Precambrian Research 399 (2023) 107227

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Precambrian Research
journal homepage: www.elsevier.com/locate/precamres

First record of trace fossils in the Ediacaran–Cambrian transition on the

northern Gondwana platform (Anti-Atlas, Morocco)
Abdelfattah Azizi a, *, Asmaa El Bakhouch a, Abderrazak El Albani b, Kalle Kirsimäe c,
Mouhssin El Halim a, Khadija El Hariri a, Mohamed Erragragui a, Ahmid Hafid a, Olev Vinn c
a
Laboratoire de Géo-ressources, Géoenvironnement et Génie civil, Département des Sciences de la Terre, Faculté des Sciences et Techniques, Université Cadi-Ayyad, BP
549, 40000 Marrakesh, Morocco
b
Université de Poitiers, UMR-CNRS, IC2MP 7285, 86022 Poitiers, France
c
Department of Geology, Institute of Ecology and Earth Sciences, University of Tartu, Ravila 14A, 50411 Tartu, Estonia

A R T I C L E I N F O A B S T R A C T

Keywords: The Anti-Atlas Mountains of Morocco preserve one of the most complete latest Neoproterozoic to Cambrian
Ediacaran stratigraphic successions worldwide, with high-resolution chemostratigraphic δ13Ccarb coverage. However, the
Cambrian exact stratigraphic position of the Ediacaran–Cambrian boundary remains unresolved. Until now, no trace fossils
Trace fossils
or body fossils have been recorded from the terminal Ediacaran strata in the Anti-Atlas Mountains. The only
Tabia Member
exception is a discoidal body fossil-like structure from the shallow-marine sandstones of the Ediacaran Tabia
Member, Adoudou Formation, in the Taroudant Group in the Igherm inlier. For the first time, we document the
occurrence of abundant trace fossils from the base of the Adoudou Formation, including Treptichnids, Mono­
morphichnus isp., Helminthoidichnites isp., Gordia isp., Palaeophycus isp., Planolites isp., and Conichnus isp. The
integration of both trace fossils and carbon isotope δ13Ccarb records suggests that the Ediacaran–Cambrian
transition in the Sous Basin can be placed at the contact between the Tifnout and Tabia Members, which che­
mostratigraphically coincides with the large negative δ13Ccarb excursion that had been previously suggested for
the Ediacaran–Cambrian boundary (BACE) in the western Anti-Atlas Mountains.

1. Introduction diversification of complex animals capable of burrowing and pene­

The Ediacaran–Cambrian boundary is one of the most interesting
transitions in the history of life, characterized by the first mass extinc­
tion of the Ediacaran biota in the fossil record (Darroch et al., 2015,
2018; Smith et al., 2017; Nance et al., 2022), the first appearance of
diverse metazoans with biomineralized skeletons (Germs, 1995), and an
exceptional development of metazoan complexity and behaviour
(Erwin, 1999; Knoll and Carroll, 1999; Narbonne, 2004, 2005). How­
ever, the exact timing of this bioevent and the stratigraphic position of
the Ediacaran–Cambrian boundary remain poorly resolved. Reaching
this goal is complicated due to an approximately 20-million-year gap
between the youngest preservation of Ediacaran body fossil assemblages
and that of Cambrian Stage 3 (Buatois et al., 2014). In contrast, the trace
fossil record across the Ediacaran–Cambrian transition is far more
diverse and continuous than that of the body fossils, providing valuable
information about this critical biological transition. One of the ecolog­
ical manifestations of the Cambrian Explosion was a rapid spread and
trating the sediment, thereby producing a dramatic increase in the depth
and intensity of bioturbation, known as the ‘Agronomic Revolution’ (e.
g., Seilacher, 1999; Bottjer et al., 2000; Seilacher et al., 2005; Buatois
et al., 2014, 2020). This bioturbation event was marked by a sudden
shift in the trace fossil style from horizontal surficial traces in the late
Ediacaran to vertically penetrative ichnofabrics in the Cambrian. The
Ediacaran–Cambrian boundary is defined by the first appearance of the
trace fossil Treptichnus pedum (Brasier et al., 1994). Secondary markers
include the last occurrences of Ediacaran-type body fossils and the first
appearance of small shelly fossils (SSFs), as well as a large negative
carbon isotope excursion, termed the BACE (BAsal Cambrian carbon
isotope Excursion) (e.g., Kimura and Watanabe, 2001; Amthor et al.,
2003; Smith et al., 2016; Nelson et al., 2022, 2023).
In Morocco, the Ediacaran–Cambrian Adoudou Formation of the
Taroudant Group is well exposed across the Anti-Atlas Mountains and
serves as a standard for the global chemostratigraphic carbonate δ13C
curve of the Ediacaran–Cambrian boundary interval (Maloof et al.,

* Corresponding author.
E-mail address: a.azizi@uca.ma (A. Azizi).

https://doi.org/10.1016/j.precamres.2023.107227
Received 11 April 2023; Received in revised form 22 September 2023; Accepted 2 November 2023
Available online 11 November 2023
0301-9268/© 2023 Elsevier B.V. All rights reserved.
A. Azizi et al. Precambrian Research 399 (2023) 107227

Fig. 1. A, simplified map of Africa showing the location of the Anti-Atlas on the northern border of the West African Craton (WAC); B, geological map of the Anti-
Atlas with locations of measured sections (redrawn and modified after Saadi et al., 1985); C, Composite section of the Taroudant and Tata Groups in the western Anti-
Atlas (δ13Ccarb data are from Maloof et al., 2005).

