From alluvial fans to lacustrine systems: trace fossils applied to paleoenvironmental analysis in the Taubaté Basin (Brazil)

Abstract

Studies integrating ichnology and sedimentology in paleoenvironmental interpretations are rare in Brazilian rift successions. This study applies an integrated ichno-sedimentological analysis to a well core from the Resende and Tremembé Formations (Cenozoic Taubaté Basin) to reconstruct paleoenvironmental conditions. Three paleoenvironments were identified: alluvial fan, braided river, and lacustrine. The alluvial settings is characterized by coarse-grained deposits displaying bioturbation intensity and sporadic trace fossils, dominated by Palaeophycus and Skolithos, reflecting opportunistic colonization. Braided river deposits consist of stacked sandstones with trough cross-bedding and a trace-fossil assemblage with Palaeophycus, Skolithos, Macanopsis, and Taenidium. The lacustrine environment is characterized by dark shale interbedded with fine-grained sandstones, including ostracod-rich white beds. Sparse beds containing Planolites indicate a complex lacustrine system with potential variations in salinity and oxygenation. Comparison with previous studies suggests that ichnofossils and paleosols reflect water table fluctuations and humid conditions in the Resende Formation. For the Tremembé Formation, ichnofossil data, palynomorphs (Botryococcus and Pediastrum), and taphonomic signatures in fishes suggest a dynamic environment with fluctuating water levels, periodic salinity increases, and decreased oxygen levels.

KEYWORDS:
continental rift; ichnology; Skolithos ichnofabric; paleosols; lacustrine system

1. INTRODUCTION

The integration of trace fossil and sedimentological analyses has enabled the inference of paleoecological and paleoenvironmental parameters in a wide range of sedimentary environments (Pemberton & MacEachern, 1995). Biogenic structures reflect the behavior of their producers, which was influenced by paleoenvironmental conditions (e.g., Seilacher, 1964; Ekdale et al., 1984; Pemberton et al., 1992; Sedorko & Francischini, 2021). The application of ichnology for interpreting marine environments has been extensively studied (e.g., MacEachern et al., 1999, 2007; MacEachern & Bann, 2008, 2020; Carmona et al., 2009; Abdel-Fattah et al., 2016; Buatois et al., 2019; Sedorko et al., 2019, 2023) compared to continental environments (Buatois & Mángano, 1995b, 2002, 2004, 2007, 2009; Bohacs et al., 2000; Genise et al., 2000; Melchor, 2004; Hunt & Lucas, 2007). Compared with marine environments, the number of studies on continental ichnology have increased after 1980 (e.g., Klappa, 1980; Bown, 1982; Sarjeant, 1983; Bown & Kraus, 1983, 1987; Genise & Bown, 1994a, b; Hasiotis & Dubiel, 1993, 1994; Buatois & Mángano, 1995a; Genise et al., 2000; Hasiotis, 2002, 2004; Hasiotis et al., 2013; Mueller & McCann, 2014; Nascimento & Guimarães Netto, 2019; Rangel et al., 2022), especially in alluvial deposits. Compared to other Brazilian basins, the Cenozoic basins are relatively less studied in terms of ichnological content (e.g., Rossetti, 2000; Guimarães Netto & Rossetti, 2003; Gandini et al., 2017; Sedorko et al., 2024, 2025). While the fossil content in the Taubaté Basin is well-documented, with records of leaf and wood impressions in the mud facies of the Resende Formation (Riccomini et al., 1987), as well as sponges (Wickert, 1974), amphibians (Riff & Bergqvist, 1999; Santos, 2020), fishes (Woodward, 1898; Schaeffer, 1947; Malabarba, 1998), birds (Alvarenga, 1982, 1985; Olson & Alvarenga, 2002; Mayr et al., 2011), and mammals (Soria & Alvarenga, 1989; Bergqvist & Ribeiro, 1998; Alvarenga et al., 2005), the occurrence of trace fossils have been only mentioned in the Resende and Tremembé Formations (e.g., Bergqvist & Ribeiro, 1998; Torres-Ribeiro & Borghi, 2007; Voltani et al., 2023), without a detailed characterization. And therefore, there are no studies using an integrated ichno-sedimentologic approach in the Taubaté Basin. In this paper, we describe the trace fossil content and the sedimentary deposits that host them. This integrated approach opens new windows to the reconstruction of paleoenvironmental conditions in the rift basins during the Cenozoic for the Taubaté Basin.

A continuous well-log enabled the description of transitions between different paleoenvironments throughout the Paleogene in this basin. Distinct ichnological patterns in these lithostratigraphic units revealed colonization behaviors controlled by high energy, oxygen deficient, and fluctuating water table levels. The comparison between trace fossils and other proxies enhanced the interpretation of paleoenvironments in the Taubaté Basin, reinforcing that ichnological analysis serves as a valuable proxy for reconstruction of palaeoenvironments.

2. GEOLOGICAL SETTING

The Taubaté Basin, located in southeastern Brazil in São Paulo state, extends NNW-SSW for 170 km in length and 20 km in width, with depths reaching 1,200 m (Fernandes, 1993; Padilha et al., 1991) (Figure 1). Its basement comprises igneous and metamorphic rocks from the Paleoproterozoic to the Neoproterozoic Ribeira Fold Belt (Hasui & Ponçano, 1978). The basin is part of the Continental Rift of Southeastern Brazil (CRSB), associated with a Paleogene distensive tectonic phase (Fernandes, 1993; Zalán & Oliveira, 2005; Riccomini et al., 2004). The sedimentary fill can be divided into two phases. The first phase, syn-tectonic to the rift, includes alluvial fans, lacustrine, and fluvial deposits, corresponding to the Resende, Tremembé, and São Paulo Formations (Riccomini, 1990; Carvalho et al., 2011). The second phase, post-diastrophic tectonics, is characterized by the deposition of alluvial deposits of the Pindamonhangaba Formation and braided fluvial deposits of the Itaquaquecetuba Formation (Melo et al., 1987; Riccomini, 1990; Carvalho et al., 2011).

