Taxis behaviour of burrowing organisms recorded in an Ediacaran trace fossil from Ukraine

Journal Pre-proof Taxis behaviour of burrowing organisms recorded in an Ediacaran trace fossil from Ukraine Alfred Uchman, Andrej Martyshyn PII:

S0031-0182(19)30515-2

DOI:

https://doi.org/10.1016/j.palaeo.2019.109441

Reference:

PALAEO 109441

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Palaeogeography, Palaeoclimatology, Palaeoecology

Received Date: 28 May 2019 Revised Date:

30 October 2019

Accepted Date: 31 October 2019

Please cite this article as: Uchman, A., Martyshyn, A., Taxis behaviour of burrowing organisms recorded in an Ediacaran trace fossil from Ukraine, Palaeogeography, Palaeoclimatology, Palaeoecology (2019), doi: https://doi.org/10.1016/j.palaeo.2019.109441. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2019 Published by Elsevier B.V.

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Taxis behaviour of burrowing organisms recorded in an Ediacaran trace fossil from Ukraine

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Alfred Uchman a, *, Andrej Martyshyn b

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a

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University, Gronostajowa 3a, PL-30-387 Kraków, Poland. E-mail: [email protected]

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b

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Kyiv 03022, Ukraine. E-mail: [email protected]

Faculty of Geography and Geology, Institute of Geological Sciences, Jagiellonian

Institute of Geology, Taras Shevchenko National University of Kyiv, 90 Vasylkivska Str.,

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ABSTRACT

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The trace fossil Archaeonassa cf. fossulata Fenton and Fenton is a bilobate, slightly

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undulating, corrugated ridge on upper bedding plains in the Ediacaran shallow marine

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siltstones of Ukraine which are no younger than 557 Ma. It was produced partly under

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microbial mats. This trace fossil shows strong orientation within a sector of 20º–40º, which is

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approximately perpendicular to the expected shoreline. The orientation points to ability of

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some Ediacaran burrowing organisms to taxis during early evolutional stages of bilaterians in

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response to physical or chemical stimuli, probably according to direction of tides. Undulations

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of A. cf. fossulata can be regarded as a record of the in-and-out burrowing in response to

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diurnal cycles of oxygen production within microbial mats. Probably, this is one of the oldest

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examples of such behaviour. In general, Archaeonassa is considered as a relatively common

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trace fossil in Upper Ediacaran deposits worldwide, especially in shallow marine deposits in

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the time interval from 560 to 550 Ma.

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Keywords: Proterozoic; ichnofossils; microbial mats; ichnotaxonomy; ichnology; evolution.

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1. Introduction

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Taxis, which is understood as a movement of organisms in response to a stimulus, can

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be observed in the geological record as oriented trace fossils, which are an evidence of in situ

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life activity of organisms. Orientation of trace fossils shows that their tracemakers were

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physically or chemically stimulated to a certain behaviour having some directional aspect in

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the space. One can ask when the ability to taxis started in the Earth history? Good examples

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of oriented trace fossils are known in shallow-marine deposits since the Cambrian through the

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whole Phanerozoic (e.g., Salter, 1856; Seilacher, 1953, 1959; Pickerill, 1995; Bromley et al.,

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2009; Pandey et al., 2014; Boyer and Mitchell, 2017; Uchman et al., 2016), rarely in deep-sea

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deposits (Simpson, 1970) and in non-marine deposits at least since the Triassic (e.g., Pollard,

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1985). Orientation of Recent traces (lebensspuren) is documented in shallow-marine or

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marginal sediments (Hohenegger and Pervesler, 1985; Pervesler and Hohenegger, 2006;

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Uchman and Pervesler, 2006) and rarely on hard rocky substrates (Cachão et al., 2011).

