Cretaceous spreite-dominated ichnoassemblages in Antarctica

Abstract

This study investigates ichnoassemblages characterized by spreite trace fossils from the Upper Cretaceous Snow Hill Island Formation on Vega Island, Antarctica. The succession reveals alternating heterolithic beds of sandy siltstones to very fine- to fine-grained sandstones, suggestive of a deltaic depositional setting influenced by fluctuating energy conditions. The dominance of spreite structures, such as Paradictyodora antarctica and Euflabella, suggests the prevalence of a colonization window for deposit- or detritus-feeding activity. Notably, the prevalence of Euflabella towards the upper levels of the succession, in more proximal and impacted depositional setting, suggests an adaptive strategy for coping with varying food resources, corroborating its role as a trophic generalist. Additionally, the absence of distinct shifts between fully marine and stressed ichnofauna challenges a clear assignation to the Phycosiphon or Rosselia ichnofacies, although in vertical succession the Euflabella dominance in upper levels is related to a lower diverse ichnoassemblage than Paradictyodora in lower levels, suggesting some freshwater impact in the top. The occurrence of horizontal and vertical sessile deposit feeding structures would align with the Rosselia ichnofacies definition, but the presence of spreite burrows and high ichnodisparity better fits into the Cruziana ichnofacies. These spreite-dominated ichonoassemblages are here attributed to Cruziana ichnofacies.

Key words Vega Island; Cape Lamb; Snow Hill Island Formation; Paradictyodora; Euflabella

INTRODUCTION

Trace fossils are widely used for paleoenvironmental reconstructions, mainly after the seminal works of Seilacher (1953, 1967),nwhen the concept of ichnofacies emerged. Since then, this paradigm has undergone successive modifications by subsequent authors, resulting in new ichnofacies and ongoing debates concerning their characterization (e.g., Buatois & Mángano 2011, Knaust & Bromley 2012, MacEachern & Bann 2020). Currently, trace fossils have been recognized in almost all depositional settings after the origin of metazoans and its distribution is a guide to interpret paleoecological parameters and paleodepositional scenarios (Mángano & Buatois 2016, Sedorko & Francischini 2020). In soft ground shallow marine depositional setting, four ichnofacies have been described: the Skolithos, the Phycosiphon, the Rosselia, and the Cruziana ichnofacies (e.g., MacEachern & Bann 2020).

The Cruziana ichnofacies is referred to as “Seilacherian or archetypical” because it was proposed with the concept of ichnofacies. It is widely recognized in shallow marine settings, above a storm wave base level, during the Phanerozoic, with a dominance of horizontal deposit and detritus feeding trace fossil, associated with variable ethological categories, high ichnodiversity, and high abundance (MacEachern & Pemberton 1992). It can be argued that the Cruziana ichnofacies, as well the Skolithos ichnofacies, are the most investigated and recurrent ichnofacies in the trace fossil literature (MacEachern et al. 2010, 2012).

The Cruziana ichnofacies in fact includes several variations, which has allowed for the identification of a proximal, archetypal, distal, and depauperated ichnoassemblages in this ichnofacies (MacEachern et al. 2005, Sedorko et al. 2018a, 2019, 2021). The boundaries between these variations are not clear, and new ichnofacies were suggested for assemblages that deviate from the archetypal model (e.g., Phycosiphon and Rosselia ichnofacies, MacEachern & Bann 2020). Ichnoassemblages dominated by spreite structures such as Zoophycos, are often attributed to the Zoophycos ichnofacies (Buatois & Mángano 2011), even though the ichnogenus Zoophycos can be also a component of the Cruziana ichnofacies, especially for Paleozoic records (Zhang et al. 2015, Sedorko et al. 2018b).

Recently, two ichnofacies were proposed for deltaic settings, the Rosselia and the Phycosiphon ichnofacies (MacEachern & Bann 2020). The Rosselia ichnofacies was proposed for sandy delta-front environments, characterized by deposit-feeding behaviors and structures that reflect high sedimentation rates and sporadic deposition. These include equilibration with spreite and escape traces, with occasional marine ichnogenera indicating shifts between stable and stressed conditions. In contrast, the Phycosiphon ichnofacies was proposed for muddy prodelta settings, marked by diverse marine trace fossils. These are interspersed with low-diversity traces, reflecting short-term alternations between stable marine conditions and physico-chemically stressed environments.

