1. Introduction
Animals have transformed the planet, transporting matter and energy through biogeochemical cycles, predation and locomotion (Logan et al. Reference Logan, Hayes, Hieshima and Summons1995; Lenton et al. Reference Lenton, Boyle, Poulton, Shields-Zhou and Butterfield2014). However, even before mobility in marine animals was widespread, they were already capable of altering their host sedimentary environments. Biomineralizing metazoans appeared in the terminal Ediacaran ca. 550 Ma, and the transport and redeposition of derived calcareous bioclasts (Warren et al. Reference Warren, Simões, Fairchild, Riccomini, Gaucher, Anelli, Freitas, Boggiani and Quaglio2013) can be interpreted as events of redistribution and potentially restructuring of matter (sensu Judson, Reference Judson2017), which generated a new sedimentary grain source in shallow marine carbonate environments (e.g., Warren et al. Reference Warren, Simões, Fairchild, Riccomini, Gaucher, Anelli, Freitas, Boggiani and Quaglio2013). Here, we present evidence for the physical and chemical substrate redistribution of bioclasts generated by Ediacaran biomineralizing metazoans in siliciclastic settings. The presence of these metazoans thus increased facies heterogeneity and complexity far earlier than previously thought, resulting in a restructuring of this sedimentary factory which persists until today.
Bioclasts in fine-grained siliciclastic rocks can indicate sedimentary processes. Ordovician subtidal bioclastic deposits with a fine matrix indicate storm-induced bioclastic reworking/winnowing of fine sediments and of bioclastic-filled gutter casts (Kreisa, Reference Kreisa1981). Bioclastic turbidites can create flow-associated sorting that in turn create more complex textures than their siliciclastic-only counterparts (Bornhold & Pilkey, Reference Bornhold and Pilkey1971).
Here, we show that Corumbella, an Ediacaran metazoan of problematic affinity, formed bioclastic deposits that record sedimentary dynamics in fine siliciclastic facies of the Tamengo Fm., Brazil. Disarticulated remains of Corumbella were reported from carbonate facies from the Tagatiyá Guazú Formation, Paraguay (Warren et al. Reference Warren, Fairchild, Gaucher, Boggiani, Poire, Anelli and Inchausti2011; Warren et al. Reference Warren, Pacheco, Fairchild, Simões, Riccomini, Boggiani and Cáceres2012; Warren et al. Reference Warren, Quaglio, Simões, Gaucher, Riccomini, Poiré, Freitas, Boggiani, Sial and Sial2017). Corumbella has a tubular form, reaching up a few centimetres in length, which has a multi-element skeleton (cataphract) built by originally aragonitic sclerites that can readily disarticulate (Osés et al. Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022) (Fig. 1).
2. Geological setting
The Corumbá Group (western Brazil) comprises a mixed siliciclastic-carbonate succession in the South Paraguay Belt, Amazon Craton (Fig. 2; Alvarenga et al. Reference Alvarenga, Moura, Gorayeb, Abreu, Cordani, Milani, Filho and Campos2001). The fossiliferous Tamengo Formation, that outcrops in the cities of Corumbá and Ladário (Mato Grosso do Sul state), comprises mudstones, sandstones and carbonates deposited across a storm-dominated carbonate ramp (Amorim et al. Reference Amorim, Afonso, Leme, Diniz, Rivera, Gómez-Gutiérrez, Boggiani and Trindade2020). Radiometric dating constrains the deposition of the Tamengo Formation in the interval ca. 555–542 million years old (Parry et al. Reference Parry, Boggiani, Condon, Garwood, Leme, McIlroy, Brasier, Trindade, Campanha, Pacheco, Diniz and Liu2017). Here, we concentrate on the siliciclastic levels that outcrop in sections along the Paraguay river (Fig. 2). Shales bearing Corumbella were previously interpreted as transgressive hemipelagic sediments (Boggiani, Reference Boggiani1997) from outer-ramp settings (Oliveira, Reference Oliveira2010; Amorim et al. Reference Amorim, Afonso, Leme, Diniz, Rivera, Gómez-Gutiérrez, Boggiani and Trindade2020; Ramos et al. Reference Ramos, Giorgioni, Walde, do Carmo, Fazio, Vieira, Denezine, Santos, Adôrno and Guida2022). Here, these fine-grained clastic facies are reinterpreted as shallow marine, as supported by previous works (Almeida, Reference Almeida1945; Almeida, Reference Almeida1965; Boggiani, Reference Boggiani1997; Antunes et al. Reference Antunes, Warren, Okubo, Freitas, Inglez, Caetano-Filho and Morais2023).
