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Oases of biodiversity: Early Devonian palaeoecology at Hamar
Laghdad, Morocco
Christian Klug, Elias Samankassou, Alexander Pohle, Kenneth De Baets, Fulvio Franchi, and
Dieter Korn
With 22 figures and 3 tables
Abstract: The mudmound locality of Hamar Laghdad (Tafilalt Platform) contains possibly the greatest
palaeobiodiversity, both horizontally in the surrounding sediments and stratigraphically through the
Devonian sedimentary succession of the Moroccan eastern Anti-Atlas. From the Ordovician to the
Devonian, sediments of the Hamar Laghdad region and their fossil contents tend to differ slightly from
those in time-equivalent strata of other parts of the Tafilalt Platform. Our research provides a descrip-
tion of the sedimentological and palaeontological record at Hamar Laghdad covering the Lochkovian
to Givetian interval. We use alpha-diversity data based on macrofossils from selected fossiliferous
strata and compare the results with the relative abundance of bioclasts in polished slabs. We show that
the palaeobiodiversity at Hamar Laghdad was similar to that of the southern Tafilalt with a normal
diversity during the Pragian, i.e. prior to mudmound initiation and growth. By contrast, the layers
covering the mudmound carbonates (i.e. when their activity had ceased) show a high diversity during
the latest Emsian with a trophic nucleus comprising 36 species. This fauna, here exemplified by the
‘Red Fauna’ (from the Red Cliff), is dominated by benthic suspension feeders (corals, brachiopods and
crinoids) and cephalopods. The shift in biodiversity is most likely related to the local occurrence of a
favourable environmental setting, characterized by a temporally and spatially varying topography. It
is assumed that this setting was influenced by sea-level fluctuations, currents and nutrient availability.
Key words: Devonian, Tafilalt Platform, alpha diversity, bioclast diversity, palaeoecology, carbonate
microfacies, carbonate buildups.
1. Introduction
Hamar Laghdad (or Hamar El Khdad; Fig. 1) is world-
renowned among geoscientists for its prominently ex-
humed, mostly conical and well-preserved mudmounds
dubbed Kess-Kess referring to the shape reminiscent of
heaps of couscous (e.g., Clariond 1934a, 1934b; roCh
1934; Choubert 1952; Choubert et al. 1952; leMaitre
& P
oueyto
1955; l
e
M
aitre
1956; h
ollard
1960, 1963,
1967, 1974; Massa et al. 1965; Gendrot 1973; MiChard
1976; Wendt 1984; Wendt & aigner 1985; braChert
1986; J
oaChiMski
1986; t
öneböhn
1991; b
raChert
et
al. 1992; MounJi et al. 1998; belka 1998; JoaChiMski et
al. 1999; aitken et al. 2002; buggisCh & kuMM 2005;
Wendt & kaufMann 2006; Cavalazzi et al. 2007;
l
eda
2010; b
erkoWski
& W
eyer
2012; C
orMinboeuf
et al. 2013; berkoWski & Zapalski 2014; franChi et
al. 2015a, 2015b; sgavetti et al. 2015). A very detailed
literature overview was recently undertaken by b
eCker
et al. (2018b); not all of these details are repeated here.
This locality is situated in a series of east-west trend-
ing folds that expose the Ordovician to Carbonifer-
ous sedimentary succession of the eastern Anti-Atlas
(MiChard 1976; Wendt 1985, 1988; bourrouilh 1987;
©2018 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de
DOI: 10.1127/njgpa/2018/0772 0077-7749/2018/0772 $ 10.00
N. Jb. Geol. Paläont. Abh. 290/1-3 (2018), 9–48 Article
Stuttgart, November 2018
E
eschweizerbart_xxx
10 C. Klug et al.
piqué & MiChard 1989; piqué et al. 1993; bourrouilh
& bourque 1995; baidder et al. 2016). In particular, a
great number of synclines, including Bou Tchrafine,
occur east of the town of Erfoud (Fig. 2). Because of
the proximity to Erfoud, their easy accessibility and
high fossil abundance, outcrops in Bou Tchrafine have
been subject of many detailed studies (beCker & house
1994, 2000; beCker et al. 2018a and reference therein).
This ridge extends eastward, interrupted by a few dry
valleys (oueds). At Hamar Laghdad, the height of this
ridge increases to reach up to 180 m in elevation ac-
cording to bultynCk & Walliser (2000) as a result of
intermittent growth of mudmounds along the platform
(e.g., aitken et al. 2002).
From the Cambrian until the Carboniferous, this
region was part of the north-western shelf of Gond-
wana (ScoteSe 1997, 2014; Golonka & Gawęda 2012)
and was covered by an epicontinental sea. This palaeo-
geographic position, along with the moderate southern
latitude, allowed the repeated establishment of diverse
ecosystems (e.g., tönebohn 1991; kaufMann 1996,
1998; klug et al. 2008; de baets et al. 2010; van roy
et al. 2010; frey et al. 2014, 2018; eiChholt & beCker
2016; JakuboWiCz et al. 2018).
Devonian mudmounds do not occur only at Hamar
Laghdad, but also in other areas of Morocco (k
auf
-
Mann
1998) and further east, with examples from the
Middle Devonian of Algeria, considered by Wendt et
al. (1993, 1997) as the “world’s most spectacular mud-
mounds”. However, in terms of accessibility and fos-
sil abundance (hence the term ‘oases’ in the title, as
suggested earlier by berkoWski 2008), those of Hamar
Laghdad can compete with the Algerian examples: the
fossil abundance is extremely high, with hundreds of
trilobite species described (e.g., alberti 1969, 1970,
1980, 1982; MorzadeC 2001; Chatterton et al. 2006;
beCker et al. 2018; Crônier et al. 2018).
Apart from trilobites, many other fossil groups
have been reported from Hamar Laghdad in previous
works and others are described in this volume: corals
(berkoWski 2004, 2006, 2008, 2012, 2018; król et al.
2016, 2018), brachiopods (HalamSki & BalińSki 2018;
Mergl 2018), rostroconchs (aMler 2018), gastropods
(unpublished; determinations by J. frýda), bivalves
(hrynieWiCz et al. 2018), cephalopods (klug 2002;
pohle & klug 2018a, 2018b), ostracods (oleMpska &
b
elka
2010), crinoids (k
lug
et al. 2014; n
ardin
2018),
crinoids associated with infesting corals (berkoWski
& klug 2012; berkoWski & zapalski 2014), cystoids
(nardin 2018; Waters & klug 2018), placoderms
(lehMan 1956, 1976, 1978; antCzak & berkoWski
2017; rüCklin & CléMent 2017; rüCklin et al. 2018),
chondrichthyans (deryCke 1992, 2017; deryCke et al.
2014), and trace fossils of actiniarians (klug & hoff-
Mann 2018).
Conceived as a combination of an introduction for
this volume about Hamar Laghdad and a palaeoeco-
logical study, the aims of this paper are (1) to provide
an overview of the stratigraphy and palaeontology of
the sediments of Lochkovian to Eifelian age hosting
the Kess-Kess mudmounds, (2) to compare palaeoeco-
logical data obtained from polished slabs with those
from macrofossil samples to reveal changes in palaeo-
ecology through the Early Devonian, (3) to put these
fossil-based data into a sedimentological context and
(4) to evaluate the interplay of environmental changes
and mudmound growth.
Fig. 1. The Hamar Laghdad ridge seen from the south. The high ridge on the left (west) is composed of the Zlíchovian crinoid
and coral limestones. The incision to the right of the inclined surface formed due to a set of faults. The hills visible towards
the right (east) are the Kess-Kess mudmounds.
eschweizerbart_xxx
Oases of biodiversity 11
2. Material and methods
We compiled a data base of palaeontological information
from the Lochkovian to Eifelian strata at Hamar Laghdad in-
cluding new unpublished facies and palaeoecology descrip-
tions. This data base has been then investigated to highlight
fluctuations in palaeoecology throughout the sequence. The
new unpublished data have been collected applying three
different methods:
1. Facies:
We cut and polished rock samples from the Lochkovian to
Eifelian succession in order to determine the carbonate fa-
cies following d
unhaM
(1962). The facies types are informa-
tive with respect to water energy and other palaeoecological
factors (flügel 2004).
2. Analyses of the bioclast content in polished slabs:
We examined the bioclast content seen in the polished slabs
semiquantitatively (explanation below). We chose this ap-
proach in order to obtain palaeoecological information fill-
ing the big gaps between the macrofossil faunules included
in the analyses of the third approach described below.
Polished slabs (50 to 100 mm wide) were used to quan-
tify the abundance of bioclast types in a defined surface area
(400 mm
2
). Bioclasts were assigned to fossil groups, usually
at a much lower systematic resolution than in the case of
approach 3. These bioclasts were, where possible, assigned
to palaeoecological guilds, but at a lower resolution than
that presented by bush et al. (2007). Bioclasts were ordered
mainly according to their tiering, i.e., from deep infaunal
as one extreme to those animals inhabiting the upper part
of the water column, from suspension feeders to predators,
and from fixed sessile to fast swimming nekton. Cumulative
histograms were generated, in which bioclast abundance is
ordered as follows: Infauna (if present) – Tabulata – Rugosa
– Brachiopoda – Gastropoda – Ostracoda – Trilobita – Cri-
noida – Dacryoconarida – Orthoceratoidea – Bactritida –
Ammonoidea – Placodermi.
Fig. 2. Maps of Morocco, the Tafilalt and the Hamar Laghdad. The two small maps show the localisation and the larger
map reveals geological and palaeontological details of Hamar Laghdad. The smaller maps were modified using maps of
klug (2002) and klug et al. (2016). The larger map is based on a detailed map published in belka (1998) and JakuboWiCz
et al. (2013, 2015).
eschweizerbart_xxx
12 C. Klug et al.
Although 400 mm
2
is a relatively small area, the number
of fossils counted in each sample varies between 20 and 280.
