinites Glaessner, 1966
total estimated diameter. Central disc 10 cm in Species Medusinites asteroides Sprigg, 1949
diameter, surrounded by an outer disc 4.5 cm in (Figure 4F)
width. Inner disc slightly raised above the outer
disc, representing ¾ of the impression size. Radial Description: A bipartite circular impression in
grooves absent. positive epirelief. Outer raised disc separated from
Material: MDJ Ed-53, found at the Contra the inner disc by a deep surrounding depression.
Fogo Farm. Radial grooves preserved in the edge of the outer
Discussion: Ediacaria flindersi is the largest disc, oriented toward the central disc. Central disc
and most common medusoid known from the various relatively smaller, with a total diameter of 10 cm.
Ediacaran assemblages (De 2006). Glaessner and Material: One specimen described and
Wade (1966) stated that radial structures may photographed in the field at the Lameirão creek.
or may not appear in the specimens and that the Discussion: Specimens with diameters of 1 to
largest complete specimen from Australia was 12 5 cm have been identified in Australia (Glaessner
cm in diameter. However, other fragments have and Wade 1966) and Africa (Bertrand-Sarfat et al.
indicated specimens much larger than 21 cm. The 1995). However, the Pacujá specimen has a diameter
phylogenetic affinity of Ediacaria has also been twice as large. Medusinites spp. are considered to be
attributed to cnidarians (McCall 2006). sedentary medusoid cnidarians (McCall 2006).
Material: One specimen described and of Parvancorina specimens and found no evidence
photographed in the field at the Contra Fogo Farm. favouring its affinity with arthropods. They related
Discussion: Gehling and Rigby (1996) the size of the specimens to the environmental depth:
described the first occurrence of this species from the greater the depth, the smaller the specimen.
the Ediacaran Member (Pound Subgroup) in the The Pacujá specimen is three times larger than the
Flinders Ranges region of southern Australia. largest specimen recorded by Ivantsov et al. (2004).
Palaeophragmodictya specimens from the Flinders In Brazil, Parvancorina was found at the Itajaí
Ranges reached 10 cm in diameter and exhibited Basin, in the state of Santa Catarina (Netto 2012).
a single central contact point that could be flat
or depressed and root-like structures (radial
Genus Pectinifrons Bamforth, Narbonne and
extensions surrounding the edge). Therefore, the
Anderson, 2008
Pacujá specimen shares the main taxobases of P.
Species Pectinifrons abyssalis Bamforth, Narbonne
reticulata, although it is slightly larger. This species
and Anderson, 2008
is the only Ediacaran organism whose affinity can
(Figure 5B)
be confidently attributed to Porifera.
and the primary supports, representing the central Discussion: Due to its morphological simplicity,
branches of the fronds, were originally composed of Palaeophycus is one of the most common ichnogenera
resistant materials that did not decompose even after associated with the Ediacaran fauna. However, most
lithification (Bamforth et al. 2008). The Pectinifrons species have millimetric diameters. Hagadorn and
specimens from Mistaken Point are associated with Waggoner (2000) described Palaeophycus specimens
Fractofusus, Charnia, Charniodiscus, Ivesheadia from the Great Basin of the United States as Ediacaran
and Aspidella discs. representatives.
be inferred for the deposits studied here. However, conditions at the time of deposition; both areas
Pectinifrons abyssalis is an intruded fossil belonging were shallower environments.
to the Avalon Assemblage, the oldest in the world. The action of microbial mats, which is consid
The age of the Ediacaran fauna contained in the ered primarily responsible for the process of
Contra Fogo Sandstone is consistent with previous fossilisation without mineralisation, is inferred to
datings for the key events in the evolution of the have been minimal or absent in the Contra Fogo
Jaibaras Basin (B.B Brito Neves and M.M.C. Neto, Sandstone fauna due to the intense bioturbation of
unpublished data; F.R.G. Novais et al., unpublished): the substrate and the sediment particle size. The
the Brasiliano Cycle (670-490 Ma), orogeny (625- ichnofossils associated (Paleophycus and Planolites)
580 Ma), the Mucambo Granite (548 + 24 Ma), the had been described in Ediacaran deposits worldwide,
metamorphism of Pacujá Formation (535 + 27 Ma) and including Brazil (Netto 2012)
the Neoproterozoic sedimentation inferred, the Meruoca The specimens of the Ediacaran assemblage from
Granite (540 + 7 Ma) and the Parapuí Suite (502 + 8 Ma). northeastern Brazil are much larger than conspecific
The presence of P. abyssalis is better understood specimens from elsewhere in the world. For example,
as a geographical intrusion, as suggested by analysis the Parvancorina minchami specimen described
of the palaeogeographical map of Veevers (2004). here is three times larger than the largest specimen
The dominance of taxa described from Russia and recorded by Ivantsov et al. (2004). Similarly, the
Australia can be attributed to the environmental Pacujá specimen of Medusinites asteroides is 10 cm in
diameter, while specimens from Australia are between the outcrops and facilitating access to the sites;
1 and 5 cm in diameter (Glaessner and Wade 1966). the Master’s student Robbyson Mendes and Prof.