Figure 1
Schematic geological map of the studied area. (A) Location of the Taubaté Basin on the southeastern Brazil; (B) Location of the drilled core in the upper middle part of the Taubaté Basin. The distribution of the geological units after Campos et al. (2021); (C) Stratigraphic chart of the Taubaté Basin (modified from Riccomini et al., 2004).

This study focuses on the Resende and Tremembé Formations. The lowermost Resende Formation comprises clast- and matrix-supported conglomerates deposited in proximal areas and mudstones in medial and distal alluvial fan areas, grading into sandy facies of medial to braided fluvial deposits (Riccomini et al., 2004; Carvalho et al., 2011). The alluvial fan deposits of the Resende Formation transition laterally and vertically into the fine-grained sediments of the Tremembé Formation (Riccomini & Coimbra, 1992; Riccomini et al., 2004). The Tremembé Formation consists of fine-grained siliciclastic rocks, primarily mudstone and shale rich in organic matter, interbedded with sandstones, marls, and dolomites (Riccomini, 1990; Riccomini et al., 1996, 2004; Mendonça Filho et al., 2010). The paleoenvironment is interpreted as lacustrine, developed in the central portion of the Taubaté Basin (e.g., Riccomini, 1993; Riccomini et al., 2004; Torres-Ribeiro & Borghi, 2007; Bergamaschi et al., 2010; Setta et al., 2012; Bergue et al., 2015).

The main debate concerns the stratigraphy and limnology of the lake. Riccomini et al. (1996) characterized the lacustrine system as a playa lake, indicating ephemeral saline water conditions, drying up annually or every few years (Renaut & Gierlowski-Kordesch, 2010). Other authors have emphasized deep water and perennial conditions (Torres-Ribeiro & Borghi, 2007; Bergamaschi et al., 2010; Setta et al., 2020). Torres-Ribeiro (2004) argued that the lake was a closed, oligomictic, eutrophic system in a semiarid climate, and therefore, could not be classified as a playa lake due to the absence of evaporitic deposits.

3. MATERIAL AND METHODS

This study presents data from a well core drilled in Pindamonhangaba municipality (45°28’22.91”W; 22°51’37.21”S), designated as 2 PD-1SP-PINDA 2. This core, which samples the Resende and Tremembé Formations, was acquired from the upper middle part of the basin. It has a diameter of 40 mm, organized in 181 boxes, and spans 715.45 m of thickness, with 34 m corresponding to the top of the well core. Macroscopic core description was conducted at a 1:12 scale, subdivided into three stratigraphic intervals, involving the recognition of lithology, texture, sedimentary structures, and bed contacts.

Trace fossil analysis and ichnotaxonomy were performed alongside sedimentary deposit descriptions using the ichnofabric approach. The degree of bioturbation was quantified using the Reineck (1963) scale, which ranges from 0 (no bioturbation) to 6 (completely homogenized by biogenic activity). Ichnofabric characterization was based on the ichnotaxobases of Bromley (1990) Paleosols were identified according to their maturity index (Retallack & Reinhardt, 1988), and the nomenclature of paleosol orders follows modern soil orders (United States Department of Agriculture, 2014).

4. RESULTS

4.1 Sedimentary deposits

The core reveals four distinct deposits across two recognized stratigraphic intervals, attributed to the Resende and Tremembé Formations (Table 1). The lower interval consists of matrix-supported conglomerates with a quartz and feldspar sandy matrix, interbedded with paleosol, and features coarse-grained deposits with medium-to-coarse sandstone exhibiting trough cross-bedding and parallel lamination, characteristic of the Resende Formation. The upper interval is predominantly dark shale, intercalated with beds containing ostracods, fishes, and plant remains, interpreted as the Tremembé Formation.

Table 1
Summary of integrated data described in the well core.

Deposits	Ichnology	Body Fossils	Paleopedology	Oxi-Redox Features	Depositional System
Fine-grained deposits	Planolites	Plants and fishes	-	-	Lacustrine
Sandstone deposits	Macanopsis, Palaeophycus, Skolithos and Taenidium	Plants and fishes	-	-	Braided river
Paleosols	Planolites	-	Entisols and Vertisols	Concentration and depletion of Fe and Mg	Alluvial fan
Coarse-grained deposits	Palaeophycus and Skolithos	-	-	Concentration and depletion of Fe	Alluvial fan

4.1.1 Coarse-grained deposits

Coarse-grained deposits account for 17% of the core, totalizing 116 m in thickness. These deposits, concentrated mainly in the lower interval above basement rocks, are composed essentially of feldspar and quartz clasts derived from metamorphic rocks, ranging from 3 to 10 mm in diameter, totalizing 40% of fabric, being 25% composed by feldspar, classified as oligomictic conglomerate. Matrix-supported conglomerates with granule- to very coarse pebble-sized clasts in a sandstone-mudstone matrix. They are poorly sorted and display massive or crudely stratified textures (Figures 22B), occasionally with Palaeophycus and Skolithos.

