RESEARCH PAPER

Rugose corals from the upper Visean (Carboniferous)of the Jerada Massif (NE Morocco): taxonomy, biostratigraphy,facies and palaeobiogeography

Markus Aretz

Received: 14 September 2009 / Accepted: 24 October 2009 / Published online: 25 November 2009

� Springer-Verlag 2009

Abstract The Visean succession of the Jerada Massif

contains a relatively diverse rugose coral fauna, which

comprises 12 genera. Only two of these (Siphonodendron

and Lithostrotion) are reported to include more than one

species. Most taxa show distinctive facies dependencies.

Small solitary corals are found in shaly environments of the

Oued Es-Sassi Formation. On the northern flank a coral

patch reef formed on an oolitic shoal in the Koudiat Es-

Senn Formation. Its core mainly consists of Lithostrotion

vorticale colonies, whereas in its surroundings literally

hundreds of specimens of Siphonophyllia samsonensis

occur. Associated with these dominant taxa occur colonial

and further large solitary taxa (Palaeosmilia, Dibunophyl-

lum). Tizraia and Pareynia are restricted to microbial-

dominated buildup facies. Their presence might be strongly

controlled by the development of this buildup type, because

further occurrences in Algeria, Morocco, and Belgium are

all in the same facies. The coral fauna at Jerada is a typical

Late Visean assemblage for the Western European Prov-

ince. The Eastern Moroccan Meseta may be an important

pathway within the province for the connection between

the Central Saharan basins and NW Europe. The bio-

stratigraphic ages of the coral fauna partly contradict ages

based on carbonate microfossils; the coral ages are slightly

older and typical Brigantian coral taxa are absent.

Keywords Visean � Rugose corals � Jerada Massif �Eastern Moroccan Meseta � Palaeobiogeography

Kurzfassung Die viseische Abfolge des Jerada-Massivs

enthalt eine relativ diverse Fauna. Diese Fauna umfasst 12

Gattungen, aber mehrere Arten sind nur in zwei dieser

Gattungen (Siphonodendron und Lithostrotion) vertreten.

Die meisten Taxa zeigen eine deutliche Fazies-

Abhangigkeit. Kleine solitare Formen sind nur aus schiefr-

igen Gesteinen der Oued Es-Sassi-Formation bekannt. Auf

dem Nordflugel des Jerada-Synklinoriums bildete sich in

der Koudiat Es-Senn-Formation an einer oolitischen Barre

ein Korallen-Patch-Riff.Sein Kern besteht im Wesentlichen

aus Lithostrotion vorticale-Kolonien. Das Riff wird von zah-

lreichen (einigen hundert) Exemplaren von Siphonophyllia

samsonensis umgeben. Assoziiert mit diesen beiden domi-

nierenden Taxa sind weitere großwuchsige solitare (Pala-

eosmilia, Dibunophyllum) und koloniale Taxa. Tizraia und

Pareynia sind nur in mikrobiell-dominierter Buildup-Fazies

zu finden. Ihr Vorkommen scheint generell auf diesen

Builduptyp beschrankt, wie weitere Vorkommen in gleicher

Fazies in Algerien, Marokko und Belgien zeigen. Die

Korallenfauna von Jerada ist typisch fur Assoziationen des

spaten Viseums innerhalb der westeuropaischen Faunen-

provinz. Die Ostliche Marokkanische Meseta stellt einen

moglichen Verbindungsweg innerhalb dieser Provinz

zwischen den Becken der zentralen Sahara und Westeuropa

dar. Die biostratigraphischen Alter der Korallen stehen in

einem gewissen Widerspruch zu den auf kalkigen Mikro-

fossilien beruhenden Altern. Die Korallen-Alter sind etwas

alter, typische Vertreter des Brigantiums fehlen.

Schlusselworter Viseum � Rugose Korallen �Jerada Massiv � Ostliche Marokkanische Meseta �Palaobiogeographie

M. Aretz (&)

Universite de Toulouse (UPS), LMTG (OMP),

14 Avenue Edouard Belin, 31400 Toulouse, France

e-mail: markus.aretz@lmtg.obs-mip.fr

M. Aretz

CNRS, LMTG, 31400 Toulouse, France

M. Aretz

IRD, LMTG, 31400 Toulouse, France

123

Palaontol Z (2010) 84:323–344

DOI 10.1007/s12542-009-0046-0

Introduction

Rugose corals of Carboniferous age are relatively poorly

studied in NW Africa. Diverse shallow water faunas have

been described from the Bechar Basin (Semenoff-Tian-

Chanksy 1974, 1985) and the Eastern Central Moroccan

Meseta (Said et al. 2007; Said and Rodrıguez 2007, 2008,

Aretz and Herbig 2009). However, limited numbers of

specimens from various regions and facies found during

reconnaissance work and mapping (e.g. Termier and

Termier 1950; Conrad 1984) clearly indicate the likelihood

of comparable abundances and diversities in NW Africa

(Fig. 1) as in contemporaneous tropical-shelf systems (e.g.

western USA: Sando 1980, Belgium: Poty 1981). The only

briefly described fauna of the Anti-Atlas (Wendt et al.

2001) shows a possible mixture of taxa of deeper and

shallow water environments. The unpublished material

of the Bechar Basin and further Algerian Basins (coll.

Semenoff-Tian-Chansky, Museum Nationale d’Histoire

Naturelle, Paris) contains numerous small undissepimented

solitary rugose corals, which probably represent deeper

water environments and environments similar to the

Anti-Atlas.

The record of Carboniferous corals in the Eastern

Moroccan Meseta is very poor. Owodenko (1946, 1976)

mentioned Caninia juddi Nicholson (? = Pseudazphren-

toides juddi), Dibunophyllum sp., Zaphrentis sp. and

Favosites parasitica d’Orbigny (? = Sutherlandia) from

the Visean succession of the Jerada Massif. However, the

depository for this material is unclear, and a revision of

those specimens is inevitable.

It is the purpose of this paper:

1. to describe newly collected rugose corals from the

upper Visean of the Jerada Massif;

2. to place the specimens in a facies context; and

3. to briefly discuss stratigraphic and palaeobiogeograph-

ic implications.

Geological setting

The Jerada Massif is part of the Palaeozoic basement of the

Eastern Moroccan Meseta deformed during the Variscan

Orogeny and today widely covered by Mesozoic–Caino-

zoic strata. It belongs to the inner zones within the tectonic

framework of the northwest African Variscan domain

(Pique et al. 1993).

The simple-structured WSW–ENE synclinorium is sit-

uated about 50 km south of the town of Oujda near the

Moroccan–Algerian border (Fig. 2). It is about 30 km in

strike and about 15 km normal to strike. The dip of the

northern flank is gentle (15–30�S), whereas the southern

flank dips steeply (80�N), and may locally be overturned.

The synclinorium contains an exclusively Carboniferous

succession (Owodenko 1946, 1976). A Visean basin fill

sequence (Herbig et al. 2006) is unconformably overlain by

a Namurian to Westphalian paralic sequence (Desteucq

et al. 1988). It ends with coal-bearing strata of Westphalian

B–C age below the Variscan unconformity.

Herbig et al. (2006) established a lithostratigraphic

subdivision of the Visean succession (Fig. 3), which

substituted the formations of Berkhli et al. (1999) defined

in repeated succession of tectonic slices. The Cafcaf For-

mation mainly consists of bedded cherts. The succeeding

Oued El Koriche Formation is lithological variegated,

which results from the deposition of debris flows, turbidites

Fig. 1 Geological map of NW Africa (after Pique 2001) with

location of the main coral occurrence. See text for main references for

coral data

324 M. Aretz

123

(siliciclastic and carbonate dominated ones), and olistoliths

in mud- to silt-dominated lower and middle slope facies.

The Oued Es-Sassi Formation comprises mainly silty

shales. The general lithology and thickness of all forma-

tions varies between northern and southern flank, but also

on the individual flanks. Probably the most striking dif-

ference between the two flanks is the composition of the

Koudiat Es-Senn Formation, which tops the Oued Es-Sassi

Formation. On the southern flank it contains a single sed-

imentary cycle, which includes microbial-sponge buildups

and small microbial-metazoan buildups (Aretz and Herbig

2008); whereas on the northern flank the formation is twice

as thick, consists of two sedimentary cycles (Berkhli et al.

1999), and only contains a single rugose coral patch reef in

the uppermost beds (Aretz et al. 2006). All sedimentary

cycles in the Koudiat Es-Senn Formation consist of a lower

siliciclastic-dominated part and a carbonate-dominated

upper part. The formation is locally topped by an ignim-

brite (Berkhli et al. 2000) indicating subaerial exposure of

the region around the Visean/Namurian boundary. Sub-

sequent Namurian strata are dominated by fine-grained

siliciclastics facies (‘‘Namurian-Westphalian shales’’).

Aretz and Herbig (2007) showed that generally more

distal facies occurs in the NE of the synclinorium, whereas

the most proximal facies is reported from the SW (Oued

Agaia area), thus indicating a NE-wards tilted basin. Hence

local and regional palaeogeography markedly affected the

sedimentation patterns in the Jerada Massif.

Materials and methods

Most corals are from limestones of the Oued El Koriche

and Koudiat Es-Senn formations, but a few records are

from the Cafcaf and Oued Es-Sassi formations (Fig. 3).

They have been collected during two field campaigns in

2004 and 2006, which focussed on mapping, buildup, and

reef facies and Mississippian basin development. Rare

single section of tabulate corals (michelinids, auloporids)

and heterocorals (Hexaphyllia), not described herein, were

found in petrographic thin sections.