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It appears that already some Ediacaran organisms are able to behave according to the

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taxis. In this paper, an oriented trace fossil (Archaeonassa) from Ediacaran shallow marine

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deposits of Ukraine is presented. It is a record of taxis, probably the oldest one, not on the

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sediment surface, but subsurface, because the tracemaker produced this trace under microbial

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mats. The aim of this paper is its presentation and interpretation. Moreover, a review of

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Ediacaran occurrences of Archaeonassa is provided.

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2. Material and methods

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The study area is located in the southwest surrounding of the Ukrainian Shield in the so-called Volyn-Podillya-Moldavia Basin. During the Ediacaran Period, it was a rifted,

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passive continental margin subjected to tectonic extension (Poprawa et al., 2018). The studied

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trace fossil occurs in the Novodnistrovs’k Quarry (Fig. 1) in the Mohyliv-Podilsky Group,

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which is subdivided into the Mohyliv Formation (Mogilev Series in the Russian literature) in

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the lower part and the Yaryshiv (Yaryshev) Formation in the upper part (Martyshyn, 2012). In

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the bottom of the quarry, Lower Proterozoic granites and migmatites of the Ukrainian Shield

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are exposed. They are covered by thin, discontinuous layer (up to a meter) of basal

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conglomerates followed by the Lomoziv (Lomozov) Member, which is dominated by

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mudstones and siltstones and represents the second sedimentary cycle in the Vendian of

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Podolia (Korenchuk and Ishchenko, 1981). It contains relatively common disc-like fossils of

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the Petalonamae group (Dzik and Martyshyn, 2017) and rare dickinsoniids (Dzik and

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Martyshyn, 2015). Above, the Yampil (Yampol) Member occurs. It is composed of thick and

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very thick, partly cross-bedded sandstone beds (up to 9 m thick) with a package (up to 3 m

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thick) of thin sandstone and siltstone beds at the top. The trace fossil studied is present in this

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package, close to the top of the Mohyliv Formation. Grazhdankin et al. (2011) and

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Grazhdankin (2014) dated the base of the overlying Yarishiv Formation to 553 Ma.

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According to new dating of bentonite layers, the layer B1, located just above the trace fossil

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horizon investigated is 556.78 ±0.18 Ma old (Soldatenko et al., 2019). Thus, the trace fossil is

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not younger than this date (roughly 557 Ma). The overlying Lyadova (Lyadov) Member of the

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Yaryshiv (Yaryshev) Formation consists of siltstone at the base (up to 2 m) followed by

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mudstone with intercalations of thin siltstone or sandstone beds (up to 14 m).

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The described trace fossil has been collected in the uppermost part of the Yampil

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Member (Fig. 1), about 1.45 and 2.75–3.00 m above the thick sandstone beds in the northern

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margin of the quarry (48º39.378'N, 027º27.939'E). Twelve collected slabs of grey laminated

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siltstone have been analyzed in detail. The surface of slab shows uneven morphology typical

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of fossil microbial mats. The siltstone contain poorly sorted, mostly quartz silt grains.

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Lamination is mostly parallel, uneven, or with isolated, small ripple lamination. The laminae

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differ in subtle grain sizes and packing. Thin, darker laminae mark the microbial mats (Fig.

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2A–D).

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Orientation of the trace fossil has been measured in the laboratory because only small

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surfaces were available in situ in the field. As a part of specimens of the trace fossil shows a

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winding course, orientation of particular segments are measured. The prevailing direction was

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adjusted to the dominant direction of the trace fossil measured in the field and determined as

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290º–300º.

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3. The oriented trace fossil

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3.1. Systematic description

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Ichnogenus Archaeonassa Fenton and Fenton, 1937

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Type ichnospecies. Archaeonassa fossulata Fenton and Fenton, 1937.

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Diagnosis. Raised, narrow traces, straight to sinuous or gently meandering, having a median

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groove flanked by rounded ridges (Yohelson and Fedonkin, 1997).