Under the Brazilian Antarctic Program (PROANTAR, Simões et al. 2023), the Project PALEOANTAR has carried out fieldwork on the Antarctic Peninsula, obtaining results regarding fossil vertebrates (e.g., Kellner et al. 2019, Souza et al. 2023), including paleohistological analyses (e.g., Brum et al. 2023), invertebrates (e.g., Pinheiro et al. 2020), fossil plants (e.g., Lima et al. 2021, Santos Filho et al. 2023), and microfossils (e.g., Santos et al. 2022). Here, the first contribution focused on trace fossils is presented, opening a new research line of this project. In this study, ichnoassemblages dominated by spreite trace fossils from a Late Cretaceous succession in the Antarctic Peninsula are analyzed, discussing implications for ichnofacies attribution. We interpret the significance of the dominance of spreite structures in the Snow Hill Island Formation on Vega Island based on an integrated sedimentological and ichnological analyses.

Geological Setting

The studied area is located in the Antarctic Peninsula (Fig. 1), specifically in the James Ross Basin (JRB) which is part of the larger Larsen Basin (Macdonald et al. 1988). The basin evolved during the Cretaceous and Paleogene due to southeast-directed subduction of the proto-Pacific Plate (Hathway 2000), resulting in the deposition of approximately 7,000 m of Cretaceous–Paleogene clastic strata (Rinaldi et al. 1978, Olivero et al. 1986, Pirrie 1989, Crame et al. 1991, Pirrie et al. 1991, Olivero 2012, Crame 2019). These strata are divided into three major depositional cycles: the Gustav Group, the Marambio Group, and the Seymour Island Group.

Figure 1
Location of the studied area. a. Position of the studied area in the Antarctic Peninsula. b. Location of Vega Island. c. Cropping out lithological units in part of James Ross Island and Vega Island; the three profiles, part of the composite sections, were located at 63°54’15.6” S 57°35’47.5”W; 63°53’10.0” S 57°33’49.22”W; and 63°53’07.5” S 57°32’27.8”W (orange arrows). Adapted from Pinheiro et al. (2020).

The Gustav Group represents deep marine facies, succeeded by the shallowing up settings of the Marambio Group, likely associated with basin filling, uplift, and sea-level changes during the Late Cretaceous and early to middle Paleogene (Hathway 2000). The depositional environment extended across a broad shelf, influenced significantly by sea level fluctuations linked to third-order eustatic sea level cycles (Olivero 2012).

The Marambio Group crops out on Vega Island, where the Cape Lamb Member of the Snow Hill Island Formation (SHIF) occurs. This unit is characterized by grayish-green massive, fine-grained muddy sandstones to sandy mudstones with carbonate concretions. Planar lamination is only visible within concretions, which also enhance the visibility of trace fossils (Roberts et al. 2014). The SHIF is overlain by the López de Bertodano Formation. This study focuses on the trace fossil assemblage of SHIF on Vega Island.

MATERIALS AND METHODS

The data acquisition considered an integrated ichno-sedimentological analysis. Trace fossils were photographed and described considering the level of occurrence and associated sedimentary facies, which also considered texture, physical sedimentary structures, composition, relations of contact, and macrofossil content. Three sedimentologic logs with trace fossil occurrences were prospected, resulting in a composite geological succession for the SHIF in the studied area (Fig. 2).

Figure 2
Facies and trace fossil distribution in studied succession. a. Geological composite log section with levels of trace fossil occurrences (Ch=Chondrites, Di=Diplocraterion, Eu=Euflabela, Ne=Nereites, Op=Ophiomorpha, Pa=Palaeophycus, Pr=Paradictyodora, Ro=Rosselia, Sk=Skolithos, Te=Teichichnus); in green the lower unit, mostly representing transitional offshore settings, and in light pink the upper unit mostly representing a prodelta reaching transitional offshore zone. b. Sandy siltstone typical of Snow Hill Island Formation interbedded in the base by a fine-grained sandstone bed. c. Level with concretions within a siltstone bed. d. Pebbles in the silty matrix. e, f. Sandstone beds with spreite-burrows, such as Teichichnus (Te) and Rhizocorallium (Rh). g. In situ Pinna sp. in a sandstone bed. Scale bar = 2cm.