A more focussed collection on corals may result in a

more diversified fauna, and would enable, in many cases,

more precise determination of intra-specific variability.

The quality of the material and thus the availability of thin

sectioned rugose corals is limited, because not all of the

material studied has been collected for this purpose.

However, at least one vertical section is always present. In

a few cases well preserved specimens could be found, but

the majority is either enclosed in limestone or outer parts

have been eroded, as typical for many African specimens

(Semenoff-Tian-Chansky 1974). The terminology of mor-

phological characters follows Hill (1981) and Poty (1981).

The attribution of genera to the higher taxonomic levels of

Hill (1981) is largely followed in this work. All material

is stored in the Institut fur Geologie und Mineralogie,

Universitat zu Koln under the numbers GIK 2156-2240.

Institutional abbreviations: BM, British Museum (Natural

History) London; DPM, Departamento de Paleontologıa,

Universidad Complutense de Madrid; HM Hunterian Museum

Glasgow; MNHNP, Museum National d’Histoire Naturelle,

Paris; NMD, National Museum of Irland, Dublin; RSM, Royal

Scottish Museum, Edinburgh; SM, Sedgwick Museum,

University of Cambridge; UMO, University Museum Oxford.

Systematic palaeontology

Systematic descriptions are presented for all taxa. Some

descriptions of well-known species are limited to key

Fig. 2 Geological map of the

Jerada synclinorium and

location of the sample localities

(P1-P10), with inset showing

the Palaeozoic massifs of the

Eastern Moroccan Meseta. Inset

after Berkhli et al. (1993)

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 325

123

points and less common described characters. To reduce

the lengths of the synonymy lists, the only references added

are those not included in recent synonymy list which the

author agrees with (cum. syn.).

Class Anthozoa Ehrenberg, 1834

Subclass Rugosa Milne-Edwards and Haime, 1850

Order Stauriida Verrill, 1865

Suborder Caniniina Wang, 1950

Family Cyathopsidae Dybowski, 1873

Genus Siphonophyllia Scouler in McCoy, 1844

Type species: Siphonophyllia cylindrica Scouler in

McCoy, 1844.

Siphonophyllia samsonensis (Salee, 1913) (Fig. 4a–f)

*1913 Caninia samsonensis Salee: 48, pl. D fig. 1.

1975 Siphonophyllia benburbensis (Lewis, 1927).—

Hoffmann et al.: pl. 4 fig. 3.

1981 Siphonophyllia samsonensis (Salee, 1913).—Poty:

52, fig. 49 pl. 25, fig. 4, pl. 26, fig. 3, 4. (cum. syn.)

?1992 Siphonophyllia cf. samsonensis (Salee, 1913).—

Rodrıguez and Falces: 190, pl. 16 fig. 1, 2.

2004 Siphonophyllia samsonensis (Salee, 1913).—Rod-

rıguez et al.: 74, fig. 2.5, figs. 3.1-2, 3.4.

2005 Siphonophyllia samsonensis (Salee, 1913).—Cozar

et al.: fig. 12.12.

2006 Siphonophyllia samsonensis (Salee, 1913).—Galla-

gher et al.: fig. 13.4.

Holotype: The specimen illustrated by Salee could not

be located by Poty (1981) in the collections at Louvain-la-

Neuve, thus it might be considered lost.

Locus typicus and stratum typicum: Nameche, Bel-

gium; lower Upper Visean.

Diagnosis: See Poty (1981)

Remark: S. benburbensis (Lewis 1927) is a synonym of

S. samsonensis (Salee 1913), but still widely used in the

literature.

Material: Four fragments from P1 (GIK 2172-2175), 16

fragments from P4; thin sectioned specimens (2156-2163),

uncut specimens (GIK 2164-2171).

Description: Literally hundreds of fragments from this

large straight cylindrical solitary coral can be found in the

two studied localities. However, younger stages of the

species are absent, as in many other outcrops (Semenoff-

Tian-Chansky 1974).

The state of preservation is very different. The better

preserved specimens show 2–3 growth rings per cm, and

well differentiated furrows and ridges (10–15 per cm).

Rejuvenesence occurs occasionally. The thin undulating

wall of most corallites is eroded, and this may also

include large parts of the dissepimentarium. Corallite

diameters, partly reconstructed, vary between 4.0 and

6.3 cm. There are 55–64 septa in two series. The major

septa are of variable length, but always leave a large

open axial space. They are straight and thickened in the

tabularium. Steroplasma may completely fill the space

between major septa. Within the dissepimentarium the

major septa become sinuous. Minor septa are mostly

restricted to septal spines within the dissepimenta-

rium, and only occur very rarely in the tabularium. A

cardinal fossula is always present. The counter septum

may be:

Fig. 3 Lithostratigraphy and simplified schematic lithological logs of

the Visean succession on the two flanks of the Jerada synclinorium

(not to scale)

326 M. Aretz

123

1. slightly longer, as the other major septa;

2. inconspicuous; or

3. in a counter fossula (very rare).

The dissepimentarium is variable in thickness. It con-

sists of an inner more regular part and an outer dissepi-

mentarium with transeptal dissepiments of both orders. In

the longitudinal section there are 4–9 rows of elongated

and variably inclined dissepiments (3–8 per cm). Maxi-

mum elongation is 1 cm. Tabulae are complete (9–12 per

cm). In the axial part they are flat and continue into a

peripheral depression. Within this depression stereoplas-

ma may be observed. Occasionally axial tabellae are

present.

Discussion: The Jerada specimens fit into the variability

described for this species, and almost perfectly match the

drawings of Dixon (1970) for specimens from Streedagh

Point (Ireland). A number of species of Siphonophyllia

are known from the uppermost Famennian to the early

Visean (e.g. S. garwoodi Ramsbottom and Mitchell 1980,

S. gigantea (Michelin 1843), whereas only a few species

are reported from younger strata. Vaughan (1915) has

already made the case that S. samsonensis is the end

member of one phylogentic linage within the genus char-

acterised by the arrangement of the dissepiments. S. siblyi

(Semenoff-Tian-Chansky 1974) differs from S. samsonensis

by its thicker major septa, the extreme reduction of the

minor septa, and a higher diameter/septa ratio. Dixon

(1970) showed the differences between S. samsonensis and

S. cylindrica Scouler in McCoy 1844, but, because of the

characteristic high morphological variability, single speci-

mens of S. samsonensis may show a habit of the second

species.

Distribution: Well known species in the middle and late

Visean of Europe and North Africa.

Family Hapsiphyllidae Grabau, 1928

Genus Zaphrentites Hudson, 1941

Type species: Zaphrentis curvulena Thomson, 1881

?Zaphrentites sp. (Fig. 4g)

Material: Three fragments from P4 (GIK 2177-2179),

one fragment from P8 (GIK 2176).

Description: Small solitary, curvated, trochoid coral.

The preservation is variable, calices and mature stages are

absent, walls are locally eroded. The maximum diameter is

15 mm. There are 34–37 major septa, which are thick and

confluent at the axial ends to close the marked cardinal

fossula above the centre. Septa in the cardinal half of the

corallite are less thick than those in the counter half. Ste-

roplasma is found between individual septa along the wall

preferentially in the counter half. Although the longitudinal

section is dominated by the thickened septa, few tabulae

rising towards the centre are seen. Dissepiments and minor

septa seem to be absent.

Discussion: The specimens belong to the group of

zaphrentoid corals. Studies of British coral workers (Vaughan

1906; Carruthers 1910; Hudson 1942) clearly showed the

need for detailed phylogenitic studies of the single coral and

of populations. This is not possible with the Jerada material

because of the small number of specimens and their bad

preservation. However, the transverse sections of these

specimens are similar to section in medial parts of Am-

plexizaphrentis enniskilleni (Milne-Edwards and Haime

1851) specimens (Lewis 1930; Hill 1940). However, the

characteristic variability of the fossula of that species can-

not be observed in the Jerada specimen, because the mature

stages are absent. The specimens are, in some respects,

provisionally assigned to the genus Zaphrentites. Further

specimens may help to elucidate their taxonomic position.

Suborder Aulophyllina Hill, 1981

Family Palaeosmiliidae Hill, 1940

Genus Palaeosmilia Milne-Edwards and Haime, 1848

Type species: Palaeosmilia murchisoni Milne-Edwards

and Haime, 1848

Palaeosmilia murchisoni Milne-Edwards and Haime,

1848 (Fig. 4h)

*1848 Palaeosmilia murchisoni Milne-Edwards and

Haime: 261.

?1999 Palaeosmilia murchisoni Milne-Edwards and

Haime.—Liao and Rodrıguez: 551, figs. 5.4–5.5.

2005 Palaeosmilia murchisoni Milne-Edwards and

Haime.—Aretz and Nudds: 172, pl. 1 fig. 5. (cum. syn.)

Holotype: specimen BM 48398, Bowerbank Collection.

Locus typicus and stratum typicum: Frome, Somerset,

UK; Visean. Doubts about the age and locality are sum-

marized in Hill (1940).

Diagnosis: See Semenoff-Tian-Chansky (1974)

Material: Three specimens from P1 (GIK 2183), P4

(GIK 2181) and P6 (GIK 2182).