Figure 2
Coarse-grained deposits. (A) Matrix-supported conglomerate with granule to coarse pebble-sized deposits and mud-sandy matrix (670.50 m); (B) Clast-supported conglomerate of very coarse pebble-sized clasts of quartz and feldspar, and scattered reddish-brown redoximorphic features.

4.1.2 Paleosols

The paleosols make up 8% of the core, totaling 56 m in thickness, and occur interbedded with conglomerates. The profiles consist of intervals of stacked compound paleosols, ranging from 90 cm to 2 m in thickness, classified as Entisols and Vertisols (Figure 3A). The Entisols display a horizon sequence of Cg and C, with boundaries ranging from sharp to gradual and wavy topography. Horizon textures are predominantly sandy, with colors including light gray (5GY 8/1), light pink (7.5YR 8/4), and red (2.5YR 5/8). Pedogenic structures are fine to medium, angular to subangular, and blocky, sometimes exhibiting redoximorphic features (Figure 3B). The Vertisols exhibit a horizon sequence of Bss and C, with gradual transition and wavy topography. Horizon textures are primarily loamy-sand, with colors ranging from light gray (5GY 8/1) to dusk red (7.5R 3/4) and red (7.5R 4/6). Pedogenic structures include fine angular blocky peds and fine to medium cuneiform structures with wedge-shaped peds, subdivided by slickenside surfaces (Figure 3C). The Bss and Cg horizons of both paleosol types contain an abundance of redoximorphic features, such as redox depletion and redox concentration (Figure 3B). Distinct clay and iron depletions along secondary pores and the matrix, ranging from fine to medium (0.1-8 cm in diameter), are distributed within these horizons and occupy 5 to 10% of the surface. Redox concentrations, such as Fe-nodules ranging from 2 to 5 mm, are particularly abundant in the Bss horizon. Another notable feature is pore linings found in root channels (rhizohaloes) and invertebrate trace fossils, showing colors varying from gray to red to yellow (Figure 3D). The trace fossils preserved in the described paleosols has low ichnodiversity, only with Planolites and rhizoliths, with a bioturbation index of 1 to 2 (4% to 8%).

Figure 3
Paleosols. (A) Succession of paleosols evidencing the occurrence of disseminated redoximorphic features marked by the alternation of red and grey colors in the well-core (618.50 m); (B) Subangular blocky structure and elongate, grey-rimmed depletion and reddish zones with concentration of iron (yellow arrow); (C) Pedogenetic structures marked by slickensides. Note the polished aspect of the paleosol matrix (yellow arrow); (D) Root cast filled with coarse-grained sand displaying an outer reddish-purple halo separated from the inner greyish core.

4.1.3 Sandstone deposits

These deposits represent 29% of the core, totalizing 205 m, and occur in the middle core interval (Figure 4). They consist of fine- to coarse-grained conglomeratic sandstone deposits, with feldspar clast ranging from 2 to 4 mm in diameter, with some gravel-sized clasts occurring locally, moderately sorted, with trough cross-bedding, and trace fossils identified as Macanopsis, Palaeophycus, Skolithos, and Taenidium. Plant remains are also present (Figures 55B). In subordinated interval, the sandstone is very fine- to fine-grained, well-sorted, and displays parallel lamination with 1 mm in thickness composed of sand (essentially by quartzs and feldspars) containing sparse Palaeophycus and plant remains (Figure 5C).

Figure 4
Sedimentological log showing the trace fossil distribution in the studied well-core 2PD-1SP-Pinda 2.

Figure 5
Sandstone deposits. (A, B) Conglomeratic sandstone with trough cross-bedding B. The red line highlights the trough cross strata (298.0 m); (C) Very fine- to fine-grained sandstone with parallel lamination.

4.1.4 Fine-grained deposits

These deposits constitute 38% of the core, totalizing 260 m in thickness, and predominate in the upper core interval. They are organized in millimeter-thick parallel lamination, black or dark gray in color, forming dark shale with trace fossils of Planolites as well as fish and plant remains. Locally, these deposits alternate between dark shales and fine-grained sandstones. In the uppermost intervals (from 95 to 34 m), fine-grained deposits include interbedded millimeter-thick white beds rich in ostracods and dark shale. The contacts between beds are clear and abrupt, marked by color changes. The ostracod-rich laminae range from 1 to 2 mm in thickness, with ostracod valves ranging from 0.7 to 1.01 mm in diameter, mostly disarticulated, slightly fragmented, and with rounded edges (Figures 6AC).

Figure 6
Fine-grained deposits. (A) Fine-grained deposits showing homogeneous aspect of the dark shale; (B) Detail of the dark shale displaying the millimeter-thick parallel lamination; (C) The uppermost interval of fine-grained deposits formed of alternations between lighter laminae rich in ostracods and darker organic laminae (38.0 m).

4.2 Trace fossils

The ichnofabric characterization allowed to identify the recurrent ichnogenera in the analyzed intervals. Five ichnogenera were identified: Skolithos, Palaeophycus, Macanopsis, Taenidium, and Planolites. These ichnogenera were described following the order of occurrence, from base to top (Figures 7 and 8).

