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Ammonites and stratigraphic succession of the middle toarcian – Lowermost

aalenian from the jebel amergou (Moulay Bouchta area, internal prerif zone,
external Rif Belt, northern Mo...

Article in Journal of African Earth Sciences · October 2023

DOI: 10.1016/j.jafrearsci.2023.105082

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Ammonites and stratigraphic succession of the middle toarcian – Lowermost
aalenian from the jebel amergou (Moulay Bouchta area, internal prerif zone,

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external Rif Belt, northern Morocco)
Mohamed Benzaggagh
University of Moulay Ismail, Faculty of Sciences, Department of Geology, BP 11.201, Zitoune, Meknes, Morocco

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ARTICLE INFO ABSTRACT

Handling Editor: DR Damien Delvaux In the Rif Belt, the Toarcian–Aalenian outcrops are rare. The most important of these outcrops are those of the
northern side of Jebel Amergou (70 km northwest of Fez). These outcrops, consisting of lumpy marlstones and
Keywords: marly limestones, are rich in ammonites. A section (JAM) located in a small ravine near Douar Amergou have
Ammonites been studied. Its stratigraphic series (20 m-thick), called the Amergou Formation, is divided into two members.
Systematics paleontology The lower member yielded in the levels 3 and 4 a rich ammonite fauna from the middle Toarcian (Bifrons and
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Toarcian–lower aalenian Gradata zones) and some ammonites from the upper Toarcian (Bonarellii and Speciosum? zones); the upper
Internal prerif
member, divide into two levels, yielded in the level 2 rare ammonites from the lower Aalenian (Opalinum Zone).
External rif
44 species of ammonites have been identified, described and illustrated for the first time from the Toar-
Morocco
cian–lower Aalenian succession of the Rif Belt. Most of these species are common to several basins of both, the
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northern and the southern margins of the Tethys and the NW European province.

1. Introduction 2. History of the previous works on the Toarcian–Aalenian

ammonites and successions from the internal prerif zone
The present work is a stratigraphic and paleontological study based
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on a rich Toarcian–lower Aalenian ammonite fauna collected from a
section located northeast of Jebel Amergou (Moulay Bouchta area), be-
longing to the Internal Prerif Zone of the western External Rif belt. Am-
monites of this time interval from the Rif belt are still unknown, never
described or figured in the previous works. Lacoste (1934), then
Bulundwe Kitongo (1987) in their unpublished theses on the geology of
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Daguin (1927a) was the first author having mentioned the pres-
ence of the Toarcian succession in the northern side of the Lower
Jurassic carbonate massif of Jebel Amergou, by the finding of some
specimens of Hildoceras during a short mission in the Moulay
Bouchta area. Lacoste (1934) in his thesis on the geology of the
southern part of the Rif Belt has described (p. 148, pl. 3) a Toar-

the southern part of the External Rif belt, have reported, mainly from
the Moulay Bouchta area, lists of Toarcian and lower Aalenian am-
monites but without any descriptions or illustrations. The collected am-
monites (44 species) complete data from these authors and allow char-
acterizing the Bifrons, the Gradata (middle Toarcian), the Bonarellii,
the Speciosum? (upper Toarcian) and the Opalinum (lower Aalenian)
zones. These species are described and illustrated for the first time from
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cian–lowermost Aalenian section located northeast of Jebel Amergou
in a small ravine near Douar Amergou. The author distinguished in
this section, three stratigraphic levels, from bottom to top.
- White silty marlstone with thin limestone beds having provided
ammonites of the middle Toarcian, Coeloceras crassum Phillips,
Coeloceras sp., Dactylioceras commune J. Sowerby, Hildoceras bifrons

the Internal Prerif Zone and the External Rif Belt, in general. Most of
them are common to several Jurassic basins of the northern and the
southern margins of the Tethys and the NW European province. The
stratigraphic and paleogeographic distributions of each species in these
different paleogeographic domains have been discussed.
Bruguière, H. grunowi Hauer, H. levisoni Simpson, H. mercati Hauer,
Hildoceras sp., Phylloceras heterophyllum J. Sowerby, Phylloceras
nilssoni Hébert, Phylloceras sp. And Rhacophylites;
- Gray limestone and white marlstone alternation containing
ammonites of the middle and upper Toarcian, Hammatoceras
insigne Schübler, Harpoceras bicarinatum (Zieten), H. wunstorfi
Monestier var. Coecilia Reinecke, Haugia grandis Buckman, H.

E-mail address: benzaggagh@gmail.com.

https://doi.org/10.1016/j.jafrearsci.2023.105082
Received 26 June 2023; Received in revised form 21 September 2023; Accepted 2 October 2023
1464-343/© 20XX

Note: Low-resolution images were used to create this PDF. The original images will be used in the final composition.
M. Benzaggagh
variabilis d’Orbigny, Haugia sp., Hildoceras emilianum Reynès,
Lytoceras germaini d'Orbigny, Lytoceras cf. Jurense Zieten,
Phylloceras aveyronnense Meneghini, Polyplectus discoides Zieten
and Pseudolioceras rivierense Monestier.
- Brown limestone and small reddish sandstone beds containing
ammonites of the lowermost Aalenian, Braunsina elegantula
Buckman, Catulloceras (Tmetoceras) scissum Benecke, Dumortieria
cf. Levesquei d’Orbigny and Ludwigella opaca Buckman.
Above this level, the lithofacies consisting of brown and reddish
schistose marlstone with ferruginous nodules and sandstone beds is typ-
ical of the Callovian–Oxfordian “Ferrysch” formation.
Bulundwe Kitongo (1987), in his thesis on the stratigraphy of the
Jurassic carbonate massifs (“sofs”) of the Internal Prerif Zone, studied
several sections of marlstone and marly limestone series lying above
the Lower Jurassic carbonate massifs. The most important of these sec-
tions are those of the northern side of Jebel Amergou. In section 3, cor-
responding to the section described by Lacoste (1934), Bulundwe
Kitongo (1987, p. 260) reported from the level called “Ammonitico-
rosso 2" (5 m-thick), ammonites of the Levisoni and Sublevisoni sub-
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zones, including Harpoceratoides, Harpoceras maghrebense Guex, Hildo-
ceras sublevisoni Fucini, Hildoceras sp. And Nodicoeloceras sp., In a lat-
eral point outcrop 2 (“affleurement ponctuel 2″, p. 261), located at the
mosque of Douar Amergou, about 100 m southeast of section 3, the au-
thor reported Hildaites striatus Guex, H. cf. Striatus Guex and Nodicoelo-
ceras sp. Characterizing the Levisoni Subzone; Hildoceras sp. Character-
izing the Bifrons Zone and Gruneria from the upper Toarcian. In section
2, located 500 m west of section 3, the author distinguished (p. 253),
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in the level “Ammonitico-rosso 2" (14 m-thick), five ammonite faunal
assemblages (called faunes A to E). According to the author, the “Faune
A″, composed of Hildaites striatus Guex, Hildaites sp., Polyplectus pluri-
costatus Haas and the “Faune B″, composed of Harpoceras maghrebense
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Guex, Hildaites sp. And Polyplectus pluricostatus Haas, characterize the
Levisoni Subzone; the “Faune C″ composed of Harpoceras submediterra-
neum (Pinna), Hildoceras sublevisoni Fucini, Mercaticeras sp., and the
“Faune D″ composed of Collina (C.) gemma Bonarelli, Hildoceras sublevi-
soni Fucini, H. semipolitum Buckman and Phymatoceras characterize the
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Sublevisoni and Bifrons subzones and the Gradata Zone; the “Faune E″
composed of Brodieia sp., Crassiceras cf. Gradatum (Merla), Hammato-
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Fig. 1. Structural map of the Rif Belt (from Suter, in: Michard, 1976; amended with data from Suter, 1980a-Suter, 1980b) and location of Moulay Bouchta area.
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Journal of African Earth Sciences xxx (xxxx) 105082
ceras costulosum Merla, Phymatoceras gr. Erbaense (V. Hauer), P. venus-
tulum (Merla), P. gr. Pulcher (Merla) and Vacekia sp. Characterizes the
Gradata Zone. The overlying level, the “Ammonitico-rosso 3" (20 m-
thick), provided Hammatoceratinae, Docidoceras sp., Graphoceras gr.
Limitatum Buckman, Graphoceras sp. And Tmetoceras gr. Scissum (Be-
necke) characterizing the lower Aalenian Opalinum–Murchisonae tran-
sition zones.
3. Geological framework of the Rif Belt
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Authors (Durand Delga et al., 1962; Andrieux, 1971; Suter, 1980a-b;
Wildi, 1983) distinguished in the Rif Belt, three large paleogeographic
and structural domains (Fig. 1): the Internal Rif and the Flysch nappes,
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to the north, and the External Rif, to the south. This latter domain,
which form the larger part of the Rif Belt consists of thick marlstone,
marly limestone and sandy marlstone sequences (4000 to 6000 m-
thick) of Triassic to Middle Miocene age folded and thrusted outwards.
Suter (1965) distinguished in the External Rif, three paleogeographic
zones: the Intrarif Zone, a large zone to the north composed of au-
tochthonous to parautochthonous units (Ketama, Tangier and Loukous
units) and allochthonous units called the “Intrariffian nappes” such as
Sanhadja, Bou Haddoud, Aknoul and Tsoul nappes, in the eastern Exter-
nal Rif; El Habt and Ouezzane nappes, in the western External Rif; the
Mesorif Zone (zone of windows) forming a narrow zone in the median
part of the Rif belt. In this zone, the Jurassic–Cretaceous sequences are
widely covered by the “Intrariffian nappes” and the Middle
Miocene–Pliocene post-orogenic deposits of the Ouerrha and Zoumi
basins.; the Prerif Zone, divided into two subzones, the Internal Prerif
Zone (zone of “Sofs”), to the north, characterized by the dominance of
the Jurassic and Lower Cretaceous outcrops and the External Prerif
Zone (zone of Hills), large zone, to the south, dominated by marlstone,
marly limestone and sandstone deposits mainly of Cenozoic age.
4. Triassic–Upper Jurassic stratigraphic sequence of the External
Rif
- Triassic–earliest Jurassic. It consists of red silty clays with
evaporates and basalt lava flows (Fig. 2) always found in
abnormal positions underline major tectonic contacts (thrusts,
M. Benzaggagh
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Fig. 2. Synthetic Triassic to Lower Cretaceous stratigraphic column of the Ex-
ternal Rif, compiled by data from Suter (1965), Andrieux (1971), Leblanc
(1979), Wildi (1981), Favre (1992), Benzaggagh (2000) and Benzaggagh et al.
(2020).
overlaps) or reworked as olistoliths into the Senonian and/or the
Miocene marlstones of the External Rif.
- Sinemurian–Pliensbachian. It corresponds to a thick carbonate
formation (ca. 500 m-thick) consisting of Sinemurian–lower
Carixian massive limestones with neritic fauna (Bulundwe
Kitongo, 1987; Papillon, 1989; Favre, 1992), and the upper
Carixian–Domerian bedded limestone with pelagic and/or benthic
fauna. Several large exposures of this formation crop out along the
south side of the Middle Ouerrha Valley (e.g., Jebels, Messaoud,
Amergou, Arechgou, Beni Ouassal and Sidi Redouane) overlapping
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Journal of African Earth Sciences xxx (xxxx) 105082
the Lower Cretaceous marlstones of the Internal Prerif Zone
(Benzaggagh, 2016; Benzaggagh et al., 2020). These limestone
blocks also crop out on the southern front of the Ketama Unit
(Andrieux, 1971; Favre, 1992; Benzaggagh et al., 2020) and within
several thrusted units such as Bou Haddoud and Senhadja nappes
(Leblanc, 1983; Papillon, 1989).
- Toarcian–Bathonian. Deposits of this time interval usually crop
out above the Lower Jurassic carbonate massifs and are
represented by condensed pelagic series (100–200 m-thick)
consisting of marlstone, marly limestone and bedded limestone
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series of Toarcian–Aalenian age; red marly limestone,
“ammonitico-rosso” type facies, of Bajocian age and marlstone
with thin platy microfolded laminated limestone often rich in
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microfilaments (Posidonomya) of Bathonian age (Suter, 1965;
Bulundwe Kitongo, 1987; Papillon, 1989; Favre, 1992), mainly the
Mjara outcrop located near Al Wahda Barrage, about 15 km
northwest of Jebel Amergou.
- Callovian–Oxfordian. It corresponds to a thick (1500 m-thick)
monotonous detrital series called the “Ferrysch” formation (Wildi,
1981) consisting of fine sandstone and silty clay alternations
forming very large outcrops in the Mesorif and the Internal Prerif
zones, the southern front of the Ketama Unit and the Senhadja and
Bou Haddoud nappes. Wildi (1981) regards it as a deep-sea fan
comparable to that of the Nile River.
- Kimmeridgian–lower Tithonian. This time interval is represented in
the whole External Rif domain by a second carbonate formation
which is strongly dismantled by the Alpine Miocene orogeny, and
forms alignments of decametric to barely kilometric blocks called
“sofs”. These “sofs” (Fig. 3) document three successive lithofacies
types (Benzaggagh and Atrops, 1997; Benzaggagh, 2000): “sofs”
with neritic facies (200–400 m) and “sofs” with pelagic facies
(10–70 m) in the Internal Prerif Zone and “sofs” with brecciated
limestone facies (10–150 m) in the Mesorif Zone.
5. Geological setting of the Moulay Bouchta area and location of
the study section
In the Moulay Bouchta area (70 km northwest of Fez city, Figs. 1
and 3) belonging to the western part of the Internal Prerif Zone, the
Mesozoic sequences are strongly scaled and overthrusted towards the
south. In this area, the Lower Cretaceous marlstones are frequently
overlapped by klippe nappes of Lower and Upper Jurassic materials,
mainly the Lower and Upper Jurassic limestones (Fig. 5A) and the
Callovian–Oxfordian detrital deposits of the “Ferrysch” formation (Fig.
3). The cross-sections, Fig. 4a-b illustrate the geological structure of this
area.
The study section, called JAM (Jebel Amergou), is located about
2 km north of the Marabout of Moulay Bouchta in a small ravine (Figs.
3 and 5B). The access to this section is easy by taking a track that leaves
towards the north from the Marabout of Moulay Bouchta.
6. Study section
The study section (Fig. 6) corresponds to the section first described
by Lacoste (1934). In this section, the Lower Jurassic massive lime-
stones of Jebel Amergou (called here the Moulay Bouchta Formation)
are surmounted with a sudden lithological change (Fig. 5A–C) by a
lumpy marlstone and marly limestone series (15-20 m-thick) called
here the “Lumpy marlstone and Marly limestone formation " or the
Amergou Formation. This formation is divided into two members:
lower and upper (Fig.. 5C and 6).
- the Lower member consists of four levels: the level 1 (20-50 cm-
thick) is composed of laminated gray marly limestone barren of
macrofauna; the level 2 (1–1.5 m-thick, Fig. 5C) consists of
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Fig. 3. Simplified geological map of Moulay Bouchta area, Middle Ouerrha, after 1/50,000 geological map of “Tafrannt de l’Ouerrha-Moulay Bouchta” (Suter, 1964),
completed by personal field data and location of the study section.
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Fig. 4. NE-SW cross-sections illustrating the geological structure of Moulay Bouchta area, south of Ouerrha Valley (Internal Prerif). A. Cross-section through Jebel
Amergou; B. Cross-section through Jebel Mekaoukine.