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Remarks. Archaeonassa fossulata derives from the early middle Cambrian of Alberta,

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Canada (Fenton and Fenton, 1937; Yochelson and Fedonkin, 1997). Archaeonassa is

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interpreted as a crawling trail produced of mostly by gastropods (Fenton and Fenton 1937;

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Buckman, 1994; Stanley and Feldmann, 1998) or crustaceans (Yochelson and Fedonkin,

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1997; Mángano and Buatois, 2003). Matz et al. (2008) presented recent, short, bilobate traces

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produced by a giant protist Groomia on the deep sea floor and suggested that production of

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bilobate traces does not necessarily needs bilaterian organisms. However, the traces are short,

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produced on the surface, where they can be eroded or smoothed after production (Gheling and

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Droser, 2009). Production of A. cf. fossulata under the microbial mats excludes protists as its

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trace maker. Bilobate trails in the Ediacaran Period can be produced also less advanced

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bilaterians similar to modern ceriantharian anemones and flatworms, which excrete mucus

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tracts trapping sediments; their traces show transitions from bilobate to unilobate morphology

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and surface corrugations (Collins net al., 2000). Retallack (2013) regarded a possibility that

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Ediacaran Archaeonassa was produced on land by “metazoan slugs or worms after rainstorms

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on land, but terrestrial habitats also open the possibility that these trails were created by slug-

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aggregating phases of slime moulds”, similarly to the interpretation of the trace fossil

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Myxomitodes stirlingensis from the Palaeoproterozoic of the south-western Australia (see

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Bengston et al., 2007). Such a possibility is also excluded in the presented case of A. cf.

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fossulata, because it was produced under the microbial mats and in association with the

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marine disc-shaped body fossil similar to Charniodiscus. Moreover, M. stirlingensis is a

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hypichnial trace showing local divergence of the lobes, and A. cf. fossulata is an epichnial

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form, without divergence of the lobes.

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Archaeonassa cf. fossulata Fenton and Fenton, 1937

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Figs. 3–6

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Material. Eleven slabs collected are housed in the Geological Museum of the National Taras

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Shevchenko University of Kyiv, Ukraine (KSU 17p197 – 17p200, KSU17p916 – 17p923).

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Two slabs (INGUJ254P1, 2) are housed in the Nature Education Centre (CEP) of the

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Jagiellonian University – Museum of Geology in Kraków, Poland.

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Description. Epichnial, discontinuous bilobate, slightly undulating ridge, mostly 3–4 mm

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wide, rarely 2–3 mm or 4–5 mm, divided by the central furrow. The width can change

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gradually within ridge by up to 20%. The ridge is wider in the elevated places than in

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depressed parts. The ridge became narrower on the ends where the ridge plunge into the bed.

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The ridge is straight, slightly curved or slightly winding. The turns are gentle, usually by 20º–

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40º. The ridge occurs in a slightly winding tract. Ridges in the tract are usually 3.5–17 mm

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long, rarely up to 55 mm long. The lobes are symmetrical (Fig. 3B) or asymmetrical (Figs.

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3D, F, 5B), corrugated, commonly covered by oblique or transverse discontinuities or bleb-

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like elevations. Some lobes look like composed of transverse pads (Fig. 3B). Lobes are

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usually wider than the median furrow. Locally, one lobe or even two lobes disappear (Fig.

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3C). In the latter case, only the median, shallow furrow is visible (Fig. 5C). Externally, the

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lobes transit to the bedding surface without any discontinuity, or rarely with some

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discontinuity, with a lobate margin (Fig. 5D). The highest part of the lobes is not in the

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middle but closer to the median furrows. The narrow median furrow of A. cf. fossulata

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represents the collapsed burrow roof, but the furrow is locally roofed and ridge become

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unilobed. The median furrow is V-shaped, usually narrower than individual lobe on the upper

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part. In rare cases, the furrow is locally roofed and the ridge became unilobed. Width and

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depth of the furrow may change along the ridge.