Description: All fragments fit well into the variability

of this species (Semenoff-Tian-Chansky 1974). The best

preserved specimen is cylindrical and the calice is well

visible, but lacks most of the apical parts and the wall. The

calicular platform (max. 2.3 cm wide) surrounds a deep

U-shaped calicular pit (3.0 cm in width), which is sur-

mounted by a gentle mounted calicular boss (\0.5 cm

high, 1.5 cm wide). The 0.5 cm deep cardinal fossula is

well visible. Maximum diameter is up to 7.0 cm. In the

mature stages there are two rows of 82–94 septa. Many

septa show a marked thickening in the dissepimentarium

when arrangements of the dissepiments become more

regular, probably corresponding to the inner boundary of

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 327

123

328 M. Aretz

123

the calicular platform. Minor septa are 1/2 to 2/3 as long as

the major septa. Septa are straight to slightly sinuous in the

dissepimentarium, whereas in the tabularium the major

septa become more sinuous and may touch each other near

the axis. In the transverse section the dissepimentarium is

three-folded, an outer zone of lonsdaleoide and sometimes

naotic dissepiments, is followed by a middle zone of

more irregular dissepiments, and finally an inner zone of

numerous rows of often elongated, regular dissepiments.

Discussion: Semenoff-Tian-Chansky (1974) showed the

large variability of the species and skipped the formerly

defined sub-species (Hill 1938–1941). The Jerada speci-

mens differ by a lower diameter/septa ratio from P. mul-

tispetata Semenoff-Tian-Chansky 1974, which might be a

more compact variant of P. murchisoni. P. ressoti Menc-

hikoff and Hsu 1935 differs by its smaller size, a more open

dissepimentarium, and a large open axial space.

Distribution: This species is well known from the lower

Visean to Serpukhovian in Europe and North Africa

(Semenoff-Tian-Chansky 1974; Mitchell 1989; Aretz 2002)

Family Aulophyllidae, Dybowski, 1873

Genus Axoclisia Semenoff-Tian-Chansky, 1974

Type species: Axoclisa cuspiforma, Semenoff-Tian-

Chansky, 1974

?Axoclisa sp. (Fig. 4i, k)

Material: Large fragment of a solitary coral from P6

(GIK 2184).

Description: The solitary coral is enclosed in a limestone

block. Its maximum diameter is 2.7 cm. There are two series

of 62 septa. Major septa are long and thickened at the base of

the tabularium. Only the cardinal septum (?) is attached to

the axial structure. The minor septa are very short and often

persist discontinuously in the outer dissepimentarium (septal

spines). The wall is thin. The axial structure (clisophyllid)

consists of a thickened axial plate, which is surrounded by

numerous lamellae and tabellae. Its maximum length is

1.0 cm. The dissepimentarium is up to 5 mm thick and very

irregular. Herringbone dissepiments are observed in the

outer rows, but also occasionally transeptal dissepiments

develop next to the wall. A longitudinal section, oblique to

the axial structure, shows numerous rows of globose to

elongated dissepiments. Tabulae are incomplete.

Discussion: The attribution of this single specimen is

not free from doubts. The axial structure is clearly cliso-

phyllid, but transeptal dissepiments are uncommon in

Clisiophyllum Dana, 1846. Semenoff-Tian-Chansky (1974)

established the genus Axoclisia for those specimens

between the two genera Clisiophyllum and Axophyllum,

thus having a clisophyllid axial structure and transeptal

dissepiments. The genus is so far monospecific and only

three specimens are known; the holotype from the Bechar

Basin (Algeria), and two specimens from SW Spain

(Rodrıguez et al. 2001b). The Jerada specimen differs from

these specimens described as Axoclisia cuspiforma Seme-

noff-Tian-Chansky 1974 in much shorter minor septa, a

higher number of septa-to-diameter ratio, less numerous

developed transeptal dissepiments, a considerably more

complicated dissepimentarium, and a larger axial structure.

A definitive attribution to the genus and/or the known

species requires more material.

Genus Dibunophyllum Thomson and Nicholson, 1876

Type species: Clisiophyllum bipartitum McCoy 1849

Dibunophyllum bipartitum (McCoy 1849) (Fig. 4l)

*1849 Clisiophyllum bipartitum McCoy: 2.

1992 Dibunophyllum bipartitum (McCoy).—Rodrıguez

and Falces: 193, pl. 17 fig. 1.

?1999 Dibunophyllum bipartitum (McCoy).—Liao and

Rodrıguez: 546, figs. 6.1–6.2.

2005 Dibunophyllum bipartitum (McCoy).—Aretz and

Nudds: 172, pl. 2 figs. 1–6. (cum. syn.)

2005 Dibunophyllum bipartitum (McCoy).—Cozar and

Somerville: figs. 12, 13.

2007 Dibunophyllum bipartitum (McCoy).—Rodrıguez

and Somerville: pl. 1 fig. 1.

2007 Dibunophyllum bipartitum (McCoy).—Aretz and

Nudds: pl. 1 fig. 2.

2008 Dibunophyllum bipartitum (McCoy).—Said and

Rodrıguez: 31, figs. 6g–i, 8.

Holotype: specimen SM A 1971, W. Hopkins Collec-

tion (chosen by Hill 1938).

Locus typicus and stratum typicum: Derbyshire, UK;

Lower Carboniferous.

Diagnosis: See Hill (1938) and Poty (1981)

Material: Single specimen from P4 (GIK 2185).

Description: The calice of this *6 cm long partly

eroded solitary coral is not preserved. Corallum curved in

early stages, upper part almost cylindrical. The maximum

diameter is 2.5 cm. A single transverse section is from the

mature stage of the corallite.

There are two series of 58 septa. The major septa are

long and reach towards the axial structure. They are

thickened at their base (forming parts of the wall?). An

open cardinal fossula is developed. Minor septa are very

Fig. 4 a–f Siphonophyllia samsonensis (Salee, 1913); a–e transverse

sections, f longitudinal section, all specimens 91, P4; a GIK 2163,

b GIK 2160, c GIK 2157, d GIK 2159, e GIK 2162, f GIK 2161.

g ?Zaphrentites sp.; transverse section, 93, GIK 2201. h Palaeosmiliamurchisoni Milne-Edwards and Haime, 1848; transverse section, 91,

P4, GIK 2181. i–k ?Axoclisia sp.; transverse sections, 92, P7, GIK

2184. l Dibunophyllum bipartitum (McCoy 1849); transverse section,

92, P4, GIK 2185. m–o Rylstonia sp.; m, n transverse sections, olongitudinal section, all sections 92, P8, GIK 2186

b

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 329

123

short, and persist only into the outer rows of dissepimen-

tarium. There are 6–8 rows of mainly irregular dissepi-

ments. The oval axial structure comprises about one-third

of the corallite diameter. It consists of a long medial plate,

five septal lamellae on each side, and numerous axial

tabellae.

Discussion: The specimen fits well into the character-

istic variability of this well-known species. Federowski

(1971) described a series of species from the same genus,

but they may either fall into the variability of D. biparti-

tum, as currently defined, or show marked differences to

the Jerada specimen in the diameter/septa ration, the

complexity of the axial structure, and the arrangement of

the dissepimentarium.

Distribution: The species is abundant in many late

Visean shallow water carbonate platforms of Europe.

Records for NW Africa are from the Bechar Basin and

Adarouch area (Semenoff-Tian-Chansky 1974; Said and

Rodrıguez 2008).

?Genus Rylstonia Hudson and Platt, 1927

Type species: Rylstonia benecompacta Hudson and

Platt, 1927

Rylstonia sp. (Fig. 4m–o)

Material: One specimen from P8 (GIK 2186).

Description: The ceratoid solitary coral is 43 mm high

and up to 19 mm in diameter. Calice, apex, and wall are

absent. There are up to 33 thick septa in two series. The

major septa are strongly thickened, especially in the car-

dinal half of the corallite. Minor septa are short. A marked

cardinal fossula is developed. The centre of the corallite is

composed of a prominent, compact axial structure to

which most major septa are fused. The axial structure is

5 mm long and 2–4 mm wide, and consists of a complex

amalgamation of an axial plate and axial lamellae. In

transverse section the dissepimentarium is 2.0–4.0 mm

thick and consists of 3–7 rows of steeply declined elon-

gated dissepiments. The tabulae are incomplete and axially

domed.

Discussion: This specimen shows similarities with the

specimens of the Bechar Basin attributed to this genus by

Semenoff-Tian-Chansky (1974). However, the character-

istic variability of the axial structure, which specimens

from the UK show, has not been observed in the Jerada

specimen.

Family Lithostrotionidae d’Orbigny, 1852

Subfamily Lithostrotioninae d’Orbigny, 1852

Genus: Lithostrotion Fleming, 1828

Type species: Lithostrotion striatum Fleming, 1828

(=Madrepora vorticale Parkinson, 1808).

Lithostrotion maccoyanum Milne-Edwards and Haime,

1851 (Fig. 5a)

*1851 Lithostrotion M’Coyanum Milne-Edwards and

Haime: 444.

2005 Lithostrotion maccoyanum Milne-Edwards and

Haime.—Aretz and Nudds: 174. (cum. syn.)

2005 Lithostrotion maccoyanum Milne-Edwards and

Haime.—Cozar and Somerville: fig. 12, 9.

Lectotype: specimen Z 87 illustrated by Milne-Edwards

and Haime (1852: pl. 17 fig. 2a), MNHNP; (chosen by

Semenoff-Tian-Chansky and Nudds 1979).

Locus typicus and stratum typicum: Oswestry, UK;

lower Carboniferous (probably Brigantian).

Diagnosis: See Semenoff-Tian-Chansky and Nudds

(1979)

Material: One colony and a fragment from P4 (GIK

2187, 2188).