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Fig. 5. Field views. A. Panoramic view of Jebel Amergou composed of Lower Jurassic massif limestones (Moulay Bouchta Formation), thrusted over the Lower Creta-
ceous marlstones of the Internal Prerif Zone, in Moulay Bouchta are; B. Study section located northwest of Jebel Amergou, showing the Amergou Formation of Toar-
cian–Aalenian age overlapped by sandstone and silty clays of Callovian–Oxfordian “Ferrysch” formation; C. Detail of the lower member of the Amergou Formation
composed of four stratigraphic levels. D. Detail of the level 2, the richest in ammonites of middle Toarcian (Bifrons Zone), showing two specimens of Hildoceras. Dot-
ted white line, abnormal contact; continuous line, slip fault.
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alternation of gray thin lumpy limestone beds and marlstone. This
level did not provide ammonites; the level 3 (1–1.5 m-thick, Fig.
5C, E), consisting of alternation of yellow to reddish lumpy
marlstone and thin lumpy limestone beds, is the richest in
ammonites of the Bifrons Zone (Sublevisoni and Bifrons subzones);
the level 4 (1.5–2 m-thick, Fig. 5C), consisting of yellow to reddish
lumpy marlstone, yielded in its lower part several ammonites of
the middle Toarcian (Gradata Zone) and some specimens of the
upper Toarcian (Bonarellii and Speciosum? zones), in its upper
part.
- Upper member consists of two levels: the level 1 (2-3 m-thick, Fig.
5D) corresponds to a series of gray limestone beds with the
intercalation of marlstone; the level 2 (5-10 m-thick) consists of
alternation of gray thin limestone beds and marlstone yielding in
its lower part rare ammonites of the lower Aalenian (Opalinum
Zone). This level is overlain in abnormal contact by the reddish
silty clay and sandstone deposits of the Callovian–Oxfordian
“Ferrysch” formation.
By comparison with data from Lacoste (1934), the “White silty marl-
stone with thin limestone beds” level of this author corresponds to the
level 3 of the lower member of the study section; the “Gray limestone
and white marlstone alternation” corresponds to the level 4 of the lower
member and the “Brown limestone and small reddish sandstone beds” cor-
responds to the upper member of the study section. The “Ammonitico-
rosso 2″ of sections 2 and 3 of Bulundwe Kitongo (1987) corresponds to
the lower member of the study section and the “Ammonitico-rosso 3″
corresponds to the upper member.
Note that Wildi (1981, p. 501) and Bulundwe Kitongo (1987, p.
261) have interpreted the lack of the Aalenian (pro-pate)–Bathonian

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Fig. 6. Stratigraphic column of the Amergou section (JAM) showing approximate distribution of ammonite species collected from the section. This distribution is
based on field data, and compilations from published works on the Toarcian ammonites.

strata in the northern side of Jebel Amergou by the emergence of the quoted in the text for morphological comparisons and/or stratigraphic
Moulay Bouchta area during the Middle Jurassic (Dogger). This inter- and paleogeographical distributions of ammonite species.
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pretation is in disagreement with the field data, since: 1, the contact be-
tween the Aalenian and the Callovian–Oxfordian strata is a clear abnor- 8. Systematic paleontology
mal contact (see Fig. 5B); 2, the Bajocian series of “ammonitico-rosso”
type facies, is well-represented on the eastern and the northern sides of Class Cephalopoda Cuvier, 1797
the Lower Jurassic carbonate massifs of Jebels Arechgou and Beni Order Ammonoidea Zittel, 1817.
Ouassal (Fig. 3); and 3, in the Mjara area, the Middle Jurassic succes- Suborder Phylloceratina Arkell, 1950.
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sion is more complete showing above the Bajocian “ammonitico-rosso” Superfamily Phylloceratoidea Zittel, 1884.
facies (20 m-thick), a Bathonian series consisting of marlstones and thin Family Phylloceratidae Zittel, 1884.
platy microfolded limestones rich in Posidonomya (Suter, 1964). Subfamily Phylloceratinae Zittel, 1884.
Genus Phylloceras Suess, 1865.
7. Methodology Type species: Ammonites heterophyllum J. Sowerby, 1812
Phylloceras heterophyllum (J.).
250 specimens of ammonites have been collected from the study Fig. 8A–C.
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section during several field missions and are numbered JAM1 to
JAM250 (Jebel Amergou) and hosted in the Department of Geology,
Faculty of Sciences of Meknes. Most of these specimens are usually
phragmocones which rarely preserve a part of the body chamber and
often have a flank well-preserved while the other one is often badly pre-
served. The collected specimens allow to identify 44 ammonite species
characterizing biozones of middle-upper Toarcian and lower Aalenian
(Opalinum Zone). Most of these specimens are not collected from lay-
ers. Consequently, the stratigraphic distribution of ammonite species
given in Fig. 6, mainly for the upper Toarcian, is based on compilation
of data from published works on the Toarcian-lower Aalenian am-
monites. Fig. 7 illustrates the Toarcian–base Aalenian ammonite bio-
zones from the Tethyan domain and their equivalents from the NW Eu-
ropean province. The list of references is restricted to the references
1820 Ammonites heterophyllus J. Sowerby, p. 119; pl. 266.
1927b Phylloceras heterophyllum (J. Sowerby) - Daguin, p. 251, pl.
11, Fig. 1.
1963 Phylloceras heterophyllum (J. Sowerby) - Zanzucchi, p. 107; pl.
13, Fig. 2, with synonymy.
1998 Phylloceras heterophyllum (J. Sowerby) - Rulleau, p. 21: pl. 1,
Figs. 1–3; pl. 2, Fig. 3, with synonymy.
2001 Phylloceras heterophyllum (J. Sowerby) - Rulleau et al. pl. 1,
Fig. 1.
2002 Phylloceras heterophyllum (J. Sowerby) - Fauré, pl. 1, Fig. 2.
2011 Phylloceras heterophyllum (J. Sowerby) - Kovács, p. 8; pl. 1,
Figs. 1–4, with synonymy.
Material: Seven specimens.

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M. Benzaggagh
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Fig. 7. Toarcian and Aalenian ammonite biozones from the Tethyan domain and their equivalent from the NW European province.
Description: Smooth, small to large-sized shell with narrow umbili-
cus and oval whorl-section higher than wide with maximum width near
the umbilicus edge; slightly convex flanks and rounded venter.
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Distribution: According to Rulleau (1998), P. heterophyllum occurs
from the Serpentinum to the Bifrons zones in Lyon region, SE France.
It is reported from the Bifrons Zone of SE England (J. Sowerby, 1812),
eastern Pyrenees, south France (Fauré, 2002, 2013); the Serpentinum
and Bifrons zones of Hungary (Kovács, 2011); the Upper Liassic of
Italy (Principi, 1915; Zanzucchi, 1963) and eastern High Atlas, Beni
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Tadjit area, Morocco (Daguin, 1927b); the middle Toarcian of Cauca-
sus, northern Russia (in Kovács, 2011) and the study area (Lacoste,
1934, and this work).
Phylloceras doderleinianum (Catullo, 1853).
Fig. 8D–G.
1853 Ammonites doderleinianus Catullo, p. 19; pl. 1, Fig. 3.
1963 Phylloceras doderleinianum (Catullo) - Zanzucchi, p. 108; pl.
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13, Figs. 1, 4 and 4a, with synonymy.
2010 Phylloceras doderleinianus (Catullo) - Galácz et al. pl. 1, Fig. 4.
2011 Phylloceras doderleinianum (Catullo) - Kovács, p. 8; pl. 1, Figs.
5–6, pl. 2, Figs. 5–6, with synonymy.
Material: Eight specimens.
Description: Smooth, small to large-sized shell with narrow and
relatively deep funnel-shaped umbilicus; oval whorl-section higher
than wide with maximum width near the mid-flank; slightly convex
flanks; moderately wide and rounded venter.
Remarks: P. doderleinianum differs from P. heterophyllum by its
slightly more convex flanks, deeper and funnel-shaped umbilicus, and
the maximum width near the mid-flank.
Distribution: According to Kovács (2011), P. doderleinianum occurs
in Hungary from the.
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Journal of African Earth Sciences xxx (xxxx) 105082
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Serpentinum to the Bifrons zones. It is reported from the Upper Lias-
sic of Italy (Principi, 1915; Zanzucchi, 1963); the middle Toarcian of
Greece, Albania, Montenegro and Austria (in Kovács, 2011) and the
study area (this work).
Subfamily Calliphylloceratinae Spath, 1927.
Genus Calliphylloceras Spath, 1927.
Type species: Phylloceras disputabile Zittel, 1869
Calliphylloceras nilssoni (Hébert, 1866).
Fig. 8H–J.
1866 Ammonites nilssoni Hébert, p. 527, Fig. 3.
1963 Calliphylloceras nilssoni (Hébert) - Zanzucchi, p. 110; pl. 13, Fi gs.
6–7, 12, with synonymy.
1969 Calliphylloceras nilssoni (Hébert) - Pinna, pl. 6, Fig. 11, with
synonymy.
1998 Calliphylloceras nilssoni (Hébert) - Alméras et al. p. 90; pl. 15, Fig.
1.
1998 Calliphylloceras gr. Nilssoni (Hébert) - Rulleau, p. 27; pl. 4, Figs.
1–2, with synonymy.
2002 Calliphylloceras nilssoni (Hébert) - Fauré, p. 694; pl. 1, Fig. 1.
2011 Calliphylloceras nilssoni (Hébert) - Kovács, p. 10; pl. 2, Figs. 3–4,
with synonymy.
2022a Calliphylloceras nilssoni (Hébert) - Benzaggagh et al. p. 9; Fig.
8B–F.
Material: 17 specimens.
Description: Smooth, mall to medium-sized shell with involute
umbilicus; compressed oval whorl-section higher than wide with
maximum width near the mid-flank; slightly convex flanks gradually
converging towards the venter bearing on the last whorl four to five
proverse constrictions arising from the umbilicus and crossing the
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Fig. 8. Toarcian ammonites from the Jebel Amergou section, Internal Prerif
Fig. 9. Toarcian ammonites from the Jebel Amergou section, Internal Prerif
Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-C. Phyllo-
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Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-B. Calliphyllo-
ceras heterophyllum (J. Sowerby, 1812), A. JAM3, B. JAM5, C. JAM6; D-G.
ceras capitanii (Catullo, 1847), A. JAM22, B. JAM23; C-G. Calliphylloceras
Phylloceras doderleinianum (Catullo, 1853), D. JAM7, E. JAM185, F. JAM9,
aff. Capitanii (Catullo, 1847), C. JAM28, D. JAM27, E. JAM25, F. JAM24, G.
G. JAM186; H-J. Calliphylloceras nilssoni (Hébert, 1866), H. JAM14, I.
JAM26; H–I. Lytoceras francisci (Oppel, 1856), H. JAM150, I. JAM30; J-K.
JAM159, J. JAM160.
Alocolytoceras coarctatum (Pompeckj, 1896), J. JAM31, K. JAM32; L. Alo-
colytoceras dorcadis (Meneghini, 1875), JAM29.
venter where are slight curved forward; venter rounded and moder-
RR

ately wide.
Description: Smooth, small to medium-sized shell with narrow um-
Distribution: According to Kovács (2011), C. nilssoni occurs in
bilics; compressed oval whorl-section; wide and weakly convex flanks
Hungary from the Serpentinum to the Bifrons zones. It is reported from
bearing six to seven shallow constrictions arising from the umbilicus
the Bifrons Zone of eastern Pyrenees (Fauré, 2002, 2013) and Lyon re-
and crossing the venter where are slightly curved forward; narrow and
gion (Alméras et al., 1998; Rulleau, 1998; Rulleau et al., 2001); the
rounded venter.
Upper Liassic of Italy (Pinna, 1969; Zanzucchi, 1963); the Levisoni
Remarks: C. capitanii differs from C. nilssoni by its more compressed
CO

Zone of the central High Atlas (Benzaggagh et al., 2022a); the Bifrons
shell and more numerous and less spaced constrictions.
and Gradata zones of the Folded Middle Atlas, Morocco (El Hammichi
Distribution: According to Kovács (2011), Calliphylloceras capi-
et al., 2008, 20,016) and the middle Toarcian of Turkey, Caucasus,
tanii occurs in Hungary from the Serpentinum to the Bifrons zones. It
Iran, Japan (in Kovács, 2011) and the study area (Lacoste, 1934, and
is reported from the Upper Liassic of Italy (Pinna, 1969; Zanzucchi,
this work).
1963); the Levisoni Zone of NW Algeria (Douas Bengoudira, 2012);
Calliphylloceras capitanii (Catullo, 1847).
the Bifrons Zone of Greece, Austria and Albania (in Kovács, 2011) and
Fig. 9A–B.
the middle Toarcian (Bifrons zone) of the study area (this work).
Calliphylloceras aff. Capitanii (Catullo, 1847).
1847 Ammonites capitanii Catullo, p. 5; pl. 12, Fig. 4.
Fig. 9C–G.
1963 Calliphylloceras capitanioi (Catullo) - Zanzucchi, p. 109; pl. 13,
Material: Five specimens.
Fig. 13, with synonymy.
Description: Smooth, small to medium-sized shell with narrow
1969 Calliphylloceras capitanioi (Catullo) - Pinna, pl. 6, Fig. 7.
umbilics; strongly compressed oval whorl-section, much higher than
2011 Calliphylloceras capitanii (Catullo) - Kovács, p. 10; pl. 2, Figs. 1–2;
wide; wide and weakly convex flanks bearing three to four substraight
pl. 3, Figs. 5–6, with synonymy.
and relatively deep constrictions arising from the umbilicus and cross-

Material: Three specimens.