The fossil content shows strong lateral variation within the
same layer or stratum. Therefore, the data obtained can only
be considered as an approximation of the bioclasts assem-
blage of the different strata. This approach cannot perfectly
reflect the composition of the bioclastic assemblage of each
stratum and this is thus considered a semiquantitative ap-
proach. Wherever possible (polished surface 800 mm2 or
larger), up to three different areas of 400 mm2 have been
investigated in order to reduce the sampling bias. The aver-
age values for both the bioclast group and the number of
fossils have been used for statistical reasons (Table 2). In the
graphical representation of the statistics, the data related to
nekton and plankton are plotted aginst a blue field, whereas
data related to benthos are plotted in a yellow field for the
sake of clarity.
Components such as isolated echinoderm and trilobite
sclerites were counted, although it cannot be ruled out that
these belonged originally to only one single specimen. This
has to be taken into account when comparing these results
to those based on macrofossils (approach 3). Additionally,
daryoconarids and ostracods were not included in the macro-
fossil analyses (approach 3), while the bioclast determination
according to approach 2 permitted determination at species
level only in a few exceptional cases.
3. Macrofossil sampling of faunas:
We collected macrofossils from distinct limestone beds and
marly intervals indicated in the sections (Tables 2, 3). All
fossils were determined as far as possible and assigned to
ecological guilds in order to quantitatively evaluate the pal-
aeoecological factors and their changes that occurred be-
tween the Lochkovian and Eifelian. For this purpose, we
collected as many macrofossils as possible from six layers
of this stratigraphic interval. It must be noted that the latest
Emsian assemblage, here dubbed Red Fauna (referring to its
red colour), was sampled during several field trips between
1996 and 2017, while the Pragian strata were sampled only
during one single field campaign in 2012. The Red Fauna
was collected at the Red Cliff (k
lug
2002; k
lug
et al. 2009,
2014); due to its peculiar palaeoecological, diagenetic history
and marly matrix, its fossil content can be extracted rela-
tively easily. Faunas of more or less the same age and similar
composition occur also elsewhere at Hamar Laghdad, but at
the Red Cliff, these uppermost Emsian strata are particularly
well-exposed and, due to the unique position of this site, this
fauna might be especially diverse (see the detailed discus-
sion further below). Nevertheless, future sampling at other
sites may reveal even higher diversities; thus, the Red Cliff
represents an example of an assemblage of latest Emsian
age, which is interesting for the reconstruction of the post-
mudmound evolution of the basin.
Because of the more intense sampling, the number of
specimens of the Red Fauna is much higher (>3000) than that
of the Pragian faunas (between 79 and 178 specimens). All
macrofossils were assigned to taxa, where possible to species
level, and specimens were counted per taxon. Taxonomic
determinations were carried out for Tabulata by J
an
k
ról
and M
ikolaJ
z
apalski
, for Rugosa by B
łazej
B
erkowSki
, for
Brachiopoda by adaM t. halaMski, andrzeJ balinski and
M
iChal
M
ergl
, for Rostroconchia by M
iChael
a
Mler
, for
Bivalvia by k
rzysztof
h
rynieWiCz
and m
icHał
j
akuBowicz
,
for non-ammonoid Cephalopoda by AP, for Ammonoidea
by CK and KDB, for Trilobita by Catherine Crônier, Mor-
gane
o
udot
and r
aiMund
f
eist
, for Echinodermata by e
lise
nardin and Johnny Waters and for placoderms by Marti n
rüCklin. The according papers are included in this volume
(aMler 2018; berkoWski 2018; Crôni er et al. 2018; feist
& belka 2018; HalamSki & BalińSki 2018; hrynieWiCz et
al. 2018; klug & HoffMann 2018; król et al. 2018; Mergl
2018; nardin 2018; pohle & klug 2018b; rüCklin et al.
2018; Waters & klug 2018; zapalski, et al. 2018; zapalski
& klug 2018).
Fossil fragments were usually considered for statistical
purposes only when at least half of the entire fossil organ-
ism was preserved. In the case of crinoids, only calyces were
counted. Trilobites also are often preserved as isolated scle-
rites and, thus, only complete specimens and cranidia were
counted. In the case of tabulate corals, we counted also sin-
gle branches. Corals are so abundant that their number can
hardly be underestimated; although counting each single co-
ralite would have been more appropriate, we counted whole
colonies and significant fragments of colonies. This method
might seem conservative because tabulate coral abundance
might become underestimated. When looking at the results,
however, corals play a very important role as visible in the
diagrams produced for the Red Fauna (Table 3).
Histograms were produced to represent taxa according to
their abundance (number of specimens in the collection), to
determine the trophic nucleus of each assemblage (for details
on the trophic nucleus-concept, see n
eyMan
1967; f
rey
et al.
2014, 2018). In a second step, each taxon was assigned to an
ecological guild following b
ush
et al. (2007). Further details
of this method are provided in frey et al. (2014). For each
assemblage, pie charts were plotted depicting the proportion
of specimens that belong to the respective guild. In order to
estimate potential biases in sampling, we rarefied our data,
using the freely available software PAST 3.19 (h
aMMer
et
al. 2001; https://folk.uio.no/ohammer/past/).
Where data from all three approaches are available, the
results were compared with each other in order to evalu-
ate the validity of the results obtained using these three
methods.
Material: Except for the material photographed in the field,
all specimens described and figured herein are housed in the
Paläontologisches Institut und Museum of the University of
Zurich (PIMUZ numbers).
3. Palaeoecological analyses
3.1. Facies changes through the Devonian
The base of the Devonian sedimentary succession is not
well-exposed at Hamar Laghdad (bultynCk & Wal-
liser 2000) except for some small outcrops in fossil
exploitation sites. In most parts, the “Scyphocrinites
Limestone” of the Silurian-Devonian boundary is cov-
eschweizerbart_xxx
Oases of biodiversity 13
ered by talus, but the surrounding sedimentary succes-
sion likely resembles that of Ouidane Chebbi (b
elka
et
al. 1999) to the east and Bou Tchrafine (hollard 1974;
beCker & house 1994, 2000) to the west of Hamar
Laghdad. In the Tafilalt region, the earliest Devonian
succession is normally represented by argillaceous to
marly sediments with orthocerids, dacryoconarids, and
graptolites (hollard 1974; belka et al. 1999; frey et
al. 2014). It is not clear to what extent the Lochkovian
sedimentary succession is recorded at Hamar Laghdad,
because most of the local Lochkovian deposits (Oumej-
joud Formation; b
eCker
et al. 2018b) consist of subma-
rine volcanites (peperites; Figs. 2-6), which formed “a
volcanic submarine high” (aitken et al. 2002; braCh-
Fig. 4. Satellite image (Google) of the eastern part of Hamar Laghdad ridge, where all mudmounds are situated. Note the
SW-NE-trending faults and the reddish erosional surface that shows the level to which the post-Visean-erosion removed
the Hamar Laghdad carbonates. The red colour is linked with dolomitization of the carbonates and hematite infiltration.
Fig. 3. Satellite image (Google) of the western part of Hamar Laghdad ridge. Note how the thick-bedded Zlíchovian crinoid
limestones of Hamar Laghdad wedge into the Deiroceras Limestone and the Erbenoceras Limestone.
eschweizerbart_xxx
14 C. Klug et al.
ert et al. 1992; franChi et al. 2015a; JakuboWiCz et al.
2015).
The interval of submarine volcanic eruptions ended
during the latest Lochkovian, when carbonate sedimen-
tation has resumed along the western edge of Hamar
Laghdad. The Lochkovian rocks consist of black lime-
stones (orthocerid floatstone), which contain current-
aligned orthocerids in rock-forming number, locally
accompanied by bivalves. Similar deposits occur char-
acteristically in most sections of the eastern Anti-Atlas
(e.g., belka et al. 1999; bultynCk & Walliser 2000;
frey et al. 2014).
As documented by several authors, including some
in the present volume (Massa et al. 1965; hollard
Fig. 5. Schematic section through the Hamar Laghdad ridge, modified after hollard (1967). A – Geological overview. B –
Main chrono- and lithostratigraphic terms used in the text.
eschweizerbart_xxx
Oases of biodiversity 15
1981; b
elka
et al. 1999; k
röger
2008; l
ubeseder
et
al. 2009; frey et al. 2014; klug et al. 2015; pohle &
klug 2018a, 2018b), carbonate content and palaeobio-
diversity increased around the Lochkovian-Pragian
boundary. The Pragian succession is poorly exposed
at Hamar Laghdad except for the western end of the
ridge (J
akuboWiCz
et al. 2013; b
erkoWski
& z
apalski
2014), where the overlying thick-bedded Emsian car-
bonates were eroded (Figs. 2, 3). These strata consist of
floatstones, wackestones and packstones rich in dacryo-
conarids, crinoids and brachiopods with subordinate
corals, gastropods, ostracods and trilobites. Through
the Pragian, facies changed from predominantly wacke-
stone to packstone and the biota from pelagic-dominat-
ed (orthoconic cephalopods) to benthic-dominated (e.g.,
crinoids; berkoWski & zapalski 2014).
The identification of the Pragian-Emsian bound-
ary is hampered by the scarcity of index fossils and
the on-going discussion about its stratigraphic position
(haude & Walliser 1998; Carls & valenzuela-ríos
2007; C
arls
et al. 2009; a
boussalaM
et al. 2015). In
this study, the Pragian interval corresponds to the “Pra-
gian Limestone” of a
boussalaM
et al. (2015), which
possibly contains the Pragian-Emsian boundary (see
also bultynCk & Walliser 2000). This is relevant be-
cause the overlying crinoid and stromatactis limestone
Fig. 6. Submarine volcanites (peperites) of Lochkovian age (Oumejjoud Formation) underlying the thick-bedded carbonates
at Hamar Laghdad. A – The Hamar Laghdad ridge as seen from the Northwest; note the thick greyish carbonates ontop,
underlain by the greenish to brownish peperites below; the arrow points at an angular unconformity of unknown origin
within the peperites. B – Carbonatic Neptunian dykes crossing the peperites. C, D – spheroidal weathering of the peperites.
eschweizerbart_xxx
16 C. Klug et al.
Fig. 7. Hollard Mound. A – Seen from the southeast. Note how the the Eifelian limestones gain in thickness towards the seep.