3. Materials and methods
Samples were collected in the Sobramil Mine (ELC VI) and in the Cacimba Ecopark (ELC VII), Tamengo Formation, in the cities of Corumbá and Ladário, state of Mato Grosso do Sul, Brazil (Fig. 2). In this study, seven polished slabs and five thin sections were analysed. Fossil samples and thin sections are housed either in the Palaeontological Collections of the Laboratory of Paleobiology and Astrobiology, University of São Carlos (codes CAP/1A and CAP/1F), or of the Institute of Geosciences, University of São Paulo (USP) (code GP/1E).
For microfacies observation, rocks were cut with a diamond saw in the Institute of Astronomy and Geophysics, University of São Paulo (USP). Slabs were serially polished either manually or with progressively finer discs in the Materials Group facilities, Engineering and Technology division of the Brazilian Centre for Research in Energy and Materials. Samples were imaged using either Olympus DSX110 microscope or Zeiss stereomicroscope with camera, respectively, in the Laboratory of Palaeobiological Studies and in the Laboratory of Petrographic Microscopy, in the Institute of Geosciences, USP. After the inspection of polished blocks, samples were selected for petrographic thin-sectioning and photographed in an Olympus petrographic microscope with camera.
CL (cathodoluminescence) analysis was done using a Cathodoluminescence Cold Cathode CITL 8200 MK3A coupled with a Nikon microscope (School of GeoSciences, University of Edinburgh). Photomicrographs were taken with a x10 objective lens, current of 900–1000 A and voltage of 25–30 V.
SR-microXRF (synchrotron radiation micro X-ray fluorescence) mapping was performed at the XRF beamline of the Brazilian Synchrotron Light Laboratory (LNLS/CNPEM), following the protocol of Osés et al. (Reference Osés, Petri, Voltani, Prado, Galante, Rizzutto, Rudnitzki, da Silva, Rodrigues, Rangel, Sucerquia and Pacheco2017). PIXE (particle-induced X-ray emission) measurement was performed at the external beam of the LAMFI (Laboratory of Materials Analysis by ionic beams) facilities (Silva et al. Reference Silva, Rodrigues, Added, Rizzutto, Tabacniks, Mangiarotti, Curado, Aguirre, Aguero, Allegro, Campos, Restrepo, Trindade, Antonio, Assis and Leite2018), at the Institute of Physics, University of São Paulo. The analysis followed the protocol described in Osés et al. (Reference Osés, Petri, Becker-Kerber, Romero, Rizzutto, Rodrigues, Galante, da Silva, Curado, Rangel, Ribeiro and Pacheco2016).
SEM (scanning electron microscopy) analyses were made at the Institute of Geosciences, University of São Paulo, using a FEI Quanta 250 microscope with an Oxford Si(Li) EDS detector. Samples were analysed uncoated. The parameters used can be found in the data bar of each micrograph.
4. Results
In the Sobramil Mine (Figs. 2 and 3), clayey/silty sandstones are associated with black calcareous skeletal fragments (Fig. 4) of Corumbella (Fig. 4g) and unidentified calcareous taxa. These bioclasts occur as discontinuous laminae, lenses and pods with erosive sharp basal contacts (Figs. 4, 5a and b). Bioclasts and massive clay form heterolithic structures of planar/undulated/wavy horizontal strata with internal plane-parallel lamination and cross-lamination with <1 cm-thick cross-sets (Figs. 4, 5a, b and e). The cross-laminae have low-angle contacts to the cross-strata set boundaries (Figs. 4a and 5e). Ripples are capped by bioclasts and/or iron oxy(hydr)oxides that also subtly follow ripple cross-lamination (Fig. 4i). Thick bioclastic crusts are cut by iron oxy(hydr)oxide partings of pyrite pseudoframboids (Fig. 5a–c). Bioclastic accumulations also form syneresis cracks (Fig. 5f–h). Bioclasts are entrapped in microbial lamination with orientated grains and pyrite pseudoframboids (Fig. 5i–l).