8
M. Benzaggagh
ing the venter where are slightly curved forward; narrow and rounded
venter.
Remarks: C. aff. Capitanii differs from C. capitanii by its deeper,
substraight and less numerous constrictions. It differs from P. nilssoni
by the less convex flanks, the less numerous and less curved con-
strictions.
Distribution: Middle–upper Toarcian of the study area (this work).
Suborder Lytoceratina Hyatt, 1889.
Superfamily Lytoceratoidea Neumayr, 1875.
Family Lytoceratidae Neumayr, 1875.
Subfamily Lytoceratinae Neumayr, 1875.
Genus Lytoceras Suess, 1865.
Type species: Lytoceras confusum Buckman, 1909
Lytoceras francisci (Oppel, 1856).
Fig. 9H–I.
1856 Ammonites francisci Oppel, p. 551.
1963 Lytoceras francisci (Oppel) - Zanzucchi, p. 113; pl. 14, Fig. 3, with
synonymy.
1968 Lytoceras francisci (Oppel) - Pinna, pl. 10, Fi gs. 9, 16, 18–19,
PR
with synonymy.
2011 Lytoceras francisci (Oppel) - Kovács, p. 14; pl. 3, Fi gs. 1–2, 4,
with synonymy.
Material: Two specimens.
Description: Smooth, medium-sized shell with fast growth whorls;
oval whorl-section higher than wide; wide and shallow umbilicus with
short and rounded umbilical wall; convex flanks and wide rounded
D
venter. Specimen Fig. 9I is compressed.
Remarks: L. francisci is characterized by its fast growth whorls and
shallow umbilicus.
Distribution: Lower–upper Toarcian of Hungary (Kovács, 2011),
TE
Austria, Greece, Albania, Caucasus (in Kovács, 2011); Toarcian
(Pinna, 1968) and Upper Liassic (Zanzucchi, 1963) of Italy and mid-
dle Toarcian (Bifrons Zone?) of the study area (this work).
Subfamily Alocolytoceratinae Spath, 1927.
Genus Alocolytoceras Hyatt, 1900.
EC
Type species: Ammonites germaini d'Orbigny, 1845 (p., 320; pl. 101,
Fi gs. 1–2, 4–5), designated by Hyatt (1900, p. 572).
Alocolytoceras coarctatum (Pompeckj, 1896).
Fig. 9J–K.
1885 Ammonites interruptus striatus Quenstedt, p. 387; pl. 48, Figs. 6
and 8.
RR
1927b Lytoceras coarctatum (Pompeckj) - Daguin, p. 252, pl 11, Fig. 3.
1998 Alocolytoceras coarctatum (Pompeckj) - Rulleau, p. 63; pl. 23,
Figs. 5–6; pl. 24, Fig. 4.
2002 Alocolytoceras gr. Germaini (d’Orbigny) - Fauré, p. 696; pl. 2, Fig.
3a-b.
2008 Alocolytoceras coarctatum (Pompeckj) - Bourillot et al. p. 601;
CO
text-Fig. 4, with synonymy.
Material: Three specimens.
Description: Smooth, small-sized shell with slow growth whorls;
subcircular whorl-section, as higher as wide; wide and shallow um-
bilicus; convex flanks bearing five constrictions on the last whorl;
moderately wide and rounded venter.
Remarks: A. coarctatum is characterized by its small size, subcircu-
lar whorl-section and well-marked transverse constrictions. Studied
specimens are close to those figured by Bourillot et al. (2008, text-Fig.
4).
Distribution: According to Bourillot et al. (2008), A. coarctatum is
reported from the upper Toarcian (Dispansum/Speciosum Zone) of
eastern Pyrenees (Fauré, 2002), Causses, southern France (Monestier,
1931; Elmi and Rulleau, 1993) and Lyon region (Rulleau, 1998); the
9
Journal of African Earth Sciences xxx (xxxx) 105082
upper Toarcian of southern Germany (Schlegelmilch, 1976) and Italy;
the Aalensis Zone of the Lusitanian Basin, Portugal, and the Folded
Middle Atlas (Benshili, 1989); the Upper Liassic of eastern High Atlas
(Daguin, 1927b); the upper Toarcian (Speciosum–Meneghinii zones) of
the central High Atlas (Bourillot et al., 2008) and the upper Toarcian?
Of the study area (this work).
Alocolytoceras dorcadis (Meneghini, 1875).
Fig. 9L.
1875 Ammonites (Lytoceras) Dorcadis Meneghini, p. 107; pl. 21, Fig. 1.
F
1969 Alocolytoceras dorcadis (Meneghini) - Pinna, p. 19; pl. 6, Fig. 5.
1972 Alocolytoceras dorcadis (Meneghini) - Guex, pl. 7, Fig. 7.
2008 Alocolytoceras dorcadis (Meneghini) - Bourillot et al. p. 600; pl. 1,
OO
Figs. 1–3; text-Fig. 3, with synonymy.
Material: Two specimens.
Description: Smooth, medium sized shell with serpenticone coil-
ing; wide and relatively deep umbilicus with rounded umbilical edge
and subvertical umbilical wall; subquadratic whorl-section, slightly
wider than high with maximum thickness on the umbilicus edge; sub-
flat flanks showing some badly marked shallow constrictions in the
inner whorls; broad and rounded venter.
Remarks: A. dorcadis is characterized by its subquadratic whorl-
section, subflat flanks, deep umbilicus with subvertical umbilical wall
and the presence of some shallow constrictions in the inner whorls.
Distribution: According to Bourillot et al. (2008), A. dorcadis
ranges from the middle to basal upper Toarcian (Bifrons–Bonarellii
zones). It is reported from Causses (Monestier, 1931), Bakony, Hun-
gary (Géczy, 1967), Italy (Dezi and Ridolfi, 1975; Venturi and Ferri,
2001), central High Atlas (Bourillot et al., 2008), Folded Middle Atlas
(Benshili, 1989), NW Algeria (Elmi et al., 1974) and the study area
(this work).
Genus Perilytoceras Rulleau, 1998
Type species: Ammonites jurensis Zieten, 1833 (p. 90; pl. 68, Fig. 1).
Perilytoceras perlaevis (Denckmann, 1887).
Fig. 10A–C.
1887 Ammonites perlaevis Denckmann, p. 45; pl. 2, Fig. 5.
1998 Perilytoceras perlaevis (Denckmann) - Rulleau, p. 54; pl. 17, Fig.
3; pl. 19, Figs. 1–2, with synonymy.
2007 Perilytoceras gr. Perlaevis (Denckmann) - Fauré, p. 40; Fi g. 6.1–3.
Material: Three specimens.
Description: Smooth, small to medium-sized shell with compressed
subtriangular whorl-section, higher than wide with maximum thick-
ness near the umbilical edge; deep and medium-sized umbilicus with
whorls covering about two-thirds of the flanks; vertical and relatively
high umbilical wall; subflat flanks; narrow and rounded venter.
Remarks: P. perlaevis is distinguished by its subflat flanks, com-
pressed subtriangular whorl-section, deep and medium-sized umbili-
cus with vertical umbilical wall. Studied specimens are close to the
specimens figured by Fauré (2007, Fi g. 6.1–3) and Rulleau (1998,
pl. 17, Fig. 3; pl. 19, Fig. 1).
Distribution: Thouarsense Zone of Lyon region (Rulleau, 1998)
and eastern Pyrenees (Fauré, 2007); upper Toarcian? Of the study
area (this work). It should be noted that this species has never been
reported from the Mediterranean domain before.
Superfamily Eoderoceratoidea Spath, 1929
Family Dactylioceratidae Hyatt, 1867.
Subfamily Dactylioceratinae Hyatt, 1867.
Genus Dactylioceras Hyatt, 1867.
Type species: Ammonites communis J. Sowerby, 1812
Subgenus Dactylioceras Hyatt, 1867.
Dactylioceras (Dactylioceras) aff. Attenuatus (.).
Fig. 10D.
M. Benzaggagh
PR
D
TE
Fig. 10. Toarcian ammonites from the Jebel Amergou section, Internal Prerif
Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-C. Perilyto-
ceras perlaevis (Denckmann, 1887), A. JAM33, B. JAM34, C. JAM35 (micro-
conch); D. Dactylioceras (Dactylioceras) aff. Attenuatus Buckman, 1909,
JAM36; E. Telodactylites eucosmus (Lippi-Boncambi, 1947), JAM37; F. Zu-
EC
godactylites cf. Braunianus (d'Orbigny, 1845), JAM38; G. Zugodactylites
sp. Indet, JAM39; H. Nodicoeloceras tuberculatum (Kottek, 1966), JAM42;
I-J. Nodicoeloceras sp. Indet, I. JAM47, J. JAM48; K-M. Collina gemma
Bonarelli, 1893, K. JAM43., L. JAM44, M. JAM45.
aff. 1926 Microdactylites attenuatus Buckman, pl. 655.
RR
aff. 2017 Dactylioceras (Dactylioceras) attenuatus Buckman - Fauré et al.
p. 55; pl. 2, Fig. 12.
Material: One specimen.
Description: Small-sized shell with compressed oval whorl-section
higher than wide; moderately evolute umbilicus; convex flanks bearing
slightly flexuous ribs, simple or bifurcated more or less high on the
CO
flanks; secondary ribs, almost of the same thickness as the primary
ones, cross the venter without interruption and slightly curved forward.
Remarks: The specimen differs from that figured by Fauré et al.
(2017, pl. 2, Fig. 12) by its slightly thicker ribs.
Distribution: Bifrons Zone of SE England (Buckman, 1909),
Quercy, SW France (Fauré et al., 2017) and middle Toarcian (Bifrons
Zone?) of the study area (this work).
Telodactylites Pinna and Levi-Setti, 1971,
Type species: Peronoceras eucosmus Lippiboncambi, 1947, desig-
nated by Venturi and Ferri (2001).
Telodactylites eucosmus (Lippi-Boncambi, 1947).
Fig. 10E.
1947 Peronoceras eucosmus Merla in Schedis - Lippi-Boncambi, p. 145;
pl. 2, Fi g. 21.
10
Journal of African Earth Sciences xxx (xxxx) 105082
1987 “Porpoceras” eucosmus (Dezi and Ridolfi) - Elmi and Benshili, pl.
2, Figs. 1–2.
2014a Telodactylites eucosmus (Lippi-Boncambi) - Kovács, p. 58; pl. 7,
Figs. 3–4, with synonymy.
Material: Two specimens.
Description: Medium-sized shell with subtrapezoidal whorl-
section, as high as wide; evolute umbilicus; flat to slightly convex
flanks bearing simple, sharp and relatively thick ribs less wide than
the intercostal spaces; one rib among two ends at a radial elongated
F
ventrolateral tubercle; from each tubercle arise two secondary ribs
crossing the venter radially and without interruption.
Remarks: Although not complete, the specimen is close to that fig-
OO
ured by Elmi and Benshili (1987, pl. 2, Figs. 1–2) under the name “Por-
poceras” eucosmus (Dezi and Ridolfi) (= Telodactylites eucosmus (Dezi
and Ridolfi, 1975) from the Gemma Subzone (middle Toarcian) of the
Folded Middle Atlas.
Distribution: Bifrons and Gradata zones of Hungary (Galácz et al.,
2010; Kovács, 2014a), Italy and France (in Galácz et al., 2010); Gra-
data Zone of the Folded Middle Atlas (Elmi and Benshili, 1987; El
Hammichi et al., 2008); middle–upper Toarcian of the study area (this
work).
Genus Zugodactylites Buckman, 1909
Type species: Ammonites braunianus d’Orbigny, 1845.
Zugodactylites cf. Braunianus (d'Orbigny, 1845).
Fig. 10F.
cf. 1845 Ammonites braunianus d’Orbigny, p. 327; pl. 104, Figs. 1–3.
cf. 1973b Zugodactylites braunianus (d'Orbigny) - Guex, pl. 3, Figs.
10–11.
cf. 2002 Zugodactylites braunianus (d'Orbigny) - Fauré, p. 709; pl. 9,
Fig. 2, with synonymy.
cf. 2014a Zugodactylites braunianus (d'Orbigny) - Kovács, p. 49; pl. 1, Fi
gs. 4–5, 9.
Material: One specimen.
Description: Compressed medium-sized shell with subrectangular
whorl-section higher than wide with maximum thickness on the um-
bilical edge; wide and moderately deep umbilicus with rounded um-
bilical edge; short, vertical umbilical wall; flat flanks slightly sloping
towards the ventrolateral edge, bearing simple, fine and radial ribs
almost of the same width as the intercostal spaces and are bifurcated
high on the flank and curved forward on the venter which is
rounded and moderately wide.
Remarks: By its flat flanks, dense ribbing and ribs bifurcated high
on the flanks, the specimen studied is close to that figured by Fauré
(2002, pl. 9, Fig. 2).
Distribution: Bifrons Zone of Lyon region (Rulleau et al., 2001),
eastern Pyrenees (Fauré, 2002, 2013), Provence, SE France (Guex,
1973b), Hungary (Kovács, 2014a) and middle Toarcian (Bifrons Zone?)
of the study area (this work).
Zugodactylites sp. Indet.
Fig. 10G.
Material: One specimen.
Remarks: It differs from Z. cf. Braunianus by its smaller size, nar-
rower flank, thick and more spaced ribs.
Distribution: Middle Toarcian (Bifrons Zone?) of the study area.
Subfamily Nodicoeloceratinae Venturi and Ferri, 2001.
Genus Nodicoeloceras Buckman, 1909
Type species: Ammonites crassoides Simpson, 1855.
Nodicoeloceras tuberculatum (Kottek, 1966).
Fig. 10H.
1966 Catacoeloceras tuberculatum Kottek, p. 135; pl. 17, Figs. 1–3, text-
fig. 65.
M. Benzaggagh Journal of African Earth Sciences xxx (xxxx) 105082

1966 Peronoceras baconicum Géczy et al., 2008, p. 438; pl. 1, Fig. 2.
2012 Nodicoeloceras tuberculatum (Kottek) - Kovács, p. 18; pl. 6, Figs.
3–4, with synonymy.
2014a Nodicoeloceras tuberculatum (Kottek) - Kovács, p. 4; Fi gs. 2–4,
6–7; pl. 5, Figs. 6–7.
Material: Three specimens.
Description: Thick medium-sized shell with cadicone whorl-
section, wider than high, with maximum thickness at the mid-flank;
wide and deep funnel-shaped umbilicus; low overlapping whorls, and
Harpoceras falciferum (J.).
Fig. 11A–D.
1820 Ammonites falcifer J. Sowerby, p. 99; pl. 254. Fig. 2.
1976 Harpoceras (Harpoceras) falciferum (J. Sowerby) - Gabilly, p. 94;
pl. 5, Figs. 3–4; pl. 9, Figs. 3–5; pl. 10,
Figs. 1–2; pl. 11, Figs. 1–3; pl. 12, Figs. 1–2; pl. 14, Figs. 1–2; pl. 15; pl.
16, Figs. 1–2, with synonymy.
1992 Harpoceras falciferum (J. Sowerby) - Howarth, p. 119; pl. 18, Fi g.
33; pl. 19, Figs. 2–4; pl. 20, Figs. 1–11; text-figs 18F. 19B, 27–34.

F
very convex flanks bearing numerous simple and regularly spaced ribs, 1997 Harpoceras falciferum (J. Sowerby) - Elmi et al. pl. 9, Figs. 9–10.
bifurcated high on the flank at a small ventrolateral tubercle and cross 2001 Harpoceras falciferum (J. Sowerby) - Rulleau et al. pl. 6, Fig. 1.
radially without interruption the venter, which is broad and rounded. 2002 Harpoceras falciferum (J. Sowerby) - Fauré, p. 717; pl. 13, Figs.