Figure 7
Biogenic structures. (A) Skolithos in coarse-grained deposits, in cross section with redoximorphic features, enhanced the contrast with the matrix; (B, C) Skolithos in conglomerate sandstone deposits, with little contrast in relation to the matrix; note are slightly irregular; (D) Skolithos in fine-grained sandstone; note the contrast with the matrix, possibly due to the presence of mud; (E, F) Palaeophycus in matrix- supported conglomerate; note the red circles in figure showing the opening burrows with little contrast with the matrix. Note also the aspects with slight roughness; (G) Palaeophycus showing opening in sandstone deposits; note slight contrast between the burrow limit and the matrix; (H) Palaeophycus in conglomerate sandstone deposits note; the slight contrast with the host rock; (I) Palaeophycus in sandstone deposits with high contrast with the host rock caused by lens of the mud in matrix.

Figure 8
Biogenic structures. (A, B) Macanopsis in sandstone deposits; note the termination of burrow with chamber in final part of structure, J- shaped; (C, D) Taenidium in coarse-grained sandstone; notes the active fill meniscate with sinuous traced with full relief; (E, F) Planolites in dark shales showing horizontal burrow and slight wall with active filled; (G) Planolites in cross section in paleosol facies; (H) Root cast with color reddish- purple rims separated from the external inner greenish and internal inner grayish core; note the fill material (coarse-grained sand).

Skolithos is characterized by straight, cylindrical, unbranched, vertically oriented unlined burrows. The diameter ranges from 1 to 2 mm, and the length varies from 1 to 4 mm (Figures 7AD). It is found in association with Palaeophycus, which features cylindrical, straight to gently sinuous (when in visible bedding planes), unbranched, and horizontally oriented burrows. Palaeophycus is more common in fine-grained sandstone deposits, with diameters ranging from 2 mm to 10 mm. The appearance of Palaeophycus compacted limits is influenced by grain size, showing higher contrast with the matrix in fine-grained sandstones and lower contrast in coarse-grained sandstones (Figures 7EI). Macanopsis is characterized as cylindrical, unbranched and unlined J-shaped burrows that are passively filled and range from vertical to inclined. The 'J' ends in a basal chamber without scratch marks. The burrow diameter is 3 mm, expanding to 5 mm in the chamber, and the total length ranges from 1 to 2 cm (Figures 88B). These burrows are associated with Taenidium and Palaeophycus. Taenidium is characterized by cylindrical, meniscated, unwalled, moderately sinuous, unbranched, and vertically oriented burrows, with diameters ranging from 3 to 4 mm and lengths from 1 to 2 cm (Figures 88D). Planolites is described as cylindrical, horizontally oriented, straight to slightly sinuous, actively filled burrows. The diameter of the circular apertures ranges from 2 to 5 mm, with lengths reaching up to 10 mm when visible. Planolites is also commonly associated with Skolithos (Figures 8EG).

Rhizoliths are subclassified into rhizohalos and root casts. Rhizohalos are characterized by zones of Fe and Mn depletions, with grayish and greenish mottled areas and reddish-purple (10Y 8/1) rims surrounding an inner gray core. These structures are vertically oriented with lateral downward branches, varying in diameter from 2 to 4 cm and lengths from 10 to 15 cm. The matrix textures of rhizohalos match the hosting paleosol, ranging from medium- to coarse-grained sandstone, with occasional gravel-sized clasts. Root casts are described as vertically oriented cylinders, ranging in length from 9 to 11 cm, with lateral branches measuring 4 to 6 cm. They are filled with coarse-grained reddish-purple sandstone, with rims separated from the inner greenish and grayish (2.5YR8/4) core. The patterns exhibit a herringbone style with lower lateral branches present (Figure 8H).

5. DISCUSSION

5.1 Distribution of trace fossils in the Taubaté Basin

The core from well 2 PD-1SP-PINDA 2 is divided into three stratigraphic intervals. The lower interval consists of matrix-supported conglomerates interspersed with paleosols, interpreted as part of the Resende Formation. The intermediate interval is characterized by sandstone deposits, also attributed to the Resende Formation. The upper interval is composed primarily of fine-grained deposits, mainly by dark shale, corresponding to the Tremembé Formation (Figure 4).

Coarse-grained deposits, including matrix-supported and clast-supported conglomerates, are typical of alluvial fan systems and are influenced by climate and bedrock type (Blair & McPherson, 1992, 1994; Nichols & Fisher, 2007). In rift basins, these deposits result from sediment gravity flow processes (Blair & McPherson, 1994). Debris flows in alluvial fans are generated by steep slopes and converging drainage networks that concentrate sediment and overland flow (Reneau et al., 1984, 1990; Reneau & Dietrich, 1987). The deposits of the Resende Formation include ortho- and paraconglomerates in proximal areas, mudstones in medial and distal fan sections, and sandstones in braided river channels (Amador, 1975; Riccomini, 1990; Riccomini et al., 2004; Ramos et al., 2005, 2006). In the studied well-core, the matrix-supported conglomerates with sandstone and mudstone matrix occurs in the lower interval and are characterized by the wide-ranging distribution of oxi-redox features (Figure 2B). Redoximorphic features form from the reduction, translocation and oxidation of Fe and Mn (Vepraskas, 2015). Fe and Mn oxides are reduced in the soil under saturation with stagnant, oxygen depleted (anaerobic) water (Vepraskas,2015). The pattern of redox pedofeatures varies depending on the movement of water and air through the soil, the duration of water saturation and anaerobic conditions, and the occurrence of organic matter (Vepraskas, 2015). In this sense, the presence of water in the system—when saturation persists long enough to reduce Fe and Mn—leads to the depletion of iron oxides throughout the groundmass. This process results in the formation of hypocoatings composed of these oxides (Vepraskas, 2015). In contrast, short duration of water saturation or short periods fluctuation of water table levels results in intrapedal Fe/Mn oxide nodules and hypocoatings (Schwertmann & Fanning, 1976; Richardson & Hole, 1979; Arshad & St. Arnaud, 1980). Accordingly, based on our observations and the features observed in coarse-grained deposits (Figure 2B), the low degree of iron reduction suggests that water and air movement through the soil occurred in brief cycles. These short-duration fluctuations were insufficient to affect the entire soil matrix, as indicated by the lacustrine context.