The trace fossil analysis considered ichnotaxonomic assignment, the tier structure, ichnodiversity, and ichnodisparity for the characterization of the ichnoassemblages (based on the categories of Buatois et al. 2017). Also, the amount of bioturbation (AB) was estimated on a linear scale (Knaust 2012), ranging from 1 (0-20%) to 5 (80-100%).

RESULTS

Three geological successions were examined, resulting in a composite section of the SHIF in the studied area (Figs. 1, 2a). It is dominated by sandy siltstone interbedded with very fine-grained sandstone (Fig. 2b) and is abundant in carbonate concretions that are frequently fossiliferous (Fig. 2c). The succession shows an upward increase in the occurrence of lenticular beds of sandstone, occasionally with basal erosive contact and containing pebbles (Fig. 2a, d), often displaying hummocky cross-stratification and spreite burrows (Fig. 2e, f). These sandstones are also fossiliferous, including bivalves among other groups (Fig. 2g). In weathered outcrops, the color tends to be rusty-brown, while in fresh expositions it appears gray. The mudstone facies typically exhibit thinly laminated beds with abundant mica flakes, occasionally preserving thin lenses of rippled or parallel-laminated sandstones. Conversely, the sandstone facies are characterized by massive or low-angle cross-stratified beds, reaching thicknesses of up to 60 cm. Climbing ripples and wave ripples are less common, but present in some sandstone beds, being more frequent to the top. The cyclic package begins with a thin, highly bioturbated siltstone, locally with lenses of wave-rippled sandstone, overlayed by lenticular sandstone locally bearing hummocky cross-stratification.

The poorly consolidated sandy siltstones are often bioturbated, but it is difficult to specifically identify the trace fossils due to the highly fractured sediment. When trace fossils are visible, they appear as simple vertical or horizontal burrows, which can be tentatively identified as Skolithos and Palaeophycus, respectively. In these beds, the siltstone occurs as moderately bioturbated, with AB = 3. Concretions are common in these siltstones, and, in addition to several macroinvertebrates, also preserve trace fossils.

Palaeophycus and Skolithos occur in the lower part of Cape Lamb Member associated with simple horizontal spreite structures, identified as Teichichnus rectus (Fig. 3a). Very small, ramified vertical to oblique structures, identified as Chondrites isp. (Fig. 3b) and “U”-shaped vertical structures bearing protusive spreite, identified as Diplocraterion can occur associated with whitish, very small meandering burrows with a lining identified as Macaronichnus (Fig. 3c, d). U-shaped horizontal burrows are identified as Rhizocorallium commune (Fig. 3e). Less common are vertical burrows with a pelleted wall, identified as Ophiomorpha (Fig. 3f), and concentrically filled vertical burrows, identified as Rosselia. This interval is defined as Paradictyodora-dominated (Fig. 2) because of the dominance of burrows bearing a vertical spreite structure with upward enlargement and prismatic-to-conical shape made of subvertical folded laminae, identified as Paradictyodora antarctica (Fig. 3g). These traces occur dominantly in moderated bioturbation amount (AB = 3), but can locally reach high intensities (AB = 4-5).

Figure 3
Trace fossils from low interval in Vega Island. a. Teichichnus rectus (Te) in vertical view. b. Chondrites isp. (Ch) in an oblique view. c. Diplocraterion isp. (Di) in vertical view. d. Diplocraterion isp. (Di) and Macaronichnus (Ma) in vertical view. e. Rhizocorallium comune (Rh) in horizontal view; f. Ophiomorpha (Op) and Skolithos (Sk) in vertical view; g. Paradictyodora antarctica (Pr) in vertical view; h. Palaeophycus striatus (Pa) in horizontal view. i. Nereites biserialis (Nb) as concave epirelief (bedding plane view). Scale bar = 2 cm.