Remarks: By their small size, cadicone whorl-section, wide and
deep funnel-shaped umbilicus, the study specimens are close to those
figured by Kovács (2014a, pl. 4, Fig. 6–) they slightly differ from them
by their denser and less spaced ribs.
Distribution: According to Kovács (2014a), N. tuberculatum is re-
ported from the Bifrons Zone of SE England and Hungary. It occurs in
the middle Toarcian (Bifrons Zone?) of the study area.
OO
1–3.
2002 Harpoceras falciferum (J. Sowerby) - Neige and Rouget, p. 769,
Fig. 3C.
2004 Harpoceras gr. Falciferum (J. Sowerby) - Morard, p. 325; pl. 16. Fi
gs 1, 3–4, 6; pl. 17, Figs. 1–6, text-fig. B.7.16.
2006 Harpoceras falciferum (J. Sowerby) - Bécaud, p. 61; pl. 19, Fig.
1a-c; pl. 20, Fig. 1a-c; pl. 24, Fig. 2a–b; pl. 26, Figs. 1,2a-c, 3a-c, 4a-

PR
Nodicoeloceras sp. Indet. c; pl. 27, Fig. 1a–b, 2; pl. 32, Fig. 2a–b, 4a-b, with synonymy.
Fig. 10I–J. 2011 Harpoceras falciferum (J. Sowerby) - Kovács, p. 28; pl. 7, Fig. 2;
Material: Three specimens. pl. 11, Fig. 1, with synonymy.
Description: Wide and shallow umbilicus, low overlapping 2023 Harpoceras falciferum (J. Sowerby) - Kovács et al. p. 225; Fi g. 5.4
whorls with cadicone whorl-section wider than high; short and con- and 7, with synonymy.
vex flanks bearing fine, simple ribs with one rib among four ends at
a ventrolateral tubercle; from each tubercle arise two secondary ribs
slightly less thick than the primaries and cross radially the broad and
D
rounded venter.
Distribution: Middle Toarcian of the study area (this work).
Genus Collina Bonarelli, 1893.
Type species: Collina gemma Bonarelli, 1893.
TE

Collina gemma Bonarelli, 1893.

Fig. 10K-M.

1893 Collina gemma Bonarelli, p. 205, text-fig.

1987 Collina gemma (Bonarelli) - Elmi and Benshili, pl. 2, Figs. 5–8.
EC

1998 Collina gemma Bonarelli - Parisi et al. pl. 3, Fig. 8.

2008 Collina gemma Bonarelli - El Hammichi et al. pl. 1, Figs. 2–3.
2010 Collina gemma Bonarelli - Galácz et al. pl. 6, Fig. 4.
2014a Collina gemma Bonarelli - Kovács, p. 59; pl. 7, Fi g. 10.11, with
synonymy.

Material: Five specimens.

RR

Description: Small to medium-sized shell with serpenticone coil-

ing and subcircular to subrectangular whorl-section slightly higher
than wide; wide and shallow umbilicus without distinct umbilical
wall; convex flanks bearing strong, simple, sharp and spaced radial
ribs almost one rib among two ends at a ventrolateral tubercle; sec-
ondary ribs, with the same thickness as the primaries cross the venter
CO

without interruption and are slightly curved forward; from each tuber-
cle arise two secondary ribs ending at an opposite ventrolateral tuber-
cle; narrow and rounded venter.
Distribution: Gradata Zone (El Hammichi et al., 2008) and
Gemma Subzone of the Folded Middle Atlas (Elmi and Benshili, 1987)
and Italy (Parisi et al., 1998); Gradata Zone (Galácz et al., 2010) and
Semipolitum to Gemma horizons of Hungary (Kovács, 2014a); Middle
Toarcian (Gradata Zone?) of the study area (Bulundwe Kitongo, 1987,
and this work).
Superfamily Hildoceratoidea Hyatt, 1867.
Family Hildoceratidae Hyatt, 1867. Fig. 11. Toarcian ammonites from the Jebel Amergou section, Internal Prerif
Subfamily Harpoceratinae Neumayr, 1875. Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-D. Harpoceras
Genus Harpoceras Waagen, 1869. falciferum (J. Sowerby, 1812); A. JAM210, B. JAM49, C. JAM, 103, D. JAM60
Type species: Ammonites falcifer J. Sowerby, 1812 (p. 99; pl. 254, (microconch), E. JAM52; E. Harpoceras pseudoserpentinum Gabilly (1976),
JAM50; F–I. Harpoceras subplanatum (Oppel, 1856), G. JAM57, H. JAM53, I.
Fig. 2), ICZN designation, Opinion 303.
JAM54.