2005, 2010). Based on carbon isotope δ13Ccarb correlations, a negative
δ13Ccarb excursion (down to δ13Ccarb values of − 6 ‰) located at the
bottom of the Tifnout Member (Tamjout bed) in the sous basin was
suggested to represent the Ediacaran–Cambrian boundary (BACE)
(Maloof et al., 2005, 2010). However, neither body fossils nor trace
fossils have yet been found, except for a discoidal fossil preserved in
shallow-marine sandstones from the Tabia Member in the Igherm inlier
that was originally interpreted as a fossil jellyfish (Houzay, 1979) and
redescribed as Aspidella discs (Maloof et al., 2005; Letsch et al., 2019).
The aims of this study are to (1) document the Adoudou Formation
ichnofauna and (2) discuss the most plausible position for the Ediacar­
an–Cambrian boundary in the western Anti-Atlas Mountains of Morocco
by using an integration of the newly available ichnological data and
carbonate δ13C chemostratigraphy.
2. Geological setting and stratigraphy
The Anti-Atlas Mountains are an ~1300 km long NE-SW-trending
belt on the northern edge of the Eburnian (c. 2 Ga) West African
Craton (WAC) (Fig. 1A). This orogen is separated from the High Atlas
Mountains to the north by the South Atlas Faults (Fig. 1B). The southern
slope of the Anti-Atlas Mountains is composed of a Palaeozoic sedi­
mentary succession that overlies the Precambrian rocks cropping out
within numerous basement inliers (Fig. 1B), providing well-exposed

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A. Azizi et al. Precambrian Research 399 (2023) 107227

Fig. 2. A lithostratigraphic section of the studied outcrop in the Zaouia locality, with associated carbonate oxygen and carbon isotope data, is presented. The
horizontal dashed line marks the Ediacaran–Cambrian boundary based on the lowest occurrence of the Fortunian trace fossils as well as the nadir of a negative
carbonate carbon isotope excursion that may correlate to the BACE (Corsetti and Hagadorn, 2000).

outcrops of crystalline and sedimentary rocks of Palaeo-, Meso- and
Neoproterozoic age (e.g., Thomas et al., 2002, 2004; Walsh et al., 2012;
Hefferan et al., 2014; Ikenne et al., 2017). The Precambrian basement
and its Palaeozoic cover were deformed in a thick-skinned fashion
during the Hercynian orogeny (Faik et al., 2001; Baidder et al., 2016).
The Palaeoproterozoic basement (referred to as PI in the regional
literature), occurring on the southwestern side of the Anti-Atlas Major
Fault (AAMF; Choubert, 1947) (Fig. 1B), is composed of schists, para­
gneisses, and migmatites that are intruded by batholiths yielding U–Pb
zircon ages ranging from ~2.2 to 2.0 Ga (e.g., Thomas et al., 2002;
Walsh et al., 2002; Gasquet et al., 2008; O’Connor, 2010). The quartzite
and limestone series of Taghdout and Jbel Lkest that overlie the Palae­
oproterozoic rocks were attributed to the Cryogenian, based on the Rb/
Sr 789 ± 10 Ma age of the contact-metamorphosed wall-rocks of mafic
dykes (Clauer, 1974). Recent studies have highlighted the existence of
Mesoproterozoic mafic dikes within the Taghdout sedimentary series,
which yield U–Pb baddeleyite ages of ~1710 Ma (Ikenne et al., 2017;
Youbi et al., 2013).
Relics of coeval oceanic crust rocks are preserved along the AAMF,
from the Siroua to Bou Azzer inliers (Fig. 1B), the Tachdamt Bleïda series
and the associated ophiolitic complex (referred to as PII in the regional
literature, e.g., Choubert, 1963). The PII units document the Neo­
proterozoic Pan-African cycle (c. 760–600 Ma), with the opening and
closing of a Cryogenian ocean basin between the WAC’s northern rifted
margin and a northern continental block. The closure of the basin was
followed by widespread intracontinental deformation and continental
and marine sedimentation (Gasquet et al., 2008; El Hadi et al., 2010;
Blein et al., 2014; Hefferan et al., 2014; Soulaimani et al., 2018; Tri­
antafyllou et al., 2016).
Both PI and PII are unconformably covered by the Ouarzazate Su­
pergroup (sensu Thomas et al., 2004) or PIII (Choubert, 1963). This unit
consists of a ~2000 m-thick pile of subaerial, volcano-sedimentary

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A. Azizi et al. Precambrian Research 399 (2023) 107227

