El Mers Group
Stratigraphic range: Bajocian-Callovian
~168–161 Ma

PreЄ

El Mers I Formation at Thaghzout river canyon

El Mers II along the El Mers river

El Mers II along the El Mers river

El Mers III Dinosaur Bearing beds

Type| Group
Sub-units| El Mers Formations 1–3
Underlies| Unconformity with Barremian or Plio-Pleistocene sediments
Overlies| Ich Timellaline–Bou Akrabene Formation
Thickness| Over 500 m (1,600 ft)
Lithology
Primary| Marl, gypsum (only in unit 3)
Other| Sandstone, limestone
Location
Coordinates|  [ ⚑ ] : 33°30′N 4°18′W﻿ / ﻿33.5°N 4.3°W﻿ / 33.5; -4.3
Paleocoordinates|  [ ⚑ ] 28°18′N 2°00′W﻿ / ﻿28.3°N 2.0°W﻿ / 28.3; -2.0
Region| Boulemane and Azilal Provinces
Country| Morocco
Extent| Middle Atlas

El Mers Group (Morocco)

The El Mers Group is a geological group in the Middle Atlas of Morocco. It is subdivided into three formations named the El Mers I, II, and III Formations, respectively. It is a marine deposit primarily consisting of marl, with gypsum present in the upper part of unit 3. It is the lateral equivalent of the terrestrial Guettioua Sandstone. Dinosaur remains are among the fossils that have been recovered from the group, most notably those of sauropods and the unusual thyreophorans Adratiklit, Thyreosaurus, and Spicomellus. This formation records a series of coastal to shallow marine settings of Middle Jurassic (with an age of Bajocian-Bathonian for El Mers 1 and a Bathonian-Callovian? for El Mers II-III), based on detailed stratigraphical biomarkers such as ammonites or Foranimifera.[1]

## Contents

* 1 Paleoenvironment
* 2 Fossils
* 2.1 Foraminifera
* 2.2 Dinoflagellates
* 2.3 Brachiopoda
* 2.4 Ostracoda
* 2.5 Xiphosura
* 2.6 Mollusca
* 2.7 Echinoderms
* 2.8 Fishes
* 2.9 Testudinata
* 2.10 Crocodylomorpha
* 2.11 Dinosauria
* 2.11.1 Theropoda
* 2.11.2 Sauropoda
* 2.11.3 Ornithischia
* 2.12 Dasycladales
* 2.13 Charophyta
* 2.14 Plantae
* 3 See also
* 4 References

## Paleoenvironment

A Modern analogue for El Mers Group is found in the coastal sequences of Coorong, South Australia.

The El Mers formations are composed primarily of marls with intermittent layers of limestone and sandstone, characterized by abundant organic matter and benthic fauna.[2] It is delineated into five distinct members, each exhibiting diverse lithological characteristics. The three members comprise a combination of layers of marls, sandstones, and limestones, marked by various unique fossils and organic materials. Sedimentary discontinuities reveal deltaic sequences that exhibit varying sedimentary structures and lithological compositions, reflecting dynamic depositional environments ranging from subtidal to continental. The facies associations within the El Mers Group are organized into elementary and mesosequences, delineating the transition from prodelta to deltaic plain environments. These sequences capture the evolution of tidal regimes and depositional dynamics.[3][4]

During the Bajocian-Bathonian transition the local faices paint a picture of a dynamic paleoenvironment, where the interplay between land and sea was pronounced.[1] This setting oscillated between a proximal continental shelf, marked by significant terrestrial input, and episodic marine incursions that suggest fluctuating sea levels.[1] The lower parts of the section hint at shallow, sometimes oxygen-deprived, stagnant waters, much like modern lagoon or restricted bays where circulation is limited. Upwards there are signs of marine influence growing stronger, particularly at the base of "Interval C", where a notable sea-level rise seems to have occurred, akin to what we might see on today's outer continental shelves, like those in the North Sea, where marine life diversity peaks, specially Bivalves and Ammonites. However, this marine dominance is not consistent, reverting back to a more coastal, land-influenced environment towards the top, similar to modern deltaic regions where riverine and marine waters mix, such as the Louisiana coast or the Coorong National Park. The environment would have experienced variable salinity, much like estuaries where fresh and marine waters blend, affecting the types of biota that could thrive.[1]

The shifts in facies and microfauna evolution in the area revelated intriguing patterns, from the establishment of initial supratidal environments characterized by charophytic marls and ostracod limestones followed by a resurgence of marine conditions, evident from the reappearance of coastal facies and the diversification of ostracod faunas, indicating a significant transgressive phase during the Upper Bathonian–Lower Callovian period. While in the Central High Atlas marine influences ceased by the Upper Bajocian–Lower Bathonian giving way to continental red layers, in the Middle Atlas marine influences persisted until the Upper Bathonian–Lower Callovian boundary, indicating a diachronic filling of the Atlas gulfs towards the Tethyan domain from west to east.[5]