Description: The maximal diameter of the colony is

25 cm and it is 10 cm in height. The shape of the colony is

two-folded into a bulbous lower half and a tabular upper

half. The epitheca is usually absent, but on the bottom of

the colony concentric growth indicates the growth centre.

The calicular pit is, on average, 1–2 mm deep, the calicular

boss consists of a single elongated axial plate. Attached

septa may rise towards the columella

Single corallites are of polygonal shape, with prefer-

ences of pentagonal to hexagonal corallites. There are 9–

12, average 10, septa in two series. The major and minor

septa are long. Major septa are often attached to the axial

plate, thus forming a spider-web structure. There are 3–7

rows of flat-lying regular to elongated relatively small

dissepiments. The diameter of the tabularium is less than

2 mm. Tabulae are incomplete and rise towards the axis.

Discussion: L. maccoyanum is the smallest cerioid

species of the genus in the West European faunal province,

and thus its recognition is relatively easy. The Jerada

specimens represent a morphotype with relatively small

corallite diameters within the variability of this species

described by Semenoff-Tian-Chansky and Nudds (1979),

Poty (1981) and Aretz (2002).

Distribution: The species is known from the late As-

bian-early Brigantian interval in NW Europe (Aretz and

Nudds 2005). In Morocco Said et al. (2007) listed it for the

Asbian of the Adarouch area.

Lithostrotion vorticale (Parkinson, 1808) (Fig. 5b)

*1808 Madrepora vorticalis Parkinson: 45, pl. 5,

figs. 3, 6.

2005 Lithostrotion vorticale (Parkinson).—Aretz and

Nudds: 175, pl. 2 fig. 8. (cum. syn.)

330 M. Aretz

123

2005 Lithostrotion vorticale (Parkinson).—Cozar et al.:

fig. 12/3

Types: The Parkinson specimens are regarded as lost. A

lectotype has not yet been chosen.

Diagnosis: See Poty (1981).

Material: One colony fragment from P1 (GIK 2189),

one colony fragment from P3 (GIK 2190), 18 colony

fragments (GIK 2191-2207) from P4, one from P5 (GIK

2208) and one from P7 (GIK 2209).

Description: A well preserved tabular colony enables

description of several outer characters. The corallum is

15 cm in diameter and 7 cm high. The growth is mainly

peripheral. The bottom surface shows a growth centre of

seven corallites. Usually the single corallites are of polyg-

onal shape, with preference of hexagonal outlines. Where

corallites are not in direct contact with each other corallite

walls become curved. This is not only seen on the bottom

surface of the corallum, but also in areas within the colony

where corallites were partly killed by sediment supply.

Especially in those latter areas corallites look like a Siphono-

dendron corallite. Fragments of further colonies show a

relatively deep calice surmounted by a thin axial structure.

There are two series of 18–21 major septa in mature

stages. Cardinal and counter septum are differentiated by

the orientation of the styliform columella, and are usually

attached to it. Further major septa are only occasionally

attached. Minor septa persist shortly into the tabularium.

Major and minor septa are more sinuous in the dissepi-

mentarium than in the tabularium. The tabularium is 3.0–

4.4 mm wide and consists of incomplete tabulae, which

increase towards the axis. The dissepimentarium consists

of 2–7 rows of globose to elongated dissepiments, which

are slightly declined. Transeptal dissepiments may occur in

the peripheral parts of corallites, especially in corners and

in offset structures.

Discussion: Based on morphometric criteria (e.g. width

of tabularium, number of septa) the Jerada specimens can

be easily placed in this species; L. maccoyanum and

L. decipiens (McCoy 1849) are smaller, L. araneum larger.

Distribution: In Europe the species extends from the

Holkerian/Livian to the Late Brigantian (Mitchell 1989,

Rodrıguez et al. 2002). In Morocco Said et al. (2007)

reported it from the Brigantian of the Adarouch area and

Semenoff-Tian-Chansky (1985) from the Visean of the

Algerian Bechar Basin.

Lithostrotion araneum (McCoy 1844) (Fig. 5c)

*1844 Astrea aranea McCoy: 187, pl. 27 fig. 6.

1981 Lithostrotion araneum (McCoy).—Poty: 20,

Figs. 10, 15, 16, pl. 4 figs. 1–4. (cum. syn.)

1992 Lithostrotion araneum (McCoy).—Rodrıguez and

Falces: 198.

2002 Lithostrotion araneum (McCoy).—Rodrıguez

et al.: 22, figs. 5, 8.

2005 Lithostrotion araneum (McCoy).—Cozar et al.:

fig. 12/4.

2005 Lithostrotion araneum (McCoy).—Cozar and

Somerville: fig. 12, 8.

2006 Lithostrotion araneum (McCoy).—Gallagher et al.:

fig. 13, 1.

Syntypes: specimen F7467/1-3, NMD. In Mitchell

(1989) specimen 50-1926 is called holotype.

Locus typicus and stratum typicum: Ireland; Lower

Carboniferous.

Diagnosis: See Poty (1981)

Material: Single specimen from P9 (GIK 2210).

Description: A fragment of a colony, which is 10 cm

wide and 2.5 cm high. Corallites are polygonal (up to

15 mm wide). There are, on average, 24 septa in two series.

The major septa are long, slightly to highly sinuous. The

may or may not attain the axial structure. In some corallites

major septa are fused next to the axial structure. Minor

septa persist a short distance into the tabularium. The axial

structure is a lenticular plate of variable thickness. The

orientation of the axial plate defines cardinal and counter

septum, which are otherwise not recognizable. The tabu-

larium is larger than 4.2 mm, and consists of incomplete

tabulae. Numerous rows of dissepiments are observed.

Discussion: This fragment has by far the largest values

for corallite diameter, number of septa, and tabularium

width of all recovered Lithostrotion specimens from

Jerada. It well fits into the morphological variability of

L. araneum described from Europe (Poty 1981, Rodrıguez

et al. 2002).

Distribution: In Europe L. araneum appears in the

middle Visean (Corphalie Member of the Lives Formation;

Livian) and continues into the Asbian. Said et al. (2007)

listed it for the upper limestone horizon of the Tizra

Formation.

Genus Siphonodendron McCoy, 1849

Type species: Siphonodendron pauciradiale (McCoy,

1844)

Siphonodendron junceum (Fleming, 1828) (Fig. 5d)

*1828 Caryophyllia juncea Fleming: 508.

2005 Siphonodendron junceum (Fleming).—Aretz and

Nudds: 176, pl. 2 figs. 9–10. (cum. syn.)

2007 Siphonodendron junceum (Fleming).—Rodrıguez

and Somerville: pl. 1 fig. 2.

Lectotype: specimen C4203, Ure Collection, HM.

Locus typicus and stratum typicum: Rutherglen,

Lanarkshire, UK; Lower Carboniferous.

Diagnosis: See Kato (1971) and Poty (1981).

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 331

123

332 M. Aretz

123

Material: Nine colony fragments from P4 (GIK 2211-

2219).

Description: Fragments of phaceloid colonies up to

15 cm in length and 10 cm in height. The largest fragment

shows the existence of a spreading centre in the middle of

the corallum, thus indicating medial growth and a bulbous

colony shape. The distances between corallites are, in

general, up to few millimetres. The length of individual

corallites can attain several centimetres. The increase is

lateral.

Mature corallites are 2.0–3.2 mm in diameter, and there

are, on average, 13–14 major septa; the maximum value is

16. Minor septa, if developed, are very short spines. The

major septa are variable in length and thickness. However,

in many cases they leave an almost open space in the

central part of the corallite, and only cardinal and counter

septum may attach to the columella. The columella is a

single axial plate, but not present in all corallites. These

diphymorphic corallites are randomly distributed. A dis-

sepimentarium is not observed. The presence of a colu-

mella defines the morphology of the tabularium.

Incomplete tabulae generally rise convex towards the

columella. A depression of the tabulae is often developed

near the wall. In diphymorphic corallites the tabulae are

complete and arranged horizontally in the corallite centre.

Discussion: The Jerada specimens fit well into the

recorded variability of that taxon, which is the most

unusual Siphonodendron species, because of the lack of

dissepiments. As typical for many occurrences the intra-

specific variability in a single outcrop is very limited.

Compared with specimens from Royseux, Belgium, and

Little Asby Scar, Britain (Aretz 2002, Aretz and Nudds

2005) the corallite sizes at Jerada are slightly smaller, and

thus the average number of septa shifts to lower values.

Distribution: This species is characteristic of the late

Asbian and Brigantian of NW Europe and SW Spain (Poty

1981, Mitchell 1989, Rodrıguez and Somerville 2007). In

Morocco Said et al. (2007) listed it for the Adarouch area.

Siphonodendron pauciradiale (McCoy, 1844) (Fig. 5e)

*1844 Lithodendron pauciradialis McCoy: 189, pl. 27

fig. 7.

2002 Siphonodendron pauciradiale (McCoy).—Poty: pl.

1 fig. 9.

2005 Siphonodendron pauciradiale (McCoy).—Aretz

and Nudds: 176. (cum. syn.)

2005 Siphonodendron pauciradiale (McCoy).—Cozar

and Somerville: fig. 12, 10.

2007 Siphonodendron pauciradiale (McCoy).—Rodrı-

guez and Somerville: pl. 1 fig. 8.

Lectotype: specimen 82-1925, Griffith Collection,

NMD.

Locus typicus and stratum typicum: Magheramore,

Tobercurry, Co. Sligo, Ireland; Glencar Limestone For-

mation, Asbian.