series deposited between ~615 and 538 Ma within alluvial and fluvial
environments (e.g., Thomas et al., 2002; Gasquet et al., 2008; Walsh
et al., 2012;). The youngest U-Pb zircon age (538 ± 6 Ma) has been
reported from the Jbel Saghro inlier in the eastern Anti-Atlas Mountains
(Baidada et al., 2018), whereas the maximum depositional age of ~541
Ma has been reported from the Bas-Draa inlier in the western Anti-Atlas
Mountains and shifts the Precambrian–Cambrian boundary down from
the Lower Adoudou Formation, as proposed by previous studies, to the
uppermost Ouarzazate Supergroup or to the hiatus between both units
(Karaoui et al., 2015). PIII rocks are, in turn, covered by a thick suc­
cession of nonmetamorphic Palaeozoic successions (Fig. 1B and C). The
Cambrian sedimentary rocks that unconformably overlie the Ouarzazate
Supergroup are dominated by shallow-marine carbonate and siliciclastic
rocks and are subdivided into two main groups (Fig. 1C). The ~1000 m
thick Taroudant Group is subdivided into two formations: the Adoudou
and Taliwine Formations (Choubert, 1952). The Adoudou Formation is
subdivided into the Tabia and Tifnout members (sensu, Maloof et al.,
2005). The Tabia Member (la série de base, sensu Choubert, 1952) is
represented by a mixed siliciclastic–carbonate succession that is sub­
divided into three units (Benssaou and Hamoumi, 2001, 2004). The
‘basal conglomerate’ unit is composed of coarse-grained sandstone and Fig. 3. Cross-plot and linear correlation cart of the δ13C and δ18O values of the
conglomerate deposited in a marine fan-delta setting dominated by al­ 45 samples from the Zaouia section.
luvial processes; the middle ‘basal limestone’ unit preserves well-
developed cyanobacterial stromatolites formed in a shallow carbonate
4. Carbonate carbon isotope systems
platform, and the overlying ‘basal siltstone’ (studied in this paper;
Fig. 1D, 2) consists of alternating transgressive–regressive depositional
Carbonate δ13C and δ18O values exhibit rather large variations
sequences, displaying typical wave-generated structures, such as hum­
(Fig. 3, Table 1). The carbonate δ13C and δ18O cross-plot does not show a
mocky cross-stratification and polygonal ripples that record episodic
distinctive positive correlation between C and O isotopic values that
storm events (Benssaou and Hamoumi, 2001, 2004). The Tifnout
would indicate diagenetic alteration (Banner and Hanson, 1990; Kauf­
Member is dominated by stromatolitic carbonate deposited in peritidal
man and Knoll, 1995), and the studied rocks can be considered well
platforms. The Adoudou Formation is overlain by the regressive deposits
preserved (but see below).
of the Taliwine Formation, consisting of fine-grained siliciclastic sedi­
The basal limestone unit shows overall higher δ13Ccarb values, typi­
ments with several carbonate interbeds. The overlying fossiliferous Tata
cally varying between − 1 and 1 ‰, but they show large variations from
Group (Cambrian stage 2) is well exposed in the Sous Basin (~1000 m
− 2.6 to 2.1 ‰ (Fig. 2) over a few metres at the contact between the basal
thick) and is dominated by variegated shale with interbedded black
limestone and overlying basal siltite, which most likely indicates a
oolitic limestone. The Tata Group records a sudden shift in the compo­
diagenetic imprint in this interval. In the Tifnout Member, the δ13Ccarb
nents of ecosystems, marked by the disappearance of microbial con­
values range narrowly between − 2.9 and 3.8 ‰ and show a slight trend
sortia (stromatolite-dominated) and the emergence of thrombolites and
from values closer to − 4 ‰ in the lower part of the unit to values ca. − 3
shelly metazoans, including archaeocyaths, trilobites, chancelloriids,
‰ in the middle and upper parts of the studied section (Fig. 2).
hyoliths and calcimicrobes (e.g., Hupé, 1960; Schmitt and Monninger,
1977; Sdzuy, 1978; Schmitt, 1979; Debrenne and Debrenne, 1995;
5. Synopsis of trace fossils
Álvaro and Clausen, 2006; Álvaro and Debrenne, 2010; Clausen et al.,
2014; Azizi et al. 2022).
Trace fossils from the Tabia Member were divided into four cate­
gories of traces: simple horizontal scratches and trails; simple actively
3. Materials and methods
filled (massive) horizontal to oblique structures; horizontal burrows
with horizontal to vertical branches and vertical plug-shaped burrows. A
The Tabia Member is well exposed along a road crossing the Alma
wide variety of ichnofossils are associated with microbially induced
inlier (Fig. 1). One representative section (Zaouia section; GPS: N
sedimentary structures (MISSs) at the tops of siltstone and sandstone
29◦56′27′′, W 8◦4′42′′), comprising the ‘Basal siltite’ unit, was logged
beds. Most of the beds containing MISSs yield well-preserved trace
(Fig. 2), and trace fossil samples were collected during field campaigns
fossils. Trace fossils are variably distributed on exposed bedding
in 2018, 2019 and 2022. Photographs were taken with a Canon EOS
surfaces.
650D. Polished slabs were scanned by a digital scanner (EPSON
EXPRESSION 10000 XL). Specimens are housed at the Faculté des Sci­
ences et Techniques, Université Cadi-Ayyad. 5.1. Simple horizontal scratches and trails
Carbonate O and C stable isotope compositions were analysed using
a Thermo Scientific Delta V Advance continuous flow isotope ratio mass 5.1.1. Ichnogenus Monomorphichnus Crimes, 1970 (Fig. 4 A–C)
spectrometer and a GasBench II preparation line connected to a Delta V This ichnogenus consists of horizontal straight to slightly sigmoidal
Advantage IRMS (Thermo Fisher Scientific) at the University of Tartu. subparallel ridges, preserved in positive epirelief. The ridges are ar­
Samples were chosen from the lithologically homogenous zones, and ca. ranged in pairs, with one ridge deeper than the other, and continual
2 mg of sample powder was dissolved in orthophosphoric acid overnight series that meet in a V. This trace fossil is unbranched, but crossovers or
at 70 ◦ C. The results of carbonate analyses are expressed in per mil de­ interpenetrations are found. The ridges are 10–13 cm long and are
viation relative to the Vienna PeeDee Belemnite (VPDB) scale for oxygen separated from each other by distances of 3–5 mm. The transverse ridges
(δ18O) and carbon (δ13C). Long-term reproducibility was better than ± are 2.0–3.0 mm wide. Central ridges are usually deeper than outer
0.2 ‰ (2σ) for δ18O and δ13C values. ridges.
These traces are tentatively assigned to Monomorphichnus Crimes,
1970. The presence of two main scratch marks at first sight makes this

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A. Azizi et al. Precambrian Research 399 (2023) 107227

Fig. 4. A, subsurface of bed with arthropod scratch marks of the Monomorphichnus type (sample AA–MO–1). B & C, Magnified view of the boxed area in (A) show the
arrangement of parallel ridges and alternation of high and deeper ridges (arrowed). Scale bars are 1 cm.