CL (cathodoluminescence) imaging of bioclastic level in Fig. 5a shows that it is formed by calcareous bioclasts with a non-luminescent core and bright luminescent calcite cement (Fig. 6a). SR-microXRF (synchrotron radiation micro X-ray fluorescence) mapping shows the association of calcium to calcareous bioclasts and to cement in these levels (Fig. 6b). Figure 6c and d indicate that the Corumbella fossil in Fig. 4g and other bioclasts have a correlation of Ca and of Sr. PIXE (particle-induced X-ray emission) spectrum of a bioclast has high counts of Ca and of Sr relative to the host rock (Fig. 6e).
SEM and EDS (energy-dispersive X-ray spectroscopy) analyses of syneresis cracks in Fig. 5g and h show details of the mineralogy (Fig. 7) and of the composition and cross-cutting relationships of both bioclastic level and clay veins (Fig. 8) forming these structures.
The Cacimba Ecopark has outcrops of dark grey silty/sandy claystone (Figs. 2 and 3). Plane/wavy discontinuous irregular bioclastic laminae/lenses of silt/fine sand forming erosive features (Fig. 9a–c) interlaminate with dark organic wavy-crinkled discontinuous lamination with clay minerals, trapped orientated elongated translucent crystals and pseudoframboids (Fig. 9d–f). V-shaped feature is putative evaporite feature in cross-section (Fig. 9g).
SEM imaging details pyrite framboid in the sample in Fig. 9d–f (Fig. 10a and b). EDS mapping further supports a pyritic composition (Fig. 10c–f). These framboids are associated with organic, microbial, laminae and scattered organic matter (Fig. 10g–k).
5. Discussion
An elemental composition consistent with a calcite mineralogy is shown for bioclasts, as evidenced by SR-microXRF mapping and by PIXE point analysis (Fig. 6b–e). These analyses indicate an association of Sr to Ca-bearing Corumbella articulated fossils (Fig. 6c and d) and to the black grains interpreted as bioclasts (Fig. 6c–e). Osés et al. (Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022) showed that Corumbella had a primary aragonitic mineralogy. The bioclasts are black since they are organic-rich (kerogenized), as shown for Corumbella fossils by Osés et al. (Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022). CL imaging of bioclastic levels shows bioclasts as grains with non-luminescent cores that are outlined by bright luminescent margins (Fig. 6a), exactly as shown for Corumbella sclerites by Osés et al. (Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022). This variation in luminescence was attributed by these authors to an increase of Mn and Fe contents towards the margins of the skeletal elements due to recrystallization. CL also shows that the bioclasts are cemented by bright luminescent calcite (Fig. 6a).
The bioclastic deposits are similar to the pre-trilobite, early ‘Cambrian shell concentrations’ (Li & Droser, Reference Li and Droser1997), as both are dominated by Small Shelly Fossils (SSFs), <10 cm-thick, and forming discontinuous thin beds and lenses. An increase of deposit thickness later in the Phanerozoic was explained by a decrease of organic content in skeletons, successful colonization of high-energy habitats and increase of carbonate production (Kidwell & Brenchley, Reference Kidwell and Brenchley1994).