## Fossils

Template:Paleobiota-key-compact

### Foraminifera

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Bullopora[5] |

* B. tuberculata

* SW part of the Skoura syncline

| Tests/Shells  | A benthic foraminiferan of the family Polymorphinidae  |
Pfenderella[5] |

* P. cf. arabica

* SW part of the Skoura syncline

| Tests/Shells  | A benthic foraminiferan of the family Pfenderininae  |
Praekurnubia[5] |

* P. crusei

* SW part of the Skoura syncline

| Tests/Shells  | A benthic foraminiferan of the family Pfenderinidae; a genus found in the Tethys between Bajocian and Oxfordian-aged layers  |
Pseudocyclammina[5] |

* P. maynci

* SW part of the Skoura syncline

| Tests/Shells  | A benthic foraminiferan of the family Hauraniidae  |

### Dinoflagellates

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Aldorfia[1] |

* A. aldorfensis

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Cribroperidinium[1] |

* C. crispum
* C. venustum

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Ctenidodinium[1] |

* C. combazii
* C. continuum
* C. cornigerum

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate, incertae sedis |
Dichadogonyaulax[1] |

* D. sellwoodii

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate, incertae sedis |
Korystocysta[1] |

* K. gochtii
* K. pachyderma

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Meiourogonyaulax[1] |

* M. caytonensis
* M. reticulata
* M. valensii

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Nannoceratopsis[1] |

* N. deflandrei

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Nannoceratopsiaceae  |
Pareodinia[1] |

* P. ceratophora

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Pareodiniaceae  |
Rhynchodiniopsis[1] |

* R. cladophora

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Trichodinium[1] |

* T. scarburghense

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Tubotuberella[1] |

* T. dangeardii

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Wanaea[1] |

* W. acollaris
* W. indotata
* W. verrucosa

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate of the family Gonyaulacaceae  |
Willeidinum[1] |

* W. spp.

* FD section, Skoura Syncline

| Cysts  | A Dinoflagellate, incertae sedis |

### Brachiopoda

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Cymatorhynchia[6][7] |

* C. quadriplicata

* El Mers

| Isolated shells  | A branchipod of the family Tetrarhynchidae  |
Daghanirhynchia[6][7] |

* D. termierae

* El Mers

| Isolated shells  | A branchipod of the family Tetrarhynchidae  |
Formosarhynchia[4] |

* F. (Rhynchonella?) dumortieri

* El Mers

| Isolated shells  | A branchipod of the family Rhynchonellidae  | Example of specimen of the genus (from Spain)
Rugitela[6][7] |

* R. praebullata

* El Mers

| Isolated shells  | A branchipod of the family Zeilleriaceae  |
Stiphrothyris[6][7] |

* S. mouterdei

* El Mers

| Isolated shells  | A branchipod of the family Lobothyrididae  |
Tubithyris[6][7] |

* T. chouberti
* T. cf. wrighti

* El Mers

| Isolated shells  | A branchipod of the family Lobothyrididae  |
Zeilleria[7] |

* Z. subbucculenta

* El Mers

| Isolated shells  | A branchipod of the family Zeilleriaceae  | Zeilleria specimens (from Spain)

### Ostracoda

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Cytherella[5] |

* C. index

* Ait Hammou

| Valves  | A freshwater/brackish ostracod of the family Cytherellidae |
Fastigatocythere[5] |

* F. aff. brentonensis

* Ait Hammou

| Valves  | A freshwater/brackish ostracod of the family Progonocytheridae  |
Kinkelinella[5] |

* K. aff. triangula

* Ait Hammou

| Valves  | A freshwater ostracod of the family Progonocytheridae  |
Metacypris[5] |

* M. spp.

* Ait Hammou

| Valves  | A freshwater ostracod of the family Limnocytheridae. Local members of the genus genus would not be confined to limnic environments, but would be euryhaline or brackish, since they disappear when faunal diversity is at its maximum.  |
Procytheridea[5] |

* P. ihopyensis

* Ait Hammou

| Valves  | A freshwater ostracod of the family Schulerideidae  |

### Xiphosura

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Selenichnites[8][9] |

* S. tesiltus
* S. isp.