form comparable to Monomorphichnus multilineatus, in which the central
ridges are deeper than the outer ridges (Gibb et al., 2017). Mono­
morphichnus can be distinguished from Dimorphichnus by the lack of
pusher marks (Crimes, 1970; Osgood, 1970; Crimes et al., 1977). Jensen
(1997) considered that both ichnogenera might represent different be­
haviours of the same producer. Monomorphichnus has been widely
documented in the Fortunian and younger strata (Mángano and Buatois,
2014, Marusin et al., 2021). Monomorphichnus-type scratch marks are
usually attributed to a trilobite tracemaker (Alpert, 1976; Sharma et al.,
2018).
5.1.2. Ichnogenus Gordia Emmons, 1844 (Fig. 5 A-D)
The ichnogenus Gordia appears as a smooth meandering, un­
branched, horizontal trace, is uniformly 1.0–1.4 mm wide, and it
commonly has overcrossings, generating repetitive, looping spirals. This
ichnogenus is preserved as concave epirelief along bedding planes in
laminated siltstone.
Gordia differs from Helminthopsis by its looped form (Pickerill and
Peel, 1990). The Tabia examples exhibit increased meandering and
crossing. Gordia is reported in various terrestrial and marine deposi­
tional environments from several localities ranging in age from the late
Ediacaran to Holocene (Mángano and Buatois, 2014; Turk et al., 2022).
Polychaetes or worm-like organisms have been proposed as possible
tracemakers (Książkiewicz, 1977). Peel (2010) reported examples of
‘Gordia’-type traces on large arthropod shields from the early Cambrian
Sirius Passet fossil lagerstaette.
5.1.3. Ichnogenus Helminthoidichnites Fitch, 1850 (Fig. 5C, D)
The ichnogenus Helminthoidichnites appears as shallow, curved to
straight, 1.3–2.5 mm wide, unbranching trails, preserved as negative

5
A. Azizi et al.
Fig. 5. Photographs and schematic tracing of the Tabia Member ichnofossils. A & B, Photograph and schematic tracing of Gordia isp. C & D, Photograph and
schematic tracing showing Helminthoidichnites isp and Gordia isp. (field photograph). Scale bars are 1 cm.
relief on the top surface of medium- to coarse-grained sandstone bed.
The trajectory of the trace is irregular. Overcrossing between several
burrows is common, and random loops are rare. The width is constant in
individual specimens.
Helminthoidichnites is one of the most common trace fossils from the
basal siltstone of the Tabia Member, and it is usually found in association
with Gordia. Häntzschel (1975) regarded Helminthoidichnites as a junior
synonym of Gordia. However, other later studies (Buatois et al., 1998)
retained this ichnogenus. Helminthoidichnites differs from Gordia by
lacking self-overcrossing and from Helminthopsis in possessing a less
winding nature (Hofmann and Patel, 1989). Helminthoidichnites range in
age from the Ediacaran to the Holocene (e.g., Narbonne and Aitken,
1990; Uchman et al., 2009; Turk et al., 2022). Vermiform animals are
suggested as potential trace makers of Helminthoidichnites-type traces
(Buatois et al., 1998; Evans et al., 2020).
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Precambrian Research 399 (2023) 107227
5.2. Simple actively filled (massive) horizontal to oblique structures
5.2.1. Ichnogenus Palaeophycus Hall, 1847 (Fig. 6A, B)
This ichnogenus has horizontal, smooth, cylindrical, straight to
slightly curved unbranched burrows that are 3–5 mm wide and up to
150 mm long, with a circular cross section. The walls are smooth and
nonornamented. The filling is the same as the host rock.
Palaeophycus is commonly interpreted as a dwelling burrow of ver­
miform animal deposit feeders, mostly polychaetes, usually moving
parallel to the sediment surface (e.g., Pemberton and Frey, 1982; Hof­
mann et al., 2012; Knaust, 2017). Palaeophycus burrows can be distin­
guished from Planolites burrows by the lined trace walls, as well as the
nature of the sediment filling. In contrast to Planolites, the filling of
Palaeophycus burrows is usually similar to the surrounding rock matrix
(Pemberton and Frey, 1982). Palaeophycus is an unbranched burrow;
however, pseudobranching is usually caused by the crossing of different
A. Azizi et al.
Fig. 6. Photographs of the Tabia Member ichnofossils. A & B, Palaeophycus isp. (field photographs) The coin diameter is 2.4 cm. C, Planolites isp (sample; AA–Pl–1).
D, vertical cross section of Planolites isp. Scale bars are 1 cm.
burrow generations, which may result in misinterpretations and confu­
sion with other similar traces, such as Thalassinoides and Planolites
(Weber et al., 2013). Palaeophycus-type traces have been reported from
the late Ediacaran to Holocene (Scott et al., 2009; O’Neil et al., 2020).
5.2.2. Ichnogenus Planolites Nicholson, 1873 (Fig. 6C-D)
The ichnogenus Planolites appears as horizontal to slightly oblique,
straight or irregularly sinuous, unbranched simple flattened cylinders,
without ornamentation. It is elliptical in cross-section and 4–10 mm in
diameter, and the maximum preserved length is 10 cm. The burrow is
filled with a material that is different from the host rock matrix; the
filling is finer-grained, dominated by quartz grains, while the sur­
rounding rock matrix is composed of quartz, feldspar, and mica flakes.
Planolites is an actively filled burrow, interpreted as a feeding trace
of vermiform deposit feeders (Pemberton and Frey, 1982, Rodríguez-
Tovar and Uchman, 2004; Pervesler et al., 2011). Worms, molluscs and
arthropods are proposed as possible tracemakers of Planolites (Bromley,
1996; Buatois and Mángano, 2002). Planolites have been described in
various continental and marine environments (e.g., Pemberton and
Frey, 1982; Rodríguez-Tovar and Uchman, 2004; Sarkar et al., 2009;
Phillips et al., 2011) and range in age from the late Ediacaran to Holo­
cene (Häntzschel, 1975; Crimes, 1992; O’Neil et al., 2020). Recent in­
vestigations have revealed an absence of Planolites in the late Ediacaran
(Mángano and Buatois, 2016; Buatois et al. 2020).
5.3. Horizontal burrows with horizontal to vertical branches; Treptichnids
(Fig. 7A–F)
Treptichnid traces are common in the upper part of the Tabia
Member (3 m below the base of the Tifnout Member), with two distinct
morphologies observed. The first trace morphology (Fig. 7A, B) consists
of individual probes that are generally oval in shape, with a maximum
diameter of 6 mm. The second morphology (Fig. 7C–F) is composed of a
subhorizontal series of individual, straight spindle-shaped probes ar­
ranged in a zigzag pattern. The shorter series consists of two segments
(Fig. 7E), whereas the longer series is composed of approximately five
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Precambrian Research 399 (2023) 107227
segments (Fig. 7F). Probes are segments that vary in length from 7.0 to
13.0 mm and in width from 2.1 to 3.0 mm. The traces change direction
at the segment junctures, but no projections are observed. Segment
junctures occur as small depressions (Fig. 7C), although rare continued
and curved junctions are observed (Fig. 7F). Angles between segments
range from 95◦ to 165◦ .
Treptichnid traces have been previously reported from terminal
Ediacaran strata, such as in Spain (Jensen et al., 2007), Newfoundland
(Gehling et al., 2001), Finnmark (Högström et al., 2013; McIlroy and
Brasier, 2017), and Namibia (Darroch et al., 2016; Jensen and Runne­
gar, 2005; Jensen et al., 2000). Short-sized segments scattered on the
bedding plane (Fig. 7A) share a similar morphology with those reported
from the Huns Member in Namibia but with more widely spaced ele­
ments (Jensen et al., 2000; Darroch et al., 2021; Nelson et al., 2022), and
those from the Cijara Formation in Spain (Jensen et al., 2007). However,
uniserial probes arranged in a zigzag pattern are similar to those
described by Jensen et al. (2007) and assigned to Treptichnid type
traces. This complex structure may possibly represent three-dimensional
burrow systems approaching the morphology of Treptichnus but is of
lesser complexity. The relationship between Treptichnus and treptichnids
is uncertain (Jensen et al., 2007). It is unclear whether both traces have
the same origin but are preserved differently or if treptichnids are
similar to Treptichnus but represent less complex trace fossils. (see Jen­
sen et al., 2000). These occurrences provide insight into the potential
evolutionary origins of complex trace fossil behaviour and can be easily
incorporated into the criteria used in recognizing and correlating the
Ediacaran–Cambrian boundary (Buatois et al., 2013; Landing et al.,
2013). The Tabia Member treptichnids represent the earliest form of
’complex’ trace fossil behaviour, albeit not as complex as that reflected
by T. pedum, the latter marking the base of the Cambrian (Brasier et al.,
1994). It is uncertain whether the simpler treptichnid burrows were
created by the same organisms as T. pedum, which is commonly thought
to have been formed by priapulid worms (Vannier et al., 2010; Kesidis
et al., 2019).
A. Azizi et al. Precambrian Research 399 (2023) 107227