B – Mass occurrence of trilobite exuviae near the vent. C – Limestones with seep-bivalve conchs arranged concave-upward;
insert: an isolated specimen of the seep-bivalve Ataviaconcha wendti from the same place. D – Vein-filling with alternations
of dark and light grey cements. E – Mound core with weathering dyke-like structures.
eschweizerbart_xxx
Oases of biodiversity 17
(mostly crinoid and coral packstones) are not of Pragian
age, at least not in the western part of Hamar Laghdad.
As far as the stratigraphy of the thick-bedded
lower Emsian carbonates (Kess-Kess Formation) is
concerned, the base appears to correlate with the Em-
sian Deiroceras Limestone of other localities in the
Tafilalt like Ouidane Chebbi (belka et al. 1999; de
baets et al. 2010), Filon 12 (frey et al. 2014; klug &
p
ohle
2018) or Bou Tchrafine (a
boussalaM
et al. 2015;
beCker et al. 2018a). aboussalaM et al. (2015, text-fig.
13) reported a massive crinoid limestone at the base
of the Deiroceras Limestone, which corroborates the
correlation used in this study. Also, when examining
the lateral continuation of the Lochkovian to Emsian
strata in satellite images (Figs. 3, 4), the thick-bedded
limestones of the Kess-Kess Formation wedge into the
Zlíchovian (Lower Emsian) Deiroceras- to Mimagoni-
atites Limestone succession (Fig. 5; compare h
ollard
1967: 223). The latter correlation is corroborated by the
presence of Mimagoniatites fecundus in the uppermost
strata of the mudmounds (k
lug
et al. 2009). Further
details of the formation of the thick-bedded carbon-
ates of the Zlíchovian and the geological history of the
mudmounds at Hamar Laghdad are discussed in detail
by belka (2018).
After the deposition of the Zlíchovian carbonates
and mudmound formation, the influence of the eus-
tatic sea-level rise of the Daleje Event caused a rapid
relative increase in clay content and a strong decrease
in carbonate sedimentation (e.g., Massa et al. 1965;
haq & sChutter 2008; babek et al. 2017). Much of the
Dalejan (late Emsian) is correspondingly represented
by marls and marly limestones (crinoid and dacryo-
conarid wackestones). This interval is of particular
interest because its sediments cover the inter-mound
area and pinch out against the mounds (Fig. 6; see also
b
erkoWski
2008, 2012). Consequently, thickness and
facies vary significantly over short distances. Current
regimes, water depth, possibly hydrothermal activities
and thus likely the ecological conditions also varied
strongly laterally (berkoWski 2008, 2012).
Near the Emsian-Eifelian boundary, the carbonate
production increased again. The upper Emsian rocks
consist of claystones, marls and limestones (mostly
dacryoconarid and crinoid wackestones) with various
cephalopods including anarcestids (e.g., aboussalaM et
al. 2015). Sedimentation during the Choteč transgres-
sion is characterized by dark nodular limestones that
contain a diverse cephalopod assemblage. This more
argillaceous interval is overlain by thick-bedded ceph-
alopod-rich limestones (dacryoconarid wackestones),
which is a characteristic feature for this interval across
the Tafilalt Platform (Wendt 1985, 1988; klug 2002;
lubeseder et al. 2010).
Above the thick-bedded limestones, some meters of
mostly nodular, more or less marly cephalopod lime-
stone follow (for the carbonate content see M
assa
et
al. 1965). The upper Emsian sediments did not fully
obliterate the topography that was created by the mud-
mounds during the early Emsian (berkoWski 2008,
Fig. 8. Givetian and Frasnian sediments and fossils. A – Trace fossils (Thalassinoides cf. paradoxicus; identified by l
othar
h. v
allon
, Faxe, Denmark) in Givetian limestones. Due to the coarser material in the burrows, hydraulic conductivity is
probably slightly higher, thus facilitating the formation of desert varnish. In turn, the latter makes the burrows more resistant
to erosion (hammer is 30 cm long). B – Frasnian limestone layer with bioturbation, a multilobate beloceratid ammonoid and
an orthocerid. Width of the image is about 20 cm. C – Detail of B showing a weathered beloceratid.
eschweizerbart_xxx
18 C. Klug et al.
2012; k
ról
et al. 2016). Accordingly, Middle Devonian
sediments continued to fill the inter-mound depressions
between the mudmound summits. Since mound growth
continued after the Emsian in the eastern part of Hamar
Laghdad (Hollard Mound/ Mound 50: Fig. 7; berkoWs-
ki
2006; J
akuboWiCz
et al. 2013, 2014, 2015; h
rynie
-
WiCz et al. 2016; belka et al. 2018), these sedimentary
unconformities continued to develop into the Late De-
vonian (e.g., h
ollard
1967). Accordingly, facies differ-
ences can be encountered across very small distances
(tens of meters) in the Emsian and more rarely from
the Eifelian to the Frasnian interval (only in eastern
Hamar Laghdad; e.g., Massa et al. 1965; töneböhn
1991). In turn, this palaeotopography likely created a
multitude of different habitats during the Emsian with
a broad range of varying physical properties includ-
ing water depth, energy, sediment accumulation and
probably also light (h
ebbeln
& s
aMankassou
2015).
Additionally, fluids advection to the seafloor continued
into the early Eifelian at the Hollard Mound, which, ac-
cording to the isotopic signatures, represents an ancient
methane seep (b
erkoWski
2006; C
avalazzi
et al. 2012;
JakuboWiCz et al. 2014a, 2014b; hrynieWiCz et al. 2017).
Reduced sedimentation rates are a characteristic
feature for most sections of the Middle and Late Devo-
nian of the Tafilalt Platform (W
endt
1985, 1988; t
öne
-
böhn 1991). This condensation is absent or less distinct
in most parts of the Lochkovian to Emsian succession.
By contrast, starting during the late Emsian, the sedi-
ment accumulation rates on the platforms and basins
in the eastern Anti-Atlas differentiated increasingly
(Massa et al. 1965; hollard 1967, 1970, 1974; Wendt
1985, 1988; kaufMann 1998; lubeseder et al. 2010).
The Eifelian succession ends with a more argilla-
ceous interval including limestone nodules (Fig. 5). In
the Givetian strata, abundant trace fossils speak for a
well-oxygenated seafloor (Fig. 8A). The more thickly
bedded Givetian limestones (dacryoconarid wacke-
stones in layers of 10 to 40 cm thickness) contain ceph-
alopods, which increase in abundance up the section.
The cephalopod limestone facies extends into the upper
Frasnian (Fig. 8B). In most parts of Hamar Laghdad,
the outcropping succession ends with the dark cepha-
lopod limestones of the Kellwasser Limestone (W
endt
Fig. 9. Gastropods from the Pragian of Hamar Laghdad. A – Palaeozygopleura sp. B – Eohormotomina restisevoluta. C –
Australonema sp. D – Oriomphalus sp. E – Paraoehlertia sp. F – Platyceratidae indet. with a rugose coral in the aperture.
Fig. 10. The latest Lochkovian to earliest Emsian succession in the west of the Hamar Laghdad ridge with the proportional
bioclast abundance (Method 2, Table 2), carbonate microfacies (Method 1) and polished rock samples from some strata.
eschweizerbart_xxx
Oases of biodiversity 19
Fig. 10.
eschweizerbart_xxx
20 C. Klug et al.
Fig. 11. The thick-bedded mudmound-carbonates (late early Emsian, latest Zlíchovian) and their fossil content. A, B –
stromatactis with branching tabulate corals. C – Large fragment of a phragmocone of the actinocerid Deiroceras hollardi.
D – The index ammonoid Mimagoniatites fecundus near the summit of a mudmound. E – Aggregation of over thirty short-
stemmed cystoids, probably more or less autochthonous.
eschweizerbart_xxx
Oases of biodiversity 21
Fig. 12. Mudmounds at Hamar Laghdad. A – The crescent shaped mud-ridge surrounding the Red Cliff seen from the
northwest; note the plateau of the Hamada du Guir at the horizon (Late Cretaceous). B – Completely exhumed conical mud-
mounds in the center of the Hamar Laghdad ridge. C – A stromatactis-like cavity in the mound carbonates with a lumachelle
of exuviae of the trilobite Scutellum. D – This mudmound is still covered by Dalejan and Eifelian sediments except for its
summit. E – Branching tabulate corals in a mound. F – A small mound or bioherm inhabited by many tabulate corals.
eschweizerbart_xxx
22 C. Klug et al.
& b
elka
1991). This outcrop situation can be explained
by the increase in argillaceous deposition after the
Frasnian. These more argillaceous Famennian and
Tournaisian sediments were eroded and subsequently
covered by scree (Figs. 3, 4).
3.2. Bioclast abundance in the Lochkovian to
Emsian sediments
As mentioned above, the Devonian sedimentary re-
cord at Hamar Laghdad begins with deposits of latest
Lochkovian age. These deposits are characterized by
nodular limestones with highly abundant orthocerids,
dominated by representatives of the genus Temper-
oceras (kröger 2008; klug et al. 2015). Additionally,
bivalves occur, along with rare gastropods (Fig. 9).
The Pragian interval is marked by a shift in bio-
clast composition from pelagic towards predominantly
benthic organisms (Fig. 10). Probably, two factors are
responsible for this change, namely a decrease in wa-
ter depth along with an improved oxygenation of the
lower part of the water column and the upper part of the
sediments (frey et al. 2014). Initially, the brachiopods
contributed strongly to the increasing benthos abun-
dance, later followed by crinoids. The latter reached
rock-forming abundance near the Pragian/Emsian
boundary (Fig. 10).