* El Mers
* Tassmante O'Moche

| Isolated crescent-shaped trace fossils  | Traces of xiphosurans, representing the second occurrence of this ichnogenus in Morocco  | Extant Horseshoe crab (in New York state)

### Mollusca

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Cadomites[5] |

* C. bremeri
* C. cf. bremeri

* East Skoura syncline

| Isolated shells  | An ammonite of the family Stephanoceratidae and indicator of the Bathonian age  | Example of specimens (from France)
Cercomya[4] |

* C. undulata

* El Mers

| Isolated shells  | A marine clam of the family Laternulidae |
Choffatia[10] |

* C. cf. urinacensis

* East Skoura syncline

| Isolated shells  | An ammonite of the family Perisphinctidae and indicator of the Callovian age  |
Corbula[4] |

* C. pectinata

* Aït Kermouss

| Isolated shells  | A marine clam of the family Corbulidae |
Clydoniceras[10][11] |

* C. discus

* Tamguert n'Tarit
* Tizi n'Juillerh

| Isolated shells  | An ammonite of the family Clydoniceratidae  |
Eomiodon[4][12] |

* E. angulatus
* E. gardeti
* E. algeriensis
* E. indicus

* El Mers
* Skoura syncline

| Isolated Shells  | A marine clam of the family Neomiodontidae  |
Isognomon[4] |

* I. (Mythiloperna) murchisoni

* El Mers

| Isolated shells  | A marine oyster of the family Malleidae | Example of extant member of the genus (from Cape Verde)
Lima[4] |

* L. (Plagiostoma) subcardiformis

* Aït Kermouss

| Isolated shells  | A marine file clam of the family Limidae | Example of extant member of the genus (from Puerto Rico)
Lopha[4] |

* L. costata

* Aït Kermouss

| Isolated shells  | A marine oyster of the family Ostreidae | Example of extant member of the genus (from Micronesia)
Modiolus[4][11] |

* M. imbricatus
* M. cf. bipartitus

* Aït Kermouss
* El Mers
* Tamguert n'Tarit

| Isolated shells  | A marine mussel of the family Mytilidae | Example of extant member of the genus (from Philippines)
Morphoceras[10] |

* M. macrescens

* East Skoura syncline

| Isolated shells  | An ammonite of the family Morphoceratidae and indicator of the Bathonian age  |
Mytilus[4] |

* M. (Falcimytilus) tumidus

* Aït Kermouss

| Isolated shells  | A marine mussel of the family Mytilidae | Example of extant member of the genus (from France)
Neomiodon[12] |

* N. skouraensis

* El Mers
* Skoura syncline

| Isolated Shells  | A marine clam of the family Neomiodontidae  |
Nucula[4] |

* N. (Paleonucula) waltoni

* El Mers

| Isolated shells  | A marine nut clam of the family Nuculanidae | Example of extant member of the genus (from Massachusetts)
Oraniceras[10] |

* O. sp.

* East Skoura syncline

| Isolated shells  | An ammonite of the family Parkinsoniidae and indicator of the Bathonian age  |
Orionoides[4] |

* O. cf. pseudorion

* East Skoura syncline

| Isolated shells  | An ammonite of the family Perisphinctidae |
Ostrea[4][10][11] |

* O. sandalina
* O. (Liostrea) hebridica

* Aït Kermouss
* El Mers
* Tamguert n'Tarit

| Isolated shells  | A marine oyster of the family Ostreidae | Example of extant member of the genus (from Croatia)
Oxyeurax[12] |

* O. fimbriata

* Aït Kermouss
* El Mers
* Tamguert n'Tarit

| Isolated shells  | A marine clam of the family Astartidae |
Paracenoceras[4] |

* P. sp.

* Aït Kermouss

| Isolated shells  | A Nautilidan of the family Paracenoceratidae | Example of specimen (from an unknown location)
Parkinsonia[5] |

* P. sp.

* Skoura syncline

| Isolated shells  | An ammonite of the family Parkinsoniidae and indicator of the Bajocian age for the lower portions of the Group  | Example of specimen (from England)
Protocardia[10][11] |

* P. tikechkachensis

* El Mers
* Tamguert n'Tarit

| Isolated shells  | A marine cockle of the family Cardiidae |

### Echinoderms

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Acrosalenia[4] |

* A. spinosa

* El Mers
* Ich Timellaline

| Isolated Individuals  | An Echinoidean of the family Acrosaleniidae  |
Hemicidaris[4] |

* H. jauberti
* H. cf. icaunensis

* Ich Timellaline

| Isolated Individuals  | An Echinoidean of the family Hemicidaridae | Example of specimen of the genus (from Switzerland)

### Fishes

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Scheenstia[13] |

* S. cf. mantelli
* S ("Lepidotes") ssp.