Fig. 7. Photographs of Treptichnids traces from the Tabia Member, all in sole view. A, Treptichnids on the sole of a sandstone bed showing individual probes oval in
shape (white arrows). B, Treptichnids on the sole of a sandstone bed showing individual probes of irregular shape (yellow arrows). C and D, Possible Treptichnids
composed of serial spindle-shaped probes arranged in a zigzagging pattern (white arrows); segment junctures occur as small depressions (yellow arrows). E and F,
Possible Treptichnids composed of serial straight probes arranged in a zigzagging pattern (white arrows); some segments are connected and show curved junctions
(yellow arrows). Scale bars are 1 cm.

5.4. Vertical plug-shaped burrows
5.4.1. Ichnogenus Conichnus Männil, 1966 (Fig. 8A, B and C)
This ichnogenus has short, conical, vertical to slightly inclined bur­
rows that are round to elliptical in transverse section. The studied
specimens show variable dimensions; the maximum diameter of the
burrows is 23 mm, and they are up to 25 mm deep, with a resulting
diameter/depth ratio of 0.92. The burrow fill is composed of fine-
grained siliciclastic–carbonate sediments. The lithological difference
between the burrow fill and the host rock is clearly marked. The central
cone is rich in siliciclastic material dominated by fine-grained quartz
grains (Fig. 8C), which is somewhat more resistant to weathering than
the surrounding part and the host sediments. Concentric laminae are
observed on the top (Fig. 8B). The basal part may be slightly rounded or
conical.
In certain aspects, the majority of these conical traces resemble to the
ichnogenera Conichnus Männil, (1966). According to Männil (1966), the
ichnogenus Conichnus comprises acuminated subcylindrical structures
oriented perpendicular to the bedding plane. Fillings may reveal
patterned internal structures, such as chevron laminae, but not radial
medusoid symmetry. Conichnus-type traces can be distinguished from
Amphorichnus by the lack of amphora-like shape and the papillate
termination. Amphorichnus also differs from Conichnus by the presence of
lateral adjustment traces (Vinn et al., 2015). Conichnus traces have been
recorded from the lower Cambrian to the Cenozoic (Pemberton et al.,
1988). Plug-shaped burrows interpreted as Conichnus are the most
common trace fossils found in the late Ediacaran strata of Namibia
(Darroch et al., 2016; Cribb et al., 2019; Darroch et al., 2021). Con­
ichnus-type traces are interpreted as the dwelling burrows of a soft-
bodied anemone-like organism (Frey and Howard, 1985, Pemberton
et al., 1988). The variously filled internal structures of Conichnus bur­
rows suggest that the trace maker moved periodically upwards as the
animal tried to keep pace with sedimentation (Frey and Howard, 1985).
5.4.2. Problematic paired plug-shaped burrows (Fig. 9 A, B and C)
Pairs of plug-shaped holes are very common at the boundary