The presence of Skolithos and Palaeophycus in continental setting is commonly associated with alluvial, fluvial meandering, and lacustrine deposits (Alpert, 1974; Mueller & McCann, 2014; Hasiotis, 2004; Buatois & Mángano, 2004; Buatois et al., 2007; Hasiotis et al., 2013) and reflects moist, subaerial to aquatic habitats, likely linked to shelter burrows and scavenging by arthropods, mostly insects (Stanley & Fagerstrom, 1974; Gierlowski-Kordesch, 1991; Knaust, 2017; Nascimento & Guimarães Netto, 2019). Palaeophycus, often found in fluvial and lacustrine deposits (Buatois & Mángano, 2004; Hasiotis, 2004; Krapovickas et al., 2010; Knaust, 2017), is interpreted as the work of semi-aquatic insects and annelids (Pemberton & Frey, 1982; Keighley & Pickerill, 1995; Krapovickas et al., 2010; Knaust, 2017). In debris flow deposits, Palaeophycus may indicate brief colonization by dwelling invertebrates during periods of low sedimentation rates (Neto de Carvalho & Baucon, 2016). Trace fossils in coarse-grained deposits, including Skolithos and Palaeophycus (Figure 4), are often interpreted as opportunistic life strategies (Levinton & Bambach, 1970; Vossler & Pemberton, 1988; McCall & Soster, 1990). High hydrodynamic energy and organic matter availability are key factors affecting colonization during deposition (DeCelles et al., 1991; Gierlowski-Kordesch, 1991). Periods of debris flow deposition may render environments unsuitable for colonizers due to coarse-grained substrates and high sedimentation rates, affecting the settlement and food availability (Dashtgard et al., 2008). Coarse-grained deposits are often overlain by mud that infills pore spaces between clasts, indicating reducing energy flows and allowing colonization (Dashtgard et al., 2008). Insects, for example, have diverse feeding strategies based on available organic matter in continental settings (Wootton, 1988), and would be able to colonize these deposits.

Paleosol development in alluvial fan systems are controlled mainly by sedimentation rates (Hartley et al., 2013). Entisols and Vertisols reflect different patterns of developments: rapid burial associated with frequent sedimentation contrasts with reduced sedimentation rates during conglomerate deposition, which indicate longer periods of exposure and, consequently, greater root colonization (Hartley et al., 2013; Golab et al., 2018).

The paleosols are composed by coarse-grained sandstone with root cast filled by same material. Furthermore, the clasts are less than 2 mm in size in the matrix (Figure 8H), however, paleosol is interbedded with coarse-grained deposits (Figure 4). In this sense, the difference in sediment composition between the paleosols and the underlying deposits suggests that their development occurred in the final stages, within braided channels located downstream of the fan system.

The presence of rhizoliths with redoximorphic features such as manganese and iron oxides reflect seasonal flooding (Stiles et al., 2001; Fiedler & Sommer, 2004; Birkeland, 1999; Hartley et al., 2013). Gray rhizohaloes with red or purple hypocoatings are characteristic of the moderately well-drained red paleosols (Kraus & Hasiotis, 2006). The colors associated with the traces are typical of redoximorphic features developed since changes in the redox conditions in response to fluctuations in degree of saturation, when the soil was saturated and relatively depleted of oxygen (O), Fe was reduced and subsequently removed from areas of the soil matrix (e.g., Duchaufour, 1982; Fanning & Fanning, 1989; Vepraskas et al., 1992; Vepraskas, 1994).

The presence of Fe concentrations in the matrix around gray depleted zones (rhizohaloes) is indicative of surface-water gleying caused by perched water tables (e.g., Fanning & Fanning, 1989; Vepraskas et al., 1992). In this case, the paleosol deposits were subject to brief flooding events, as evidenced by the small red rims and limited bleaching, which reflect a low intensity of gleying. Possibly, the paleosol deposits in alluvial final portion have started development with high humidity levels of the system and likely experienced a sudden and permanent rise in water table (Kraus & Hasiotis, 2006), associated with deposition of debris flow deposits drowning the vegetation. The sparse occurrence of Planolites in these deposits corroborates harsh conditions for colonization of the paleosoil.

The sandstone deposits are predominantly fine- to coarse-grained conglomeratic sandstones with poorly defined trough cross-bedding (Figure 5A). In rift basins, these deposits are part of a braided river system, indicating a gradual reduction in hydrodynamic energy (Sadler & Kelly, 1993; Riccomini et al., 2004; Nichols & Fisher, 2007). The transition from alluvial fan to braided river is marked by decrease in the clast size and increase in the proportion of overbank facies (Nichols and Fisher, 2007). In alluvial fans, conglomeratic sandstones in proximal zones show cross-bedding and indicate braided streams near the basin margin (Graham, 1983; MacCarthy, 1990; Sadler & Kelly, 1993). Proximal river channels are bedload-dominated, while further into the basin, braided rivers create sandy bars (Collinson, 1996).