* Ait Hammou
* Djmila
* Oued El Mers
* Tizi n'Juillerh

| Partial specimen wirth cranial material; Referred cranial material, isolated teeth and scales  | A marine/brackish bony fish of the family Lepidotidae | Reconstruction

### Testudinata

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Testudinata[10][11] |

* Indeterminate

* Tamguert n'Tarit
* Tizi n'Juillerh

| Very numerous debris: escutcheons, plates, vertebrae, ribs, long bones  | Turtle remains of uncertain affinity  |

### Crocodylomorpha

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Hatcherichnus[9] |

* H. ispp.

* Inzar O'Founass

| Footprints  | Indeterminate Crocodrylomorph Tracks  |

"Megalosaurus"[10][11]

* "M." mersensis

* Botane
* Tizi n'Juillerh

Isolated teeth and vertebra

A marine crocodylomorph of the family Teleosauridae. Mistaken as Theropod remains in the past.[14]

Steneosaurus?[10]

* S.? spp.

* Bou Iferaoun
* Botane
* Darak
* Djmila
* Oued Tamemecht
* Tamguert r'Tane
* Tirardine
* Tissenfelt

Isolated teeth, cranial and postcranial material, both isolated and semi-articulated

A marine crocodylomorph of the family Teleosauridae. The referral to the genus Steneosaurus should be taken with caution, as this genus was overlumped.

### Dinosauria

#### Theropoda

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Carmelopodus[9] |

* cf.C. isp.

* Tassmante O'Moche

| Footprints  | Theropod tracks of uncertain affinity, suggested to come from Ceratosauria members  |
Grallator?[9] |

* G.? isp.

* Ifri N'Tfrane

| Footprints  | Theropod Tracks, referred usually to small-medium sized Genera  |
Hispanosauropus?[9] |

* H.? isp.

* Ifri N'Tfrane

| Footprints  | Theropod tracks of uncertain affinity, different from Megalosauripus in the larger metatarsophalangeal region  |
Megalosauripus[15][9] |

* M. ispp.
* cf.M. ispp.

* El Mers
* Tasra
* Tasra Westbank
* Tassmante O'Moche
* Oued Tamghilt

| Footprints  | Traces of theropods; members of the ichnofamily Eubrontidae, incertae sedis within Theropoda |
Averostra[10][15] |

* Indeterminate

* Oued Botane

| Isolated Teeth  | Isolated teeth of unidentified or undescribed/studied theropods. Teeth associated with the Teleosaur "Megalosaurus" mersensis have recently been suggesting to come from indeterminate Averostrans. Other remains need revision.[16][17] |
Theropodipedia[9][18][15] |

* Indeterminate

* El Mers
* Inzar O'Founass
* Ifri N'Tfrane
* Laach O'Medda
* Oulad Ali
* Tasra
* Tasra Westbank
* Tassmante O'Moche

| Footprints  | Theropod tracks of uncertain affinity, suggested to come from "allosaurs or megalosaurs"[18] |

#### Sauropoda

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Breviparopus[15][9] |

* B. isp.

* El Mers
* Oued Tamghilt

| Footprints  | Traces of sauropods typical of the ichnofamily Parabrontopodidae, incertae sedis within Sauropodomorpha. Includes traces with pes similar to Diplodocoidea.[15] |

Cetiosaurus[10][19]

* "C." mogrebiensis

* Aîn ou N'Jourh
* Taghrout
* Tamguert r'Tane
* Tamguert n'Tarit
* Tich Niouine
* Oued Botane

Complete skeleton and referred co-types of partially complete and incomplete specimens

A eusauropod of the family Cetiosauridae, currently awaiting revision and likely not a member of the genus Cetiosaurus.[20]

Eusauropoda[21][22] |

* Indeterminate

* Aït Tamjout
* Boulahfa
* 1 km east of Taguelft

| More than 45 pieces: 2 long femora, large vertebrae and other remains  | The remains include gigantic bones of an animal of very large dimensions: a femur measures 236 cm.[22] |
Sauropodina[9][18][23] |

* Indeterminate

* El Mers
* Tasra
* Tasra Westbank
* Oulad Ali

| Footprints  | Sauropod tracks of uncertain affinity. There are both small and enormous sauropod tracks, including some among the largest known worldwide, with a pes track length of up to 130 cm.[9] |

Turiasauria[24]

* Indeterminate

* Boulahfa

USMBA 002–004, three isolated teeth

Teeth suggesting the oldest known African turiasaur

#### Ornithischia

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Adratiklit[25] |

* A. boulahfa

* Boulahfa

| NHMUK PV R37366 (A dorsal vertebra), three cervical vertebrae (NHMUK PV R37367; R37368), a dorsal vertebra (NHMUK PV R37365), and a left humerus (NHMUK PV R37007)  | A dacentrurine stegosaur  | Adratiklit boulahfa Spicomellus afer Thyreosaurus atlasicus

* Indeterminate

* Boulahfa

| USMBA 001, a partial left femur  | The oldest known cerapodan dinosaur
Ornithopodichnus?[9] |

* O.? ispp.