In the basal Paradictyodora-dominated interval, ten ichnogenera are present and correspondingly categories of architectural designs (sensu Buatois et al. 2017). In these architectural categories, Chondrites is defined in the category of “a burrow with shaft or bunch with downwards radiating probes”; Diplocraterion as “a vertical single U- or Y-shaped burrows”; Ophiomorpha “as maze and boxwork burrows”; Macaronichnus as “a simple actively filled (massive) horizontal to oblique structure”; Palaeophycus as “a passively filled horizontal burrow”; Paradictyodora as “a burrow with complex vertically oriented spreite”; Rhizocorallium as “a burrow with horizontal spreite”; Rosselia as “a vertical concentrically filled burrow”; Skolithos as “a vertical unbranched burrow”; and Teichichnus as “a horizontal burrow with simple vertically oriented spreite”.

In the upper part of the succession, in the informal “upper unit of Camp Lamb Member”, fine-grained sandstones are more frequent. In these sandstones, simple horizontal burrows, identified as Palaeophycus isp. and Palaeophycus striatus (Fig. 3h) are common. It also occurs an alternating sequence of paired depressions flanked on either side by a diffuse, irregular to lobed surrounding area, identified as Nereites biserialis (Fig. 3i). In addition, there are common Paradictyodora, but this interval is dominated by horizontal to oblique burrows with single or multiple spreite, including laminae with curved or sigmoidal backfill connected with an oblique to subvertical tube at the base, identified as Euflabella. Two morphologies can be recognized for Euflabella: one presents a single spreite, typically consisting of folded, subhorizontal to slightly oblique laminae, diverging from a common tube and expanding laterally, referred to as Euflabella singulari (Fig. 4a-b), and the other shows a radial arrangement of spreite, with single or folded laminae extending in subhorizontal to oblique directions, identified as Euflabella radiata (Fig. 4c-d). Associated to Euflabella, it is also present vertical structures with relatively long spreite, identified as Teichichnus (Fig. 4e-f). Paradictyodora antarctica is subordinated (Fig. 4g) while Euflabella and Teichichnus dominate in these sandstones from the upper informal member (Fig. 2). Where Euflabella is present, the amount of bioturbation is usually high (AB = 4-5) (Fig. 4h).

Figure 4
Spreite-dominated beds in upper interval of studied section. a-b. Euflabella singulari (Es) in bedding plane view. c-d. Euflabella radiata (Er) in bedding plane to slight oblique view. e-f. Teichichnus rectus (Te) in vertical view. g. Paradictyodora antarctica (Pr) in horizontal views. h. High bioturbated level with Euflabella singulari (Es) in bedding plane view. Scale bar = 2 cm.

In the basal levels of the Euflabella-dominated interval, five ichnogenera are present and represent four categories of architectural designs (as defined by Buatois et al. 2017). Both Euflabella and Paradictyodora are in the category of “burrows with complex vertically oriented spreite”; Teichichnus is “a horizontal burrow with simple vertically oriented spreite”; Nereites is “a complex actively filled horizontal structure”; and Palaeophycus is “a passively filled horizontal burrow”.

DISCUSSION

The composite section is dominated by sandy siltstone interbedded with very fine- to fine-grained sandstones, locally with hummocky cross-stratification in the upper part of the section. The coarsening upward arrangement and lenticular bodies suggest a lobate form typical of deltaic environments (Macellari 1988, Pirrie 1989, Scasso et al. 1991). The erosion surface at the base of sandstone packages (Fig. 2b, e) along with the presence of wave ripples and their distribution towards the top of the succession, indicates shifting and abandonment of active mouth bar deposition sites. During abandonment phases, wave action reworked previous sandstone beds, forming erosion surfaces and basal sandstone layers (Bhattacharya 2006). The lack of fluvial delta plain deposits between successive packages suggests a permanent subaqueous deposition (Bhattacharya 2006). Thus, the alternating beds of bioturbated siltstones and sandstones suggests the interplay of depositional processes in a deltaic setting, as inferred from the cyclical stacking pattern observed within the formation (Olivero et al. 2008). Additionally, there is a rich fossil assemblage within concretionary horizons indicating marine conditions, including ammonites, bivalves, gastropods, echinoderms, and decapod crustaceans (Roberts et al. 2014). Noteworthy among these are the ammonites Gunnarites antarcticus and bivalves such as Lahillia, Modiolus, and Pinna (Fig. 2g). Pinna is very common in these beds (Pirrie et al. 1991, Olivero 2012).