8
A. Azizi et al.
Fig. 8. Trace fossil specimens of the Conichnus type from the boundary between the Tabia and Tifnout Members are shown. A, a field photograph of Conostichus in
sandy carbonate mixed sediments (sample: AA–Co–1). B, a top view of Conostichus showing concentric laminae (arrowed) (sample: AA–Co–2). C, a vertical view of
Conichnus displaying a conical shape. Scale bars are 1 cm.
between the Tabia and Tifnout members and may occur in association
with most of the other traces described above. They are usually pre­
served on bedding planes as a pair of holes and look superficially like the
Arenicolites-type trace, but a “U” shape has not been detected in cross-
section, and this type of trace does not exceed 4 mm in depth. The di­
ameters of the holes are 4–12 mm, and the spacing between the holes of
each pair varies between 4 and 10 mm.
Paired plug-shaped burrows have been previously described within
the E-C boundary interval in the Nama Group, Namibia (Darroch et al.,
2016; Cribb et al., 2019; Darroch et al., 2021). As mentioned by Darroch
et al. (2021), the neighbouring holes seem to be alternating or aligned,
suggesting affinities with treptichnids. Similar plug-shaped forms are
interpreted as Brooksella by Crimes and Germs (1982). However, the
association of burrows in pairs remains mysterious.
6. Discussion
The trace fossil record provides a key piece of evidence to evaluate
the timing of diversification and ecosystem construction during the
terminal Ediacaran–early Cambrian. Ediacaran shallow marine deposits
worldwide contain simple horizontal trace fossils reflecting the activity
of nonspecialized microbial mat grazers (Buatois and Mángano, 2012;
Darroch et al., 2021). The Cambrian radiation of complex animal life
induced a dramatic increase in the depth and complexity of bioturbation
in the seafloor sediment, known as the ‘agronomic revolution’ (AR). The
AR marks the shift from matgrounds to bioturbated mixgrounds (Buatois
et al., 2014). The timing of this geobiological event is still a matter of
debate, whereas significant diversification of trace fossils and the advent
of bioturbation occurred ca. 20 Myr earlier than the “Cambrian Explo­
sion” of body fossils (Buatois and Mangano, 2016).
The transition between the Tabia and the Tifnout members in the
9
Precambrian Research 399 (2023) 107227
Zaouia section includes abundant horizontal burrows and trails, prob­
ably produced by priapulids (Treptichnids) and arthropods (Mono­
morphichnus), as well as by anemone-like organisms (Conichnus). The
ichnotaxonomic composition and stratigraphic distribution of the trace
fossils of this part of the Adoudou Formation are consistent with For­
tunian ichnoassemblages (Mángano and Buatois, 2017).
The Ediacaran–Cambrian boundary is marked by the first appear­
ance datum (FAD) of T. pedum, the first three-dimensionally branching
burrows (Narbonne et al., 1987; Brasier et al., 1994; Landing, 1994;
Buatois et al., 2013), as well as other biostratigraphic ichnofossils that
made their first appearance at or immediately above the boundary, such
as Arenicolites sp., Conichus conicus, Monomorphichnus spp., Phycodes sp.,
or Helminthopsis tenius (Jensen, 2003; Landing et al., 2013; Babcock
et al., 2014).
The FAD of T. pedum roughly corresponds to the base of the
Cambrian worldwide, including the Mackenzie Mountains of Canada
(Carbone and Narbonne, 2014); Sonora, Mexico (Sour-Tovar et al.,
2007); Death Valley, Eastern California (Jensen et al., 2002); Flinders
Ranges of South Australia (Jensen et al., 1998); eastern Finnmark,
Norway (Føyn and Glaessner, 1979); and Nama Group, Namibia (Lin­
nemann et al., 2019; Darroch et al., 2021), among many other regions.
Nevertheless, the selection of T. pedum as a proxy for the base of the
Cambrian period has also been heavily criticized (Babcock et al., 2014;
Zhu et al., 2019). On the one hand, the three-dimensional structure of
T. pedum remains the subject of debate (Geyer, 2005; Seilacher, 2007;
Vannier et al., 2010; Chen et al., 2013; Meyer et al., 2014). According to
Seilacher (2007), the trace fossil T. pedum is indicative of a sophisticated
trace of undermat mining rather than the vertical bioturbation proposed
by Geyer and Uchman (1995) and Geyer (2005). Undermat mining
forms are reported in the Ediacaran Shibantan Member of the Dengying
Formation in South China (Meyer et al., 2014), as well as in the lower
A. Azizi et al.
Fig. 9. Trace fossil specimens of the paired vertical plug-shaped burrows from the boundary between the Tabia and Tifnout Members. A and B, slab specimens
showing numerous pairs of holes (sample; AA–PH–1). B & C, vertical cross–section of the holes showing in (A), the position of holes is marked by white arrows. Scale
bars are 1 cm.
Member of the Wood Canyon Formation in the Great Basin, United
States (O’Neil et al., 2020). In addition, the morphology of Treptichnids is
not yet well understood, making it difficult to make a definitive taxo­
nomic determination of them in relation to the ichnogenus Treptichnus.
Jensen et al. (2000) described Treptichnids in more detail in the base of
the Huns Member of the Nama Group and identified them as Treptichnus
isp.
On the other hand, the facies dependence of T. pedum has been
regarded as a major complication in its use for global correlation
(Gehling et al., 2001; Geyer, 2005; Babcock et al., 2014; Zhu et al.,
2019). Generally, these trace fossils are found predominantly in silici­
clastic rocks, and although they are known to occur in a range of lith­
ofacies (Landing et al., 2013; Buatois et al., 2013; Buatois, 2018),
carbonate facies are usually not included. This limitation reduces the
application of T. pedum as a proxy for the base of the Cambrian, as this
interval often consists of carbonate facies in numerous key regions, such
as Kazakhstan, China, Mongolia, and Siberia. Facies dependence may
explain the delayed appearance of T. pedum in some
10
Precambrian Research 399 (2023) 107227
Ediacaran–Cambrian sections (Buatois et al., 2013; Shahkarami et al.,
2017; Buatois, 2018; Devaere et al., 2021). The Fortune Head stratotype
is a coarse siliciclastic-dominated sequence that has undergone meta­
morphism and lacks a chemostratigraphic, magnetostratigraphic, or
other non-biostratigraphic record that would enable correlation with
other sections regionally or globally (Geyer, 2005). The FAD of T. pedum
is inferred to lie above the interval that contains mostly Ediacaran fossils
and below the interval containing mostly Cambrian fossils. This section
has not been proven to be the stratotype; however, it is strongly sup­
ported by numerous studies around the world (Geyer, 2005; Maloof
et al., 2005, 2010; Landing et al., 2013). It should also be considered that
Ediacaran-type macrofossils have rarely been reported from lower
Cambrian strata. For example, Ediacaran-type fronds were reported
from the Uratanna Formation in the Angepena syncline, northern Flin­
ders Ranges, above the Ediacaran–Cambrian transition (Jensen et al.,
1998). Simple discoidal fossils and frond-like soft-bodied fossils are
described from the Cambrian strata of the Salt Spring Hills and the White
Mountains (Hagadorn et al., 2000). In addition, Cloudinomorphs and
A. Azizi et al. Precambrian Research 399 (2023) 107227