* Ifri N'Tfrane

| Footprints  | Ornithichian tracks, referred to short-toed forms such as Gilmoreosaurus
Spicomellus[26] |

* S. afer

* Boulahfa

| NHMUK PV R37412, a partial rib bearing four co-ossified spines; USMBA 84, a partial skeleton including dermal armour.[27] | An early ankylosaurian
Thyreosaurus[28] |

* T. atlasicus

* Boulahfa North

| HIIUC-BN00, a partial postcranial skeleton that includes nine dorsal vertebrae and 21 dorsal rib remains, a limb bone (fibula?), and six dermal elements  | A dacentrurine stegosaur with unusual recumbent osteoderms

### Dasycladales

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Heteroporella[5] |

* H. lusitanica

* SW part of the Skoura syncline

| Calcareous imprints  | A dasycladacean algae of the family Dasycladaceae |
Sarfiatella[5] |

* S. dubari

* SW part of the Skoura syncline

| Calcareous imprints  | A dasycladacean algae, associated in adjacent regions with Aalenian–Bajocian faunas  |

### Charophyta

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Porochara[5] |

* P. hians

* Ait Hammou

| Oogonia  | A characean algae of the family Porocharaceae; charophytes are abundant in lower levels, associated with freshwater ostracods  | Example of modern characean algae

### Plantae

Genus  | Species  | Stratigraphic position  | Material  | Notes  | Images

|
|
|
|
|

Araucariacites[1] |

* A. australis

* FD section, Skoura Syncline

| Pollen  | Affinities with Araucariaceae inside Coniferae.  | Example of extant Araucaria cones
Callialasporites[1] |

* C. dampieri
* C. segmentatus
* C. turbatus
* C. spp.

* FD section, Skoura Syncline

| Pollen  | Affinities with Araucariaceae inside Coniferae.
Classopollis[1] |

* C. (Corollina) spp.

* FD section, Skoura Syncline

| Pollen  | Affinities with Cheirolepidiaceae inside Coniferae.  |
Cyathidites[1] |

* C. ssp.

* FD section, Skoura Syncline

| Spores  | Affinities with the family Cyatheaceae inside Cyatheales. Arboreal Fern Spores  | Modern Cyathea, Cyathidites come probably from similar genera
Densoisporites[1] |

* D. spp.

* FD section, Skoura Syncline

| Spores  | Affinities with the Selaginellaceae in the Lycopsida.  | Modern Selaginella. Densoisporites probably come from a similar Plant
Lycopodiumsporites[1] |

* L. austroclavatidites
* L. spp.

* FD section, Skoura Syncline

| Spores  | Affinities with the family Lycopodiaceae in the Lycopodiopsida. Lycopod spores, related to herbaceous flora common in humid environments.  | Modern Lycopodium. Lycopodiumsporites probably come from a similar plant
Protocupressinoxylon[29] |

* P. choubertii

* Skoura syncline

| Fossil wood  | Conifer wood of the family Hirmeriellaceae, considered to be a potential synonym of the genus Brachyoxylon | Example of petrified wood trunk of the same genus (from an unknown location)
Protopodocarpoxylon[30] |

* P. teixeirae

* Skoura syncline

| Fossil wood  | Conifer fossil wood of the family Hirmeriellaceae  |

## See also

* Tafraout Group (Toarcian–Aalenian)
* Azilal Formation (Toarcian-Aalenian)
* Guettioua Formation (Bathonian–?Callovian)
* Anoual Formation (Bathonian)
* Ksar Metlili Formation (Tithonian–?Berrasian)