The absence of many sedimentary features in non-Corumbella-hosting clastic sediments of the Tamengo Fm. suggests that an interplay of bioclasts, mud and sedimentary dynamics creates sedimentary complexity. The cataphract organization of Corumbella (Osés et al. Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022) when disarticulated and fragmented yields bioclasts/crystals that reach very coarse sand grade, up to 2 mm in diameter (Figs. 1 and 4). The predictable disarticulation behaviour in Fig. 1 (Osés et al. Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022) illustrates how progressively smaller unities were formed. Bioclast thickness, microstructure and organic content influence strength, which in turn controls fragmentation (Zuschin et al. Reference Zuschin, Stachowitsch and Stanton2003). A thin skeleton with a possibly laminar microfabric (Osés et al. Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022) may have decreased strength in Corumbella, presumably in turn increasing fragmentation (Zuschin et al. Reference Zuschin, Stachowitsch and Stanton2003). The multi-element skeleton disarticulation/fragmentation of Corumbella yielded variations in grain size and shape that interacted with flow dynamics influencing the geometry, thickness, internal fabrics and sedimentary structures of the deposits. Fine sand-sized bioclasts form <1 mm-thick horizontal discontinuous laminae (Fig. 4e, g and i), thin lenses (Figs. 4i and 5e) and low-angle cross-lamination (Figs. 4a, c and 5e). Flow energy limits the maximum particle size that is transported, and these fine-grained thin deposits formed by lower-energy conditions. The well-sorting of bioclasts forming cross-lamination is explained by current winnowing during ripple formation/migration. Coarse sand-sized bioclasts form ≥2 mm lenses and pods only (Fig. 4a and b). These are local erosive lag deposits – cut-and-fill structures – formed after erosion and flow deceleration (Baas et al. Reference Baas, Best and Peakall2015). The red films enveloping bioclastic levels bear pyrite pseudoframboids (Fig. 5b and c), representing films that have likely resulted from the decay of biofilms around the deposits.
The heterolithic alternation of bioclasts and massive clay forms by current flow in an upper plane bed regime, which creates sediment ‘bursting’, followed by deposition of large, heavy bioclasts and clay settling, yielding bioclast/clay fining-upward couplets (Fig. 4d–g) (Cheel & Middleton, Reference Cheel and Middleton1986). Turbulence explains bioclasts subvertical to the lamination (Fig. 4e). Unequivocal Corumbella sclerites (Fig. 4g) have a convex-up configuration and form ‘house-of-cards’-like imbrication (Fig. 4e and f), which might indicate tractive flow action (Quaglio et al. Reference Quaglio, Warren, Anelli, dos Santos, Rocha-Campos, Gazdzicki, Strikis, Ghilardi, Tiossi and Simões2014). The convex-up configuration indicates deposition in high-energy flows (McFarland et al. Reference Mcfarland, Westrop and Cheel1999; Yao et al. Reference Yao, Aretz, Li and Wang2016). The variation of thickness of bioclastic-clay couplets might point to periods of longer high-energy conditions – possibly storms with deposit cannibalism (Einsele, Reference Einsele1992) – followed by also long intervals of low-energy deposition (Fig. 4i) vs rapid events (Fig. 4e and g).
Polygonal features with similar orientation and variable sizes have bioclastic accumulations and are here interpreted as syneresis cracks (Fig. 5f). The surface separating part and counterpart – positive epirelief and negative hyporelief – possesses a reddish iron oxy(hydr)oxide parting (Fig. 5f). In cross-section, the features manifest as an irregular discontinuous thick level of silt/sand-sized anhedral calcite crystals with jigsaw texture (bioclasts), that is interlaminated with clay, and cut by vertical sinuous clay mineral veins that taper downwards (Figs. 5g, h, 7 and 8).
There are several hypotheses for the polygonal features. Load structures can be ruled out since deformation of underlaying clay strata is not observed (Fig. 5g). A MISS origin is undermined by the absence of microbial lamination, organic matter or pseudoframboids associated with the structures (Figs. 5g, 7 and 8). The polygonal features are similar to polygonal desiccation cracks, but no evidence of subaerial exposure has been observed. Molar-tooth structures can also be ruled out since these are more irregular in plane-view and possess ductile and brittle features and vertical crumpled veins infilled with calcite microspar (Pratt, Reference Pratt1998a).