Table 1 Ediacaran–Cambrian transition to be somewhat speculative in practice.

Carbon and oxygen isotope data. Nonetheless, the first appearance of typical Fortunian trace fossils allows
Lab. ID ID1 ID2 δ13C(‰V-PDB) δ18O(‰V-PDB) us to constrain the Cambrian age for the base of the Tifnout Member.
This is also supported by the carbon isotopic δ13Ccarb trend in the studied
0948-22 Ti 1 − 3,75 9,40
succession, showing δ13Ccarb values in the basal limestone unit aver­
−
0949-22 Ti 2 − 3,82 − 9,38
0950-22 Ti 3 − 3,70 − 9,50 aging 0 ‰ but staying rather stable at − 3 to − 4 ‰ in carbonates of the
0951-22 Ti 4 − 3,50 − 9,33 Tifnout Member (Fig. 2). A large negative δ13C excursion, the BACE, has
0952-22 Ti 5 − 3,47 − 8,59 been recorded worldwide postdating the last appearance of Ediacaran-
0953-22 Ti 6 − 3,34 8,03
type fossils and/or predating the first occurrence of T. pedum (Zhang
−
0954-22 Ti 7 − 3,26 − 8,83
0955-22 Ti 8 − 3,20 − 9,19 et al., 1997; Corsetti and Hagadorn, 2000; Amthor et al. 2003; Smith
0956-22 Ti 9 − 3,07 − 9,04 et al., 2016; Chen and Feng, 2019). The BACE is known from Mongolia
0957-22 Ti 10 − 3,17 − 8,51 (Brasier et al., 1996; Smith et al., 2016), the Siberian platform (Kaufman
0958-22 Ti 11 − 3,38 9,00
−
and Knoll, 1995; Bartley et al., 1998; Kouchinsky et al., 2017), north­
0959-22 Ti 12 − 3,23 − 8,40
0960-22 Ti 13 − 3,05 − 9,60 west Canada (Narbonne et al., 1994), Iran (Kimura et al., 1997), the
0961-22 Ti 14 − 3,07 − 9,26 Great Basin (Corsetti and Hagadorn, 2000; Corsetti and Kaufman, 2003;
0962-22 Ti 15 − 3,20 − 9,44 Smith et al., 2016), Kazakhstan (Stammeier et al., 2019), South China
0963-22 Ti 16 − 2,97 − 10,22 (Ishikawa et al., 2008, 2014; Li et al., 2009; Steiner et al., 2020), Mexico
0964-22 Ti 17 − 3,00 10,39
(Loyd et al., 2012, 2013; Hodgin et al., 2021) and Morocco (Maloof
−
0965-22 Ti 18 − 3,22 − 10,29
0966-22 Ti 19 − 3,09 − 9,69 et al., 2005, 2010; and herein). Therefore, this negative excursion is
0967-22 Ti 20 − 2,95 − 10,98 thought to mark the Ediacaran–Cambrian transition and has been linked
0968-22 Ti 21 − 3,10 − 10,14 to the perturbation of the carbon cycle and, possibly, a mass extinction
0969-22 Ti 22 − 3,07 12,01
−
(Amthor et al., 2003; Darroch et al., 2018; Hodgin et al., 2021);
0970-22 Ti 23 − 2,78 − 10,51
0971-22 Ti 24 − 3,18 − 11,80 nevertheless, its absolute age has not been clearly defined. Geochrono­
0972-22 Ti 25 − 3,44 − 11,30 logical dating of the Ediacaran–Cambrian excursion comes from the Ara
0973-22 Cb 1 − 0,81 − 7,12 Group in Oman, where the U–Pb ages obtained from a volcanic ash bed
0974-22 Cb 2 − 1,01 − 8,25 that coincides with the negative excursion yield 542.0 ± 0.3 Ma
0975-22 Cb 3 − 0,78 6,85
(Amthor et al., 2003) and are recalibrated to 541.00 ± 0.13 Ma by
−
0976-22 Cb 4 − 1,67 − 5,34
0977-22 Cb 5 − 0,89 − 8,41 Bowring et al. (2007). Recent U-Pb ages obtained for a volcanic ash bed
0978-22 Cb 6 − 0,85 − 8,78 located three metres above the BACE in southern China have yielded
0979-22 Cb 7 − 0,39 − 6,85 540.7 ± 3.8 Ma (Chen and Feng, 2019), providing further support for
0980-22 Cb 8 0,29 6,59
−
the validity of the overall excursion of the Ediacaran–Cambrian
0981-22 Cb 9 − 0,08 − 7,16
0982-22 Cb 10 − 0,33 − 7,40 boundary.
0983-22 Cb 11 0,18 − 7,77 The composite carbon isotope curve of the Sous Basin shows two
0984-22 Cb 12 0,02 − 7,31 negative ‘W’-shaped excursions at the base of the Adoudou Formation
0985-22 Cb 13 − 0,05 − 7,53 (e.g., Maloof et al., 2005, 2010), where the lower anomaly (~− 4 ‰) is
0986-22 Cb 14 − 0,25 7,43
recorded in the Basal limestone unit of the Tabia Member, and the upper
−
0987-22 Cb 15 0,93 − 7,55
0988-22 Cb 16 1,32 − 8,47 excursion (~− 6 ‰) oscillates in the lower fifty metres of the Tifnout
0989-22 Cb 17 − 0,97 − 6,64 Member. Identical ‘W’-shaped δ13Ccarb signatures have been recorded in
0990-22 Cb 18 0,32 − 8,64 the Siberian Platform (Brasier et al. 1996) and SW Mongolia (Smith
0991-22 Cb 19 − 0,25 6,63
−
et al., 2016). In the Zaouia area, this upper excursion is recorded in the
0992-22 Cb 20 − 0,23 − 8,20
0993-22 Cb 21 − 0,39 − 8,07
base of the Tifnout Member and regarded as marking the Ediacar­
0994-22 Cb 22 − 2,61 − 6,50 an–Cambrian transition of the Sous Basin (e.g., Maloof et al., 2005,
0995-22 Cb 23 2,22 − 9,37 2010). The carbonate isotope values in the studied profile suggest,
0996-22 Cb 24 1,62 − 9,94 however, that basal limestone with nearly normal marine values is
chemostratigraphically below the ‘W’-shaped excursion, and the sam­
Erniettomorphs are found in the Nomtsas Formation in Namibia, and they ples from the Tifnout Member carbonates overlying the basal siltite unit
are dated to 538.56 ± 0.09 Ma and stratigraphically overlap complex with the observed ichnofossils show values and a rather flat stratigraphic
bilaterian trace fossils, such as Archaeonassa, Psammichnites, Para­ trend corresponding to the lower δ13Ccarb anomaly at the beginning of
psammichnites, and treptichnids (Nelson et al., 2022). Moreover, dis­ the ‘W’-shaped excursion. Therefore, based on the C isotope character­
coidal fossils were previously described from the Tabia Member in the istics and ichnofossil occurrence, the Ediacaran–Cambrian boundary can
Warmandaz region, which neighbours the Igherm inlier and is located be placed at the base of the Tifnout Member and below the most nega­
approximately 30 km eastwards from the studied section (Houzay, 1979; tive part of the BACE. This conclusion is further justified by the last
Maloof et al., 2005, 2010; Letsch et al., 2019). These are 1–13 cm wide occurrences of Treptichnids and Ediacaran-type fossils in the Tabia
and preserved both as positive epireliefs on the tops of beds and corre­ Member.
sponding positive hyporelief casts on the soles of beds. Following the In addition, the Ediacaran–Cambrian transition coincides with a
argumentation of Letsch et al. (2019), the Warmandaz specimens share major unconformity observed in many sections: Mackenzie Mts., Canada
numerous characteristics with the genus Aspidella, which is described in (Narbonne et al., 1994); Death Valley, United States of America (Corsetti
many Ediacaran strata around the world (Tarhan et al., 2015). Never­ and Kaufman, 1994; Corsetti and Hagadorn, 2000); and the Ara Group,
theless, it is evident that the decline of Ediacaran fauna is coincident Oman (Amthor et al., 2003). Additionally, this major discontinuity is
with the rise of bioturbation, also hinting that sediment disturbance may located below and not far from the BACE in Lena-Aldan, Siberia (Brasier
have played a prominent role in the decline of the soft-bodied Ediacaran et al., 1994; Pelechaty et al., 1996), and SW Mongolia (Brasier et al.,
organisms (Mángano and Buatois, 2014; Darroch et al., 2015). 1996; Smith et al., 2016). Similarly, a major discontinuity in the Sous
Therefore, the facies dependence of T. pedum, as well as the strati­ Basin is located at the base of the Tabia Member (Fig. 1C), which is in
graphic overlaps of complex bilaterian trace fossils with Ediacaran body agreement with the C isotope and fossil data.
fossils cause both the localization and the correlation of the