Diagnosis: See Poty (1981).

Material: Single fragment from P3 (GIK 2220).

Description: The fragment of a phaceloid colony is

strongly weathered. The upper surface of the sample shows

well-separated cylindrical corallites, whereas the transverse

thin section shows many offsets and connections between

individual corallites. Corallites have an average diameter

of 4.0 mm (3.7–4.5 mm). There are two series of 20 major

septa (rarely 19–21). Major septa are variable in length.

Cardinal and counter septum may be attached to columella.

The minor septa are short in the tabularium. The columella

is an axial plate which may be thickened. There is one row

of globose dissepiments. The incomplete tabulae increase

towards the axis.

Discussion: The Jerada specimen is a rather large rep-

resentative within the reported species variability. The

single row of dissepiments is a sufficient character to dif-

ferentiate the Jerada specimens from small specimens of S.

irregulare and S. intermedium, which may have similar

number of septa and diameters, but have two rows of

dissepiments.

A badly preserved fragment from P4 (GIK 2221) shares

many characteristics of S. pauciradiale, but has inferior

corallite diameters and, often, a number of septa around the

lower limit for this species. However, because at least one

row of dissepiments is present and the largest and most

complete corallites are clearly within the normal size for the

species, the specimen is determinate as S. cf. pauciradiale.

Distribution: S. pauciradiale appears in Europe in the

early Asbian and persists into the Brigantian. Semenoff-

Tian-Chansky (1985) mentioned it from the late Visean of

the Bechar Basin, and Said et al. (2007) reported it from the

Brigantian of the Adarouch area.

Siphonodendron irregulare (Phillips, 1836) (Fig. 5f, g)

*1836 Lithostrotion irregulare Phillips: 202, pl. 2,

figs. 14, 15.

1852 Lithostrotion irregulare Phillips.—Milne-Edwards

and Haime: 198, pl. 11 fig. 1a–e.

Fig. 5 a Lithostrotion maccoyanum Milne-Edwards and Haime,

1851; transverse section, P4, GIK 2187. b Lithostrotion vorticale(Parkinson, 1808); transverse section, P3, GIK 2190. c Lithostrotionaraneum (McCoy 1844); transverse section, P9, GIK 2210. dSiphonodendron junceum (Fleming, 1828); transverse section, P4,

GIK 2212. e Siphonodendron pauciradiale (McCoy 1844); transverse

section, P3, GIK 2220. f Siphonodendron irregulare (Phillips, 1836);

transverse section, P4, GIK 2222. g Siphonodendron irregulare(Phillips 1836); transverse section, P2, GIK 2227. h Siphonodendronmartini (Milne-Edwards and Haime, 1851); transverse section, P4,

GIK 2229. All scale bars = 5 mm

b

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 333

123

e.p. 1940 Lithostrotion pauciradiale McCoy.—Hill: 169.

?1962 Lithostrotion cf. pauciradiale (McCoy).—Cald-

well and Charlesworth: 377, pl. 14 fig. 4.

1981 Siphonodendron irregulare (Phillips).—Poty: 30,

pl. 12 figs. 1–4.

1986 Siphonodendron irregulare (Phillips).—Herbig:

202, Abb. 4, Abb. 11 figs. 1, 5.

1992 Siphonodendron irregulare (Phillips).—Rodrıguez

and Falces: 203, pl. 19, figs. 1–3.

1997 Siphonodendron martini ecotype 2; Nudds and

Day: 96, fig. 1.

2002 Siphonodendron irregulare (Phillips).—Aretz:

112, pl. 1 fig. 8, pl. 5 fig. 4, pl. 10 fig. 4.

2002 Siphonodendron irregulare (Phillips).—Rodrıguez

et al.: 26, fig. 13a.

2002 Siphonodendron irregulare (Phillips).—Poty: pl. 1

fig. 8.

2005 Siphonodendron sp.—Aretz and Nudds: 178, pl. 3

fig. 1.

2007 Siphonodendron sp.—Aretz and Nudds: pl. 1

fig. 4.

Type material: The specimens of Phillips are regarded

as lost. Poty (1993) described topotypic material from the

locality Ashfell, Cumbria.

Locus typicus and stratum typicum: There is some

confusion on this issue, because Phillips did not indicate a

single locality, and those named contain different Si-

phonodendron species (Hill 1940). If the definition of Poty

(1981, 1993) is accepted, the Middle Visean exposed in the

road cut next to Ashfell, Cumbria, is the type area.

Diagnosis: See Poty and Hannay (1994).

Material: Single colony fragment from P2 (GIK 2222),

and six fragments from P4 (GIK 2223-2228).

Description: Phaceloid corallum with long cylindrical

corallites. Two types of corallite sizes can be differentiated.

The smaller type has corallite diameters of 4.6–6.2 mm and

there are two series of 23–24 septa. The larger type com-

prises specimens with corallite diameters 5.8–6.5 mm and

septal numbers of 21–23.

Both types have a thin wall. The major septa are thin and

straight or sinuous. Some attain a simple axial plate. Its

orientation distinguishes the cardinal and counter septum.

The minor septa are short and persist into the tabularium.

The dissepimentarium consists of one or two rows of

regular dissepiments. In the specimen from P2 diphymor-

phic stages are observed.

Discussion: Aretz and Nudds (2005) discussed the main

problems of the correct specific affiliation of Siphonoden-

dron specimens between the two well defined species

S. pauciradiale (McCoy 1844) and S. martini (Milne-Edwards

and Haime 1851). In this study S. irregulare (Phillips, 1836) is

used as defined by Poty and Hannay (1994).

In the case of the small type it differs from S. martini by

its smaller size, from S. pauciradiale by its larger size and

the second rows of dissepiments, and from S. intermedium

Poty, 1981 by the long major septa attaining the axial plate.

The larger type has the corallite diameter of typical S.

martini, but the number of septa is much lower. The Jerada

specimens do not show an isolated axial plate as commonly

observed in S. intermedium.

Distribution: Because of the confusion on the correct

specific affiliations, the precise occurrence of these species

is uncertain. In Europe and North Africa the name has been

used for specimens of middle and late Visean age.

Siphonodendron martini (Milne-Edwards and Haime

1851) (Fig. 5h)

*1851 Lithostrotion martini Milne-Edwards and Haime:

436.

2005 Siphonodendron martini (Milne-Edwards and

Haime).—Aretz and Nudds: 178, pl. 3 fig. 2. (cum. syn.)

Lectotype: specimen E 1446, Phillips Collection, UMO.

Locus typicus and stratum typicum: Yorkshire, UK,

possibly Teesdale or Ribblesdale; Lower Carboniferous.

Diagnosis: See Semenoff-Tian-Chansky and Nudds

(1979).

Material: One colony fragment from P4 (GIK 2229).

Description: Phaceloid corallum with long cylindrical

corallites. The corallite diameter is 6.0–7.3 mm. There are

23–25 septa in two series. Major septa are long, cardinal

and counter septum attain the styliform columella. The

latter is a single axial plate thickened in its central part. The

minor septa are 1/5 to 1/2 as long as the major septa, often

only entering the tabularium. The septa are straight to

slightly sinuous. There are 2–3 rows of regular globose to

elongated dissepiments. Tabulae are tent-shaped, incom-

plete and in two series.

Discussion: The Jerada specimen is a relatively small

representative within the variability of this species (Se-

menoff-Tian-Chansky and Nudds 1979, Poty 1981).

However, it also shares many similarities with specimens

of S. irregulare from the same locality, except its corallite

size and size-related characters.

Distribution: S. martini is perhaps the most common

species of Siphonodendron in the middle and late Visean of

Europe as Aretz and Nudds (2005) stated. Records in NW

Africa are so far from the late Visean (Semenoff-Tian-

Chansky 1985, Said et al. 2007).

Subfamily Diphyphyllinae Dybowski, 1873

Genus Tizraia Said and Rodrıguez, 2007

Type species: Tizraia berkhlii Said and Rodrıguez,

2007

334 M. Aretz

123

Tizraia berkhlii Said and Rodrıguez, 2007 (Fig. 6a–d)

?2005 Lublinophyllum sp.—Cozar and Somerville:

figs. 12, 6–7.

*2007 Tizraia berkhlii.—Said and Rodrıguez: 27,

figs. 3–8.

Holotype: specimen DPM-TIZ2/5-6.

Locus typicus and stratum typicum:, Adarouch area,

Morocco; upper beds of the Tizra Formation, upper Visean

(Brigantian).

Diagnosis: See Said and Rodrıguez (2007).

Material: Two fragments, one from P7 (GIK 2230), one

from P10 (GIK 2231).

Description: The fragments are from phaceloid colonies.

The increase is parricidal and may occur axial and lateral.

Mature corallites are 7–12 mm in diameter (mean 8.8 mm),

and there are 30–32 major septa; maximum value is 33.

The variably thick corallite wall is generally undulose, but

may also be smooth. The lengths and thickness of the

septa are variable, but they always leave an open space in

the axis of the corallite. Sinuosity of septa is higher in the

dissepimentarium with no difference between major and

minor septa. In general there is a tendency of thickened

septa in the dissepimentarium and/or at the limit of the

dissepimentarium/tabularium. However, in single corallites

there is no thickening at all or it occurs in the tabularium.

Lengths of major septa are 1/3 to 3/4 of the corallite diam-

eter. Minor septa are comparable, variable, and, in general,

1/3 to 1/2 of the major septa long. The dissepimentarium is

variable. It mainly consists of regular dissepiments, but

transeptal dissepiments of both orders regularly occur.