11
M. Benzaggagh
Material: Eight specimens.
Description: Compressed, small to medium-sized shell with oval
whorl-section, higher than wide with maximum thickness near the
umbilical edge; moderately evolute umbilicus with short subvertical
umbilical wall; slightly convex flanks gradually joining the short ven-
trolateral edge and bearing falcoid ribs, simple or grouped in pairs or
bundles on the lower third of the flank and inflected forward on the
outer part of the flank, rounded venter showing or no trace of a me-
dian keel bordered by two narrow and smooth spaces.
Remarks: The species is characterized by its moderately involute
umbilicus with short subvertical umbilical wall.
Distribution: H. falciferum, a species of wide geographic distribu-
tion, is reported from the uppermost lower Toarcian and the middle
Toarcian (Sublevisoni Subzone) of SE England (Howarth, 1992), NW
France (Bécaud, 2002, 2006; Gabilly, 1976; Morard, 2004; Neige and
Rouget, 2002), eastern Pyrenees (Fauré, 2002); uppermost lower Toar-
cian (Serpentinum Zone) of Hungary (Kovács, 2011; Kovács et al.,
2023), Bulgaria (Metodiev, 2008), Lyon region (Elmi and Rulleau,
1991; Elmi et al., 1997) and the middle Toarcian (Sublevisoni Zone?)
of the study area (this work).
PR
Harpoceras pseudoserpentinum Gabilly (1976).
Fig. 11E.
1973 Harpoceras pseudoserpentinum Gabilly, p. 79; pl. 7, Fi gs. 1, 4–5;
pl. 8. Fi g. 1; pl. 9, Fi gs. 1–2.
1976 Harpoceras pseudoserpentinum Gabilly, p. 90; pl. 7, Fi gs. 1, 4–5;
pl. 8, Fi g. 1; pl. 9, Fi gs. 1–2, with synonymy.
1997 Harpoceras pseudoserpentinum Gabilly - Elmi et al. pl. 9, Fi g. 8.
D
2006 Harpoceras (Harpoceras) pseudoserpentinum Gabilly - Bécaud, p.
60; pl. 16, Fig. 1a-c; pl. 17, Fig. 1a-c, with synonymy.
Material: One specimen.
TE
Description: Medium-sized shell with compressed oval whorl-
section higher than wide with maximum thickness near the umbilical
edge; moderately evolute umbilicus with short subvertical umbilical
wall; slightly convex flanks gradually joining the short ventrolateral
edge and bearing fine falcoid ribs, simple or grouped in bundles on the
EC
inner part of the flank and inflected forward on the outer part; the
lower third of the flank is smooth on the last whorl and showing some
small spiral depressions; narrow and rounded venter with a high keel
bordered by two smooth and narrow spaces.
Remarks: H. (H.) pseudoserpentinum is characterized by its rela-
tively wide umbilicus, smooth inner part of the flank, and the presence
of spiral depressions.
RR
Distribution: Serpentinum Zone of NW France (Bécaud, 2002,
2006; Gabilly, 1976) and Lyon region (Elmi et al., 1997); lowermost
middle Toarcian? Of the study area (this work).
Harpoceras subplanatum (Oppel, 1856).
Fig. 11F–I.
CO
1856 Ammonites subplanatus Oppel, p. 244.
1968 Harpoceras subplanatum (Oppel) - Pinna, p. 42; pl. 5, Fig. 3; pl. 6,
Fig. 4.
1984 Harpoceras subplanatum (Oppel) - Riegraf et al. p. 137; pl. 8, Fig.
9.
1992 Harpoceras subplanatum (Oppel) - Howarth, p. 136; pl. 22, Figs.
4–7; pl. 23, Figs. 1–3; text-figs 18H, 29, 35–36, with synonymy.
1998 Harpoceras subplanatum (Oppel) - Alméras et al. p. 48; pl. 6, Fig.
4; pl. 9, Fig. 1.
2006 Harpoceras (Harpoceras) subplanatum (Oppel) - Bécaud, p.65; pl.
18, Fig. 3; pl. 25, Fig. 1; pl. 31, Fig. 2.
2001 Harpoceras subplanatum (Oppel) - Rulleau et al. pl. 7, Figs. 1–2.
2013 Harpoceras cf. Subplanatum (Oppel) - Fauré, pl. 3, Figs. 2–3, Fig.
12.
12
Journal of African Earth Sciences xxx (xxxx) 105082
2022a Harpoceras subplanatum? (Oppel) - Benzaggagh et al. p. 13; Fig.
10F–H.
Material: Nine specimens.
Description: Compressed, medium to large-sized shell with invo-
lute umbilicus and short subvertical to slightly undercut umbilical
wall; broad and slightly convex flanks gradually joining the short
ventrolateral edge and bearing fine falculiform ribs strongly project-
ing forward on the outer part of the flank, showing or no some small
spiral depressions on the inner part of the flanks; venter narrow and
F
rounded showing trace of a low median keel bordered by two nar-
row and smooth spaces.
Remarks: H. subplanatum differs from the other species of Har-
OO
poceras by its more closed umbilicus and fine ribbing.
Distribution: Bifrons Zone of SE England (Howarth, 1992), south-
ern Germany (Riegraf et al., 1984), NW France (Bécaud, 2006; Fauré,
2013) and Lyon region (Alméras et al., 1998; Rulleau et al., 2001);
lower and middle Toarcian (Levisoni and Bifrons zones) of the central
High Atlas (Benzaggagh et al., 2022a); middle Toarcian (Bifrons
Zone?) of North America (Jakobs, 1992); lower Toarcian of Italy
(Pinna, 1968) and middle Toarcian (Bifrons Zone?) of the study area
(this work).
Genus Taffertia Guex, 1973b
Type species: Taffertia taffertensis Guex, 1973b (p. 503; pl. 2, Fig. 6;
pl. 14, Fig. 7).
Taffertia taffertensis Guex, 1973b
Fig. 12A–B.
1973a Taffertia taffertensis Guex, p. 503; pl. 2, Fig. 6; pl. 10, Fig. 7; pl.
14, Fig. 7; pl. 15, Fig. 13.
1992 Taffertia taffertensis Guex - Jakobs, p. 258; pl. 1, Fig. 2.
Material: Three specimens.
Description: Compressed, small to medium-sized shell with subo-
val whorl-section higher than wide, with maximum thickness on the
umbilical edge; moderately evolute umbilicus with short subvertical
umbilical wall; slightly convex flanks, joining gradually a short ven-
trolateral edge and bearing straight to slightly flexuous rounded ribs
strong and spaced on the lower half of the flank, becoming arched and
inflected forward on the upper part of the flank and mostly bifurcated
in the inner part of the flank; ribs are more or less blunt on the body
chamber; venter narrow and sharp with a well-developed keel bor-
dered by two narrow sulci.
Remarks: The three specimens differ from those figured by Guex
(1973a, pl. 2, Fig. 6 and pl. 10, Fig. 7) by their less thick ribs.
Distribution: Levisoni Zone of the Folded Middle Atlas (Guex,
1973a) and North America (Jakobs, 1992); lowermost middle Toar-
cian? Of the study area (this work).
Genus Polyplectus Buckman, 1909
Type species: Ammonites discoides Zieten, 1831 (p. 21; pl. 16, Fig. 1).
Polyplectus appenninicus Haas, 1913.
Fig. 12C.
1881 ammonites discoides Zieten - Meneghini, p. 21 (pars).
1968 Polyplectus appenninicus Haas - Pinna, p. 17; pl. 5, Fig. 4, with
synonymy.
2003–2004 Polyplectus appenninicus Haas - Pallini et al. pl. 3, Fig. 1.
Material: Two specimens.
Description: Medium-sized oxycone shell with involute umbilicus;
short vertical umbilical wall; slightly convex flanks joining the venter
without a distinct ventrolateral edge, and bearing numerous fine and
flexuous ribs; narrow and sharp venter, showing trace of a median keel
without smooth surfaces on either side.
M. Benzaggagh
PR
D
TE
Fig. 12. Toarcian ammonites from the Jebel Amergou section, Internal Prerif
Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-B. Taffertia
taffertensis Guex, 1973b, A. JAM61, B. JAM250; C. Polyplectus appenninicus
(Hass, 1913), JAM62; D-E. Polyplectus discoides (Zieten, 1831), D. JAM63, E.
JAM 155; F-G. Hildoceras apertum Gabilly, 197, F. JAM67, G. JAM71; H–K.
EC
Hildoceras bifrons (Bruguière, 1789), H. JAM67, I. JAM191, J. JAM68, K.
JA192.
Distribution: Lower Toarcian (Pinna, 1968) and Bifrons Zone
(Pallini et al., 2003-2004) of Italy; middle Toarcian (Bifrons Zone?) of
the study area (this work).
RR
Polyplectus discoides (Zieten, 1831).
Fig. 12D–E.
1831 Ammonites discoides Zieten, p. 21; pl. 16. Fig. 1.
1927b Polyplectus discoides (Zieten) - Daguin, p. 253, pl. 11, Fig. 7.
1968 Polyplectus pluricostatus (Haas) - Pinna, p. 46; pl. 6, Figs. 5, 8 and
10.
CO
1975 Polyplectus discoides (Zieten) - Guex, p. 115, pl. 7, Fig. 8.
1992 Polyplectus discoides (Zieten) - Howarth, p. 153; pl. 28, Figs. 6–8;
text-Fi g. 37B, with synonymy.
1996 Polyplectus discoides (Zieten) - Bernad and Martínez, p. 10; pl. 2,
Fig. 2.
2002 Polyplectus discoides (Zieten) - Fauré, p. 718; pl. 13, Fig. 11
2004 Polyplectus apenninicus Haas - Myczyński, p. 53, Fi gs. 24.3 and
32.1, with synonymy.
2006 Polyplectus discoides (Zieten) - Rulleau, p. 68; Figs. 7–8.
2006 Polyplectus discoides (Zieten) - Bécaud, p. 55; pl. 4, Fig. 2, with
synonymy.
Material: Two specimens.
Description: Small to medium-sized oxycone shell with narrow
umbilicus; slightly convex flanks joining the venter without a distinct
13
Journal of African Earth Sciences xxx (xxxx) 105082
ventrolateral edge and bearing fine and blunt falcoid ribs inflected
forward on the outer part of the flank; acute venter with a sharp keel.
Remarks: P. discoides differs from P. apenninicus by its sharper ven-
ter and more closed umbilicus.
Distribution: Dispanum Zone of SE England (Howarth, 1992) and
NW France (Bécaud, 2006); Crassicosta–Hillebrandti zones (mid-
dle–upper Toarcian?) of North America (Jakobs, 1992); Thouarsense
and Insigne zones of Causses (Guex, 1975); Thouarsense and Fallacio-
sum subzones of eastern Pyrenees (Fauré, 2002); Toarcian of Iberian
Range (Bernad and Martínez, 1996); Thouarsense Zone of Poland
F
(Myczyński, 2004); lower and middle Toarcian of Italy (Pinna, 1968);
upper Toarcian of eastern High Atlas (Daguin, 1927b); lowermost up-
per Toarcian (Bonarellii Zone?) of the study area (Lacoste, 1934, and
OO
this work).
Subfamily Hildoceratinae Hyatt, 1867.
Genus Hildoceras Hyatt, 1867.
Type species: Hildoceras bifrons (Bruguière, 1789) in Buckman,
1909) (pl. 114).
Hildoceras apertum Gabilly (1976).
Fig. 12F–G.
1973 Hildoceras apertum Gabilly, p. 157–160; pl. 26, Fi gs. 1–6, 9–11.
1974 Hildoceras apertum Gabilly - Elmi et al. pl. 2. Fi g. 10.
1976 Hildoceras apertum Gabilly, p. 153; pl. 26, Fi gs. 1–6, 9–11, with
synonymy.
1992 Hildoceras bifrons (Bruguière) - Howarth, pl. 37, Fig. 7.
2002 Hildoceras apertum Gabilly - Fauré, p. 720; pl. 15, Fig. 3.
1997 Hildoceras apertum Gabilly - Elmi et al. pl. 10. Fig. 3.
2002 Hildoceras apertum Gabilly - Neige and Rouget, p. 773; Fig. 5E, F,
H.
2006 Hildoceras apertum Gabilly - Rulleau, p. 71, pl. 21, Fig. 5.
2006 Hildoceras apertum Gabilly - Bécaud, p. 90; pl. 28, Fig. 3a-c; pl.
46, Fig. 1a–b, 3a-c, 4a-b, with synonymy.
Material: Three specimens.
Description: Medium-sized shell with shallow and moderately
evolute umbilicus; subrectangular whorl-section, with maximum
thickness on the umbilical edge; short, slightly sloping umbilical wall;
flat flanks slightly converging towards the ventrolateral edge showing
on the inner quarter a shallow spiral groove, separated from the um-
bilicus by a smooth band; the outer part of the flank bears relatively
strong arcuate ribs regularly spaced and slightly less wide than the in-
tercostal spaces; ribs start from the spiral groove or immediately
above; venter flat, relatively wide, bisulcate and tricarinate.
Remarks: H. apertum differs from H. bifrons by its less marked spiral
groove, located lower on the flank.
Distribution: Middle Toarcian, Sublevisoni Subzone of SE England
(Howarth, 1992), NW France (Bécaud, 2002, 2006; Gabilly, 1973,
1976; Neige and Rouget, 2002), eastern Pyrenees (Fauré, 2002), Lyon
region (Elmi et al., 1997; Rulleau et al., 2001), NW Algeria (Elmi et al.,
1974) and the study area (this work).
Hildoceras bifrons (Bruguière, 1789).
Fig. 12H–K.
1789 Ammonites bifrons Bruguière, p. 40.
1963 Hildoceras bifrons (Bruguière) - Zanzucchi, p. 120; pl. 14, Fig. 10;
pl. 15, Fi gs. 1–2, 4, 9–10; pl. 16, Fig. 3; pl. 17, Fi gs. 8–9, 11, with
synonymy.
1972 Hildoceras bifrons (Bruguière) - Guex, pl. 6, Fig. 7; pl. 7, Fig. 8.
1976 Hildoceras bifrons (Bruguière) - Gabilly, p. 157; pl. 26, Figs. 7–8;
pl. 27, Figs. 1–7; pl. 28, Figs. 1–2; pl. 29, Figs. 1–2.
1984 Hildoceras (Hildoceras) bifrons (Bruguière) - Riegraf et al. p. 119;
pl. 6, Fig. 6, with synonymy.
1992 Hildoceras bifrons (Bruguière) - Howarth, p. 184; pl. 36, Fig. 4; pl.
37, Figs. 1–10; pl. 38, Figs. 3 and 7.
M. Benzaggagh
1994 Hildoceras bifrons (Bruguière) - Ouahhabi, p. 246; pl. 8, Fig. 13;
pl. 9, Figs. 1–6.
1996 Hildoceras bifrons angustisiphonathum Prinz - Bernad and
Martínez, p. 13; pl. 3, Fig. 1.
2002 Hildoceras bifrons (Bruguière) - Fauré, p. 720; pl. 15, Figs. 5–6.
2006 Hildoceras bifrons (Bruguière) - Bécaud, p. 90; pl. 47, Figs. 1 and
3.
2008 Hildoceras bifrons (Bruguière) - Metodiev, Fi g. 4t-u.
2016 Hildoceras bifrons (Bruguière) - El Hammichi et al. pl. 4, Fig. 7.
2016 Hildoceras bifrons (Bruguière) - Jattiot et al., 2017, Fig. 4I.
2022 Hildoceras bifrons (Bruguière) - Benzaggagh, p. 15; Fig. 10D–F,
with synonymy.
2022a Hildoceras bifrons (Bruguière) - Benzaggagh et al. p. 16; Fig.
12F–J.
Material: Six specimens.
Description: Small to medium-sized shell with moderately evolute
umbilicus, and rounded umbilical edge; short, slightly sloping umbili-
cal wall; subrectangular to subtrapezoidal whorl-section, higher than
wide, with maximum thickness near the umbilical edge; flat flanks
PR
slightly sloping towards the ventrolateral edge, bearing smooth and
well-marked spiral groove, located at the lower third of the flank; the
inner part of the flank is smooth; the outer one, above the spiral
groove, bears simple, arcuate ribs, almost of the same width as the in-
tercostal spaces; venter moderately broad, flat to slightly convex, bisul-
cate and tricarinate, with a strong median keel.
Distribution: H. bifrons, a species of wide geographic distribu-
tion, has been reported from the Bifrons Subzone of several regions
D
in the world, among others: SE England (Howarth, 1992), southern
Germany (Riegraf et al., 1984), NW France (Bécaud, 2006; Gabilly,
1976; Jattiot et al., 2016; Neige and Rouget, 2002), Luxembourg
(Delsate and Weis, 2010), eastern Pyrenees (Fauré, 2002, 2013),
TE
Lyon region (Alméras et al., 1998; Elmi et al., 1997; Rulleau et al.,
2001), Causses (Guex, 1972), Iberian Range (Bernad and Martínez,
1996), Italy (Pelosio, 1968; Pinna, 1968; Principi, 1915; Zanzucchi,
1963), South Riffian Ridges (Benzaggagh, 2022), Beni Snassen, NE
Morocco (Alméras et al., 1988; Ouahhabi, 1986), Folded Middle At-
EC
las (El Hammichi et al., 2008, 2016; Elmi et al., 1989), central High
Atlas (Benzaggagh et al., 2022a; Wilmsen et al., 2002), NW Algeria
(Douas Bengoudira, 2012; Elmi et al., 1974) and the study area
(Lacoste, 1934, and this work).
Hildoceras crassum Mitzopoulos, 1930.
Fig. 13A–C.
RR
1867 Ammonites levisoni Simpsum Reynès, pl. 7, Figs. 2–3.
1976 Hildoceras crassum Mitzopoulos - Gabilly, p. 143; pl. 23, Figs.
1–5; pl. 24, Figs. 1–2; pl. 25, Figs. 1–2.
1994 Hildoceras crassum Mitzopoulos - Ouahhabi, p. 240; pl. 6, Figs.
4–8; pl. 7, Figs. 1–3.
1996 Hildoceras crassum Mitzopoulos - Bernad and Martínez, p. 13; pl.
CO
3, Fig. 2.
2001 Hildoceras crassum Mitzopoulos - Rulleau et al. pl. 10, Fig. 1.
2006 Hildoceras crassum Mitzopoulos - Bécaud, p. 87; pl. 42, Fig. 3; pl.
43, Fig. 2.
2016 Hildoceras crassum Mitzopoulos - El Hammichi et al. pl. 3, Fig.
10; pl. 4, Fig. 1.
Material: Seven specimens.
Description: Medium-sized shell with trapezoidal whorl-section,
and maximum thickness on the umbilical edge; shallow and moder-
ately evolute umbilicus; narrow and smooth periumbilical band, with-
out a distinct spiral groove; moderately thick ribs; bisulcate and tricari-
nate venter.
14
Journal of African Earth Sciences xxx (xxxx) 105082
Remarks: H. crassum differs from H. bifrons by the absence of the
spiral groove and from H. sublevisoni by a smooth, well-marked perium-
bilical band.
Distribution: Bifrons Subzone of NW France (Bécaud, 2006;
Gabilly, 1976), Lyon region (Rulleau et al., 2001), Iberian Range
(Bernad and Martínez, 1996), Folded Middle Atlas (El Hammichi et al.,
2016); Beni Snassen (Ouahhabi, 1986) and the study area (this work).
Hildoceras semipolitum Buckman, 1909
Fig. 13D–E.
F
1902 Ammonites semipolitum Buckman, p. 4.
1968 Hildoceras semipolitum Buckman - Pelosio, p. 153; pl. 18, Figs. 2,
6, 8 and 11; pl. 23, Fi gs. 5–6, 8, with synonymy.
OO
1972 Hildoceras semipolitum Buckman - Guex, pl. 6, Fi gs. 1, 4–5; pl. 7,
Fi g. 5.
1974 Hildoceras semipolitum Buckman - Elmi et al. pl. 3, Figs. 5 and 8.
1976 Hildoceras semipolitum Buckman - Gabilly, p. 165; pl. 24, Figs.
7–8; pl. 28, Figs. 3–4; pl. 29, Figs. 3–4.
1984 Hildoceras (Hildoceras) semipolitum Buckman - Riegraf et al. p.
122; pl. 6, Figs. 7-8a; pl. 7, Fig. 1, with synonymy.
1989 Hildoceras semipolitum Buckman - Benshili, pl. 19, Fig. 5.
1992 Hildoceras semipolitum Buckman - Howarth, p. 187; pl. 38, Fi gs.
1–2, 4–6, 8, with synonymy.
1994 Hildoceras semipolitum Buckman - Ouahhabi, p. 250; pl. 9, Figs.
7–16, with synonymy.
2002 Hildoceras semipolitum Buckman - Fauré, p. 720; pl. 15, Figs. 8–9.
2006 Hildoceras semipolitum Buckman - Bécaud, p. 93; pl. 18, Fig. 1; pl.
47, Figs. 2 and 4; pl. 48, Fig. 2, with synonymy.
2008 Hildoceras semipolitum Buckman - Metodiev, Fi g. 4r-s.
2010 Hildoceras semipolitum Buckman - Delsate and Weis, pl. 2, Fig. 3
2021 Hildoceras semipolitum Buckman - Arp et al. Fi g. 12.4–5.
2022 Hildoceras semipolitum Buckman - Benzaggagh, p. 16; Fig. 10D–F.
2022a Hildoceras semipolitum Buckman - Benzaggagh et al. p. 16; Fig.
13E–G.
Material: Two specimens.
Description: Two incomplete specimens showing a relatively fast
growth whorls, a relatively narrow umbilicus and a well-marked spiral
groove, located at the mid-flank.
Remarks: H. semipolitum differs from H. bifrons by the less evolute
umbilicus, the more compressed whorl-section and the spiral groove lo-
cated at the mid-flank.
Distribution: H. semipolitum, a species of wide geographic distrib-
ution, has been reported from the topmost Bifrons and lowermost
Gemma subzones of several regions in the world, among others, SE
England (Howarth, 1992), southern Germany (Arp et al., 2021;
Riegraf et al., 1984), NW France (Bécaud, 2006; Gabilly, 1976), east-
ern Pyrenees (Fauré, 2002, 2013), Luxembourg (Delsate and Weis,
2010), Lyon region (Elmi et al., 1997; Rulleau et al., 2001), Causses
(Guex, 1972), Italy (Pelosio, 1968; Pinna, 1969), Bulgaria (Metodiev,
2008), South Riffian Ridges (Benzaggagh, 2022; Faugères, 1978),
Beni Snassen (Alméras et al., 1988; Ouahhabi, 1986), Folded Middle
Atlas (Benshili, 1989; Elmi et al., 1989), central High Atlas
(Benzaggagh et al., 2022a), NW Algeria (Elmi et al., 1974) and the
study area (Bulundwe Kitongo, 1987, and this work).
Hildoceras sublevisoni (Fucini, 1919).
Fig. 13F–H.
1874 Ammonites levisoni (Simpson) - Dumortier, p. 49; pl. 9, Figs.
3–4.
1919 Hildoceras sublevisoni Fucini, p. 182.
1974 Hildoceras sublevisoni (Fucini) - Elmi et al. pl. 2, Figs. 4 and 5.
1968 Hildoceras sublevisoni Fucini - Pelosio, p. 147; pl. 18, Fi gs. 5, 7,
9–10, 12; pl. 23, Fig. 9, with synonymy.
1973a Hildoceras sublevisoni Fucini - Guex, pl. 6, Fig. 4; pl. 10, Fig. 3.
M. Benzaggagh Journal of African Earth Sciences xxx (xxxx) 105082

F
OO
PR
D
TE
EC
RR
CO

Fig. 13. Toarcian ammonites from the Jebel Amergou section, Internal Prerif Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-C. Hildoceras cras-
sum Mitzopoulos, 1930, A. JAM181, B. JAM181, C. JAM70; D-E. Hildoceras semipolitum Buckman, 1909, D. JAM72, E. JAM73; F–H. Hildoceras sublevisoni
(Fucini, 1919), F. JAM167, G. JAM76, H. 172; I. Hildoceras snoussi Elmi et al., 1997, JAM85; J-O. Hildoceras tethysi Géczy, 1967, J. JAM80, K. JAM77, L.
JAM75, M. JAM171, N. JAM197, O. JAM84.