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Remarks. The trace fossil belongs to the ichnogenus Archaeonassa Fenton and Fenton, 1937,

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which is typified by A. fossulata Fenton and Fenton, 1937. It is diagnosed as “raised, narrow

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traces, straight to sinuous or gently meandering, having a median groove flanked by rounded

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ridges” (Yohelson and Fedonkin, 1997, p. 1213). The original material of A. fossulata derives

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from the middle Cambrian of British Columbia (Fenton and Fenton, 1937). Archaeonassa is

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interpreted as a crawling trail produced mostly by gastropods (Fenton and Fenton 1937;

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Buckman, 1994; Stanley and Feldmann, 1998) or crustaceans (Yochelson and Fedonkin,

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1997; Mángano and Buatois, 2003).

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Compare to the type material of Archaeonassa fossulata (see and Fenton, 1937;

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Yochelson and Fedonkin, 1997; Figs. 6–7), the described A. cf. fossulata is of a comparable

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size (3–4 mm), but it is less continuous, with a narrower median furrow (always V-shaped but

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U-shaped in the some traces of the type material) and with more corrugated, locally

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asymmetric and uneven lobes. The epichnial stretches of A. cf. fossulata are generally more

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elevated in respect to the surface of bed than in the type material of A. fossulata, which is

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more incised in the bed. It is possible that the differences resulted from pushing of sediment

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under the microbial mat and deformation (dragging) of the plastic microbial mat by the trace

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maker of A. cf. fossulata. Probably, burrows of the type material of A. fossulata from the

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Cambrian deposits were produced beyond microbial mats on the sediment surface and hence

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they are smoother. As the considerations on the similarities and differences are partly

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interpretative and ichnotaxa in general should be based on morphological differences

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(Bertling et al., 2006), determination of the described trace fossil in the open nomenclature as

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A. cf. fossulata is proposed as an appropriate solution at this stage of research.

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4. Discussion

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4.1. Environment and ethology

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The laminated silty sediments, with rough lamination surfaces, locally with the

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elephant skin structures, suggest intensive microbial activity in microbial (probably

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cyanobacterial) mats. The structures can be considered as an example of the microbially

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induced sedimentary structures (MISS; Noffke et al., 2001). Such structures are common in

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intertidal or shallow subtidal, low energy settings (see Noffke, 2010). In Ediacaran,

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Archaeonassa and similar traces commonly occur below MISS (Buatois & Mángano, 2012,

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2016; Chen et al., 2013; Meyer et al., 2014; Arrouy et al., 2016). The most abundant traces in

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Ediacaran strata belong to mat grazers, (Gehling, 1999; Seilacher et al., 2005; Jensen et al.,

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2006; Fedonkin et al., 2007; Buatois and Mángano 2012). A. cf. fossulata can be ascribed to

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this category. On several slabs, the petalomean Charniodiscus-like discs are present (see Dzik

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and Martyshyn, 2017), including surfaces with the described trace fossil (Fig. 3A). Moreover,

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several oval, elliptical, or irregular, flat elevations can be observed in the trace-fossil baring

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surfaces (Figs, 5A, B, 8). Probably, they are “ghosts” of body fossils covered by the microbial

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mats. Nevertheless, any interactions between these structures and trace fossils are observed.

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This suggests that such places have not been nutritionally attractive for the trace maker.

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Morphology of Archaeonassa cf. fossulata suggests that the tracemaker moved under

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the microbial mat along a winding and undulating path (Fig. 4). Sediment was pushed away

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along and partly through the body digestive tract and formed the lobes. In elevations of the

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undulating path, the mat was locally snagged and ploughed and even some fragments of the

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mat was unwrapped outward (Fig. 5D). Collapsed and stretched mat behind the tracemaker

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body formed the median furrow. The change from bilobate to unilobate trail and surface

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corrugations are present in some modern traces of creeping ceriantharian anemones and

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flatworms, which produce mucus tracts trapping sediments; they are invoked as analogues of

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some Ediacaran traces (Collins net al., 2000). The presence of similar morphological features

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in A. cf. fossulata may suggest that the trace maker produced a lot of mucus, but similar effect

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can be also caused by wet microbial mats.