The remaining hypothesis is that the polygonal features are syneresis cracks. Syneresis cracks manifest as lenses or lines with varied shapes (e.g., straight, curved and sigmoidal), with distinct orientations and connection degrees, commonly yielding polygonal shapes and two population sizes at a single bed ‒ ca. 1–10 mm wide and ca. 1–30 cm long (Pratt, Reference Pratt1998). In cross-section, they are vertical/subvertical relative to the horizontal strata, V-shaped, tapering downwards or upwards, and are infilled with silt and sand from overlaying and/or underlaying beds (Pratt, Reference Pratt1998; McMahon et al. Reference Mcmahon, Hood and Mcilroy2016, and references therein). Syneresis cracks are hosted in marine and non-marine thin strata, finer than fine sand, with clay, silt, sand, and carbonate contribution. They are formed sporadically, underwater, intrastratally in shallow depth, at low-energy settings (Pratt, Reference Pratt1998). Earthquakes trigger sediment dewatering, yielding fluid scape and the formation of distorted dikelets (Pratt, Reference Pratt1998). Syneresis cracks have morphological and dimension similarities with the features herein described, though the latter are shorter, possess clay-infilled veins, and is a mixture of sand-sized bioclasts and cohesive clay deposited under high-energy conditions. The structures herein interpreted as syneresis cracks occur associated with likely bioclastics levels (as observed but not shown by Amorim et al. Reference Amorim, Afonso, Leme, Diniz, Rivera, Gómez-Gutiérrez, Boggiani and Trindade2020, for the boundary between their silty-shale and calcimudstone), sporadically, both in our bioclastic and microbial microfacies (Figs. 4, 5 and 9), suggesting that bioclasts would have had a role in the formation of such structures. As highlighted by McMahon et al. Reference Mcmahon, Hood and Mcilroy(2016), ‘subaqueous sedimentary cracks’ need adhesion to yield cracks and these form after some source of stress. In the facies here studied, adhesion could be achieved either by the mixture of clay sediment, organic-rich bioclasts, and water, and/or by the attraction between clay particles (McMahon et al. Reference Mcmahon, Hood and Mcilroy2016) infilling voids among bioclasts, or even the early cementation (McMahon et al. Reference Mcmahon, Hood and Mcilroy2016) of calcite around aragonite bioclasts (Osés et al. Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022). Additionally, stress could result from the deposition of bioclasts themselves in a stormy context. Amorim et al. (Reference Amorim, Afonso, Leme, Diniz, Rivera, Gómez-Gutiérrez, Boggiani and Trindade2020) attributed syneresis cracks in the Tamengo Formation to a storm-generated origin. A model claimed that rapid deposition over a clay bed would cause dewatering of this deposit, yielding syneresis cracks (see discussion in Pratt, Reference Pratt1998). Here, a model adapted from the model of Pratt (Reference Pratt1998) is hypothesized: the turbulent rapid deposition of sand-sized bioclasts mixed with clay during storms caused a seismic shaking-like effect that triggered fissuring (dikelets) and dewatering and fluidization of clay that subsequently filled distorted dikelets. Clay levels below the bioclastic deposit are bent and clay minerals are locally chaotically organized, indicating disturbance. Indeed, the model of Pratt (Reference Pratt1998) predicts that the cohesion of clay under the forces of a seismic event yields bending and disruption of clay. It has been proposed that the proportion of swelling clay contribution (smectite) would control the potential of syneresis crack formation (Pratt, Reference Pratt1998). Indeed, syneresis cracks are found in smectite-bearing (Fazio et al. Reference Fazio, Guimarães, Walde, Do Carmo, Adorno, Vieira, Denezine, Silva, Viana, Borges and Pinho2019) siliciclastic beds of the Tamengo Formation. In sum, bioclastic contribution in the Tamengo Formation siliciclastic settings could have been an important component of crack structures formed in predominantly cohesive (clayey) sediments.
Ripples are capped by fine sand/silt bioclasts (Fig. 4i). The aragonitic bioclasts (density=2.947 g/cm3) behaved like heavy minerals (density >2.87 g/cm3), being swept on an active plane bed, yielding internal dispersive pressures that moved bioclasts to the top of the flow. This creates sorted bioclast concentrations (‘lags’) over ripple crests (Cheel & Middleton, Reference Cheel and Middleton1986). Such ‘capping’ may have prevented underlying bedform erosion.