This interpretation is supported by our description with stacks of sandstone with trough cross bedding and with structure marked by feldspar clasts (Figure 5B). Furthermore, in the Taubaté Basin, these deposits are interpreted as the Resende Formation representing axial braided river channels (Riccomini, 1990; Sant’Anna, 1999; Riccomini et al., 2004; Ramos et al., 2005, 2006). These river beds increase in frequency and thickness towards the interior of the basin, where traction flows become more dominant than gravitational flows (Ramos, 2003). Deposits commonly associated with these gravel sets are low-angle-bedded gravel or sand include planar cross bedded (Blair & McPherson, 1994) as observed in Figure 5C, furthermore, cross-bedding and the preservation of bar forms suggest deposition in braided rivers (Collinson, 1996).

The predominance of trough cross bedding with sandstone matrix and an ichnofabric composed essentially by Palaeophycus and Skolithos suggests opportunistic colonizers during relatively lower hydrodynamic activity, but still under high energetic conditions in braided settings (Sedorko et al., 2025). This scenario is briefly interrupted by development of a lake system, with sparse beds containing Planolites, plants and fishes (around 510 m in the well core; Figure 4). Besides Palaeophycus and Skolithos, the sandstone deposits are characterized by trace fossils such as Macanopsis, and Taenidium (Figure 4). Taenidium also indicate sediment moisture and cohesion during colonization (Buatois & Mángano, 2002, 2007; Mikuś & Uchman, 2013). Taenidium is associated with detritus-feeding, locomotion, and dwelling behaviors, with menisci indicating backfilling on softground (Neto de Carvalho & Baucon, 2016). Insects can create meniscated burrows to avoid desiccation during water table fluctuations (Hasiotis, 2000). The presence of Macanopsis suggests water table fluctuations commonly observed in fluvial to marginal fluvial settings (Bown & Kraus, 1983; Hasiotis, 2004; Buatois & Mángano, 2002, 2007; Fernandes & Carvalho, 2006; Nascimento et al., 2021). This affirmation is corroborated by redoximorphic features in paleosol deposits, indicating fluctuations of the water level in the system with moments of flooding and subaerial exposure. This dynamic reflected by paleosols is also suggested by the pattern of colonization of the substate. J-shaped burrows of Macanopsis have been linked to various trace makers. Casertano et al. (1998) identified Macanopsis-like burrows produced by scarab beetles. Bown & Kraus (1983) suggest spiders, wasps, or beetles as trace makers, while Hembree & Hasiotis (2007) attribute Macanopsis to dung beetles. Ground beetles and scarab beetles are commonly considered trace makers for continental Macanopsis, though spiders, millipedes, and frogs are also noted (Mikuś & Uchman, 2013; Nascimento et al., 2021, 2024)

According to Mikuś & Uchman (2013), beetle activity is controlled by groundwater levels, avoiding flood inundation. Meniscate ichnotaxa, however, are influenced by substrate moisture and cohesiveness, which change due to progressive drying after flooding (Neto de Carvalho & Baucon, 2016). Morrissey & Braddy (2004) suggested that a Beaconites-Taenidium ichnofabric reflects colonization of subaerially exposed sediment in response to seasonal desiccation, with animals excavating to aestivate or mold at water table levels. This impoverished assemblage of meniscate forms suggests temporary communities in ephemeral water bodies and longer dry periods (Minter et al., 2007; Neto de Carvalho & Baucon, 2016). On the other hand, Macanopsis represents a period when sandy bars were exposed and water tables were lower, allowing colonization probably by arthropods, considering the size of the structures (Nascimento et al., 2021). Thus, the ichnofabric is dominated by Palaeophycus and Skolithos in sandstones, and the sparse Macanopsis and Taenidium associated with trough-cross-stratified sandstones deposited corroborates a colonization mostly of tops of sand bars (Mueller & McCann, 2014). The rapid migration of sand bars precludes extensive colonization and the predominance (Wang et al., 2014).

Overall, the trace fossils from Resende Formation, featuring Skolithos, Palaeophycus, Macanopsis, and Taenidium, shows two intervals: the lower with coarse-grained alluvial deposits containing Skolithos and Palaeophycus and paleosoils, and the middle with sandstone deposits hosting Skolithos, Palaeophycus, Macanopsis, and Taenidium (Figure 9), locally with ephemeral lake. High-energy coarse-grained deposits have few vertical structures like Skolithos, with Palaeophycus predominating. Gravel deposits typically show minimal bioturbation by Skolithos (e.g., Dashtgard et al., 2008). The recurrence of meniscated burrows would aligns with Scoyenia ichnofacies, typical of fluvial and lacustrine systems undergoing progressive desiccation (Seilacher, 1967), but the dominance of simple vertical and horizontal structures, associated with low intensities and low ichnodiversity across intervals suggest opportunistic colonization in the context of ephemeral water bodies (Buatois & Mángano, 2002, 2009; Buatois et al., 2007; Mueller & McCann, 2014; Neto de Carvalho & Baucon, 2016). In this sense, we do not attribute these ichnofabrics to any ichnofacies.

Figure 9
Paleoenvironmental evolution of the Taubaté Basin (modified of Riccomini et al., 1987). (A) Initial moment with development of alluvial fan filled the basement of the Taubaté Basin; (B) Intermediate moment with reworking of alluvial fan by braided rivers system compounding Resende Formation characterized as Skolithos ichnofabric; (C) Final moment with development of lacustrine system above Resende Formation, recover all deposits with lacustrine facies, recognize as Tremembé Formation dominated by Mermia ichnofabric.