Thick-bedded limestones dominate the basal Em-
sian succession. The fossil content changed from cri-
noid-dominated at the base to a more variable biofacies
at the top. The surface of this limestone is very well
exposed (Figs. 1, 3, 4, 6A). Consequently, small-scale
differences in biofacies are easily detectable. In many
places, this bedding plane exposes a stromatactis-rich
limestone as it is characteristic for mudmounds with
abundant tabulate corals and crinoid remains (Fig.
11A, C). Locally, small tabulate bioherms formed small
positive reliefs with diameters ranging between half a
meter and a few meters (Fig. 11F). Remains of large
actinocerids (Deiroceras hollardi; Fig. 11C) reaching
lengths of over 1.5 meters were reported (klug et al.
2015; p
ohle
& k
lug
2018a, 2018b). Remarkably, the
cystoid genus Eucystis formed small meadow-like ac-
cumulations with some tens of individuals (Fig. 11E).
On the same bedding plane, numerous Neptunian dykes
with banded infills (cements and internal sediments)
occur, interpreted to testify for hydrothermal activity
at that time (belka 1998; belka et al. 2018).
The sedimentary record of the Daleje transgression
and the following highstand (b
eCker
& h
ouse
1994;
aboussalaM et al. 2015; babek et al. 2017) is well-ex-
posed above the thick-bedded mound limestones (Fig.
12), while in many other localities of the Tafilalt, this
part is covered by debris from younger sediments or
the Mimagoniatites Limestone (klug 2017). In this
context, it has to be stressed that especially the oldest
sediments deposited during this transgression tend to
wedge out at the lower parts of the mound flanks (Fig.
12A, D). At the Red Cliff (k
lug
2002), the layers above
the mound strata are rich in pelagic bioclasts, particu-
larly dacryoconarids (Figs. 12A, 13). This changed rap-
idly in the younger sediments, and benthonic bioclasts
become increasingly important (Figs. 13, 14). Initially,
there were less bioclasts of benthic than of pelagic or-
ganisms but during the latest Emsian, benthic organ-
isms became much more important. Near the Emsian-
Eifelian boundary, planktonic and nektonic bioclasts
dominate again (Figs. 14, 15). Above the Choteč Event
level, large bivalves appear to become abundant in
Hamar Laghdad, although we have not quantified this
since our main focus was the Lochkovian to Emsian
interval.
Fig. 13. Succession dated from the early late Emsian (basal Dalejan) succession in the center of the Hamar Laghdad ridge
(Red Cliff) with the proportional bioclast abundance (Table 1), carbonate microfacies and polished rock samples from some
strata. Note the interval that yielded the Red Fauna.
eschweizerbart_xxx
Oases of biodiversity 23
Fig. 13.
eschweizerbart_xxx
24 C. Klug et al.
Fig. 14. Relative proportions of pelagic versus benthic fauna in all investigated samples (Method 2, Table 1). Age assign-
ments and possible correlations are listed. The error bars are the 95 % binomial confidence intervals (following raup 1991
and d
e
b
aets
et al. 2012) calculated using the binom.confint function of the Binom Package in R (using the exact approach).
Fig. 15. The latest Emsian (late Dalejan) to Eifelian sequence in the central part of the Hamar Laghdad ridge (Red Cliff)
with proportional bioclast abundance (Method 2, Table 1), carbonate microfacies and polished rock samples from some
strata. Layers exploited for trilobites are marked in pink.
eschweizerbart_xxx
Oases of biodiversity 25
Fig. 15.
eschweizerbart_xxx
26 C. Klug et al.
Fig. 16.
eschweizerbart_xxx
Oases of biodiversity 27
3.3. Alpha diversity fluctuations during the Early
Devonian
Four macrofossil samples from the Pragian and one as-
semblage of latest Emsian age have been analysed for
alpha diversity. Based on data from Hamar Laghdad,
the overall diversity appears to have been low in the
latest Lochkovian with dominating orthocerid cepha-
lopods (Fig. 16A). This holds true for other localities
in the Tafilalt as well (b
elka
et al. 1999; f
rey
et al.
2014), assuming a similarity in biofacies for these sites.
Benthic forms occurred there in low numbers. With the
beginning of the Pragian stage, sea level fell (h
aq
&
s
Chutter
2008)
and, probably, oxygenation of the water
near the sediment surface increased, as reflected in the
rising abundance and diversity of benthic forms (Figs.
14, 17). Alternatively, this might be linked to changes
in other physical properties of the sediment, such as
grain size, sorting, and cohesiveness and early or late
lithification; the missing infaunal macrofauna makes
Fig. 17. Faunal composition of four Pragian macrofossil associations (A to D, Method 3, Table 2) and of cross sections
counted in 4 cm2-surfaces of polished rock samples (sample numbers 4, 6, 10, 12, 16; Method 2, Table 1) from roughly the
same stratigraphic interval. The temporal order of the samples is indicated by the grey arrows. Sample size is indicated at
the top left of each pie-chart. Planktonic forms are represented by segments in blue colours. Note the strong decrease in
proportion of planktonic organisms between the latest Lochkovian and the earliest Emsian. The number of ecological guilds
sensu bush et al. (2007) stayed more or less constant.
Fig. 16. Histograms and bean curves (rarefaction analyses) of the Pragian macrofossil assemblages (Method 3, Table 2). The
percentage of specimens is given on the vertical axis. Note the low variation in the number of species forming the trophic
nucleus. It is unclear to which degree the trophic nucleus is so much smaller than in the Red Fauna due to strong differences
in sampling effort (for each fauna, a rarefaction analysis was carried out; the according diagrams are shown next to each
histogram with a grey arrow). In any case, the degree of ecological change during the Pragian was possibly low and maybe
driven by a successive reduction in water depth.
eschweizerbart_xxx
28 C. Klug et al.
this seem a less likely explanation, which is further fal-
sified by trace fossils (e.g. the patchy appearance of the
sediment in the sections in Fig. 10). Also, water depth
and changes in light availability might have played a
pivotal role in improving the ecological conditions for
benthos. The Pragian faunules A and B (Figs. 16, 17)
are still dominated by macro-remains of pelagic organ-
isms (which coincides with the bioclast composition in
samples 4 to 10), both in specimen numbers and in spe-
cies; the maximum in both values was found in faunule
B (Table 2). Faunules C and D contain less than 50%
fossils of pelagic organisms, although both layers still
yielded more than 10 cephalopod species. The overly-
ing mound carbonates are then more strongly domi-
nated by benthic organisms such as crinoids and corals.
The number of benthic species increases from 16
in faunule A to the maximum of 22 in B (see examples
in Fig. 9). Subsequently, the diversity of benthic organ-
isms slightly decreases to 18 in faunule C and 17 in
faunule D (Figs. 13, 14). Remarkably, the number of eco-
logical guilds occupied by members of these faunules
hardly varies through the Pragian. Neither remains of
fully infaunal nor of nektonic species were identified in
the Pragian faunules. It has to be pointed out, though,
that the samples are small (further sampling would, for
example, likely reveal remains of Machaeracanthus,
which would proove the presence of the nektonic guild).
The basal Emsian assemblages were not studied in
detail. However, investigations in the mound carbonates
(Kess-Kess Formation) did not reveal any fully infaunal
or nektonic forms. As expected, epifaunal organisms
such as crinoids, tabulates, rugosans and cystoids domi-
nate the fauna (Fig. 11). Small cavities (Alberti 1982;
feist & Chatterton 2015; FranChi et al. 2015a; belka
et al. 2018) were used by scutellid trilobites for moult-
ing (Fig. 12C). Both at the base and on the top of the
mudmound limestones, cephalopods occur in moderate
numbers. At the top, cephalopods are represented by
large fragments of the actinocerid Deiroceras hollardi
(Fig. 11C) and rare specimens of the ammonoid species
Mimagoniatites fecundus (Fig. 11D).
The sediments overlying the mudmound carbonates
were only sampled for alpha diversity below the Em-
sian and Eifelian boundary. In spite of this incomplete
sampling, the information obtained from bioclast oc
-
currences and relative macrofossil abundances provides
some insight into diversity changes. There are a some
common trends that became evident from the bioclast
analyses and the alpha-diversity in the latest Emsian:
At the Red Cliff, the first layers above the thick-bed-
ded limestones are dominated by pelagic organisms
(dacryoconarids). Although the proportions fluctuate,
the abundance of benthic forms increases until the
development of the Red Fauna during the latest Em-
sian, which will be described below (Figs. 13-15). The
Red Fauna comprises a very broad range of ecological
guilds sensu bush et al. (2007), ranging from fully nek-
tonic (placoderms) to semi- or shallow infaunal (some
bivalves and rostroconchs).
4. Palaeoecological changes during the
Early Devonian
The main changes in palaeoecology from the late
Lochkovian to the Eifelian depicted in the present
study correspond well with those reported by frey
et al. (2014) for the southern Tafilalt Platform west of
Taouz. In the southern Tafilalt, the Lochkovian succes-
sion is dominated by dark claystones and marls with
few limestone intercalations poor in benthic organisms
and rich in plankton such as orthocone cephalopods,
dacryoconarids and graptolites. This reflects moder-
ately deep marine shelf conditions with poorly oxygen-
ated conditions in the bottom waters. In the southern
Tafilalt, towards the base of the Pragian succession, the
sediments changed as shown by the shift from from
dark, carbonate-poor claystones to light grey marls and
limestones. The increasing abundance and diversity of
benthic organisms might also point to rising oxygen
levels (cf. edWards et al. 2017; Wood & erWin 2017).
Taking these facts together, a slow sea-level fall appears
to be the most likely driver of improved life conditions
at the sea-floor, affecting both faunal compositions and
lithofacies (Massa et al. 1965; alberti 1981; belka et
al. 1999; b
ultynCk
& W
alliser
2000; k
röger
2008;
frey et al. 2014).