11
A. Azizi et al.
7. Conclusion
The Ediacaran–Cambrian boundary has long been recognized as an
important junction in Earth history. Despite this, it is still a poorly un­
derstood geological transition due to limited geochronological data and
the difficulty in correlating stratigraphic sections regionally and
globally.
The first occurrence of the Cambrian trace fossils from the base of the
Tifnout Member allows us to suggest at least a Fortunian age for the host
strata. Further evidence of Treptichnid traces found 3 m below the base
of the Tifnout Member supports a late Ediacaran age for the Tabia
Member.
The integration of trace fossils with carbon isotope δ13C records
suggests that the Ediacaran–Cambrian transition of the Sous Basin may
have occurred at the contact between the Tifnout and Tabia Members,
chemostratigraphically corresponding to the large negative δ13Ccarb
excursion that was previously suggested for the Ediacaran–Cambrian
boundary (BACE) in the western Anti-Atlas Mountains.
CRediT authorship contribution statement
Abdelfattah Azizi: Conceptualization, Formal analysis, Funding
acquisition, Investigation, Project administration, Writing – original
draft, Writing – review & editing. Asmaa El Bakhouch: . Abderrazak El
Albani: Writing – review & editing. Kalle Kirsimäe: Writing – review &
editing. Mouhssin El Halim: . Khadija El Hariri: Writing – review &
editing. Mohamed Erragragui: . Ahmid Hafid: . Olev Vinn: Writing –
review & editing.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
No data was used for the research described in the article.
Acknowledgments
The authors are grateful to anonymous reviewers for their
constructive comments. We thank Cadi Ayyad University, Académie
Hassan II des Sciences et Techniques and the Region Nouvelle Aquitaine
for their financial support. The authors are grateful to Gabriela
Mángano, Luis Buatois, E.H. Bouougri and Ursula Toom for their help in
trace fossils identification. Idir Elhabib, and Ibrahim Ouargaga are
thanked for their assistance during the field work. The analyses at the
University of Tartu were supported by the Estonian Centre of Analytical
Chemistry.
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