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The trace maker fed on organic matter and appeared rhythmically on or close the surface when the trace was formed, alternatively with slightly deeper burrowing phase when

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the trace is not visible on the bedding plane. Probably, the rhythmicity can be coincided with

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tides and related to a more effective respiration of oxygen in shallower level within sediment

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and microbial mats. Water containing oxygen infiltrated the mats and its percolation through

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the mat enabled respiration. Xiao et al. (2019) presented a tubular trace fossil Yichnus levis

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Xiao et al., 2019 from the Late Ediacaran of China (less than 551 Ma), which shows regularly

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undulating course in respect to the bedding. This was interpreted as a record of surfing in and

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on microbial mats (“in-and-out behaviour”), foremost for oxygen, less for food, in response to

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diurnal rhythms of oxygen production within the microbial mats (see also Meyer et al., 2014)

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and in the water column. Such rhythmicity with high oxygen content during the day and low

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oxygen content during the night is proven by experiments on modern microbial mats

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(Canfield and Des Marais, 1993; Gingras et al., 2011). The in-and-out burrowing behaviour is

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considered as an Ediacaran evolutionary innovation of bilaterians organisms (Xiao et al.,

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2019).. Other or additional possibility is that the tracemaker responded to diurnal temperature

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changes. Archaeonassa cf. fossulata can be regarded as an example of the in-and-out

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burrowing behaviour sensu Xiao et al. (2019) and probably one of the oldest example of its

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occurrence (no younger than 557 Ma), as the Chinse examples are younger than 551 Ma

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(Meyer et al., 2014; Xiao et al., 2019) and the examples from Namibia (Jensen et al., 2000;

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Jensen and Runnegar, 2005) are even more younger.

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4.2. Orientation

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The most intriguing is the strong orientation of Archaeonassa cf. fossulata (Fig. 9).

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Most of the ridges are confined to a sector which angle is 20º–40°. Only small percentage of

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traces is oriented beyond this range. One dominant direction is usually associated with one or

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two accessory directions deflected by 20º–40° from the dominant azimuth. The dominant

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direction is roughly coincided with the SE-NW direction (azimuth 290º–300º), generally

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perpendicularly to the expected shore of the main land masses located to the north, and

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parallel to the direction of tide currents.

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The described orientation suggests that the tracemakers followed water mases moved

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by tides, but no signs which would allow determination if it moved with or against the water

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flow. Whatever is the cause of the orientation, this is an evidence of taxis of an Ediacaran

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bilaterian organism. Rheotaxis in the Ediacaran epibenthic organism Parvancorina is

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presented by Paterson et al. (2017). However, this case in not obvious as body fossils can be

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oriented by current. The case of Archaeonassa cf. fossulata. shows that also some organisms

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burrowing under microbial mats were able to taxis already in the Ediacaran Period (in the case

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at least 557 Ma old) and that they were able to respond to physical or chemical stimuli related

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probably to tides. Trace fossil record of the taxis excludes non-behavioural orientation (e.g.

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by currents), which is possible in the case of body fossils.

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4.3. Archaeonassa in the Ediacaran Period

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Archaeonassa ranges since the Late Ediacaran Period since 565 Ma (Table 1; see also

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Grazhdankin, 2014; Muscente et al., 2019), and it continues through the Phanerozoic (e.g.,

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Buckman, 1994; Yochelson and Fedonkin, 1997; Stanley and Feldmann, 1998; Hanken et al.,

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2016; Wang et al., 2019). In the Ediacaran Period, it occurs in shallow and deep sea

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environments (Buatois and Mángano, 2016), but mostly in shallow marine settings on several

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palaeocontinents.