The upper core interval features dark shale interbedded with fine-grained sandstone and ostracodite at the top (Figure 6). Thick, fine-grained deposits in the rift basin are formed mainly by lacustrine systems controlled by climate and tectonics, as seen in the Taubaté Basin (Bohacs et al., 2000; Torres-Ribeiro & Borghi, 2007). The dark shale layers reflect reduced sediment influx during periods of intermittent tectonic reactivation (Galloway & Hobday, 2012). Bohacs et al. (2000) propose that lake controls, such as accommodation changes and sediment-water supply balance, influence lake occurrence and character. Fine-grained deposits lacking evaporites, mud cracks, or tractive facies, and with sparse trace fossils, suggest low-energy, deep-water conditions with anoxic sedimentation (Talbot & Allen, 1996; Torres-Ribeiro & Borghi, 2007; Mendonça Filho et al., 2010; Voltani et al., 2023). These deposits, attributed to the Tremembé Formation, are microclastic units comprising siltstones, mudstones, shales, marl, limestone, and sandstone (Riccomini, 1990; Fernandes, 1993; Molinari, 2003; Torres-Ribeiro, 2004; Torres-Ribeiro & Borghi, 2007).

Oxygen deficiency in the lake, controlled by mixing frequency, primary production, and water chemistry (Bohacs et al., 2000), enhances organic preservation by limiting scavenger activity and bacterial respiration (Demaison & Moore, 1980). Allochthonous organic material from deltas and fluvial input increases food supply for infaunal deposit-feeding and grazing communities (Torres-Ribeiro & Borghi, 2007). Lakes typically support a diverse range of benthic organisms, such as annelids, semi-aquatic insects, crustaceans, and mollusks (Cohen, 2003; White & Miller, 2008). Biota in lacustrine systems is influenced by factors like oxygen, salinity, energy, nutrient supply, and substrate consistency (Buatois & Mángano, 2007, 2009). Balanced-fill lakes, depending on sediment and water supply, affect biogenic activity, with closed lakes becoming strongly stratified and oxygen-depleted, leading to high organic matter preservation and reduced biotic activity (Bohacs et al., 2000; Scott et al., 2012).

Planolites, preserved mainly in the upper interval (Figure 4), are produced by deposit-feeders actively processing sediment (Pemberton & Frey, 1982; Gierlowski-Kordesch, 1991; Keighley & Pickerill, 1995; Knaust, 2017). In lacustrine systems, Planolites indicates periods of organic material deposition (Scott et al., 2012). The high abundance of monospecific associations dominated by Planolites reflects oxygen depletion, allowing organic matter preservation (Talbot and Allen, 1996; Magyar et al., 2006; Uchman et al., 2007; Scott et al., 2012; Voltani et al. 2023). Buatois & Mángano (1995b) proposed the Mermia ichnofacies, characterized by horizontal grazing and feeding traces, subordinate locomotion traces, and moderate ichnodiversity in very soft submerged substrates. The absence of typical grazing traces precludes this attribution and, similar to the lower interval, we do not attribute this ichnofabric to one ichnofacies. The recurrence of Planolites ichnofabric increases upward due to reduced lake stress (Melchor, 2004; Torres-Ribeiro & Borghi, 2007; Scott et al., 2012; Voltani et al., 2023).

5.2 Controls of trace fossil distribution

Few studies mentioned the occurrence and distribution of trace fossils in the Resende Formation. The formation is characterized a conglomerate with a mud-sandy matrix deposited under a humid climate, as supported by Lima & Amador (1985), Blair & McPherson (1994), and Bergue et al. (2015). Trace fossils such as Skolithos and Palaeophycus in coarse-grained deposits indicate colonization by opportunistic fauna under relatively high-energy conditions in fluvial settings, but are post depositional in alluvial settings. The trace fossils of the Tremembé Formation have attracted more attention in the literature, but only with mention to its occurrences (e.g., Torres-Ribeiro & Borghi, 2007; Santana et al., 2023; Voltani et al., 2023). Regarding the depositional interpretations, Riccomini (1990) and Sant’Anna (1999) proposed the Tremembé Formation as a playa-lake system, an under-filled lake based on Bohacs et al. (2000). Such lakes are characterized by evaporitic facies and increased salinity (Bohacs et al., 2000). High salinity in profundal deposits typically limits biodiversity, leading to the absence of Mermia ichnofacies (Buatois & Mángano, 2007; Scott et al., 2012). Saline conditions also affect lacustrine faunas, potentially causing low oxygen levels (Scott et al., 2012). Deeper saline lakes are often stratified due to salinity or temperature differences (Renaut & Gierlowski-Kordesch, 2010), which would typically inhibit biotic development. Trace fossils characteristic of restricted environments, however, suggest that this lake had deep conditions and lower salinity levels, as supported by Torres-Ribeiro & Borghi (2007), Santana et al. (2023), and Voltani et al. (2023). Mendonça Filho et al. (2010) identified palynomorphs Botryococcus and Pediastrum in Tremembé Formation, indicating variations in the water column. Botryococcus, associated with high luminosity and salinity, reflects low water levels, while Pediastrum, indicative of fluvial input with high nutrients and low luminosity, suggests higher water levels. The alternation between these palynomorphs points to fluctuations in lake base levels driven by more humid climatic conditions (Mendonça Filho et al., 2010).