This larger size may have been a deciding factor Regina Raick for support during fieldwork; the
in the preservation of the organisms as moulds. Bio Conselho Nacional de Desenvolvimento Científico
turbation would have increased the oxygenation of e Tecnológico (CNPq) for the Master’s program
the environment and thus decreased the chances of fellowship granted to author 1 and for the funds
fossilisation. Smaller individuals may have been present provided through project 401781/2010, coordin
but would have been bioturbed soon after burial. ated by author 2. We also thank the Museum for
housing the specimens and for providing free access
CONCLUSIONS to the Palaeontology Collection; and LAGESE
and LABOPALEO for their support during the
These rocks, which contain Ediacaran fossils, preparation of this paper.
may represent a new geological event between the
Pacujá Formation and the Aprazível Formation of RESUMO
the Jaibaras Basin.
Este estudo relata a primeira ocorrência da fauna Ediacarana
Field analyses suggest a meandering fluvial
no Nordeste do Brasil (Município de Pacujá, noroeste do
to estuarine environment for the Contra Fogo
Estado do Ceará) e apresenta interpretações preliminares de
Sandstone. The high abundance and diversity of the
seu significado. A correlação regional indica que os fósseis
fossil assemblage, indicate that the depth, salinity,
se originaram na Bacia do Jaibaras e que podem representar
oxygenation and light conditions of this environment
um novo sistema geológico. O ambiente de deposição pode
were appropriate for faunal development.
ser atribuído a um sistema transicional. Nove espécies
The ichnofossiliferous assemblage confirms
Ediacaranas podem ser identificadas, incluindo membros
the presence of Arenicolites, Palaeophycus and
de grupos pandêmicos (ex. Charniodiscus arboreus
Planolites in the Ediacaran.
Glaessner, 1959; ?Charniodiscus concentricus Ford, 1958;
The predominance of discs representing frond
Cyclomedusa davidi Sprigg, 1947; Ediacaria flindersi
bases rather than full fronds is probably related to high
Sprigg, 1947; e Medusinites asteroides Sprigg, 1949) e
environmental oxygenation and to intense bioturbation,
grupos endêmicos (ex. Kimberella quadrata Glaessner &
which may have restricted the activity of the microbial
Wade, 1966; Palaeophragmodictya reticulata Gehling &
mats that would have preserved the more fragile portions.
Rigby, 1996; Parvancorina minchami Glaessner, 1958;
Pectinifrons abyssalis, previously recorded
e Pectinifrons abyssalis Bamforth, Narbonne, Anderson
only in Newfoundland, is also present among the
2008). Três icnogêneros também estão presentes: Areni
ichnofossils of the Contra Fogo Sandstone.
colites Salter, 1857; Palaeophycus Hall, 1987; e Planolites
Based on the predominance of representatives
Nicholson, 1873. A idade relativa dos depósitos está
from the White Sea Assemblage, the inferred age of the
entre o ?Ediacarano e Cambriano, e a fauna assemelha-
deposits is at least 560 Ma, but the ichnofossiliferous
se a Assembleia White Sea. A bioturbação apresenta
species extend these deposits to Cambrian. This date
típicos icnogêneros Ediacara não ramificados com pouca
is consistent with the ages of the key events in the
profundidade no substrato. Esta ocorrência até então
Jaibaras Basin, including the Pacujá Formation.
desconhecida da fauna de Ediacarano reforça a importância
ACKNOWLEDGMENTS do Estado do Ceará para a paleontologia brasileira e global.
We thank the director of the Pacujá Museum, Palavras-chave: Brasil, estado do Ceará, fauna de Edi
Alancardec Leopoldino for guiding the authors to acara, ambiente transicional, Assembleia White Sea.
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