Although sufficiently well preserved and diverse,
the Pragian faunules A and B (Figs. 10, 16, 17) did not
reach the high species-level diversity reported in frey
et al. (2014) in the southern Tafilalt; when rarefied to
150 specimens, there are 32 species in Pragian faunule
B compared 51 to species in the Pragian faunule of
frey et al. (2014). It is not clear whether this discrep-
ancy is real (ecological and primary) or originates from
differences in sampling effort (certainly important),
results of a taphonomic bias or represents the effect
of condensation. Regardless, the composition of the
Pragian faunules described here appears to be fairly
similar to that reported in frey et al. (2014) from the
Pragian of the southern Tafilalt with several species in
common.
eschweizerbart_xxx
Oases of biodiversity 29
Close to the Pragian-Emsian boundary, the simi-
larity of biofacies and lithofacies of Hamar Laghdad
and other parts of the Tafilalt Platform significantly
decreases due to the presence of thick-bedded crinoid-
and coral-rich limestones, and, further up in the section,
the mudmounds. The only sedimentary unit compara-
ble to the thick-bedded limestones at Hamar Laghdad
(Kess-Kess Formation) in age and – to a lesser degree
– in facies is the thick Deiroceras Limestone (which is
also rich in crinoid remains) that crops out throughout
much of the Tafilalt Platform (klug et al. 2013).
At the base of the Kess-Kess Formation, crinoids
occur in rock-forming quantity. This phenomenon is
known from several mudmound localities in the Anti-
Atlas (kaufMann 1998) and elsewhere (pratt 1995).
kaufMann (1998: 44) stated “Slightly elevated tem-
peratures may have stimulated the benthic fauna, es-
pecially crinoids, forming flat in situ lenses, which in
turn served as substrates for microbial colonization.”
At Hamar Laghdad, this lens reached an important size
with a lateral extension (E-W) of over five kilometres
and 180 m thickness (Massa et al. 1965; bultynCk &
Walliser 2000). According to belka (1998), only the
southern edge of this lens is preserved and the centre
was 1.5 km north of Hamar Laghdad. Assuming a di-
ameter of 3 km (as estimated by belka 1998; possibly
Fig. 18. Estimation of the maximum number of mudmounds based on data and a map of belka (1998). The dark lines indi-
cate the faults documented by belka (1998), which were extrapolated to the north to identify the approximate center of the
carbonates of the Kess-Kess Formation. The two outer concentric circles (3 and 5 km), indicate the potential original extend
of the Kess-Kess Formation; in the 3 km-circle, 40 mudmounds are shown per 1 km2. This would correspond to more than
700 mudmounds.
eschweizerbart_xxx
30 C. Klug et al.
reaching 5 km diameter), we suggest that the lens (as-
suming that it was circular and correspdonds in surface
extension to the area of the laccolith intrusion) would
have covered between 7 and 20 km2. Such an accumu-
lation of crinoids likely formed because of the favour-
able conditions provided by the submarine elevation
on the Tafilalt Platform resulting from the Lochkovian
submarine volcanites and the late volcanic hydrother-
malism that provided nutrients. This underwater relief
may also have promoted a high-diversity association
dominated by filter feeders (cf. roberts et al. 2006,
2009; de Clippele et al. 2018).
The cause of the transition from an elevation popu-
lated by crinoids to the nucleation and growth of large
mudmounds remains poorly understood. The sea-level
rise in the early Emsian, during which the so-called
Erbenoceras-Metabactrites Shale was deposited in
the basins of the eastern Anti-Atlas (aboussalaM et
al. 2015), may be responsible for the reduced crinoid
growth, perhaps simultaneously fostered carbonate
production around the main hydrothermal vents. As-
suming that (1) mudmounds were distributed evenly on
this crinoid limestone lens (20 to 40 per km2), (2) only
10 to 20 % of the crinoid limestone lens is preserved
today, (3) the crinoid lens had an approximately circular
shape, and (4) we can speculate that there were possibly
as many as 100 (for a ratio of 20 mounds per km2 and
an area of 7 km2) to over 700 (for a ratio of 40 mounds
per km2 and an area of 20 km2) mudmounds at Hamar
Laghdad (Fig. 18).
As evidenced by the plethora of fossils embedded in
the mudmound limestones including soft-bottom dwell-
ers such as many tabulate and rugose corals, cystoids
and crinoids, the palaeogeographic and palaeoecologi-
cal setting provided favourable living conditions for
these organisms. Most of these organisms were sus-
pension feeders, which likely profited from the wealth
of micro-organisms that, in turn, thrived because of the
minerals and nutrients provided from the hydrothermal
fluids (e.g., berkoWski
2006, 2008, 2012; frey et al.
2014; JakuboWiCz et al. 2014b; hrynieWiCz et al. 2017,
2018; k
lug
& H
offMann
2018). Currents bringing in
nutrients and food particles were likely also important
for the fauna dwelling at Hamar Laghdad, but with the
insufficient data at hand, the respective role of these
two factors can hardly be discriminated. Also, the
previously mentioned scutellid trilobites continued to
find caves for moulting (f
ranChi
et al. 2014; f
eist
&
C
hatterton
2015; b
elka
et al. 2018; f
eist
& b
elka
2018). The occurrence of the giant actinocerids (Fig.
8C) in the Kess-Kess Formation can probably be ex-
plained by their supposed preference of moderately
shallow environments (cf. kröger et al. 2009; Manda
& fryda 2010); during the Early Devonian, the area of
Hamar Laghdad probably represented an elevation on
the northern Tafilalt Platform where water depth was
shallower compared to its surroundings (but similar to
other elevated areas such as towards the west at Jebel
Mdouar or the south near Taouz), but it was still in
a pelagic setting with the sediment surface below the
euphotic zone (Wendt 1985, 1988; belka 1998; kauf-
Mann
1998; k
lug
& k
orn
2002; b
erkoWski
2008; h
eb
-
beln & saMankassou 2015; król et al. 2016).
Cessation of growth of the majority of the mud-
mounds coincides with the Daleje transgression. The
cause for their demise appears to be directly linked
to the mode of carbonate deposition (inorganic or or
-
ganic), a topic discussed in the contribution of b
elka
et al. (2018).
Although the cause of the decline in mudmound
growth is still under debate, there is no doubt that the
biofacies changed significantly with the beginning of
the late Emsian (Figs. 13, 14). The imprint of the global
sea-level rise is reflected at Hamar Laghdad in the shift
from predominantly benthic communities replaced by
predominantly pelagic organisms (Figs. 13, 14). None-
theless, the benthic forms are dominant again in the
latest Emsian communities, which populated a highly
diversified ecosystem consisting of many different
habitats that became established (Figs. 14, 15).
In the early Eifelian, another global sea-level rise
(haq & sChutter 2008) inverted the trend in diversi-
fication of benthic organisms. Much of the Eifelian suc-
cession is dominated by ammonoids (klug 2002) and
orthocerids (p
ohle
& k
lug
2018b), although some lay-
ers are rich in bivalves of the genus Panenka (hrynie-
WiCz et al. 2018).
Fig. 19. The Red Cliff (A) and some geological features found around it. Favositid corals are extremely abundant in the Red
Fauna (B). North of the Red Cliff, many siliceous pebbles document the former presence of the Cenomaninan transgression
conglomerate on the plateau that corresponds to the pre-Cenomanian erosional level (C). Some levels near the Emsian-Eifelian
boundary contain Chondrites-like burrows (D). In the lower part of the Dalejan succession, bivalves are highly abundant (E).
eschweizerbart_xxx
Oases of biodiversity 31
Fig. 19.
eschweizerbart_xxx
32 C. Klug et al.
The late Eifelian Kačák transgression again altered
the ecosystems of the Hamar Laghdad area. In this
interval, ammonoids are much reduced in abundance and
diversity. By contrast, most of the Givetian to Frasnian
limestones are rich in trace fossils (Fig. 8). During
these two stages, cephalopods are common but their
abundance and diversity fluctuated rapidly (e.g., beCker
et al. 2018a, 2018b). Similarly, the relative abundance of
ammonoids versus other cephalopods changed.
5. The Red Cliff and the Red Fauna
The term “Red Cliff” was introduced by klug (2002)
for an erosional remnant (a butte or mesa-like hill) at
Hamar Laghdad, referring to the red colour of the de-
posits (Fig. 19A). Accordingly, the late Emsian fauna
described in this volume are referred to as the “Red
Fauna”. Similar or even richer faunas may be present
elsewhere in Hamar Laghdad. Therefore, the case re-
ported in the present paper can be considered repre-
sentative for the diverse faunas occurring in the Em-
sian strata covering the majority of the exposed mud-
mounds.
5.1. Taphonomy
Facies, palaeoecology and taphonomy at the Red Cliff
are controlled by two main factors, namely the palaeo-
topography and the proximity to the erosional surface
that formed until the Cenomanian (Fig. 19A, C). As
described by klug et al. (2009), the fauna from the
Dalejan marls, which lie close to the erosional surface
(Fig. 19A, C), was subject to intense weathering lasting
from the Viséan to the Cenomanian (i.e., during a time
interval as long as about 240 Ma). In the course of this
process, aragonitic shells were dissolved while calcitic
shells got partially silicified. Also, microscopic cavi-
ties in fossils were infilled by hematite. This alteration
along with partial silicification explains why many fos-
sils of latest Emsian to early Eifelian age weathered out
of these carbonates and why many of those display a
yellowish to reddish colour (Fig. 19B). Due to silicifica-
tion and later weathering many fossils stand prominent-
ly out of the marls and limestones; in turn, this creates
a sampling bias allowing to collect a high number of
often excellently preserved fossils, thus facilitating the
documentation of the overwhelming palaeobiodiversity
of this stratigraphic interval at the Red Cliff. By con-
trast, the layers in the last meters underneath the un-
conformity show a very strong alteration including both
silicification and dolomitisation. Weathered limestones
on top of the Red Cliff superficially resemble oolites
where the virtual ooids are actually dolomite crystals
that are macroscopically visible.
5.2. Composition of the Red Fauna
The Red Fauna has been extensively sampled, leading
to a collection of over 3000 specimens. This collection
comprises at least 140 species of macrofossils. This
number is probably still an underestimate, although the
curves obtained by a rarefaction analysis (Fig. 20) are
fairly flat at a sample size of 2000.