These transeptal dissepiments rarely form a complete circle

in the outer tabularium. There are mature corallites without

any trace of transeptal dissepiments and others with only one

or two transeptal dissepiments of 2nd order.

In the longitudinal section there are 3–4 rows of small

globose to elongated dissepiments forming a relatively thin

dissepimentarium. The tabularium mainly consists of

dome-shaped incomplete tabellae and outer inclined

smaller peripheral tabellae.

Discussion: The Jerada specimens slightly enlarge the

species variability described by Said and Rodrıguez (2007).

Especially the inconsistency of the transeptal dissepiments

and the variability of the septal shapes are noteworthy. This

clearly points to the need for detailed comparison of the

variabilities in Tizraia and Lublinophyllum Khoa, 1977,

because at least some characters used to differentiated

these two genera (Said and Rodrıguez 2007) may fall into

the intra-specific variability. Kubassophyllum Dobrolyub-

ova in Dobrolyubova et al. (1966) should also be included,

because it has many similarities with the genera under

question.

Distribution: Only the type material from the Brig-

antian of the Adarouch area and the two Jerada specimens

of this study can so far be surely assigned to Tizraia

berkhlii. Said and Rodrıguez (2007) mentioned the exis-

tence of specimens in the Jerada Basin in the Djerada

Formation (Eastern Morocco). The term Djerada Forma-

tion might be deduced from older work concerning the

Jerada Coal Basin (Owodenko 1946, 1976), but conse-

quently it must be used for the productive coal seams of

the Westphalian age, where no carbonates are recorded.

Carbonates and also corals are restricted to the interval

Oued El-Koriche Formation to Koudiat Es-Senn Forma-

tion (= upper Visean). Hence, it can be concluded that

Said and Rodrıguez (2007) intended to indicate this

stratigraphic interval.

Subfamily Aulininae Hill, 1981

Genus Aulokoninckophyllum Sando, 1976

Type species: Campophyllum carinatum Carruthers,

1909.

Aulokoninckophyllum carinatum (Carruthers, 1909)

(Fig. 6e, f)

*1909 Campophyllum carinatum.—Carruthers: 150, pl.

1 figs. 3–6.

1998 Aulokoninckophyllum carinatum (Carruthers,

1909).—Niko and Yamagiwa: 139, fig. 7.

2001 Aulokoninckophyllum carinatum (Carruthers,

1909).—Rodrıguez et al.: 90, figs: 3, 4. (cum. syn.)

Holotype: syntypes 1954.6.8-12, RSM (according to

Semenoff-Tian-Chansky 1974).

Locus typicus and stratum typicum: Cap Chernyi,

Nowaja Semlja, Russia; upper Visean.

Diagnosis: see Rodrıguez et al. (2001a)

Material: Four specimens from P4 (GIK 2232-2235),

one specimen from P8 (GIK 2236).

Description: Fragments trochoid solitary corals, which

lack apex, calice, and a large part of the wall. Maximum

height is 3.0 cm, maximum diameter is 18 mm. Areas of

intact wall show the presence of 1 mm thick horizontal

bands possibly growth bands. There are 36–47 septa of two

series. Major septa are up to 7 mm long, and thus leave an

open central space. The minor septa are 3–5 mm long.

They persist some mm or as short septal spines into the

tabularium. All septa are sinuous, carinate (zigzag), and

thickened in the dissepimentarium. A cardinal fossula is

developed; the cardinal septum is about 3/4 as long as the

other major septa. The outer part of the dissepimentarium

is irregular and rarely contains transeptal dissepiments. In

the longitudinal section there are numerous rows of glo-

bose to slightly elongated dissepiments. The tabulae are

incomplete. An aulos is discontinuously developed. In the

axial area/aulos they are almost horizontal, and axial

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 335

123

336 M. Aretz

123

tabellae occasionally occur; in the peripheral zone they are

replaced by declined or curved tabellae.

Discussion: The Jerada specimens fit well into the

variability described by Rodrıguez et al. (2001a). The

largest specimen has a diameter/septa ratio comparable

with that of the monotypic A. amarensis (Semenoff-Tian-

Chansky 1974), but the latter differs by fewer axial tabellae

and a narrower dissepimentarium.

Distribution. The species is known from the late Vis-

ean—Serpukhovian of Europe, Asia, and North Africa,

possibly also Queensland (Hill 1981).

Suborder Lonsdaleiina Spasskiy, 1974

Family Axophyllidae Milne-Edwards and Haime, 1851

Genus Axophyllum Milne-Edwards and Haime, 1850

Type species: Axophyllum expansum Milne-Edwards

and Haime, 1850

Axophyllum aff. pseudokirsopianum Semenoff-Tian-

Chansky, 1974 (Fig. 6g, h)

Material: One specimens from P6, GIK 2237.

Description: A fragment of a solitary corals, the rela-

tively thick wall is only partly preserved. The diameter is

about 2.5 cm. There are two series of 42 septa. The major

septa are long and slightly sinuous. The minor septa are not

more than half as long as the major septa, and only persist

few mm into the tabularium. The axial structure comprises

about one-third of the corallite. It consists of a slightly

sinuous partly thickened axial plate, several thickened,

curved axial lamellae, and somewhat thinner but also

curved axial tabellae. The dissepimentarium comprises

large transeptal dissepiments of two orders, which are

elongated and declined in the longitudinal section. Tabulae

are incomplete.

Discussion: The Jerada specimen is a typical larger

sized Axophyllum. It differs from A. pseudokirsopianum

Semenoff-Tian-Chansky, 1974 by its shorter minor septa,

and a slightly larger dissepimentarium. A large dissepi-

mentarium is characteristic for A. parkinsoni (Ryder,

1930), but that of the Jerada specimen is less thick. A

similar length of minor septa is known from A. kirsopia-

num (Thomson, 1880), but naotic septa have not been

observed as in the latter.

Genus Pareynia Semenoff-Tian-Chansky, 1974

Type species: Pareynia splendens Semenoff-Tian-

Chansky, 1974

Pareynia splendens Semenoff-Tian-Chansky, 1974

(Fig. 6i, k, l)

*1974 Pareynia splendens splendens Semenoff-Tian-

Chansky: 241, Figs. 93, 94, 96, pl. 62 fig. 7, pl. 63 figs. 1,

2, pl. 64 figs. 1–4, pl. 65 figs. 2, 3, pl. 73 figs. 1, 2.

?1974 Pareynia gangamophyllides Semenoff-Tian-

Chansky: 248, Fig. 95, 96, pl. 63 fig. 3, pl. 65 fig. 4,

pl. 66 fig. 1-6, pl. 70 fig. 1, 2, pl. 74 fig. 1-4.

cf. 1978 Pareynia splendens Semenoff-Tian-Chansky.—

Poty in Kimpe et al.: pl. 6 fig. 6.

1981 Pareynia splendens Semenoff-Tian-Chansky.—

Poty: 64, pl. 31 figs. 1, 2.

?1998 Pareynia? sp. indet.—Niko and Yamagiwa: 141,

fig. 8.

Holotype: specimen PAR 382/93. Semenoff-Tian-

Chansky collection, MNHNP.

Locus typicus and stratum typicum: Piton a l’Est du

Meharez el Kebir, Bechar Basin, Algeria; Ioucha 21

Member, upper Visean.

Diagnosis: (Semenoff-Tian-Chansky 1974 and Poty

1981). Pareynia with large dissepimentarium, which may

comprise up to the half of the corallite; short, but well

developed minor septa; variable axial structure, especially

the thickness of the axial plate; thin to medium-thick wall

present; variable corallite diameter throughout growth

(common rejuvenesence?); lateral increase possible.

Material: One specimens from P6 (GIK 2238), two

from P7 (GIK 2239-2240).

Description: The large solitary corals are completely

surrounded by calcareous matrix, thus the external char-

acters of the specimens are not observed. The maximum

diameter is 5.6 cm, however during growth it strongly

varies between zone of extension and contraction, resulting

in a very irregular corallite shape. A thin to medium-thick

undulating wall is present, but may be less visible in the

areas where the growth style changes; additionally these

areas are preferentially eroded.

There are two series of up to 55 septa. The major septa

are long and slightly sinuous. In the outer dissepimentari-

um they are interrupted by transeptal dissepiments of 1st

order. The maximum length of minor septa is half that of

the major septa. The axial structure is about 1 cm in

diameter. There are numerous tabellae and strongly curved

axial lamellae. The thickness of the central axial plate is

variable, thus resulting in some sections in a gangamo-

phyllid appearance of the axial structure. The dissepi-

mentarium can be separated in a vast outer zone of

Fig. 6 a–d Tizraia berkhlii Said and Rodrıguez, 2007; a, b transverse

sections, GIK 2230, c, d longitudinal sections, all scale bars = 5 mm,

P7, GIK 2231. e, f Aulokoninckophyllum carinatum (Carruthers,

1909); transverse sections, 92, P8, GIK 2236. g, h Axophyllum aff.

pseudokirsopianum Semenoff-Tian-Chansky, 1974; transverse sec-

tions, 92, P6, GIK 2237. i, k, l Pareynia splendens Semenoff-Tian-

Chansky, 1974; i, l transverse sections, 92, I: GIK 2239, l GIK 2240,

k longitudinal section, 91, P7, GIK 2240

b

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 337

123

transeptal dissepiments of both orders, and an inner dis-

sepimentarium of 3–5 rows of regular dissepiments.