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Archaeonassa is most common in the second ichnozone (ca. 558–550 Ma) among the

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three Ediacaran ichnozones of Crimes (1987); for the concept of three Ediacaran ichnozones

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see also Jensen (2003) and Jensen et al. (2006), but the problem of dating and correlation of

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the ichnozones remains unsolved (McDonald et al., 2014). In the concept of two Ediacaran

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ichnozones, Archaeonassa is a characteristic component of the lower Ediacaran zone (560–

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550 Ma), together with other simple grazing trace fossils, such as Helminthoidichnites,

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Helminthopsis, Gordia, Epibaion and Kimberichnus (Shahkarami et al., 2017). The described

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occurrence of A. cf. fossulata fits to this zone. The time range of this zone roughly

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corresponds to the Belomorian (559–550 Ma) regional stage of the East European Platform,

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which beginning marks initiation of bioturbation in low-energy shelves, before expansion of

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burrowing organisms to higher energy environments after its end (see Grazhdankin, 2014).

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5. Conclusions

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The trace fossil Archaeonassa cf. fossulata from the Ediacaran Mohyliv Formation

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(Ukraine) shows strong orientation perpendicular to the expected shore, probably in response

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to tides. This points that some bilaterian organisms, no younger than 557 Ma, which burrowed

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at least partly under microbial mats were able to reacted to some physical or chemical stimuli

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in the same way. This is probably the oldest example of taxis among burrowing organisms.

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Moreover, A. cf. fossulata course undulates regularly in respect to the bedding plane and

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fossil microbial mats. This is interpreted as a record of the in-and-out burrowing in response

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to diurnal cycles of oxygen production within the microbial mats, being one of the oldest

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example of such behaviour. The ichnogenus Archaeonassa is a typical trace fossil of the Late

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Ediacaran Period worldwide, produced on the sediment surface or under microbial mats,

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especially in shallow-marine environments between 560 and 550 Ma.

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276 277

Acknowledgements

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Field work in Ukraine by A.U. was partly sponsored by the Jagiellonian University (DS

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funds). Mark Florence (Smithsonian National Museum of Natural History, Washington, D.C.,

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U.S.A.) provided a photograph of the holotype of Archaeonassa fossulata. Waldemar

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Obcowski (Institute of Geological Sciences, Jagiellonian University, Kraków) prepared the

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laser scan image of slab INGUJ254P1. Two anonymous reviewers provided critical remarks,

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which helped to improve the paper.

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479

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480

Figure captions

481 482

Fig. 1. Location map and stratigraphic column of the Ediacaran deposits in the study area.

483

Red arrows indicates positon of the trace fossils studied.

484

Fig. 2. Trace fossil bearing slabs in cross sections. A, B. Scan images of thin sections

485

perpendicular to the bedding planes. C. Ripple and parallel cross bedding. Position of

486

Archaeonassa cf. fossulata indicated by arrow (Ar). D. Parallel and ripple cross bedding.

487

Fig. 3. Archaeonassa cf. fossulata on upper surface of slab INGUJ254P1. A. General view of

488

the slab with indication of parts shown in B–F. Ch – Charniodiscus-like disc-shaped body

489

fossil; several, poorly preserved disc-shaped structures in other parts. B. A single burrow. C–

490

F. Details of A.

491

Fig. 4. Archaeonassa cf. fossulata on upper surface of other two slabs. A. General view of the

492

slab KSU 17p197 with indication of parts shown in B and C. B, C. Details of A. D. Fragment

493

of slab KSU 17p198. Numerous oriented Archaeonassa cf. fossulata together with many

494

small Charniodiscus sp.

495

Fig. 5. Archaeonassa cf. fossulata on upper surface of slab INGUJ254P2. A. General view of

496

the slab with indication of parts shown in B and C. B, C. Details of A. D. A. cf. fossulata on

497

upper surface of slab KSU 17p923.

498

Fig. 6. Comparison of Archaeonassa cf. fossulata (based on the investigated material from

499

Late Ediacaran of Ukraine) and the type material of Archaeonassa fossulata from the

500

Cambrian of Canada (for the latest see Figure 7).