In heterolithic facies from the Sobramil Mine, microbial mats bearing bioclasts manifest as pyrite pseudoframboids/rare framboids irregular accumulations and orientated elongated translucent crystals (Figs. 5j–l), as these are features of microbial lamination (Schieber, Reference Schieber1999; Noffke et al. Reference Noffke, Gerdes, Klenke and Krumbein2001; Noffke, Reference Noffke2009). Organic-rich, microbial laminae bear pyrite pseudoframboids (Figs. 9d–f and 10g–k). Microbially laminated facies are documented from the Cacimba Ecopark (Fazio et al. Reference Fazio, Guimarães, Walde, Do Carmo, Adorno, Vieira, Denezine, Silva, Viana, Borges and Pinho2019), and similar lamination occurs lacking bioclasts when interlaminates with erosive features infilled with bioclasts in this locality (Fig. 9). These structures form by current erosion and are filled with sediment and bioclasts during waning energy flow (Pérez-López & Pérez-Valera, Reference Pérez-López and Pérez-Valera2012).
Bioclastic heterolithic deposits formed in cyclic, fair-weather conditions, under episodic storm disturbance, are considered tide-influenced facies. Lenticular laminae of fine sand interlayered with clay (Fig. 5d) likely formed when sand and clay deposited, respectively, by tidal currents and slack-water (Posamentier & Walker, Reference Posamentier and Walker2006). Evidence for evaporites (Fazio et al. Reference Fazio, Guimarães, Walde, Do Carmo, Adorno, Vieira, Denezine, Silva, Viana, Borges and Pinho2019) and desiccation features (Ramos et al. Reference Ramos, Giorgioni, Walde, do Carmo, Fazio, Vieira, Denezine, Santos, Adôrno and Guida2022) in calcimudstones points to shallow, inter-tidal to supra-tidal deposition. The presence of vendotenids indicates settings with light (Amorim et al. Reference Amorim, Afonso, Leme, Diniz, Rivera, Gómez-Gutiérrez, Boggiani and Trindade2020). Fazio et al. (Reference Fazio, Guimarães, Walde, Do Carmo, Adorno, Vieira, Denezine, Silva, Viana, Borges and Pinho2019) also described evidence of evaporites in the Cacimba Ecopark, and we show a putative evaporite feature (Fig. 9g) in the dark microbial claystone. Taken together the evidence of this paragraph, but considering that we have not observed desiccation evidence, we consider that this facies represents deposition in a microbial-dominated, occasionally evaporite protected setting with occasional storm influence, in mid-ramp, subtidal conditions. Previous works associated Corumbella with deeper, outer-ramp settings (Oliveira, Reference Oliveira2010; Amorim et al. Reference Amorim, Afonso, Leme, Diniz, Rivera, Gómez-Gutiérrez, Boggiani and Trindade2020; Ramos et al. Reference Ramos, Giorgioni, Walde, do Carmo, Fazio, Vieira, Denezine, Santos, Adôrno and Guida2022).
Corumbella is found in shallow facies worldwide. In the Tagatiyá Guazú Formation, Paraguay, disarticulated remains of Corumbella occurs in thrombolytic mudstones (Warren et al. Reference Warren, Fairchild, Gaucher, Boggiani, Poire, Anelli and Inchausti2011; Warren et al. Reference Warren, Quaglio, Simões, Gaucher, Riccomini, Poiré, Freitas, Boggiani, Sial and Sial2017) and in grainstones and massive oolitic grainstones with plane-parallel stratification and wave ripples (Warren et al. Reference Warren, Pacheco, Fairchild, Simões, Riccomini, Boggiani and Cáceres2012). The facies from the Tagatiyá Guazú Fm. are interpreted to have been deposited in a shallow carbonate platform, with protected environments, probably in an evaporitic lagoon setting (Warren et al. Reference Warren, Pacheco, Fairchild, Simões, Riccomini, Boggiani and Cáceres2012). Corumbella is also found in siltstones of the lower member from the Wood Canyon Formation, USA, together with Gaojiashania and Conotubus (Smith et al. Reference Smith, Nelson, Tweedt, Zeng and Workman2017). These shales were deposited in a shallow marine setting (Hagadorn & Waggoner, Reference Hagadorn and Waggoner2000; Smith et al. Reference Smith, Nelson, Tweedt, Zeng and Workman2017). The Kushk Series of central Iran consists of two facies associations ‒ carbonates deposited in a shallow subtidal setting and shales that represent a deep, outer-ramp environment (Vaziri & Laflamme, Reference Vaziri and Laflamme2018). The latter comprises sub-unit 6, consisting of argillaceous grey shales that bear Corumbella and other taxa. Corumbella was recently reported in fine-grained sandstones of the Huns Member, Urusis Formation, Nama Group (Turk et al. Reference Turk, Maloney, Laflamme and Darroch2022).