Tabulae are incomplete, clinotabulae are developed. Ele-

ments of the axial structure are much thicker than dissep-

iments and tabulae. Tabellae rise towards the axis.

Discussion: The genus Pareynia is rarely reported and

thus the morphological variabilities are poorly known. The

irregular growth observed in the Jerada specimens combined

with partly eroded areas of the corallites may explain some of

the difficulties with interpretation of this genus. However, the

Jerada specimens significantly increase the likelihood that

the axial structure is more variable, especially the thickness

and shape of the axial plate, as currently stated for the genus.

Hence P. gangamophylloides may fall in the variability of

P. splendens. For the same reason, use of subspecies defined

by Semenoff-Tian-Chansky (1974) is questionable.

Occurrence: Upper Visean of Algeria, Belgium and

probably Japan.

Discussion

Biostratigraphic position

Owodenko (1946, 1976) dated the studied interval as late

Visean based on brachiopods and ammonoids. Vachard and

Berkhli (1992) used carbonate microfossils and concluded

an almost exclusive Brigantian age with a possible Asbian

age for the base of the succession, and a Serpukhovian age

for a 10 m thick limestone package in the Oued Agaia area

(SW end of synclinorium). The corals collected in this

study are used to discuss these ages and to indicate pos-

sibilities for refinements (Fig. 7).

The basal volcanic rocks cannot be dated by biostrati-

graphic tools; the primitive carbonate microfossils

(Pachysphaerina and Diplosphaerina) found in thin pack-

ages of intercalated calcareous shales (Vachard and Berkhli

1992, Berkhli et al. 1999) do not have a biostratigraphic

value. The Siphonodendron irregulare colony recovered

from the lower third of the Cafcaf Formation on the

northern flank indicates a maximum age of middle Visean

(Livian/Holkerian) and a minimum age of late Visean.

Most corals recorded from the succeeding formations have

limited biostratigraphic potential. They are all known from

late Visean coral faunas, but first appearance dates (FADs)

are often uncertain. It seems to be reasonable to assume at

least a middle Visean age for the Cafcaf Formation,

especially because the accumulation of bedded cherts in

that thickness (up to 40 m), requires a substantial amount

of time for deposition. The middle/late Visean boundary is

hard to trace and might still be placed in the bedded chert

facies.

The age of the fauna of the Oued El-Koriche Formation

is not unequivocal. The biostratigraphic range of Litho-

strotion vorticale is rather large (Holkerian–Brigantian).

Lithostrotion araneum is known in the Holkerian–Asbian,

but not from the Brigantian. Tizraia is so far only known

from the Brigantian (Said and Rodrıguez 2007). Thus, in

any case, the biostratigraphic range of Lithostrotion ara-

neum or Tizraia has to be extended.

The fauna of the Oued Es-Sassi Formation contains

Aulokoninckophyllum carinatum, which is late Asbian/

Brigantian in age. The composition of the coral faunas in

the entire Koudiat Es-Senn Formation indicates at least an

Asbian age. A clear late Asbian FAD has Dibunophyllum

bipartitum found in the upper Koudiat Es-Senn Formation

Fig. 7 Distribution of rugose

corals in the Visean succession

and resulting stratigraphic

divisions, with comparison with

ages based on carbonate

microfossils (Vachard and

Berkhli 1992; Berkhli et al.

1999). HOL, Holkerian; LAS,

lower Asbian; UAS, upper

Asbian; AS, Asbian; LBR, lower

Brigantian; UBR, upper

Brigantian; ARN, Arnsbergian

(E2)

338 M. Aretz

123

(Aretz and Nudds 2005). Pareynia is a taxon, which is

generally thought to appear ‘‘high’’ in the late Visean

(Brigantian) (Poty 1981), but its FAD is very uncertain

because of its rarity. Two species recovered from the

youngest levels in the Koudiat Es-Senn Formation,

Siphonodendron junceum and Lithostrotion maccoyanum,

have a late Asbian FAD and both persist into the Brigan-

tian. L. maccoyanum is unknown from the upper Brigantian

in NW Europe (Aretz and Nudds 2005). Note that Rodrıguez

and Somerville (2007) do not differentiate the Brigantian in

their distribution pattern. Thus if the temporal distribution of

these two species in NW Africa is the same as in NW Europe,

the top of the Koudiat Es-Senn Formation is not older than

late early Brigantian.

The interpretation of these coral data is critical, because

diagnostic taxa are rare (Fig. 7). However, it is obvious that

taxa characteristic for many Brigantian faunas (e.g. colo-

nial axophyllids, Palastrea, Corwenia; Mitchell 1989,

Rodrıguez and Somerville 2007) do not occur in the faunas

collected in the Jerada Basin. Their absence might be best

explained by three possibilities:

1. so far these taxa have been overlooked;

2. the succession is mainly older than the Brigantian; or

3. an impoverished fauna has been recovered, which

lacks the typical marker.

None of these three arguments can definitely be ruled

out, but it must be stressed that those taxa indicating a

Brigantian age, for example Tizraia and Pareynia, are both

only reported from few localities. Additionally their

stratigraphic range is strongly questionable, because their

occurrence might be strongly facies-controlled (see below).

Hence a conservative age model based on the corals

(Fig. 7) indicates as possible age for the Cafcaf Formation

the Middle Visean and possibly basal late Visean, early

and/or, late Asbian for the Oued El-Koriche and Oued Es-

Sassi formations, and late Asbian and/or early Brigantian

age for the Koudiat-Es-Senn Formation. The hiatus around

the Visean/Serpukhovian boundary would at least comprise

the upper Brigantian and the basal Serpukhovian.

This interpretation contradicts the data of Vachard and

Berkhli (1992) and Berkhli et al. (1999), and thus either the

postulated ranges for corals or carbonate microfossils have

to be reconsidered (Fig. 7). This is of special importance

because Jerada is the key region for the Brigantian (Cfm 7)

in the Moroccan biozonation of carbonate microfossils

(Vachard in Berkhli et al. 1993).

New ammonoid data of Korn and Ebbighausen (2008)

from pelites in the Chebket el Hamra area, in the NE of the

Jerada synclinorium, partly help to elucidate this problem.

These data are from a somewhat deeper and almost car-

bonate-free part of the basin. A basal late Asbian assem-

blage is followed by four assemblages of early and late

Brigantian age, thus showing that at least in deeper parts of

the basin sedimentation continued into the later Brigantian.

The stratigraphic correlations into the shallow parts of the

basin are so far not well constrained, because of tectonic

complication and the partial absence of good lithostrati-

graphic and biostratigraphic marker horizons. However, the

succession studied by Korn and Ebbighausen (2008) is

lithologically similar to the interval Cafcaf to Oued

Es-Sassi formations, the absence of limestone beds makes

the Koudiat Es-Senn Formation unrecognizable.

The coral fauna recorded from the Adarouch area (Said

et al. 2007, Said and Rodrıguez 2007, 2008) is more

diverse than those at Jerada, but especially the assemblages

from the lower part of that succession (Tizra Formation)

show a comparable composition. Said et al. (2007) indi-

cated an Asbian age for most of the Tizra Formation (the

uppermost beds are Brigantian in age), and thus these

results possibly strengthen the coral-based interpretation at

Jerada. It is important to mention that L. maccoyanum is

restricted to the Asbian (Said et al. 2007), and that the late

Visean-Serpukhovian succession of the Adarouch area

contains similar depositional environments as those of the

Jerada Massif.

However, all these considerations have to take into

account that the Visean succession of the Jerada Massif

comprises very different depositional environments. Facies

dependencies of the studied organisms are inevitable, and

may largely control the biostratigraphic interpretation.

The composition of the coral faunas recovered from the

individual outcrops clearly shows these facies dependen-

cies (Table 1), and these must also be suspected for groups

such as carbonate microfossils and ammonoids.

Facies

The smallest coral individuals, Aulokoninckophyllum,

Rylstonia and ?Zaphrentitis, have been found in shales of

the Oued Es-Sassi Formation (P8). A specialised fauna,

including Pareynia, Tizraia, Palaeosmilia and ?Axoclisia,

is known from the microbial buildup environments on the

southern flank of the synclinorium (P6, 7, 10). So far all

known occurrences of Tizraia are restricted to this partic-

ular facies. The same can be suspected for Pareynia. In its

type area, Bechar Basin, it was described (Semenoff-Tian-

Chansky 1974) from a region dominated by large sponge-

fenestellid bryozoan buildups (Bourque et al. 1995) with

abundant microbial boundstones, and also the Belgian

specimens (Poty 1981) are very likely to be from the same

kind of environment (Aretz and Chevalier 2007).

A marked differentiation in the composition of a coral

fauna is known from the uppermost Koudiat Es-Senn

Formation (P4) on the northern flank. Here, mass occur-

rences of Siphonophyllia samsonensis are only found in the

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 339

123

vicinity of a coral patch reef which is mainly formed by

Lithostrotion vorticale. Associated is a diverse coral fauna,

which comprises several species of Siphonodendron and

Lithostrotion, and larger solitary corals as Palaeosmilia

and Dibunophyllum. The small solitary corals (Au-

lokoninckophyllum and ?Zaphrentites) recovered from P4

are most probably associated with calcareous shales locally

intercalated and also present some meters below the patch

reef, hence confirming the observations from the Oued Es-

Sassi Formation. The patch reef formed next to an oolitic

shoal which contains Siphonodendron pauciradiale and

Lithostrotion decipiens (P3). The composition of the fauna

in P1 has some resemblance to that of P4, especially the

dominance of S. samsonensis and L. decipiens, but fewer

specimens were collected, and (thus) the fauna is less

diverse. However, the presence of a further coral patch reef

of the type found at P4 may be envisaged for P1.