Most faunal elements of the Red Fauna belong to
the epifaunal benthos, nektoplankton or plankton. The
scarcity of infaunal elements and vertebrate nekton is
likely primary because, due to the protected situation
of the Red Cliff (Figs. 4, 11A, B, 20A), it is unlikely
that faunal elements were washed into or out of this
palaeodepression between the surrounding ridge and
the mudmound circle. Probably, the low abundance of
true endobenthos roots in physical properties of the
sediment during deposition. The scarcity of placoderms
and the absence of other gnathostomes are comparable
to that of other outcrops dating from the late Emsian
(r
üCklin
et al. 2018). However, it is still remarkable
because, taking the high local palaeobiodiversity into
account, one would expect that larger predators would
also be reasonably common. Possibly, it was the very
restricted spatial extent of this habitat between the
mudmounds, which did not allow a greater number of
large predators to thrive.
Sessile benthic organisms, stalked or not, make up
the majority of the local diversity, namely about three
quarters of the macroscopic species (Table 3). As far
as the number of species is concerned, the corals are
the most diverse with 34 tabulates and 12 rugose taxa,
which occur in coeval strata across Hamar Laghdad
(berkoWski 2018; król et al. 2018). Brachiopods are
also very common and so far, 24 species were docu-
mented from the Red Fauna (Mergl 2018).
Among the vagile forms, trilobites are represented
by at least 24 species (Crônier et al. 2018). Howev-
er, the diversity was likely higher than that listed in
C
rônier
et al. (2018), because a much greater number of
species are listed in the monographs of alberti (1969,
1970; see beCker et al. 2018a for a review). This abun-
dance of vagile forms underlines their great ecological
importance.
eschweizerbart_xxx
Oases of biodiversity 33
As far as the swimming organisms are concerned,
cephalopods are dominant (Table 3). Three bactritid
species, nine ammonoid species and at least 15 nauti-
loid species occur in moderate numbers (pohle & klug
2018b). Gnathostomes are represented by only a few
bone fragments probably belonging to two species of
placoderms (rüCklin et al. 2018).
5.3. Palaeoecology of the Red Fauna
The seafloor with the numerous, up to 50 meter high
mounds (Fig. 5) apparently created a broad range of
habitats at Hamar Laghdad (e.g., berkoWski 2004,
2008; b
elka
& b
erkoWski
2005; C
avalazzi
et al. 2007;
franChi et al. 2015a; król et al. 2016; hrynieWiCz et
al. 2017). This high local palaeobiodiversity (Figs. 20-
Fig. 20. Palaeoecological analyses of the macrofossils of the Red Fauna (Method 3, Table 3). Top left: Pie chart of the
composition of the Red Fauna; of the 14 ecological guilds (sensu b
ush
et al. 2017), surficial benthic organisms dominate
the fauna. The banana plot (rarefaction, top right) shows that sampling is not complete but already quite good. This might
partially root in the strongly varying preservation due to the differences in knowledge of forms with calcitic and aragonitic
shells. The histogram at the bottom contains only those taxa that form the trophic nucleus. Tabulate and rugose corals,
brachiopods, cephalopods and trilobites dominate the fauna. Percentage of specimens is given on the vertical axis; note that
there is some uncertainty reflected in taxa given in open nomenclature (some unidentified taxa and thus higher diversity
might be hidden here).
eschweizerbart_xxx
34 C. Klug et al.
22) likely established owing to a combination of fa-
vourable ecological factors: (1) the small-scale spatial
ecological compartimentalisation on a rather small area
roots in the irregular distribution of carbonate build-
ups of varying sizes with steep slopes, vents, cavities
(belka et al. 2018) and protected depressions between
the mounds. (2) Water depth was likely moderate, but
the seafloor of the Tafilalt Platform was probably still
below the euphotic zone as reflected in the presence of
trilobites with large eyes on the one side (compare rust
et al. 2016) and the absence of green algae on the other
side. The algae reported in the literature failed to show
morphological features providing an unambiguous at-
tribution to green algae (braChert et al 1992; Wendt
2001; discussions in h
ebbeln
& s
aMankassou
2015;
J
akuboWiCz
et al. 2018). t
öneböhn
(1991) reported rare
occurrences of corallinacean algae, which, however,
can occur in depths up to 250 m (discussion in hebbeln
& s
aMankassou
2015). It remains difficult to assess
palaeobathymetry and light availability in the geolog-
ic past. The fossils provide a somewhat contradictory
picture and detailed information on palaeobathymetry
are impossible to extract; nevertheless, most fossil evi-
dence points at maximal depths within or just below the
Fig. 21. Reconstruction of a late Emsian fauna (like the Red Fauna), which lived after the main phase of mudmound forma-
tion. Taxon names see Fig. 22.
eschweizerbart_xxx
Oases of biodiversity 35
photic zone (up to 250 m) for at least some part of the
time. (3) A wealth of nutrients was probably available
owing to the ongoing hydrothermalism (b
elka
1998;
buggisCh & kruMM 2005; berkoWski 2006, 2008; Cav-
alazzi et al. 2007; oleMpska & belka 2010; franChi
et al. 2015a, 2015b). (4) Low sediment accumulation
rates, partially due to enduring winnowing (l
ubeseder
et al. 2010) possibly led to constant conditions and fa-
voured suspension feeders such as crinoids. Conceiv-
ably, moderate currents carried food particles at a rate
favourable for the abundant suspension feeders present
between the mounds. Additionally, the low sediment
accumulation rates had a certain condensation effect
in off-mound strata, depending on the position in rela-
tion to the mound structures. (5) The palaeogeographic
setting of the Anti-Atlas in moderate latitudes during
the Emsian (sCotese 1997) and the epicontinental sea
situation also contributed to this ecologically-sustained
stability.
In the case of the fauna of late Emsian age of the
Red Cliff, scenario (1) is essential because it might ex-
plain the particularly high diversity at this special set-
ting, although other outcrops with high diversities at
Hamar Laghdad were not included in this study. The
Fig. 22. Taxa of the late Emsian fauna shown in Fig. 21.
eschweizerbart_xxx
36 C. Klug et al.
Red Cliff is situated in a roughly L- to crescent-shaped
carbonate buildup (Fig. 4), which is reminiscent of
mud-ridges of the Ahnet Basin in Algeria (W
endt
et al.
1993). In the case of the Red Cliff, this mound structure
has a diameter of about 300 meters (e.g., aitken et al.
2002, text-fig. 3A). At its northern end, it is truncated
by the erosional surface; thus, this structure was pos-
sibly originally even longer. Additionally, another large
mudmound is situated to the east. Taken together, this
topographic setting created a depression that likely pro-
duced a particularly protected and stable environment.
Possibly, a great number of hydrothermal sources in the
carbonate buildups surrounding the Red Cliff, but it is
unclear when they ceased to be active. Alternatively, all
of Hamar Laghdad had such a high palaeobiodiversity
during the latest Emsian, and the high diversity can
exclusively be explained by a combination of preserva-
tional (taphonomic) and sampling biases. Testing these
alternative hypotheses in detail requires a considerable
sampling effort and is a worthwhile task for the future.
6. Discussion and conclusions
The submarine elevation upon which the mounds of
Hamar Laghdad grew was likely situated in a palae-
ogeographic position suitable for nutrient input, e.g.,
favoured through oceanic currents. This is supported
by the high biodiversity biota including numerous sus-
pension feeders. The distribution of modern carbonate
mounds in the northern Atlantic displays such patterns
(expedition sCientists 2005; Wheeler et al. 2007; ei-
sele et al. 2008; dorsChel et al. 2010).
Numerous clustered carbonate mounds were report-
ed from continental slopes of the Rockall and Porcu-
pine banks (57°N) and the Porcupine Seabight (51°N;
W
hite
& d
orsChel
2010). In these mound provinces,
the mound-sediments reach a thickness exceeding 250
m and a diameter of up to three kilometres (freiWald
2002; roberts et al 2003; Wheeler et al. 2005). These
dimensions are fairly comparable to Hamar Laghdad,
where individual mounds reach 30-50 m, with com-
plexes as thick as 180 m covering a surface with a
diameter of up to five kilometers (M
assa
et al. 1965;
b
ultynCk
& W
alliser
2000). These occurrences are
also comparable with respect to the moderate to high
(palaeo-)latitudes.
It is assumed that surface productivity over the
Rockall and Porcupine banks, driven by high nutrient
levels there, play a significant role in the growth of
mounds at these specific locations (White et al. 2005).
Most likely, the productivity played a key role in the
Hamar Laghdad case as well. To date no unambiguous
evidences for such high productivity have been provid-
ed. Further investigation may shed light on the nutrient
level during the formation of Kess-Kess mounds (see
discussion in hebbeln & saMankassou 2015).
Water depth is poorly constrained for the Hamar
Laghdad mudmounds, but the estimated 100-200 mud-
mounds and mound complexes covering areas as large
as minimum 520 m x 130 m (braChert et al 1992)
compare to modern examples from the NE Atlantic.
W
hite
& d
orsChel
(2010) identified 1013 individual
carbonate mounds clustered in seven coral carbonate
mound provinces in the Rockall Trough and Porcu-
pine Sea Bight, NE Atlantic, located in water depths
between 525 and 1650 m. The epicontinenntal setting
and fossil content of the Dalejan sediments indicate a
lower water depth for Hamar Laghdad.
The distribution patterns of these Recent carbonate
mound provinces are primarily controlled by oceanic
circulation (driving nutrient supply) rather than by hy-
drothermalism, although the latter has been proposed
as the main driving factors in the cases of the Moroccan
mudmounds (b
elka
1998; b
erkoWski
2004; f
ranChi
et
al. 2015a, 2015b). This topic, out of the scope of the
present paper, holds a high scientific potential to con-
tribute to our understanding of the peculiar occurrences
of carbonate mound provinces in the sedimentary rock
record, in Morocco and elsewhere.