Afonso et al. (Reference Afonso, Pereira, de Faria, Romero, Amorim, Basso and Trindade2024) showed that the distribution of Cloudina calcareous bioclasts in carbonate beds from the Tamengo Fm. are related to sedimentary dynamics. They described two types of shallowing-upward cycles, Type I encompassing proximal mid-ramp deposits with poorly sorted reworked bioclasts and Type II consisting of better-preserved fossils deposited in lower energy conditions at distal mid-ramp and at outer-ramp settings. Afonso et al. (Reference Afonso, Pereira, de Faria, Romero, Amorim, Basso and Trindade2024) and our data reinforce that metazoan bioclasts reflect sedimentary dynamics in the Tamengo Fm., both in carbonate and siliciclastic settings.
The input of calcareous skeletons into siliciclastic sediments also yielded a continuum of dissolution/recrystallization of aragonite as well as decay of skeletal organics. These processes might have led to an increase in carbonate cements and to preservational microbial pathways yielding authigenic mineral precipitation (e.g., pyrite and clay minerals). Osés et al. (Reference Osés, Wood, Romero, Prado, Bidola, Herzen, Pfeiffer, Stampar and Pacheco2022) showed carbonate cements in close association with Corumbella.
6. Conclusions
The composition and morphology of the biomineralized metazoan Corumbella in the Tamengo Fm. created calcareous, sand-sized sediment in fine siliciclastic beds that in turn added sedimentary complexity. This led to a new interpretation for Corumbella depositional settings. Diverse Phanerozoic-style sedimentary features would have hardly been recorded in fine siliciclastic facies without the sand-sized clasts. The shallow marine clastic factory, therefore, underwent this step change in the Late Ediacaran, far earlier than previously thought, with the expansion of biomineralizers to shallow siliciclastic settings. The palaeoenvironmental and palaeobiogeographic distribution of Corumbella therefore favours this metazoan as a sedimentary indicator in Late Ediacaran fine siliciclastic facies. Further research could shed light on the geochemical impacts of bioclastic input in siliciclastic sediments in the Late Ediacaran.
Acknowledgements
GLO thanks the Coordination for the improvement of Higher Education Personnel – Brazil (CAPES) – Financing Code 001. GLO thanks the São Paulo research Foundation (FAPESP) for the grants #2017/21584-1, #2021/07007-7, #2016/06114-6, #2022/11586-5, #2023/04501-6, #2022/06485-5, and #2023/14250-0, and the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) grant #141115/2017-3. GLO thanks the Program of Post-Graduation in Ecology and Natural Resources (São Carlos Federal University), the Program of Post-doctorate and Pesquisador Colaborador, Institute of Physics, University of São Paulo. RW thanks the UKRI NERC Grant NE/T008458/1. We thank the Brazilian Synchrotron Light Laboratory (LNLS/CNPEM) for support during SR-microXRF analyses (proposals 20170831 and 20180422), and the Laboratory of Materials Analysis by ionic beams (LAMFI), Institute of Physics, University of São Paulo. We thank Bruno Becker-Kerber, Esthella Silva, Thibault Le Guelvout and Alessandro Marques for fieldwork support. We thank technical support from Juliana Leme, Ivone Gonzales, Sonia Alonso, Osmar Bagnato, Daniel Kakizaki, Cleber Diniz, Luiz Anelli, Renato Carvalho, Gergely Szabo, Renato Moraes, Amy Shore, John Craven, Douglas Galante, Gustavo Prado, Carlos Pérez and Isaac Sayeg. We thank Júlia d’Oliveira for the illustrations.
Competing interests
The author(s) declare none.