In P2, P5, and P8 all corals are reworked and represent

more shallow facies than the depositional environment they

are embedded in. Their distribution pattern is random and

not controlled by any ecological preferences.

Overall, the compositions of assemblages, in particular

facies, show that large solitary corals and most colonial

taxa have a preference for shallow and turbulent water

whereas smaller corals represent unfavourable conditions

(siliciclastic input, deeper water?).

Palaeobiogeography

The Jerada fauna comprises taxa similar to the coral faunas

described from the late Visean of the Eastern Central

Meseta, which display comparable depositional environ-

ments (Said et al. 2007; Aretz and Herbig 2009). Differ-

ences mainly result from the total number of samples for

individual localities and slightly different ages. Although

the fauna of the Bechar Basin (Semenoff-Tian-Chansky

1974, 1985) is only partly well studied, the available data

indicate a substantial overlap with the Jerada fauna. The

later might be characterised as an impoverished Bechar

fauna. A new aspect for Morocco is the presence of

Pareynia and ?Axoclisa in the Jerada fauna, which have

been described in NW Africa so far only from the Bechar

Basin (Semenoff-Tian-Chansky 1974). Thus, in the late

Visean the Jerada fauna can be interpreted as a link

between the coral faunas of the Central Sahara basins and

the region north of the Central Atlas Fault in Morocco.

Table 1 Overview of the

distribution of rugose corals in

the different late Visean

environments of the succession

in the Jerada Massif

Environment Coral Taxa Formation

Coral patch-reef (core) Lithostrotion vorticale possibly

further lithostrotionids

Koudiat Es-Senn

Formation (P 4)

Coral patch-reef (surroundings and off-reef

facies)

Siphonophyllia samsonensis

Palaeosmilia murchisoni

Dibunophyllum bipartitum

Lithostrotion maccoyanum

Lithostrotion vorticale

Siphonodendron junceum

Siphonodendron irregulare

Siphonodendron martini

Koudiat Es-Senn

Formation (P1, 4)

Oolitic shoal Siphonodendron pauciradiale

Lithostrotion vorticale

Koudiat Es-Senn

Formation (P3)

Silty shale Aulokoninckophyllum carinatum

Rylstonia sp.

Zaphrentites sp.

Oued Es-Sassi

Formation (P8)

Microbial-sponge buildup (core and capping

beds) and microbial-metazoan buildup

Palaeosmilia murchisoni

?Axoclisia sp.

Lithostrotion vorticale

Tizraia berkhlii

Axophyllum aff.

pseudokirsopianum

Pareynia splendens

Koudiat Es-Senn

Formation (P6, 7)

Oued El-Koriche

Formation (P10)

Calciturbidite Lithostrotion vorticale

Lithostrotion araneum

Oued El-Koriche

Formation (P5, 9)

Bedded chert (turbiditic layer?) Siphonodendron irregulare Cafcaf Formation

(P2)

340 M. Aretz

123

Comparison with regions such as Anti-Atlas, western

Central Meseta, and Jebilet are so far impossible because

of the absence of solid data from these regions.

The Jerada fauna comprises many taxa, which are (well-)

known in NW Europe north of the Variscan Orogen. Its

composition clearly indicates its belonging to the West

European faunal province which includes North Africa

(Sando 1990). Comparison of the faunas from NW Europe,

SW Spain, and NW Africa shows that the Eastern Moroccan

Meseta links NW Europe with the central Saharan basins.

Hence the appearance of ‘‘exotic’’ taxa in NW Europe, for

example Pareynia in the Vise area (Poty 1981) or Axoclisia

in SW Spain, might be explainable by migration along a

pathway through the Eastern Moroccan Meseta.

The Jerada material cannot help elucidation of the

relationships of the West-European province to palaeoge-

ographic units further east (circum-Mediterranean, Minor

Asia), because contemporaneous faunas of these regions

are unknown or too poorly studied. However, contempo-

raneous European faunas from the southern side of the

Variscan Orogen (Herbig 1986; Aretz 2002) have a dif-

ferent composition, and contain taxa, for example Kizilia,

which are literally absent from NW Africa. Thus, in the late

Visean an important palaeobiogeographic barrier hampered

faunal exchange between NW Africa and regions south of

the Variscan Orogen, today neighboured as the Betic

Cordillera. The present geographic position may be partly a

result of Late Variscan tectonic activity along important

shear zones (Vai 1991).

Conclusions

1. The lithologically variegated succession of the Late

Visean of the Jerada Massif contains a diversified

rugose coral fauna. It comprises well-known and

geographically widely distributed taxa, for example

Palaeosmilia murchisoni or Siphondendron martini,

but also rare taxa, for example Pareynia splendens and

Tizraia berkhlii.

2. The fauna of the Adarouch area is similar to those of

the herein studied fauna from the Jerada Basin with

regard to composition and age. However, the Jerada

fauna is less diversified, which may be partly the result

of significantly fewer collected specimens.

3. The biostratigraphic ages of the Visean succession

(Cafcaf–Koudiat Es-Senn formations) are difficult to

establish because of missing or poor fauna and facies

changes and dependencies. Carbonate microfossils and

possibly also ammonoids indicate a Brigantian age at

least for parts of the succession. However, the coral-

based biostratigraphic interpretation shows some

differences from these ages. Typical Brigantian taxa,

for example colonial axophyllids and Palastrea, are

absent, and, consequently, coral ages are slightly older.

4. Overall an improved and more detailed biostrati-

graphic framework is needed for NW Africa to

precisely date the first appearance dates of coral taxa.

The facies dependencies described in this study clearly

demonstrate possible complications, but also may help

to explain differences between specific regions in NW

Africa.

5. The appearance of most taxa can be correlated with

distinctive facies. Small solitary corals occur in silty

shales, locally calcareous, whereas larger solitary

corals are found in limestones surrounding a coral

patch reef and on top of large microbial-sponge

buildups. The massive colonial coral Lithostrotion

vorticale is the main framework builder of that small

coral patch reef, which is a relatively unique feature in

Visean reefs. Overall, colonial corals occur in very

different environments. Their higher abundance in

buildup facies at Jerada is linked with the predomi-

nance of this facies as the main carbonate facies in

many parts of the Jerada Basin.

6. Palaeobiogeographically, the fauna recovered at Jerada

is part of the Western European coral province (Sando

1990), which comprises NW Africa. The presence of

taxa, for example Pareynia, Tizraia, and ?Axoclisia,

which are commonly not found in NW Europe,

indicate the position and importance of the Eastern

Moroccan Meseta as a pathway between the Central

Saharan basins and NW Europe. However, more coral

data from NW Africa are needed to reveal the precise

timing and directions of migration patterns.

Acknowledgments This study benefited from the help of H.-G.

Herbig (Koln), D. Korn (Berlin), V. Ebbighausen (Odenthal), and

E. Poty (Liege). Special thanks to Sebastian, Stefan, Arne, and Florian

(all Koln) for the corals they collected during their mapping projects.

A. Mihli and J. El Mokhtari (Direction Regional du Ministere de

l’Energie et des Mines a Oujda) are thanked for permission to work in

the Jerada Basin in the autumns of 2004 and 2006. All officials from

the ‘‘Eau et Foret’’ authority in Jerada are greatly acknowledged for

their hospitality and help during our field campaigns. R. Baumler

(Koln) prepared the thin sections for this study.

Appendix

Location of sample localities based on the topographic

maps (1/50,000) of the region (Jerada, Wad Al Himar) (see

Fig. 2 for approximate positions).

A: Jerada Synclinorium, northern flank:

P1. Eastern slope of Jbel Bou Keltoum, off-patch reef

facies in the upper part of Koudiat Es-Senn Formation;

34�21042N; 2�03023W.

Rugose corals from the upper Visean (Carboniferous) of the Jerada Massif 341

123

P2. Valley flank of the creek east of abandoned barite

mine, lower third of Cafcaf Formation; 34�23015N;

2�00027W.

P3. Eastern side of the valley east of Koudiat Es-Senn;

upper Koudiat Es-Senn Formation; 34�22014N;

2�00014W.

P4. Coral patch reef on the southern slope of the hill east

of Koudiat Es-Senn; upper Koudiat Es-Senn; 34�22015N;

2�00009W.

P5. SW of Small hill (1022 m); Oued El Koriche

Formation; 34�23025N; 1�56023W.

B: Jerada Synclinorium, southern flank:

P6. Top and capping beds of the most western microbial-

sponge buildup and microbial-metazoan buildups in the

Koudiat Es-Senn Formation, west of P7; 34�17058N;

2�03021W.

P7. Large microbial-sponge buildup in the Koudiat

Es-Senn Formation west of the road Oujda-Ain Beni

Mathar and Oued Defla; 34�17059N; 2�03009W.

P8. Silty shales of the Oued Es-Sassi Formation south of

P7; west of the road Oujda-Ain Beni Mathar and Oued

Defla; 34�17055N; 2�03009W.

P9. Calciturbidites within the Oued El Koriche Forma-

tion east Oued Defla, west of P 10; 34�17058N;

2�03001W.

P10. Large olistolite mainly consisting of microbial-

sponge buildup facies east of the road Oujda-Ain Beni

Mathar and Oued Defla; 34�17056N; 2�02057W.

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