501

Fig. 7. The holotype of the trace fossil Archaeonassa fossulata Fenton & Fenton, 1937, early

502

middle Cambrian, Mt. Whyte Formation, Alberta, Canada. Slab USNM No. 489678, formerly

503

Princeton University No. 46972, Smithsonian National Museum of Natural History,

504

Washington, D.C., U.S.A.

21

505

Fig. 8. Laser scanning image of surface of slab INGUJ254P2 showing several flat elevations,

506

which may be “ghosts” body fossils covered by the microbial mats.

507

Fig. 9. Orientation of Archaeonassa cf. fossulata in six slabs.

508

22

509

Table 1

510

Occurrences of Ediacaran trace fossils ascribed to Archaeonassa and their palaeoenvironment. Determination and reference Archaeonassa fossulata; Jensen, 2003; Jensen et al., 2005; Buatois and Mángano, 2016 ?Archaeonassa;

Formation and location

Age

Palaeoenvironment

Ediacara Member,

556–550

Shoreface to

Rawnsley Quartzite; Flinders Ranges, S Australia

upper offshore

Mistaken Point Formation; Newfoundland, Canada

565 My

Deep-water deposits

Unnamed;

Miette Group, Windermere

>542

Deep water

Hofmann et al., 1991; Hofmann and Mountjoy,

Supergroup; Rocky Mountains, British Columbia, Canada

Liu et al., 2014; see also Liu and McIlroy, 2015; see Buatois and Mángano, 2016

2010 Archaeonassa; Jensen, 2003

Verkhovka, Formation; White Sea, NW Russia

558–550 Ma

Shallow marine

Archaeonassa isp.;

Ust'-Pinega Formation; NW Russia

<555 Ma

Offshore under influence of a deltaic system

Archaeonassa; Grazhdankin Zimnegory Formation; and Maslov, 2009 White Sea, NW Russia

558–555 (553– 550) Ma

Middle to upper shoreface

Archaeonassa; Grazhdankin Erga Formation (lower and Maslov, 2009; part); White Sea, NW Grazhdankin, 2014 Russia

550–547 Ma

Middle to upper shoreface

Archaeonassa isp.; Grazhdankin and Maslov, 2015, and references therein

East European Platform (not 560–540 specified) Ma

Shallow marine

Unnamed; Chen et al., 2013

Dengying Formation; S China

551–542 Ma

Shallow marine

Archaeonassa cf. fossulata; this paper

Upper part of the Yampil Member, Mogyliv Formation; Ukraine

>557 Ma

Tidal flat

cf. Archaeonassa;

Cerro Negro Formation; E Argentina

<565 Ma

Shallow marine subtidal

cf. Archaeonassa; Warren et al., 2011

Itapucumi

Shallow marine

Group; Paraguay

600–550 Ma

Archaeonassa; Warren et al., 2014

Sete Lagoas Formation, Bambuí Group; E Brazil

550–542 Ma

Shallow marine limestones and dolomites

Jensen, 2003; Jensen et al., 2005, 2006

Arrouy et al., 2016

23

Nereites sp.; Crimes and Germs, 1982; considered as Archaeonassa by Jensen, (2003) and Jensen and Runnegar (2005)

Nudaus Formation; Namibia 547–545 Ma

Tidal flat

?Archaeonassa; Hageman and Miller, 2016

Unicoi Formation (middle part); Appalachians, USA)

Coastal plain occasionally inundated by sea

511

24

Probably terminal Ediacara n

Highlights

The trace fossil Archaeonassa cf. fossulata is a bilobate, epichnial ridge. A. cf. fossulata occurs in the Ediacaran (<557 Ma) shallow marine siltstones, Ukraine. A. cf. fossulata is oriented within a sector of 20–40º, perpendicular to the shoreline. A. cf. fossulata records ability of some Ediacaran burrowing organisms to taxis.

Declaration of interest

None.