1976 Hildoceras sublevisoni Fucini - Gabilly, p. 128; pl. 21, Fig. 5; pl.
22, Figs. 1–2.
1984 Hildoceras (Hildoceras) sublevisoni Fucini - Riegraf et al. p. 117;
pl. 6, Fig. 5, with synonymy.
1994 Hildoceras sublevisoni Fucini - Ouahhabi, p. 234; pl. 5, Figs. 1–8.
1996 Hildoceras sublevisoni Fucini - Bernad and Martínez, p. 12; pl. 4,
Fig. 2.
1998 Hildoceras sublevisoni Fucini - Parisi et al. pl. 3, Fig. 1.
2002 Hildoceras sublevisoni (Fucini) - Fauré, p. 719; pl. 14, Figs. 7–8.
2004 Hildoceras sublevisoni (Fucini) - Morard, 333; pl. 20, Figs. 6–7; pl.
21, Figs. 1–4.
2006 Hildoceras sublevisoni Fucini - Bécaud, p. 85; pl. 33, Fig. 2; pl. 42,
Fig. 2.
2009 Hildoceras sublevisoni Fucini - Myczyński and Jach, p. 35, Fig. 5G-
O
2016 Hildoceras sublevisoni Fucini - Jattiot et al., 2017, Fig. 4A
2022 Hildoceras sublevisoni Fucini - Benzaggagh, p. 16; Fig. 10K–J.
2022a Hildoceras sublevisoni Fucini - Benzaggagh et al. p. 16; Fig.
13H–J.

15
M. Benzaggagh Journal of African Earth Sciences xxx (xxxx) 105082

Material: 14 specimens. Description: Medium-sized shell with shallow and moderately

Description: Small to medium-sized shell with evolute umbilicus
and subquadratic to subrectangular whorl-section, higher than wide
with maximum thickness on the umbilical edge; short and sloping
umbilical wall, with rounded umbilical edge; flat, subparallel flanks,
converging slightly towards the ventrolateral edge; arched ribs, al-
most as wide as the intercostal spaces, forming short folds above the
umbilical edge and without spiral groove; venter flat, relatively wide,
bisulcate and tricarinate with a strong median keel.
Remarks: H. sublevisoni differs from the other species of the genus
evolute umbilicus; subrectangular whorl-section; short and slightly
sloping umbilical wall; flat flanks slightly converging towards the
ventrolateral edge and showing on the inner quarter a very shallow
spiral groove or a smooth band; the outer part of the flank bears ar-
cuate ribs, regularly spaced and slightly less wide than the inter-
costal spaces; ribs start from the spiral groove or immediately above
forming a well-marked short folds above the umbilical edge; flat
bisulcate and tricarinate venter.
Remarks: H. tethysi differs from H. crassum by its narrower smooth

F
Hildoceras by the absence of the spiral groove and from H. crassum by umbilical band and more pleated ribs above the umbilical edge.
more evolute umbilicus. Distribution: Bifrons Zone, Sublevisoni Subzone of SE England
Distribution: H. sublevisoni, a species of wide geographic distribu- (Howarth, 1992), NW France (Bécaud, 2006; Gabilly, 1976), eastern

tion, has been reported from the Sublevisoni Subzone of several re-
gions in the world, among others: southern Germany (Riegraf et al.,
1984), NW France (Bécaud, 2002, 2006; Gabilly, 1976; Jattiot et al.,
2016; Neige and Rouget, 2002), Lyon region (Alméras et al., 1998;
Elmi et al., 1997; Rulleau et al., 2001), eastern Pyrenees (Fauré,
2002, 2013), Causses (Guex, 1972; Morard, 2004), Iberian Range
(Bernad and Martínez, 1996), Italy (Parisi et al., 1998; Pelosio, 1968;
OO
Pyrenees (Fauré, 2002), Causses (Guex, 1972; Morard, 2004), Lyon
region (Elmi et al., 1997), Italy (Zanzucchi, 1963), Hungary (Géczy,
1967), Beni Snassen (Ouahhabi, 1986), and the study area (this
work).
Family Phymatoceratidae Hyatt, 1867.
Genus Furloceras Elmi and Rulleau, 1995.
Type species: Hildoceras (Lillia) chelussii Parisch and Viale, 1906,

PR
Zanzucchi, 1963), Poland (Myczyński and Jach, 2009), South Riffian designated by Elmi and Rulleau (1995).
Ridges (Benzaggagh, 2022), Beni Snassen (Alméras et al., 1988; Furloceras iserensis (Oppel, 1856).
Ouahhabi, 1986), Folded Middle Atlas (Benshili, 1989; Elmi et al., Fig. 14A–B.
1989; Guex, 1973a), central High Atlas (Benzaggagh et al., 2022a),
NW Algeria (Douas Bengoudira, 2012; Elmi et al., 1974) and the
study area (Bulundwe Kitongo, 1987, and this work).
Hildoceras snoussi Elmi et al., 1997
Fig. 13I.
D
1977 Hildoceras snoussi Elmi, p. 80, pl. 4, Fig. 3.
1997 Hildoceras snoussi Elmi - Elmi et al. pl. 28, Fig. 2.
2008 Hildoceras snoussi Elmi - El Hammichi et al. pl. 1, Fig. 6.
TE

Material: Two specimens.

Description: Fragment of very compressed trapezoidal section,
with maximum thickness on the umbilicus edge; smooth flanks con-
verging towards the ventrolateral edge; smooth spiral groove located at
EC

the lower third of the flank; narrow and acute venter with a high keel
bordered by two smooth and shallow sulci.
Remarks: Studied specimens are close to the specimen H. snoussi
Elmi, figured by Elmi et al. (1997), and that figured by El Hammichi et
al. (2008, pl. 1, Fig. 6).
Distribution: Topmost Bifrons and lowermost Gemma subzones of
the Folded Middle Atlas (El Hammichi et al., 2008); middle Toarcian
RR

(Bifrons Zone?) of the study area.

Hildoceras tethysi Géczy, 1967.
Fig. 13J-N.

1886b Harpoceras (Hildoceras) bifrons Bruguière - Gemmellaro, p. 190.

1963 Hildoceras bifrons (Bruguière) - Zanzucchi, p. 120; pl. 15, Fig. 9b;
CO

pl. 16, Fig. 3a.

1967 Hildoceras bifrons (Bruguière) tethysi Geczy, p. 126; pl. 4, Fig. 4.
1976 Hildoceras tethysi Geczy - Gabilly, p. 137; pl. 21, Figs. 1–4; pl. 23,
Figs. 6–7, with synonymy.
1992 Hildoceras lusitanicum Meister - Howarth, pl. 35, Fig. 4a-b.
1994 Hildoceras tethysi Geczy - Ouahhabi, p. 238, pl. 6, Figs. 1–3; pl.
10, Fig. 2; pl. 11, Fig. 4; pl. 13, Figs. 6 and 10, with synonymy.
1997 Hildoceras tethysi Geczy - Elmi et al. pl 9, Fig. 15.
2002 Hildoceras tethysi Geczy - Fauré, p. 719; pl. 14, Fig. 9.
2004 Hildoceras tethysi Geczy - Morard, p. 334; pl. 21, Fig. 5.
2006 Hildoceras tethysi Geczy - Bécaud, p. 87; pl. 44, Fig. 2a-b; pl. 45, Fig. 14. Toarcian ammonites from the Jebel Amergou section, Internal Prerif
Fig. 2a-b. Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-B. Furloceras is-
erensis (Oppel, 1856), A. JAM100, B. 215; C-D. Furloceras chelussii (Parisch
Material: Seven specimens. and Viale, 1906), C. JAM213, D. JAM84; E-F. Crassiceras latum Merla, 1932, E.
JAM91, F. JAM88; G-H. Crassiceras gradatum (Merla, 1932), G. JAM90, H.
JAM89.