Voltani et al. (2023) supported the idea of a deep lake by noting biostratinomic characteristics of fish, such as partial flotation, which indicate water stratification and anoxic conditions at the bottom. Mass fish deaths could result from mixing of anoxic hypolimnion with the epilimnion, causing anoxic events. The presence of partially floating carcasses suggests a lake depth greater than 10 m (Voltani et al., 2023). Ostracodite in the uppermost intervals (Figure 9C) also reflects environmental stress. Bergue et al. (2015) described ostracod assemblages in the Tremembé Formation and found that high concentrations of Heterocypris correspond to shallowing phases, while Limnocythere-Cypretta assemblages indicate deep lake phases. This suggests variations in water levels and bottom oxygenation, as supported by sparse Planolites.

Overall, the Tremembé Formation aligns with Bohacs et al.’s (2000) model of a deep lake with low oxygen exchange rates, creating a stratified water column with low or absent oxygen at the lake bottom. This is corroborated by analyses of organic matter, taphonomy, palynomorphs, and ichnology (Torres-Ribeiro & Borghi, 2007; Mendonça Filho et al., 2010; Bergue et al., 2015; Setta et al., 2020; Voltani et al., 2023). When the lake was open, sediment and water input diluted the salinity, providing oxygen and allowing for colonization by opportunistic organisms, as evidenced by sparse Planolites. Conversely, when the lake was closed, increased salinity and anoxic conditions were indicated by Botryococcus, Heterocypris, and well-preserved organic matter (e.g., Voltani et al., 2023).

6. CONCLUSIONS

This study provides a comprehensive analysis of trace fossils and sedimentological characteristics in the Resende and Tremembé Formations within the Taubaté Basin, revealing distinct ichnological and sedimentological patterns. In the Resende Formation, trace fossils such as Skolithos and Palaeophycus described in coarse-grained deposits suggest colonization by opportunistic fauna under relatively high-energy conditions in fluvial settings, or to post-depositional colonization in alluvial settings, both corresponding to Skolithos and Palaeophycus. This result is in accordance with previous interpretations of the depositional setting for this unit, formed under a humid climate. The limited documentation of trace fossils in this formation (Palaeophycus, Skolithos, Macanopsis, and Taenidium) reflects its challenging depositional conditions for faunal activity.

Conversely, the Tremembé Formation evidences a complex paleoenvironmental history. Initial interpretations of the Tremembé Formation as a playa lake system, dominated by evaporitic facies and high salinity, were revisited. Evidence from trace fossils, supported by available data on palynomorphs, and taphonomic characteristics suggest that the lake exhibited restricted conditions with variable salinity. Palynomorphs such as Botryococcus and Pediastrum indicate fluctuating water levels and salinity, supporting the hypothesis of a dynamic lake environment influenced by climatic changes. The presence of trace fossils suggesting anoxic settings (Planolites), along with biostratinomic evidence of fish flotation and anoxic conditions, underscores the lake’s stratified nature and episodic oxygenation. During the phases of open lake conditions, it supported opportunistic colonization as indicated by sparse Planolites. In contrast, during closed periods characterized by elevated salinity, the lake experienced more severe anoxic conditions, as reflected for the high degree of organic matter preservation.

This study highlights the significant impact of climatic and environmental fluctuations on the ichnological and sedimentological records of the Taubaté Basin. The data supports a model of dynamic lake systems with alternating periods of high salinity, anoxia, and variable water levels, influencing faunal activity and preservation. These insights present implications for reconstructing past climatic and ecological conditions.

ACKNOWLEDGEMENTS

This study is a contribution to the project “Icnologia das bacias de Resende e Taubaté no contexto do Rift Continental do Sudeste do Brasil” (Jovem Cientista do Nosso Estado – FAPERJ 281340; 200.131/2023 - SEI-260003/000688/2023 - BBP). VC thanks CAPES for the master`s grant. DS thanks the National Council for Scientific and Technological Development (CNPq) for the research grant (CNPq 306493/2022-5), and the Alexander von Humboldt-Stiftung for the postdoc fellowship (CAPES/Humboldt Program nº 14/2022 - process nº 23038.004870/2021-69).

• Manuscript ID: BJGEO-2024-0062.R2.
• How to cite:
  Cambria, V., Menezes, M. N., Dal’ Bó, P. F., Borghi, L., & Sedorko, D. From alluvial fans to lacustrine systems: trace fossils applied to paleoenvironmental analysis in the Taubaté Basin (Brazil). Braz. J. Geol. (2026), 56:e20240062. https://doi.org/10.1590/2317-4889e20240062
• Financial support:
  Icnologia das bacias de Resende e Taubaté no contexto do Rift Continental do Sudeste do Brasil” (Jovem Cientista do 519 Nosso Estado – FAPERJ 281340; 200.131/2023 - SEI-260003/000688/2023 - BBP). CAPES - Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - 88887.834516/2023-00. National Council for Scientific and Technological Development (CNPq) for the research grant (CNPq 306493/2022-5).
• Data availability statement:
  All data supporting this study are included in the article.

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Edited by

• SCIENTIFIC EDITOR:
  Carlos Henrique Grohmann https://orcid.org/0000-0001-5073-5572

• ASSOCIATE EDITOR:
  Lucas Warren https://orcid.org/0000-0002-2050-6514

Data availability

All data supporting this study are included in the article.

Publication Dates

• Publication in this collection
  06 Feb 2026
• Date of issue
  2026

History

• Received
  01 Nov 2024
• Accepted
  21 Oct 2025