## References

1. ↑ 1.00 1.01 1.02 1.03 1.04 1.05 1.06 1.07 1.08 1.09 1.10 1.11 1.12 1.13 1.14 1.15 1.16 1.17 1.18 1.19 1.20 1.21 1.22 Khaffou, Hanane; Hssaida, Touria; Oukassou, Mostafa; Maatouf, Wafaa; Afenzar, Abdelkrim; Zafaty, Omar; Chakir, Sara; Jaydawi, Soukaina et al. (2023). "Bajocian–Bathonian dinoflagellate cyst assemblages from the Middle Atlas, Morocco: Palynostratigraphic and paleoenvironmental implications". Review of Palaeobotany and Palynology 312. doi:10.1016/j.revpalbo.2023.104862. ISSN 0034-6667.
2. ↑ Oukassou, M.; Boumir, Kh.; Benshili, Kh.; Ouarhache, D.; Lagnaoui, A.; Charrière, A. (June 2019). "The Tichoukt Massif: a Geotouristic Play in the Folded Middle Atlas (Morocco)". Geoheritage 11 (2): 371–379. doi:10.1007/s12371-018-0287-y. ISSN 1867-2477. Bibcode: 2019Geohe..11..371O.
3. ↑ Soufiani, N.; Fedan, B. (2002). "Les dépôts bathoniens du synclinal d'El Mers (Moyen Atlas, Maroc): unités formationnelles, analyse et cadre géodynamique". Bulletin de l'Institut scientifique 24 (6): 1–14. http://www.israbat.ac.ma/wp-content/uploads/2015/03/01- Soufiani et al. (01-14).pdf.
4. ↑ 4.00 4.01 4.02 4.03 4.04 4.05 4.06 4.07 4.08 4.09 4.10 4.11 4.12 4.13 4.14 4.15 Dresnay, Renaud du (1963-01-01). "Donnees stratigraphiques complementaires sur le Jurassique moyen des synclinaux d'El Mers et de Skoura (Moyen-Atlas, Maroc)". Bulletin de la Société Géologique de France S7-V (6): 883–900. doi:10.2113/gssgfbull.S7-V.6.883. ISSN 0037-9409. https://pubs.geoscienceworld.org/sgf/bsgf/article-abstract/S7-V/6/883/518424/Donnees-stratigraphiques-complementaires-sur-le.
5. ↑ 5.00 5.01 5.02 5.03 5.04 5.05 5.06 5.07 5.08 5.09 5.10 5.11 5.12 5.13 5.14 Charrière, André; Dépêche, Françoise; Feist, Monique; Grambast-Fessard, Nicole; Jaffrezo, Michel; Peybernès, Bernard; Ramalho, Miguel (1994). "Microfaunes, microflores et paléoenvironnements successifs dans la formation d'El Mers (Bathonien-?Callovien) du synclinal de Skoura (Moyen-Atlas, Maroc)". Geobios 27 (2): 157–174. doi:10.1016/s0016-6995(94)80002-2. ISSN 0016-6995. Bibcode: 1994Geobi..27..157C.
6. ↑ 6.0 6.1 6.2 6.3 6.4 Rousselle, L. (1965). "Rhynchonellidae, Terebratulidae et Zeilleriidae du Dogger marocain (Moyen-Atlas septentrional, Hauts-plateaux, Haut-Atlas).". Notes Mém. Serv. géol. Maroc, Rabat 187: 1-168. https://www.sudoc.abes.fr/cbs/DB=2.1/SRCH?IKT=12&TRM=281640688.
7. ↑ 7.0 7.1 7.2 7.3 7.4 7.5 Alméras, Y.; Elmi, S.; Mekahli, L.; Ouali-Mehadjii, A.; Sadki, D.; Tlili, M. (1994). "Biostratigraphie des Brachiopodes du Jurassique moyen dans le domaine atlasique (Maroc, Algérie). Contraintes environnementales et relation avec l'évolution verticale des peuplements d'ammonites.". Miscellanea del Servizio Geologico Nazionale 5 (5): 219-241. https://www.researchgate.net/profile/Abdelkader-Ouali-Mehadji/publication/273118588_Biostratigraphie_des_Brachiopodes_du_Jurassique_moyen_dans_le_domaine_atlasique_Maroc_Algerie_Contraintes_environnementales_et_relation_avec_l'evolution_verticale_des_peuplements_d'ammonites/links/54f70afa0cf2ccffe9d95da8/Biostratigraphie-des-Brachiopodes-du-Jurassique-moyen-dans-le-domaine-atlasique-Maroc-Algerie-Contraintes-environnementales-et-relation-avec-levolution-verticale-des-peuplements-dammonites.pdf.
8. ↑ Oukassou, Mostafa; Charrière, André; Lagnaoui, Abdelouahed; Gibb, Stacey; Michard, André; Saddiqi, Omar (April 2016). "First occurrence of the Ichnogenus Selenichnites from the Middle Jurassic Strata of the Skoura Syncline (Middle Atlas, Morocco); Palaeoecological and palaeoenvironmental context". Comptes Rendus Palevol 15 (5): 461–471. doi:10.1016/j.crpv.2015.09.013. Bibcode: 2016CRPal..15..461O.