16
M. Benzaggagh
1856 Ammonites iserensis Oppel, Jura formation, p. 249, lectotype in
Gallitelli-Wendt, 1969) (pl. 5, Fig. 6).
1963 Phymatoceras erbaense (Oppel) - Zanzucchi, p. 136; pl. 10, Fig. 9.
1969 Phymatoceras iserense (Hauer) - Pinna, pl. 3, Fig. 8.
1968 Phymatoceras (Phymatoceras) iserense (Oppel) - Pelosio, p. 160;
pl. 19, Fig. 8, with synonymy.
Material: Two specimens.
Description: Wide umbilicus; subquadratic whorl-section; strong
flexuous simple ribs regularly spaced; broad venter bearing a rounded
median keel bordered by two deep sulci.
Remarks: The specimen is close to that figured by Pelosio (1968, pl.
19, Fig. 8).
Distribution: Upper Toarcian (Pelosio, 1968) and middle-upper
Toarcian (Zanzucchi, 1963) of Italy; lowermost upper Toarcian? Of the
study area (this work).
Furloceras chelussii (Parisch and Viale, 1906).
Fig. 14C–D.
1906 Hildoceras (Lillia) chelussii Parisch and Viale, p. 156; pl. 11, Figs.
PR
10–11.
1968 Phymatoceras (Chartronia) chelussii Parisch and Viale - Pelosio, p.
174; pl. 21, Figs. 3 and 9; pl. 22, Fig. 12; pl. 23, Fig. 12, with
synonymy.
2014b Furloceras chelussii (Parisch and Viale) - Kovács, p. 19; pl. 2,
Figs. 1 and 5, with synonymy.
Material: Two specimens.
D
Description: Compressed, medium-sized shall, with subrectangu-
lar to subtrapezoid whorl-section; shallow, wide and evolute umbili-
cus; short vertical to slightly sloping umbilical wall and rounded
umbilical edge; subflat flanks slightly sloping towards the ventrolat-
TE
eral edge and bearing thick flexuous ribs, less wide than the inter-
costal spaces, simple or bifurcate at the umbilical edge from a small
elongated tubercle; venter moderately wide, subflat, bisulcate and
tricarinate.
Remarks: The two specimens are close to those figured by Kovács
EC
(2014b, pl. 2, Figs. 1, 5a-b).
Distribution: According to Kovács (2014b), the species is known
from the middle-upper Toarcian of Spain, Portugal, France, Greece,
Bulgaria, Albania and Caucasus. It occurs from the upper Toarcian of
Italy (Pelosio, 1968); the Gradata to Thouarsense zones of Hungry
(Kovács, 2014b) and the lowermost upper Toarcian? Of the study area
(this work).
RR
Subfamily Phymatoceratinae Hyatt, 1867.
Genus Mouterdeiceras Elmi and Rulleau, 1995.
Type species: Mouterdeiceras dubourgi Elmi and Rulleau (1995).
Mouterdeiceras dubourgi Elmi and Rulleau (1995).
Fig. 15A.
CO
1967 Phymatoceras tirolense (Hauer) - Géczy et al., 2008, p. 141; Fi g.
23, pl. 8.
1987 Phymatoceras evolutum Merla - Elmi and Benshili, Figs. 4–5.
1995 Mouterdeiceras dubourgi Elmi and Rulleau, p. 89, text-figs 1.1–3,
13–14, text-fig. 2.1–2; pl. 10, Figs. 1–8; pl. 11, Figs. 7–8.
2010 Mouterdeiceras dubourgi Elmi and Rulleau - Galácz et al. p. 328;
pl. 3, Fig. 3; pl. 4, Fig. 1.
2014b Mouterdeiceras dubourgi Elmi and Rulleau - Kovács, p. 22; pl. 4,
Figs. 2 and 4, with synonymy.
Material: One specimen.
Description: Large-sized specimen with subrectangular to sub-
trapezoid whorl-section, slightly higher than wide and with slow
growth whorls; wide shallow and evolute umbilicus with short,
rounded umbilical wall; subparallel to slightly convex flanks bearing
17
Journal of African Earth Sciences xxx (xxxx) 105082
strong, straight to slightly flexuous, ribs regularly spaced and slightly
inflected forward on the ventrolateral edge; presence of some bifur-
cate ribs at the umbilical margin in the inner whorls; flat to slightly
rounded venter showing a median keel bordered by two shallow and
narrow sulci.
Remarks: By its great size, the general feature of the coiling and rib-
bing, the specimen is similar to those figured by Galácz et al. (2010, pl.
3, Fig. 3; pl. 4, Fig. 3).
Distribution: Thouarsense Zone of Lyon region, Normandy and
Center-West France (Elmi and Rulleau, 1995); Gradata and
F
Thouarsense zones of Hungary (Galácz et al., 2010; Kovács, 2014b);
Gradata Zone of the Folded Middle Atlas (Elmi and Benshili, 1987) and
middle Toarcian (Gradata Zone?) of the study area (this work).
OO
Subfamily Mercaticeratinae Guex, 1973b
Genus Crassiceras Merla, 1932
Type species: Crassiceras latum Merla, 1932
Crassiceras latum Merla, 1932
Fig. 14E–G.
1932 Crassiceras latum Merla. Amm. App. Cent., p. 43, pl. 5, Fig. 10.
1963 Crassiceras latum Merla - Zanzucchi, p. 140; pl. 20, Fig. 13.
Material: Eight specimens.
Description: Medium-sized specimens of subrectangular, to sub-
quadratic whorl-section, with maximum thickness on the umbilical
edge; moderately rapid growth whorls and moderately evolute umbili-
cus; rounded umbilical edge; short subvertical to slightly sloping um-
bilical wall; subparallel to weakly convex flanks, slightly converging
towards the ventrolateral edge and bearing thick, flexuous, regularly
spaced ribs, as wide as the intercostal spaces, single or grouped in pair
at an elongated umbilical tubercle and strongly curved forward on the
outer part of the flank; wide venter with high median keel bordered by
two deep sulci.
Remarks: By their relatively thick shells with slightly open umbili-
cus, the study specimens are close to the specimen Crassiceras latum
figured by Zanzucchi (1963, pl. 20, Fig. 13). They differ from it by rel-
atively less thick ribs.
Distribution: Toarcian of Italy (Zanzucchi (1963); middle Toarcian
(Gradata? zones) of the study area (this work).
Crassiceras gradatum (.).
Fig. 14G–H.
1867-81 Ammonites comensis Meneghini. Mongraphie. p. 29 (pars), pl.
7, Fig. 1.
1931 Ammonites comensis V. Buch, p. 3, pl. 2, Fig. 3.
1932 Brodiceras gradatum Merla, p. 36, pl. 4, Figs. 6–8.
1963 Brodieia gradata (Merla) - Pinna, p. 86; pl. 11, Fi gs. 12, 17–19;
pl. 12, Fig. 10, with synonymy.
1969 Brodieia gradata (Merla) - Pinna, pl. 2, Fig. 4.
1968 Brodieia gradata (Merla) - Barbera, p. 303; pl. 7, Fig. 7.
Material: Two specimens.
Remarks: C. gradatum differs from C. latum by a more open umbili-
cus and less rapid growth whorls. The study specimens are close to the
specimen Brodieia gradata figured by Pinna (1969, pl. 2, Fig. 4).
Distribution: Toarcian of Italy (Barbera, 1968; Pinna, 1963, 1969);
middle Toarcian (Gradata? zones) of the study area (Bulundwe Kitongo,
1987, and this work).
Genus Pseudomercaticeras Merla, 1932
Type species: Pseudomercaticeras parvilobum Merla, 1932; Arkell,
1957).
Pseudomercaticeras rotaries Merla, 1932
Fig. 15B–C.
1867-81 Ammonites comensis Meneghini, p. 21; pl. 8, Fig. 3.
M. Benzaggagh
PR
D
TE
Fig. 15. Toarcian ammonites from the Jebel Amergou section, Internal Prerif
Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A. Mouterde-
iceras dubourgi Elmi and Rulleau (1995), JAM109; B–C. Pseudomercaticeras
EC
rotaries Merla, 1932, B. JAM98, C. JAM99.
1963 Pseudomercaticeras rotaries Merla - Pinna, p. 82; pl. 11, Fi gs. 1–4,
6, with synonymy.
1969 Pseudomercaticeras rotaries Merla - Pinna, p. 82; pl. 3, Fig. 4.
1963 Pseudomercaticeras rotaries Merla - Zanzucchi, p. 139; pl. 19, Fi
RR
gs. 2, 6–7, with synonymy.
2015 Pseudomercaticeras rotaries Merla - Kovács, pl. 2, Figs. 1–2.
Material: Tow specimens.
Description: Small-sized shell with shallow and wide evolute um-
bilicus; subrectangular to subquadratic whorl-section; rounded umbili-
cal edge; flat to slightly convex flanks bearing strong, mainly simple,
CO
slightly flexuous ribs, regularly spaced and less wide than the inter-
costal spaces, showing thickening at the ventrolateral edge; venter flat
relatively broad with a median keel bordered by two sulci.
Remarks: The two specimens are close to the specimens figured by
Zanzucchi (1963, Fig. 6) and Kovács (2015, pl. 2, Figs. 1–2).
Distribution: According to Kovács (2015), the genus Pseudomercat-
iceras is known from the middle Toarcian (Bifrons and Gradata/Vari-
abilis zones) of Italy, Austria, Greece, SE and NE Spain, Portugal, North
Africa, SE France, Romania, Oman, North America and Hungary. The
species occurs in the middle Toarcian (Gradata? zones) of the study
area.
Subfamily Grammoceratinae Buckman, 1909
Genus Pseudogrammoceras Buckman, 1901
Type species: Ammonites struckmanni Denckmann, 1901 (p. 72, pl. 3,
Fig. 1), designated by Donovan (1958) (p. 52).
18
Journal of African Earth Sciences xxx (xxxx) 105082
Pseudogrammoceras mediterraneum García Gómez and Rivas,
1980.
Fig. 16A.
1980 Pseudogrammoceras mediterraneum García Gómez and Rivas, p.
269; pl. 2, Figs. 1–5.
2002 Pseudogrammoceras mediterraneum García Gómez and Rivas, p.
177; pl. 1, Fig. 11.
Material: Two specimens.
F
Description: Compressed, small-sized shell with elliptical whorl-
section, much higher than wide; shallow and moderately evolute um-
bilicus; short and rounded umbilical wall; slightly convex flanks bear-
OO
ing fine, simple flexuous ribs, strongly inflected forward on the outer
part of the flank; narrow, rounded venter showing a low median keel
bordered by two narrow, flat spaces to very shallow sulci.
Distribution: Thouarsense Zone of Bulgaria (Metodiev, 2002), and
Betic Cordillera (García Gómez and Rivas, 1980); upper Toarcian? Of
the study area (this work).
Pseudogrammoceras subregale Pinna (1968).
Fig. 16B–C.
1968 Pseudogrammoceras subregale Pinna, p. 51; pl. 4, Figs. 9–11; pl.
1 nt, Fi gs. 18 and 20; pl. 2 nt, Fi g. 36.
1974 Pseudogrammoceras subregale Pinna - Elmi et al. pl. 4, Fig. 5.
1998 Pseudogrammoceras subregale Pinna - Parisi et al. pl. 3, Fig. 11.
2002 Pseudogrammoceras subregale Pinna - Metodiev, p., 174, pl. 1, Fig.
9.
2006 Pseudogrammoceras subregale Pinna - Géczy et al., 2008, pl. 9,
Figs. 3–4.
2020 Pseudogrammoceras subregale Pinna - Kovács et al. p. 147; pl. 1,
Figs. 5–6.
Material: Three specimens.
Description: Compressed, medium-sized shell with evolute umbili-
cus and subrectangular whorl-section with maximum thickness on the
umbilical edge; subvertical to slightly sloping umbilical wall; flat and
subparallel flanks slightly sloping towards the ventrolateral edge, bear-
ing simple, flexuous ribs inflected forward on the outer part of the
flank; narrow venter with a keel bordered by two narrow flat spaces to
very shallow sulci.
Distribution: Thouarsense Zone of Bulgaria (Metodiev, 2002); up-
per Toarcian of Italy (Parisi et al., 1998; Pinna, 1968); Gradata Zone of
Hungary (Kovács et al., 2020), NW Algeria (Elmi et al., 1974) and low-
ermost upper Toarcian of the study area (this work).
Genus Podagrosites Guex, 1973b
Type species: Pseudogrammoceras podagrosum Monestier, 1931.
Podagrosites pseudogrunowi Guex (1975).
Fig. 16D–E.
1874 Ammonites grunowi Hauer - Dumortier, p. 67; pl. 14, Figs. 6–7; pl.
15, Figs. 1–2.
1975 Podagrosites latescens var. Pseudogrunowi Guex, p. 105; pl. 3, Fi
gs. 4–5, 7–8, 11–13; pl. 4, Figs. 1 and 3.
1989 Podagrosites pseudogrunowi Guex - Rulleau, p. 71; pl. 9, Fig. 1; pl.
10, Figs. 3–6, with synonymy.
1993 Podagrosites pseudogrunowi Guex - Rulleau, pl. 1, Figs. 1–2.
2001 Podagrosites pseudogrunowi Guex - Rulleau et al. pl. 13, Fig. 5.
2002 Podagrosites pseudogrunowi Guex - Fauré, p. 725; pl. 17, Fi g. 18.
2002 Podagrosites aff. Pseudogrunowi Guex - Metodiev, pl. 5, Figs. 6–7.
2007 Podagrosites pseudogrunowi Guex - Elmi et al. p. 2, Fig. 1.
Material: Two specimens.
Description: Compressed, evolute shell with subtrapezoidal
whorl-section with maximum thickness on the umbilical edge; smooth
M. Benzaggagh
and relatively high subvertical umbilical wall; subflat flanks, slightly
sloping towards the ventrolateral edge, bearing strong, simple sig-
moid ribs strongly inflected forward on the outer part of the flank;
venter flat bearing a high keel bordered by well-marked sulci.
Distribution: Thouarsense and Insigne zones of Causses (Guex,
1975); Thouarsense Zone of eastern Pyrenees (Fauré, 2002) and Bul-
garia (Metodiev, 2002); upper Toarcian of Lyon region (Rulleau, 1989,
1993; Rulleau et al., 2001) and Lusitanian basin, Portugal (Elmi et al.,
1997) and lowermost upper Toarcian of the study area (this work).
Family Graphoceratidae Buckman, 1909
Subfamily Leioceratinae Spath, 1936.
Genus Leioceras Hyatt, 1867.
Type species: Nautilus opalinus Reinecke, 1818.
Leioceras opalinum (Reinecke, 1818).
Fig. 16F–G.
1818 Nautilus opalinus Reinecke, Figs. 1–2.
1981 Leioceras opalinum (Reinecke) - Wierzbowsk, p. 214; pl. 3.3–5; pl.
4.3–4.
1985 Leioceras opaliniforme (Buckman) - Ureta, p. 198; pl. 1, Figs. 4–6,
PR
with synonymy.
1989 Leioceras (Leioceras) opalinum (Reinecke) - Benshili, pl. 19, Fig.
10.
1995 Leioceras opalinum (Reinecke) - Rulleau, pl. 16, Figs. 1–5, pl. 17,
Fi gs. 1–2, 5–6.
1997 Leioceras opalinum (Reinecke) - Elmi et al. pl. 12, Fig. 1.
2001 Leioceras (Leioceras) opalinum (Reinecke) - Rulleau et al. pl. 19,
Figs. 4–6; pl. 20, Fig. 4.
D
2002 Leioceras opalinum (Reinecke) - Fauré, p. 333; pl. 24, Figs. 4–5.
2003–2004 Leioceras opalinum (Reinecke) - Pallini et al. p. 11, with
synonymy
2004 Leioceras opalinum (Reinecke) - Myczyński, p. 73; Fi g. 26.8 and
TE
Fi g. 27.5, with synonymy.
2008 Leioceras opalinum (Reinecke) - Seyed-Emami, p. 252; Fig. 6C,
with synonymy.
2008 Leioceras (Leioceras) opalinum (Reineck) - El Hammichi, pl. 2, Fig.
9.
EC
Material: Two specimens.
Description: Very compressed shell with oval elongated whorl-
section; shell smooth showing traces of fine ribs in the inner whorls;
broad and subflat flanks, with fast growth whorls; narrow, moderately
evolute umbilicus, with overlap of about three-quarter of flanks; verti-
cal umbilical wall in inner whorls, becoming sloping on the last whorl;
RR
narrow, rounded venter showing traces of a median keel.
Distribution: Lower Aalenian (Opalinum Zone) of eastern Pyre-
nees (Fauré, 2002), Lyon region (Rulleau, 1995; Rulleau et al., 2001),
West Carpathians (Myczyński, 2004), Spitsbergen, northern Norway
(Wierzbowski et al., 1981), Italy (Pallini et al., 2003-2004), Iran
(Seyed-Emami et al., 2008), central High Atlas (Sadki, 1996); upper
CO
Toarcian and Aalenian of the Folded Middle Atlas (Benshili, 1989; El
Hammichi et al., 2008) and the study area (this work).
Leioceras comptum (Reinecke, 1818).
16H–I.
1818 Nautilus comptus Reinecke, 57, pl. 1, Figs. 5–6.
1997 Leioceras comptum (Reinecke) - Elmi et al. pl. 12, Fig. 4.
2001 Leioceras (Cypholioceras) comptum (Reinecke) - Rulleau et al. pl.
19, Fig. 7; pl. 20, Figs. 1–2.
2003–2004 Leioceras (Cypholioceras) comptum (Reinecke) - Pallini et
al. p. 11; pl. 7, Figs. 1–3; pl. 15, Fig. 16, with synonymy.
2004 Leioceras comptum (Reinecke) - Myczyński, p. 78; Fi gs. 27.2, 6–7,
with synonymy.
2004 Leioceras aff. Comptum (Reinecke) - Seyed-Emami et al., p. 85, pl.
2, Figs. 3 and 5.
19
Journal of African Earth Sciences xxx (xxxx) 105082
2008 Leioceras comptum (Reinecke) - Seyed-Emami et al., p. 254; Fig.
6G, I.
Material: Two specimens.
Description: Small-sized shell with oxycone coiling and involute
umbilicus; oval whorl-section, with maximum thickness slightly below
the mid-flank; short subvertical to slightly sloping umbilical wall and
rounded umbilical edge; flanks slightly convex, gradually joining the
venter, without a distinct ventrolateral edge, and bearing broad blunt
falcoid ribs, simple or grouped in pair, they are provers on the lower
F
part of the flank, then inflected forward on the outer flank; sharp venter
bearing a sharp keel.
Remarks: L. comptum differs from L. opalinum by its more involute
OO
umbilicus, more convex flanks, with broad falcoid ribs and acute ven-
ter.
Distribution: Lower Aalenian (Opalinum Zone) of Lyon region
(Elmi et al., 1997; Rulleau et al., 2001); Italy (Pallini et al., 2003-2004),
West Carpathians (Myczyński, 2004), Iran (Seyed-Emami et al., 2004,
2008), central High Atlas (Sadki, 1996), and the study area (this work).
Family Tmetoceratidae Spath, 1936.
Subfamily Tmetoceratinae Spath, 1936.
Genus Tmetoceras Buckman, 1892
Type species: Ammonites scissus Benecke, 1865, designated by
Buckman (1909).
Tmetoceras scissum (.).
Fig. 16J-N.
1865 Ammonites scissus Benecke, p. 170; pl. 6, Fig. 4
1998 Tmetoceras scissum (Benecke) - Alméras et al. p. 104; pl. 17, Fig.
3.
1999 Tmetoceras scissum (Benecke) - Fernández-López et al., p. 94;
Figs. 2–3, 8 A-N, 9 F-Q, with synonymy.
2001 Tmetoceras scissum (Benecke) - Rulleau et al. pl. 16, Fig. 4.
2004 Tmetoceras scissum (Benecke) - Myczyński, p. 7; Fi gs. 26.4-6,
with synonymy.
2003–2004 Tmetoceras scissum (Benecke) - Pallini et al., 2003-2004 p.