Overall, we hope that the present volume underlines
the uniqueness of Hamar Laghdad. In this context, we
support the suggestion by henriet et al. (2014: 103)
to include Hamar Laghdad as a geological-palaeon-
tological highlight in the “mound Heritage Route in
Morocco. The route segment circling the High Atlas
encompasses outstanding examples of Paleozoic to
Mesozoic mound sites.” As such, we think that Hamar
Laghdad is a perfect candidate for a UNESCO World
Heritage Site.
Acknowledgements
We greatly appreciate the support provided by the Swiss
National Science Foundation (projects 200020_132870
and 200021_ 160019). Working permits and sample export
permits were kindly provided by the Ministère de l’Energie,
des Mines, de l’Eau et de l’Environnement (Direction du
Développement Minier, Division du Patrimoine, Rabat,
Morocco). bernd kaufMann (Bremen) proof-read an earlier
version of the manuscript and provided some valuable and
stimulating comments. Taxonomic determinations were car-
ried out by J
an
k
ról
(Poznan), M
ikolaJ
z
apalski
(Warsaw),
eschweizerbart_xxx
Oases of biodiversity 37
Błazej BerkowSki (Poznan), adaM t. halaMski (Warsaw),
a
ndrzeJ
b
alinski
(Warsaw), M
iChal
M
ergl
(Prague), M
i
-
Chael aMler (Cologne), krzysztof hrynieWiCz (Warsaw),
j
iří
F
rýda
(Prague), m
icHał
j
akuBowicz
(Poznan), C
athe
-
rine
C
rônier
(Lille), M
organe
o
udot
(Lille), r
aiMund
f
eist
(Montpellier), e
lise
n
ardin
(Montpellier), J
ohnny
W
aters
(Boone), Martin rüCklin (Leiden), and lothar h. vallon
(Faxe). We acknowledge the constructive reviews by B
łazej
berkoWski (Poznan) and two anonymous reviewers.
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Manuscript received: May 14th, 2018.
Revised version accepted by the Stuttgart editor: July 2nd,
2018.
Addresses of the authors:
Christian klug, Paläontologisches Institut und Museum,
Universität Zürich, Karl Schmid-Strasse 4, CH-8006 Zürich,
Switzerland;
e-mail: chklug@pim.uzh.ch
elias saMankassou, Département des Sciences de la Terre,
Université de Genève, CH-1205 Genève, Switzerland ;
e-mail: Elias.Samankassou@unige.ch
alexander pohle, Paläontologisches Institut und Museum,
Universität Zürich, Karl Schmid-Strasse 4, CH-8006 Zürich,
Switzerland;
e-mail: alexander.pohle@pim.uzh.ch
kenneth de baets, GeoZentrum Nordbayern, Fachgrup-
pe PaläoUmwelt, Friedrich-Alexander-Universität Erlan-
gen-Nürnberg, Loewenichstr. 28, 91054 Erlangen, Germany;
e-mail: kenneth.debaets@fau.de
fulvio franChi, Earth and Environmental Science Depart-
ment, Botswana International university of Science and
Technology, Private Bag 16, Palapye, Botswana;
e-mail: franchif@biust.ac.bw
d
ieter
k
orn
, Museum für Naturkunde Berlin, Invaliden-
straße 43, 10115 Berlin, Germany;
e-mail: dieter.korn@mfn-berlin.de
eschweizerbart_xxx
44 C. Klug et al.
Table 1. Faunal composition in % based on bioclast counts in polished samples from the Pragian of Hamar Laghdad (Method
2). For the stratigraphic position of samples 1 to HL58, see Figs. 10, 13, 15.
Sample
Placodermi
Arthrophyllum
Temperoceras
Orthocerida
Dacryoconarida
Crinoida
Trilobita
Ostracoda
Gastropoda
Brachiopoda
Rugosa
Tabulata
Nr of fossils
200100 00 000000020
30000.022 0.525 0.179 0.109 00.0113 0.142 0.011 089.5
40000.028 0.830 0.019 0.019 000.085 0.009 0.009 106
600.009 000.692 0.044 0.026 0.01 00.207 0.04 0113.5
80000.007 0.852 0.036 0.014 00.0049 0.077 0.007 0.004 281
900.006 00.01 0.667 0.122 0.006 00.006 0.183 00156
10 00000.543 0.172 0.06 00.017 0.207 00116
12 0000.009 0.237 0.237 0.082 00.010 0.415 0.010 0104
16 0000.01 0.101 0.416 0.01 00.010 0.455 00104.5
18 00000.281 0.193 0.193 0.07 0.015 0.252 0067.5
HL01 000 0.98 00.015 0.005 0323
HL02 00000.79 0.058 0.022 0.04 00.116 00.051 69
HL06 00000.557 0.021 0.113 000.289 0.021 045.5
HL10 00000.5 0.0165 0.06 0.12 0.044 0.236 0.011 0.011 91
HL14 0000.837 00.053 000.110 00113.5
HL18 00000.088889 0.622 0.067 000.044 0.015 0.163 67.5
HL22 00000.65 00.05 0.15 00.15 0040
HL29 0000.051 0.59 0.026 0.038 0.06 0.128 0.103 0039
HL31 0.023 000.022 0.195 0.356 0.138 0.11 0.046 0.103 0043.5
HL33 00000.292 0.3 0.236 0.03 00.111 0.028 036
HL35 00000.595 0.095 0.071 0.07 0.024 0.143 0042
HL36 00000.469 0.143 0.122 0.04 0.041 0.143 00.041 24.5
HL39 00000.625 0.166 0.073 000.135 0086.33
HL43 00000.253 0.660 0.029 000.039 0.019 051.5
HL52 00000.459 0.144 0.245 000.144 0.008 093.75
HL58 00000.249 0.613 0.023 0.02 0.012 0.081 0086.5
Ort.-
lst. 0000.575 0.402 00.023 0000043.5
eschweizerbart_xxx
Oases of biodiversity 45
Table 2. Faunal composition of faunules from the Pragian of Hamar Laghdad. For the stratigraphic position of samples A to
D (Figs. 10, 13, 15). Abbreviations of modes of life: c – coprophagous; er – erect; ff – freely fast; fs – freely slow; fu – facul-
tative unattached; g – grazer; na – non-motile and attached; p – pelagic; pr – predators; s – suspension feeder; su – surficial.
Taxonomic determinations were carried out for Tabulata by J. król and M. zapalski, for Rugosa by B. berkoWski, for Bra-
chiopoda by A. T. h
alaMski
, A. b
alinski
and M. M
ergl
, for Rostroconchia by M. a
Mler
, for Bivalvia by K. h
rynieWiCz
and
M. JakuboWiCz, for Gastropoda by jiří Frýda, for non-ammonoid Cephalopoda by AP, for Ammonoidea by CK and KDB,
for Trilobita by C. Crônier, M. oudot and R. feist, for Echinodermata by E. nardin and J. Waters, and for placoderms by
M. rüCklin. * new taxa described in pohle & klug (2018b).
Species A B C D Life mode
Echinodermata
Crinoida sp. A 1 3 3 3 er, na, s
Crinoida sp. B 2 2 1 er, na, s
Anthozoa
Rugosa 415 611 er, na, s
Favositida sp. A 1 1 3 2 er, na, s
Favositida sp. B 1 2 er, na, s
Thamnoporidae 1 1 9 er, na, s
Micheliniidae 2 5 er, na, s
Trilobita
Odontochile 3 su, ff, pr
Reedops 4 1 su, ff, pr
Crotalocephalina gibbus 1 su, ff, pr
Otarion 1 2 su, ff, pr
Phacopidae 1 7 2 su, ff, pr
Paralejurus 2 su, ff, pr
Scutellum 2 su, ff, pr
Rhynchonellata
Protathyris 2 4 7 7 su, na, s
?Aulacella eifeliensis 1 su, na, s
Cingulodermis 1 1 3 su, na, s
Bivalvia
Panenka obsequens 7 5 2 1 si, fu, m
Hercynella paraturgescens 2 su, fs, g
Bivalvia 1su, fa, s
Hyolitha
Orthotheca 1 3 su, nu, s
Amphigastropoda
Belerophontidae 1 su, fs, g
Vetigastropoda
Platyceras 3 2 3 4 su, fu, c
Oriomphalus multiornatus 2 1 1 su, fs, g
Gastropoda 1 1 1 su, fs, g
Palaeozygopleura 2 2 2 su, fs, g
Species A B C D Life mode
Paraoehlertia 3 1 su, fs, g
Eohormotomina restisevoluta 1 4 su, fs, g
Planitrochus 1 su, fs, g
Australonema 1 4 su, fs, g
Cephalopoda
Discosorida
Pseudendoplectoceras lahcani 1 p, fs, pr
Pseudendoplectoc. sp. n.* 1 p, fs, pr
Oncocerida
Tafilaltoceras adgoi 1 p, fs, pr
Lituitida
Arthrophyllum vermiculare 2 4 p, fs, pr
Pseudorthocerida
Spyroceras patronus 1 3 1 p, fs, pr
Spyroceras cyrtopatronus 1 p, fs, pr
Cancellspyroceras loricatum 1 2 2 p, fs, pr
Suloceras longipulchrum 2 1 p, fs, pr
Subdoloceras atrouzense 3 9 7 4 p, fs, pr
Subdoloceras engeseri 48 510 p, fs, pr
Geidoloceras sp. n.* 131 510
Geidoloceras ouaoufilalense 1 p, fs, pr
? Pseudorthoc. indet. A 1 p, fs, pr
Orthocerida
Mimogeisonoceras sp. 4 p, fs, pr
Pseudospyroceras reticulum 2 1 2 p, fs, pr
Arkonoceras sp. 1 p, fs, pr
? Orthocycloceras sp. 2 p, fs, pr
Tibichoanoceras tibichoanum 8 6 p, fs, pr
Dawsonoceratidae indet. 1 p, fs, pr
Orthoceratidae gen. et sp. n.* 3 1 p, fs, pr
Temperoceras migrans 14 4 p, fs, pr
Orthocerida indet. 27 25 9 p, fs, pr
Number of specimens 74 202 99 86
eschweizerbart_xxx
46 C. Klug et al.
Table 3. Faunal composition of the Red Fauna (latest Emsian in age) of Hamar Laghdad. For the stratigraphic position, see
Fig. 15. Abbreviations of modes of life: c – coprophagous; er – erect; ff – freely fast; fs – freely slow; fu – facultative unat-
tached; g – grazer; na – non-motile and attached; p – pelagic; pr – predators; s – suspension feeder; su – surficial. Taxonomic
determinations were carried out for Tabulata by J. k
ról
and M. z
apalski
, for Rugosa by B. b
erkoWski
, for Brachiopoda
by A. T. h
alaMski
, A. b
alinski
and M. M
ergl
, for Rostroconchia by M. a
Mler
, for Bivalvia by K. h
rynieWiCz
and M.