9. ↑ 9.00 9.01 9.02 9.03 9.04 9.05 9.06 9.07 9.08 9.09 9.10 Amzil, Mustapha; Oukassou, Mostafa; Lallensack, Jens N.; Klein, Hendrik; Zafaty, Omar; Saber, Hafid; Charrière, André; Meyer, Christian et al. (2024). "New dinosaur tracks from the Middle Jurassic red beds of the Middle Atlas (Morocco): Application of photogrammetry to ichnology and conservation of geological heritage". Proceedings of the Geologists' Association 22 (3): 12–31. doi:10.1016/j.pgeola.2024.06.004. https://www.sciencedirect.com/science/article/pii/S0016787824000373.
10. ↑ 10.00 10.01 10.02 10.03 10.04 10.05 10.06 10.07 10.08 10.09 10.10 Lapparent, A. F. d. (1955). "Étude paléontologique des vertébrés du Jurassique d'El Mers (Moyen Atlas) [Paleontological study of the Jurassic veretebrates from El Mers (Middle Atlas)"]. Notes et Mémoires du Service Géologique du Maroc 124 (1): 1–36. https://search.worldcat.org/es/title/Etude-paleontologique-des-vertebres-du-jurrassique-d'El-Mers-(Moyen-Atlas)/oclc/17293965.
11. ↑ 11.0 11.1 11.2 11.3 11.4 11.5 Termier, H.; Gubler, J.; Lapparent, A. F. (1940). "Reptiles et Poissons du Bathonien d'El-Mers (Moyen-Atlas marocain) [Reptiles and fishes from the Bathonian of El Mers (Moroccan Middle Atlas)."]. Comptes Rendus de l'Académie des Sciences de Paris 210 (3): 768–770. https://naturalhistory.si.edu/sites/default/files/media/translated_publications/Termier&_40a.pdf.
12. ↑ 12.0 12.1 12.2 Mongin, D. (1967). "Les Mollusques du Bathonien saumâtre du Moyen Atlas.". Notes et mémoires du Service Géologique du Maroc 200 (1): 39-95. https://www.marinespecies.org/aphia.php?p=sourcedetails&id=427877.
13. ↑ Beltan, Laurence (1957-01-01). "Etude d'un neurocrane de Lepidotes du Bathonien du Maroc". Bulletin de la Société Géologique de France S6-VII (8–9): 1091–1106. doi:10.2113/gssgfbull.s6-vii.8-9.1091. ISSN 1777-5817.
14. ↑ Chabli, S. (1986). "Données nouvelles sur un «Dinosaurien» jurassique moyen du Maroc: Megalosaurus mersensis Lapparent 1955, et sur les Megalasaurides en Sénégal". Les dinosaures de la Chine à la France 1 (2): 65–72. https://pascal-francis.inist.fr/vibad/index.php?action=getRecordDetail&idt=8370587.
15. ↑ 15.0 15.1 15.2 15.3 15.4 Hadri, M.; Lorente, F. P. (2012). "Historia de yacimientos con huellas de dinosaurio, desde su descubrimiento hasta su primer estudio: alrededores de El Mers (Marruecos)". Zubia 30 (6): 93–147. https://dialnet.unirioja.es/descarga/articulo/4118092.pdf.
16. ↑ "Non-theropods". https://theropoddatabase.github.io/Non-theropods.htm#Megalosaurusmersensis.
17. ↑ "Neotheropoda". https://theropoddatabase.github.io/Neotheropoda.htm#Averostra.
18. ↑ 18.0 18.1 18.2 Jenny, Jacques; Le Marrec, Alain; Monbaron, Michel (1981). "Les empreintes de pas de Dinosauriens dans le Jurassique moyen du Haut Atlas central (Maroc): Nouveaux gisements et precisions stratigraphiques". Geobios 14 (3): 427–431. doi:10.1016/s0016-6995(81)80186-6. ISSN 0016-6995. Bibcode: 1981Geobi..14..427J.
19. ↑ Monbaron, Michel (1983). "Dinosauriens du Haut-Atlas Central (Maroc): état des recherches et précisions sur la découverte d’un squelette complet de grand Cetiosaure [Dinosaurs of the Central High Atlas (Marocco) : state of research and details of discovery of complete skeleton of large Cetiosaur"]. Actes de la Société jurassienne d'émulation 86: 203. doi:10.5169/seals-684897. ISSN 1011-2820. https://www.e-periodica.ch/digbib/view?pid=asj-006:1983:86::288.