11; pl. 5, Figs. 1–2, with synonymy.
2007 Tmetoceras scissum (Benecke) - Rulleau, 2006, p. 93; pl. 52, Fig.
7.
2010 Tmetoceras scissum (Benecke) - Kovács, p. 363; pl. 5, Figs. 2 and
4, with synonymy.
Material: Seven specimens.
Description: Small to medium-sized shell with subcircular to
slightly oval whorl-section, higher than wide; wide and shallow umbili-
cus; rounded umbilical wall; convex flanks bearing sharp simple,
straight to slightly flexuous ribs less wide than the intercostal spaces
and showing a slight thickening at the ventrolateral edge where are in-
flected forward; narrow, flat to slightly rounded venter showing a rela-
tively narrow and shallow median sulcus.
Distribution: Opalinum Zone of Lyon region (Alméras et al.,
1998; Rulleau et al., 2001, 2007), Hungary (Kovács, 2010), West
Carpathians (Myczyński, 2004) and Spain (Fernández-López et al.,
1999); lowermost Aalenian of the study area (Bulundwe Kitongo,
1987; Lacoste, 1934, and this work) (see also Fernández-López, 1999).
Superfamily Hammatoceratoidea Schindewolf, 1964
Family Hammatoceratidae Buckman, 1887
Subfamily Hammatoceratinae Buckman, 1909
Genus Crestaites Rulleau et al., 2001.
Type species: Hammatoceras meneghinii Bonarelli (after Meneghini,
1867–1881, pl. 13, Fig. 1). Holotype refigured in Merla, 1932) (p. 10;
pl. I, Fig. 1).
Crestaites meneghinii (Bonarelli, 1899).
Fig. 17C–D.
M. Benzaggagh
PR
D
TE
Fig. 16. Toarcian and lowermost Aalenian ammonites from the Jebel Amergou
section, Internal Prerif Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral
views. A. Pseudogrammoceras mediterraneum García Gómez and Rivas, 1980,
JAM105; B–C. Pseudogrammoceras subregale Pinna (1968), B. JAM102, C.;
EC
JAM101; D-E. Podagrosites pseudogrunowi Guex (1975), D. JAM104, E.
JAM205; F-G. Leioceras opalinum (Reinecke, 1818), F. JAM86, G. JAM87;
H–I. Leioceras comptum (Reinecke, 1818), H. JAM107, I. JAM108; J-N.
Tmetoceras scissum (Benshili, 1989), J. JAM113, K. JAM120, L. JAM115, M.
JAM116, N. JAM117.
1867–1881 Ammonites insignis Schübler - Meneghini, p. 59; pl. 13, Fig.
RR
1.
1899 Hammatoceras Meneghinii Bonarelli, p. 208.
1991 Hammatoceras meneghinii Bonarelli - Elmi and Rulleau, p. 323; Fi
g. 2.1, pl. 2, Fi gs. 1–2.
2008 Crestaites meneghinii (Bonarelli) - Géczy et al. pl. 3, Fig. 4.
2008 Crestaites cf. Meneghinii (Bonarelli) - El Hammichi et al. pl. 4, Fig.
CO
6.
2009 Crestaites meneghinii (Bonarelli) - Kovács, p. 20, Figs. 10–11, with
synonymy.
2020 Crestaites meneghinii (Bonarelli) - Kovács et al. p. 154, pl. 7, Fig.
1.
Material: Two specimens.
Description: Large-sized compressed shell with oval whorl-
section, higher than wide; moderately wide and shallow umbilicus;
short rounded umbilical wall; slightly convex flank gradually joining
the rounded ventrolateral edge; flanks bearing strong and radiate pri-
mary ribs emerging from the umbilicus, without tubercules, and bi-
furcate near the mid-flank; secondary ribs bend forward from the fur-
cation points; narrow and rounded venter showing a smooth median
band.
20
Journal of African Earth Sciences xxx (xxxx) 105082
Distribution: Speciosum Zone of Hungary (Kovács et al., 2020);
upper Toarcian of the Folded Middle Atlas (Benshili, 1989; El Arabi et
al., 1999; El Hammichi et al., 2008) and the study area (this work) (see
also, Kovács, 2009).
Crestaites victorii (Bonarelli, 1895).
Fig. 17E.
1867–1881 Ammonites insignis Schübler - Meneghini, p. 56; pl. 14, Fig.
2.
1899 Hammatoceras victorii Bonarelli, p. 209.
F
1915 Hammatoceras victorii Bonarelli - Principi, p. 447; pl. 16, Fig. 2.
1991 Hammatoceras victorii Bonarelli - Elmi and Rulleau, pl. 3, Figs.
3–4.
OO
2009 Crestaites victorii (Bonarelli) - Kovács, p. 25, Figs. 14–15.
2020 Crestaites gr. Victorii (Bonarelli) - Kovács et al. p. 156; pl. 7, Fig.
5; pl. 8, Figs. 2–3; pl. 9, Figs. 1 and 3.
Material: One specimen.
Description: This species differs from C. meneghinii by its coarse
and more spaced ribs, mainly on the last whorl and more convex flanks.
Remarks: By its coarse ribs, the studied specimen is close to Ham-
matoceras victorii figured by Principi (1915, pl. 16, Fig. 2) and Crestaites
victorii figured by Kovács (2009, p. 25, Fig. 14).
Distribution: Upper Toarcian of Italy (Principi, 1915), Speciosum
and Meneghinii zones of Hungry (Kovács, 2009; Kovács et al., 2020);
Meneghinii-Aalensis zones of the Folded Middle Atlas (El Hammichi et
al., 2008) and upper Toarcian of the study area (this work) (see also
Kovács, 2009).
Genus Geczyceras Martínez, 1992.
Type species: Hammatoceras speciosum Janensch, 1902 (pl. 10, Fig.
1), designated by Martínez (1992).
Geczyceras costulosum (.).
Fig. 17A–B.
1934 Hammatoceras costulosum Merla, p. 13; pl. 1, Figs. 3–4.
1995 Geczyceras costulosum (Merla) - Goy, 1974. p. 100; pl. 12, Fig. 1.
2011 Geczyceras costulosum (Merla) - Sandoval et al. pl. 10, Fig. 1.
2015 Geczyceras costulosum (Merla) - Martínez et al. p. 17; pl. 6, Fig. 6.
2020 Geczyceras costulosum (Merla) - Kovács et al. p., 149; pl. 4, Fi gs.
3, 4–5.
Material: Two specimens.
Description: Compressed, medium to large-sized shell with oval
whorl-section, higher than wide; moderately wide and shallow umbili-
cus; short subvertical to slightly sloping umbilical wall, and rounded
umbilical edge; slightly convex flank gradually joining the rounded
ventrolateral edge; flanks bearing strong, radiate primaries ribs arising
from a more or less well-developed umbilical tubercle and bifurcate or
trifurcate near the mid-flank; secondary ribs bend forward from the
furcation points; narrow and rounded venter showing a smooth me-
dian band.
Remarks: The study specimens differ from those figured by
Martínez et al. (2015, pl. 6, Fig. 6) and Kovács et al. (2020 pl. 4, Fi gs. 3,
4–5.) by their less convex flanks and subvertical umbilical wall.
Distribution: Speciosum Zone of Hungary (Kovács et al., 2020);
Meneghinii Zone of Iberian Range and Betic Cordillera (Sandoval et al.,
2011); upper Toarcian of the study area (this work) (see also Kovács et
al., 2020; Martínez et al., 2015).
9. Biostratigraphy and paleobiogeography
The study ammonite specimens were collected during several field
missions. Most of them were not collected from layers, which makes it
difficult to characterize with precision the different zones and subzones
of the Toarcian–lower Aalenian stages and their specific assemblages.
M. Benzaggagh
PR
D
TE
Fig. 17. Upper Toarcian ammonites from the Jebel Amergou section, Internal
EC
Prerif Zone. 1. Lateral view, 2 and 3. Frontal and/or ventral views. A-B
Geczyceras costulosum (Merla, 1932), A. JAM110, B. JAM112, C-D.
Crestaites meneghinii (Bonarelli, 1899), C. JAM111, D. 216; E. Crestaites vic-
torii (Bonarelli, 1895), JAM207.
Nevertheless, several of the identified species have short stratigraphic
RR
ranges and are considered in the Mediterranean and the Submediter-
ranean domains as index species of biozones or species accompanying
the index species and therefore make it possible to confirm the presence
in the study succession, the most ammonite biozones of the mid-
dle–upper Toarcian and the lowermost Aalenian. Thus, the Sublevisoni
and Bifrons subzones have been characterized by their index species, in-
cluding, Hildoceras sublevisoni, H. apertum and Hildoceras tethysi, for the
CO
first subzone, and Hildoceras bifrons, H. crassum, H. semipolitum and H.
snoussi, for the second subzone. These two subzones also contain,
Dactylioceras (D.) aff. Attenuatus, Harpoceras falciferum, H. pseudoser-
pentinum, H. subplanatum, Nodicoeloceras tuberculatum, Taffertia taffer-
tensis and Zugodactylites cf. Braunianus. The Gradata zone is defined on
the basis of the species Merlaites gradata (Merla) which was not found in
the study section. However, this biozone is well characterized by ac-
companying species, such as Crassiceras latum, Crassiceras gradatum,
Furloceras iserensis, F. chelussii, Mouterdeiceras dubourgi, Pseudomercat-
iceras rotaries and Collina gemma. Lacoste (1934) reported from the
same section typical ammonites of the Gradata zone, including Haugia
variabilis and H. grandis Buckman; the Bonarelli Zone, equivalent to the
NW European Thouarsense Zone, can be characterized by accompany-
ing species, such as, Pseudogrammoceras mediterraneum, P. subregale,
Podagrosites pseudogrunowi, Alocolytoceras coarctatum, A. dorcadis, Peri-
21
Journal of African Earth Sciences xxx (xxxx) 105082
lytoceras perlaevis and Polyplectus discoides; the Speciosum? Zone can be
characterized by Crestaites meneghinii, C. victorii and Geczyceras costulo-
sum and the Opalinum Zone by Leioceras opalinum, Leioceras comptum
and Tmetoceras scissum. The Polymorphum and Levisoni subzones may
correspond to the level 1 and 2, of the lower member of the studied sec-
tion which did not provided ammonites. However, the second subzone
seems well-represented in the study area, as Bulundwe Kitongo (1987)
reported from his lateral point outcrop 2, two typical ammonite species
of the Levisoni Subzone, Hildaites striatus Guex and Hildaites sp. The
specimens of Calliphylloceras capitanii, C. aff. Capitanii, C. nilssoni, Lyto-
F
ceras francisci, Nodicoeloceras sp. Indet, Phylloceras heterophyllum, P. do-
derleinianum, Polyplectus appenninicus, Zugodactylites sp. Indet., and
Telodactylites eucosmus are mainly from the Bifrons and Gradata zones.
OO
Among the listed ammonites, several species such as: Calliphyllo-
ceras capitanii, C. aff. Capitanii, Collina gemma, Crassiceras gradatum, C.
latum, Crestaites meneghinii, C. victorii, Furloceras iserensis, Hildoceras
snoussi, Geczyceras costulosum, Lytoceras francisci, Phylloceras doderleini-
anum, Polyplectus appenninicus, Pseudogrammoceras mediterraneum and
P. subregale seem to be restricted to the Mediterranean areas (Italy, SE
Spain, eastern Europe, and north Maghrebian margin of the Tethys);
two species, Calliphylloceras aff. Capitanii and Hildoceras snoussi seem to
be restricted to the Moroccan Jurassic basins and Taffertia taffertensis is
knowns only in Morocco and North America; while species Alocolyto-
ceras coarctatum, A. dorcadis, Calliphylloceras nilssoni, Dactylioceras (D.)
aff. Attenuates, Furloceras chelussii, Harpoceras falciferum, H. pseudoser-
pentinum, H. subplanatum, Hildoceras apertum, H. bifrons, H. crassum, H.
semipolitum, H. sublevisoni, H. tethysi, Leioceras opalinum, L. comptum,
Mouterdeiceras dubourgi, Nodicoeloceras tuberculatum, Perilytoceras per-
laevis, Phylloceras heterophyllum, Podagrosites pseudogrunowi, Polyplectus
discoides, Pseudomercaticeras rotaries, Telodactylites eucosmus, Taffertia
taffertensis, Tmetoceras scissum and Zugodactylites cf. Braunianus are
common to the Mediterranean areas and the NW European province
(southern Germany, SE England, Luxembourg, France and North Amer-
ica).
The occurrence of the most listed ammonites, both in the Tethyan
and the NW European domains, mainly, the middle Toarcian species,
such as species of the genera Hildoceras and Harpoceras, attests to ma-
rine communications between the NW European province and the dif-
ferent basins of the Tethys allowing migration of marine macrofauna
especially from the NW European province to the Tethys, but also in the
opposite way.
10. Paleogeography
In a paleogeographic framework, the abundance of outcrops rich in
ammonites of middle Toarcian, and to a lesser degree, of upper Toar-
cian-Aalenian, in several Jurassic basins of the Tethyan and the NW Eu-
ropean domains, evokes the installation, at that time, over large marine
areas, of favorable conditions to the proliferation of ammonite faunas,
mainly the benthic and nektobenthic forms, which tolerate marine en-
vironments of moderate depth such as external platforms.
In the Moroccan Jurassic basins, two types of marine environments
can be distinguished: deep and subsiding furrows corresponding to the
axial zones of the central High Atlas (Benzaggagh et al., 2022a, 2022b;
Ettaki, 2003; Ettaki and Chellai, 2005), the Folded Middle Atlas
(Benshili, 1989), the Central unit of the South Riffian Ridges
(Benzaggagh, 2020, 2022) and the eastern foreland of the Rif Belt
(Hervouët, 1985). In these sectors, whit very thick Toarcian marlstone
and marly limestone series (up to 500 m-thick), ammonites are very
rare.
Whereas, in the External Rif, the Beni Snassen and some shallow
marine areas of the Folded Middle Atlas and the central High Atlas
basins, the Toarcian Stage is represented by condensed marlstone and
limestone series (few tens of meters), characterizing moderately deep
marine environments. In these sectors ammonites are often abundant.
M. Benzaggagh
11. Conclusions
The Toarcian and Aalenian outcrops in the Internal Prerif Zone and
the External Rif belt are rare. The most important of these outcrops are
those of the northern side of Jebel Amergou (western part of the Inter-
nal Prerif). A section (JAM) located at the northwest extremity of Jebel
Amergou have been studied. Its Toarcian–base Aalenian series (20 m-
thick) called the Amergou Formation, consists of alternation of lumpy
marlstone and thin limestone beds and is divided into two members.
The lower member is composed of four levels; the levels 1 and 2, which
did not yield ammonites, are attributed to the lower Toarcian; the level
3 is rich in ammonites of the middle Toarcian (Sublevisoni and Bifrons
subzones); the level 4 provided ammonites of the middle Toarcian (Gra-
data Zone) and upper Toarcian (Bonarellii and Speciosum? zones); The
upper member is divided into two levels, the level 2 yielded in its lower
part rare ammonites of the lower Aalenian (Opalinum Zone). The
Bifrons and Bonarellii zones are the richest in ammonites. 44 species
have been identified, described and illustrated for the first time from
the middle Toarcian–base Aalenian of the Internal Prerif Zone and the
External Rif belt.
PR
Ammonite assemblages of the defined biozones are similar to those
reported for the same stratigraphic interval from several basins of the
Tethyan and the NW European domains. The study ammonites show
that some species are restricted to the basins of the northern and the
southern margins of the Tethys and are of Mediterranean origin; while
numerous other species are common to the Mediterranean and the NW
European domains. These common species attest to marine communica-
tions between these two paleogeographic domains during the Toar-
D
cian–lower Aalenian time interval. The abundance of outcrops rich in
ammonites, particularly those of the middle Toarcian, in several Juras-
sic basins of the Tethys and the NW European domains, evokes favor-
able conditions (bathymetric and nutrient) over wide marine areas, al-
TE
lowing the proliferation of benthic and nektobenthic ammonite faunas.
In Morocco, the Toarcian–Aalenian stages are represented in the ax-
ial zones of the most Jurassic basins (central High Atlas, Folded Middle
Atlas, eastern foreland of the Rif Belt, Central unit of the South Riffian
Ridges) by very thick marlstone and marly limestone series evoking
EC
deep and subsiding furrows. In these sectors ammonites are very rare.
In the External Rif, the Beni Snassen and in some localities of the Folded
Middle Atlas, and the central High Atlas, Toarcian series, consisting of
marlstone and marly limestone alternation with reduced thickness,
evoke moderately deep marine environments. In these series, ammonite
faunas are often abundant.
RR
Declaration of competing interest
The author declares that he has no known competing financial inter-
ests or personal relationships that could have appeared to influence the
work reported in this paper.
CO
Data availability
No data was used for the research described in the article.
Acknowledgments
The author is very grateful to the reviewers, Dr. Philippe Fauré
(Toulouse) and Dr. Zoltán Kovács (Hungary) for their numerous valu-
able comments and constructive remarks which highly increased the
quality scientific of the manuscript. He also thanks Dr Damien Delvaux,
editor-in-chief of the Journal of African Earth Sciences for his remarks
on the first version of the manuscript.
22
Journal of African Earth Sciences xxx (xxxx) 105082
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