In the basal informal unit, a transitional offshore setting in a prodelta depositional scenario thus predominates, with a lesser influence of sandy bodies (Fig. 5). In these beds, the ichnoassemblage is dominated by the spreite burrows Paradictyodora antarctica and Teichichnus rectus, with the subordinated presence of Chondrites, Diplocraterion, Macaronichnus, Ophiomorpha, Palaeophycus, Rosselia, Rhizocorallium, and Skolithos. The mixture of shallow-, mid-, and deep-tier trace fossils, associated with different ethological categories supports the assignment for this ichnoassemblage to the Cruziana ichnofacies, as has been previously done (e.g., Olivero & Cabrera 2013). The presence of ten categories of architectural designs (sensu Buatois et al. 2017), characterizing a high ichnodisparity, corroborates this attribution.

Figure 5
Reconstruction of depositional environment represented in the Snow Hill Island Formation by dominance of Paradictyodora in the base and Euflabella to the top of the unit, composing a Cruziana ichnofacies. Paradictyodora morphology inspired in Olivero et al. (2004). FWWB = Fair-weather wave base; SWB = Storm wave base.

The overlying beds of the informal upper unit of Cape Lamb Member, in contrast, are dominated by sandy bodies representing a prodelta to delta front, where mainly characterized by Euflabella and a lower diverse ichnoassemblage (Fig. 5). An unconformity was proposed for this interval, placed in a conglomerate bed that would separate the informal lower and upper units (Olivero 2012). In these layers, Teichichnus, Nereites, Palaeophycus, and Paradictyodora are also present. The behavior associated to biserial arrangement in Nereites biserialis is unexplained. It likely aids in feeding in nutrient-rich sediments, possibly requiring extra space in the tunnel for waste disposal. Sideways movement may also aid in respiration (Olivero & Cabrera 2016). These ichnogenera are here representing four categories of architectural designs (sensu Buatois et al. 2017), what might be assumed as depauperated expressions of the Cruziana ichnofacies (MacEachern et al. 2010).

Euflabella and Paradictyodora are uncommon trace fossils, being reported for few localities. In Tierra del Fuego, Euflabella singularis and E. radiata are prevalent in shelfal fine-grained silty sandstones, indicative of relatively low-energy conditions within the depositional environment. Particularly, E. singularis dominates in organic-rich, storm-influenced to offshore deposits (Olivero & Cabrera 2013). On the other hand, E. radiata is prominent in fine-grained sandstones interpreted as event beds in a shelf setting or at the base of prodelta parasequences, occasionally found in prodelta thin-bedded alternations or distal subtidal settings (Olivero & Cabrera 2013). Conversely, Paradictyodora in Tierra del Fuego mostly occurs in fodinichnia-dominated ichnocoenosis within basinal fan delta systems characterized by sporadic low energy conditions and low sedimentation rates, possibly featuring disaerobic or poorly oxygenated bottoms. In the Santa Marta Formation, however, this ichnogenus occurs in deposits interpreted as delta slope and distal subaqueous delta platform (Olivero et al. 2004).

Recently, two new ichnofacies were proposed for delta settings, the Phycosiphon and the Rosselia ichnofacies (MacEachern & Bann 2020). The Phycosiphon ichnofacies is characterized by diverse marine trace-fossil associations intercalated with low-diversity suites, reflecting recurrent shifts between stable marine conditions and physico-chemically stressed environments triggered by fluctuations in fluvial influence and other shallow marine agents (wave, storms, tides). The Rosselia ichnofacies also reflects recurrent alternations between stable marine conditions and physico-chemically stressed environments, but with a prevalence of deposit feeding and equilibrium ethologic categories in sandy substrates (MacEachern & Bann 2020).