JakuboWiCz, for Gastropod by jiří Frýda, for non-ammonoid Cephalopoda by AP, for Ammonoidea by CK and KDB, for
Trilobita by C. Crônier, M. oudot and R. feist, for Echinodermata by E. nardin and J. Waters, and for placoderms by M.
rüCklin. * new taxa described in pohle & klug (2018b). ** new taxa described in Mergl (2018). *** new taxa described
in Crônier et al. (2018).
Taxon Number Ecology
CNIDARIA
Actiniaria
Conichnus conicus 9 si, na, pr
Tabulata
Favosites bohemicus 41 su, na, s
Favosites goldfussi saourensis 2 su, na, s
Crenulipora difformis 2 su, na, s
Striatopora cf. obliqua 23 su, na, s
Striatopora marsupia 16 su, na, s
Taouzia chouberti 37 su, na, s
Taouzia sp. 8 su, na, s
? Thamnoptychia sp. 15 su, na, s
Parastriatopora crassimuralis 5 su, na, s
Aulopora sp. 33 su, na, s
Bainbridgia sp. 10 su, na, s
Aulocystis wendti 21 su, na, s
Aulocystis cf. alectiformis reptata 89 su, na, s
Chia ramosa 3 su, na, s
Dualipora preciosa 51 su, na, s
?Senceliaepora sp. 1 su, na, s
Cladochonus sp. 20 su, na, s
Aulocystis sp. 6 su, na, s
? Crenulipora sp. 4 su, na, s
? Dendropora sp. 6 su, na, s
? Dualipora sp. 3 su, na, s
? Favosites sp. 2 su, na, s
Favositida indet. 32 su, na, s
? Michelinia sp. 2 su, na, s
? Parastriatopora sp. 2 su, na, s
Striatopora sp. 1 5 su, na, s
Striatopora sp. 2 3 su, na, s
Striatopora sp. 3 4 su, na, s
? Striatopora 2 su, na, s
? Hyostragulum sp. 2 su, na, s
Cleistopora cf. geometrica 7 su, na, s
? Cleistopora sp. 10 su, na, s
? Coenites sp. 1 su, na, s
Taxon Number Ecology
? Thamnopora sp. 2 su, na, s
Auloporida indet 7 su, na, s
Indet. Tabulata coral 80 su, na, s
Rugosa
Enterolasma eisenmanni 2 su, na, pr
Catactotoechus instabilis 3 su, na, pr
Syringaxon smithioides 36 su, na, pr
Syringaxon exiguus 19 su, na, pr
Syringaxon sp. (juv.) 31 su, na, pr
Sutherlandinia anna 22 su, na, pr
Sutherlandinia gotlandica 2 su, na, pr
Schindewolfia sp. 11 su, na, pr
Marocaxon sp. 26 su, na, pr
Adradosia sp. 1 su, na, pr
Amplexus sp. 2 su, na, pr
Indet. solitary Rugosa 89 su, na, pr
Hydrozoa
trachypsammid hydrozoan? 1 su, na, s
POLYCHAETA
Hicetes 2 su, na, s
MOLLUSCA
Rostroconchia
Hoareicardia sp. 4 smi, nu, s
Bohemicardia sp 1 smi, nu, s
Hyolithida
Orthotheca sp. 38 su, nu, s
Bivalvia
Panenka sp. 17 si, fu, m
Cypricardinia sp. 9 si, fu, s
Patrocardia sp. 3su, fu, s
Bivalvia indet. 3su, fu, s
Jahnia sp. 3su, fu, s
Hercynella sp. 2su, fu, s
Gastropoda
Platyceratida indet. 23 su, fu, c
Pleurotomarioidea indet. A 3 su, fs, g
Pleurotomarioidea indet. B 2 su, fs, g
eschweizerbart_xxx
Oases of biodiversity 47
Taxon Number Ecology
Loxonematoidea indet. A 6 su, fs, g
Loxonematoidea indet. B 2 su, fs, g
Gastropoda indet. 13 su, fs, g
CEPHALOPODA
Oncocerida
Oncocerida indet. 2 p, fs, pr
Lituitida
Arthrophyllum vermiculare 16 p, fs, pr
Pseudorthocerida
? Spyroceras sp. 2 p, fs, pr
? Suloceras sp. 1 p, fs, pr
Subdoloceras tafilaltense 19 p, fs, pr
? Diagoceras sp. 50 p, fs, pr
? Pseudorthocerida indet. 3 p, fs, pr
Orthocerida
Orthocycloceras tafilaltense 7 p, fs, pr
Infundibuloceras sp. n. * 10 p, fs, pr
Astoceras sp. n.* 1 p, fs, pr
? Geisonoceratidae indet. 22 p, fs, pr
Orthocerida indet. A 117 p, fs, pr
Orthocerida indet. B 2 p, fs, pr
? Orthocerida indet. C 8 p, fs, pr
? Orthocerida indet. D 2 p, fs, pr
Bactritida
Lobobactrites ellipticus 13 p, fs, pr
Devonobactrites emsiense 8 p, fs, pr
Devonobactrites obliquiseptatum 1 p, fs, pr
Ammonoidea
Amoenophyllites doeringi 29 p, ff, pr
?Gyroceratites sp. 1 p, ff, pr
Anarcestes lateseptatus 5 p, ff, pr
Anarcestes latissimus 18 p, ff, pr
Anarcestes plebeius 29 p, ff, pr
Anarcestes sp. 10 p, ff, pr
Latanarcestes latisellatus 21 p, ff, pr
Latanarcestes noeggerathi 67 p, ff, pr
Paranarcestes chalix 7 p, ff, pr
BRACHIOZOA
Brachiopoda
Lochkothele sp. 4 su, na, s
Schizotreta sp. 1 su, na, s
Plectodonta (Dalejodiscus) sp. 1 su, na, s
Novellinetes cf. novellus 1 su, na, s
Taxon Number Ecology
Zlichopyramis (?) sp. 1 su, na, s
Dalejina sp. n. ** 23 su, na, s
Lysigypa sp. 15 su, na, s
Leviconchidiella sp. 1 su, na, s
Clorinda cf. exarmata 11 su, na, s
Glossinulus mimicus 1 su, na, s
Carolirhynchia (?) sp. 39 su, na, s
Quasidavisonia tenuissima 5 su, na, s
Kaplicona sp. 7 su, na, s
Holynatrypa sp. 129 su, na, s
Radimatrypa sp. 30 su, na, s
Rhynchatrypa thetis 49 su, na, s
Trigonatrypa sp. n. ** 114 su, na, s
Merista passer 21 su, na, s
Amoenospirifer foedus 285 su, na, s
Pinguispirifer sp. n.** 50 su, na, s
Echinocoelia strobilata 2 su, na, s
Cingulodermis sp. n. ** 137 su, na, s
Eoreticularia indifferens 493 su, na, s
Cyrtina sp. 1 su, na, s
Phoronida
Hederellidae gen. et. sp. indet. 5 su, na, s
ARTHROPODA
Trilobita
Morocops granulops 30 su, ff, pr
Morocops sp. n.*** 67 su, ff, pr
Phacopine sp. J 3 su, ff, pr
Destombesina cf. tafilaltensis 3 su, ff, pr
Psychopyge cf. elegans 1 su, ff, pr
Hollardops cf. mesocristata 1 su, ff, pr
Acastid sp. H 1 su, ff, pr
Acastid sp. I 1 su, ff, pr
Cyphaspis sp. G 5 su, ff, pr
Cyphaspides sp. 6 su, ff, pr
Gerastos sp. 1 7 su, ff, pr
Gerastos sp. 2 18 su, ff, pr
Gerastos sp. 3 2 su, ff, pr
Gerastos sp. 4 3 su, ff, pr
? 1 su, ff, pr
Dohmiella? sp. E 1su, ff, pr
Cornuproetus cf. midas amlanensis 5 su, ff, pr
Diademaproetus sp. vel Cornupr. sp. 6 su, ff, pr
? 1 su, ff, pr
eschweizerbart_xxx
48 C. Klug et al.
Taxon Number Ecology
Harpes sp. n.*** 13 su, ff, pr
Ceratocephala aff. vesiculosa 3 su, ff, pr
Leonaspis sp. 1 su, ff, pr
Acanthopyge sp. 3 su, ff, pr
Scabriscutellum aff. georgei 4 su, ff, pr
ECHINODERMATA
Edrioasteroidea
Rhenopyrgus 2 si, na, s
Blastoidea
Pentremitidea pailleti 1 er, na, s
Cystoidea
Cystoid sp. A 8 er, na, s
Cystoid sp. B 6 er, na, s
Cystoid indet. 6 er, na, s
Crinoida
Crinoid sp. A 15 er, na, s
Crinoid sp. B (Pisocrinus sp.) 22 er, na, s
Crinoid sp. B 5 er, na, s
Crinoid sp. D 2 er, na, s
Elicrinus? weyeri 1 er, na, s
Tiaracrinus jeanlemenni 3 er, na, s
3096
eschweizerbart_xxx