20. ↑ Läng, E. (2008). "Les Cétiosaures (Dinosaura, sauropoda) et les sauropodes du Jurassique moyen: revision systématique, nouvelles découvertes et implications phylogénétiques". (Doctoral dissertation, Paris, Muséum national d'histoire naturelle): 1–223.
21. ↑ Termier, Henri (1942-01-01). "Donnees nouvelles sur le Jurassique rouge a dinosauriens du Grand et du Moyen-Atlas (Maroc)". Bulletin de la Société Géologique de France S5-XII (4–6): 199–207. doi:10.2113/gssgfbull.s5-xii.4-6.199. ISSN 1777-5817.
22. ↑ 22.0 22.1 Charroud, M.; Fedan, B. (1992). "Données préliminaires sur la découverte du gisement de Boulahfa à dinosauriens (SW de Boulemane, Moyen Atlas central)". Notes et Mémoire du Service géologique du Maroc 366 (5): 448–449. https://naturalhistory.si.edu/sites/default/files/media/translated_publications/Charroud&Fedan_92.pdf.
23. ↑ Meyer, C. A.; Thüring, B. (2005). "Mind the Middle Jurassicgap-bone versus track record in dinosaurs". Abstracts 3rd Swiss Geoscience Meeting Zurich 3 (1): 59–60. https://citeseerx.ist.psu.edu/document?repid=rep1&type=pdf&doi=a981e7967716fc9a9e1cb27c47ad5ca308a1efae.
24. ↑ Woodruff, D. C.; Barrett, P. M.; Ouarhache, D.; El Khanchoufi, A.; Boumir, K.; Ech-Charay, K.; Oussou, A.; Butler, R. J. et al. (2025). "Teeth from the Middle Jurassic of Morocco reveal the oldest turiasaurian sauropods from Africa". Acta Palaeontologica Polonica 70 (3): 411–420. doi:10.4202/app.01214.2024. https://app.pan.pl/article/item/app012142024.html.
25. ↑ Maidment, Susannah C.R.; Raven, Thomas J.; Ouarhache, Driss; Barrett, Paul M. (January 2020). "North Africa's first stegosaur: Implications for Gondwanan thyreophoran dinosaur diversity". Gondwana Research 77: 82–97. doi:10.1016/j.gr.2019.07.007. Bibcode: 2020GondR..77...82M.
26. ↑ Maidment, Susannah C. R.; Strachan, Sarah J.; Ouarhache, Driss; Scheyer, Torsten M.; Brown, Emily E.; Fernandez, Vincent; Johanson, Zerina; Raven, Thomas J. et al. (2021-09-23). "Bizarre dermal armour suggests the first African ankylosaur" (in en). Nature Ecology & Evolution 5 (12): 1576–1581. doi:10.1038/s41559-021-01553-6. ISSN 2397-334X. PMID 34556830. Bibcode: 2021NatEE...5.1576M. https://www.nature.com/articles/s41559-021-01553-6.
27. ↑ Maidment, Susannah C. R.; Ouarhache, Driss; Ech-charay, Kawtar; Oussou, Ahmed; Boumir, Khadija; El Khanchoufi, Abdessalam; Park, Alison; Meade, Luke E. et al. (2025-08-27). "Extreme armour in the world's oldest ankylosaur" (in en). Nature: 1–6. doi:10.1038/s41586-025-09453-6. ISSN 1476-4687. https://www.nature.com/articles/s41586-025-09453-6.
28. ↑ Zafaty, O.; Oukassou, M.; Riguetti, F.; ((Company)), J.; Bendrioua, S.; Tabuce, R.; Charrière, A.; Pereda-Suberbiola, X. (2024). "A new stegosaurian dinosaur (Ornithischia: Thyreophora) with a remarkable dermal armour from the Middle Jurassic of North Africa". Gondwana Research 131 (In press): 344–362. doi:10.1016/j.gr.2024.03.009. Bibcode: 2024GondR.131..344Z.
29. ↑ Philippe, M.; Bamford, M.; McLoughlin, S.; Alves, L.S.R.; Falcon-Lang, H.J.; Gnaedinger, S.; Ottone, E.G.; Pole, M. et al. (2004). "Biogeographic analysis of Jurassic–Early Cretaceous wood assemblages from Gondwana". Review of Palaeobotany and Palynology 129 (3): 141–173. doi:10.1016/j.revpalbo.2004.01.005. ISSN 0034-6667. Bibcode: 2004RPaPa.129..141P.
30. ↑ du Dresnay, R. (1975-01-01). "Donnees topographiques, stratigraphiques et paleontologiques concernant une ammonite citee dans la Formation d'El-Mers (Moyen-Atlas, Maroc), et leurs consequences sur l'age attribue a cette formation". Bulletin de la Société Géologique de France S7-XVII (6): 1144–1146. doi:10.2113/gssgfbull.S7-XVII.6.1144. ISSN 0037-9409.

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