In the studied succession, it is not clear if there is an alternation between fully marine and impoverished ichnofauna, as expected for the Phycosiphon and Rosselia ichnofacies (MacEachern & Bann 2020), probably because the visualization of trace fossils was not favored in the siltstone facies (e.g., Savrda 2007). The overall dominance of spreite burrows in the Cape Lamb Member at Vega Island, however, suggests a colonization window from detritus- or deposit-feeding activity in this deltaic paleoenvironment, mainly for upper beds dominated by Euflabella, where suspension-feeding habits are absent. This scenario could fit in the definition of the Rosselia ichnofacies, however, the absence of fugichnia/equilibrichnia structures precludes this attribution. Additionally, the dominance of spreite burrows is not part of the definition of Rosselia ichnofacies (MacEachern & Bann 2020).

Another point is that the vertical replacement observed in the prograding section, transitioning from a Paradictyodora-dominated to an Euflabella-dominated succession, appears to be driven by the slight differing ecological strategies employed by these tracemakers. Paradictyodora likely dominates in a more stable, deeper-water prodeltaic setting where sedimentation rates were relatively constant. Euflabella seems to be exhibiting a more opportunistic behavior suited to exploit proximal, impacted deltaic settings. Euflabella, with its diverse spreite burrows extending in subhorizontal to oblique directions, appears to have adapted to the dynamic conditions of shallower, proximal deltaic environments, where fluctuating river discharge and sediment supply create a patchy, resource-rich landscape. Furthermore, the increase in Euflabella occurrences towards the upper levels of the Cape Lamb Member supports the interpretation that Euflabella represents a trophic generalist tracemaker, adapted for deposit- or detritus-feeding activity. This adaptation allows it to prospect the substrate amidst varying food resources (Olivero & Cabrera 2013).

CONCLUSIONS

The proposal of the Rosselia and Phycosiphon ichnofacies reflects ongoing efforts to refine our understanding of deltaic settings and their trace fossil associations. In the Upper Cretaceous succession of the Antarctic Peninsula’s Snow Hill Island Formation, the dominance of spreite burrows suggest the prevalence of a colonization window for detritus- or deposit-feeding activity, but it is part of the Cruziana ichnofacies. The vertical replacement in a prograding section, from a Paradictyodora dominated to an Euflabella dominated succession seems to be related to the more opportunistic behavior reflected by the latter, which would better exploit proximal, impacted deltaic paleoenvironments.

ACKNOWLEDGMENTS

DS thanks the Alexander von Humboldt-Stiftung for the postdoc fellowship (CAPES/Humboldt Program nº 14/2022 - process nº 23038.004870/2021-69), and the Conselho Nacional de Desenvolvimento Científico e Tecnológico (grant CNPq 306493/2022-5). The Programa Antártico Brasileiro (PROANTAR – OPERANTAR XLII) is being thanked for the operational facilities that allowed the fieldtrip in Vega Island. This study is a contribution to the Project PALEOANTAR “Paleobiologia e Paleogeografia do Gondwana Sul: Inter-relações entre Antártica e América do Sul”, financed by CNPq (PROANTAR #442677/2018-9 and #440902/2023-1 to AWAK). Authors also want to thank CNPq for founding INCT PALEOVERT #406902/2022-4; #308707/2023-0 to AWAK; #309245/2023-0 to JMS. This research is also partially founded by Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) #E-26/201.095/2022 to AWAK and #E-26/210.066/2023; E-26/210.734/2024 to JMS. We thank for the help in fieldwork of Joseline Manfroi, Paula Sucerquia, Taluany Nascimento, Thales A.R. Schaefer, Millena L. Pitanguy, and Denni Moraes. The authors thank Dirk Knaust and an anonymous reviewer which suggestions improved the quality of this manuscript.

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Publication Dates

• Publication in this collection
  13 Dec 2024
• Date of issue
  2024

History

• Received
  11 June 2024
• Accepted
  16 Oct 2024