Cyrtocrinids (Cyrtocrinida,Crinoidea) and other associated crinoids from the Jurassic (Kimmeridgian–Tithonian)–Cretaceous (Berriasian–Barremian) of the Carpathian Foredeep basement (western Ukraine)

Upper Jurassic (Kimmeridgian–Tithonian) and Cretaceous (Berriasian–Barremian) strata of the Ukrainian part of the Carpathian Foredeep basement are rich, at least locally, in crinoid remains. Crinoids belonging to cyrtocrinids (Cyrtocrinida) are represented by whole cups, isolated remains of disarticulated cups, brachial plates and columnals. They are assigned to the following taxa: Cyrtocrinida indet., Eugeniacrinites cf. cariophilites (von Schlotheim), Lonchocrinus sp., Phyllocrinus stellaris Zaręczny, Ascidicrinus pentagonus (Jaekel), Gammarocrinites sp., Psalidocrinus armatus (Zittel), Psalidocrinus sp., and Hemibrachiocrinidae gen. indet. Cyrtocrinids are associated with other stalked (isocrinids, Isocrinida and millericrinids, Millericrinida) and stemless (saccocomids, Roveacrinida) crinoids. Columnals, pluricolumnals, brachial plates, and cirrals of isocrinids are assigned to Balanocrinus sp., Isocrinina fam. et subfam. indet., and columnals of millericrinids to Millericrinida indet. Free-living roveacrinids are assigned to Saccocoma sp. and Crassicoma sp. Knowledge on Jurassic and Cretaceous crinoids formerly described from Ukraine is discussed. Although majority of crinoids described herein seems to be allochthonous, autochthonous forms were also found with certainty in some intervals. These include some cyrtocrinids, which dominate in shallow-water environments of the Ukrainian Carpathian Foredeep basement. Isocrinids are also common in this shallow marine environment, whereas sessile saccocomids are assigned to low-energy, mud-supported bottom, open marine, outer-platform/upper slope, and relatively deep environments.     

First record of non-marine ostracods from the Paleogene “hamadian deposits” of Méridja area,west of Bechar (southwestern Algeria)

A new non-marine ostracod fauna from the Paleogene “hamadian deposits” outcropping west of Bechar (southwestern Algeria) has been recovered from lacustrine to fluvial deposits of the Oued Méridja section and fluvial deposits on the southern edge of the Hamada de Méridja section. Recently, these sections have been dated as late Thanetian – early Ypresian (latest Paleocene to earliest Eocene) and Ypresian – earliest Lutetian (early to earliest middle Eocene), respectively, based on charophytes. The associated ostracod fauna recovered consists of relatively mostly moderately to badly preserved specimens and comprises 14 taxa, none of which could be identified to species level in view of its poor state of preservation; we have nevertheless been able to identify and describe the following taxa: Herpetocypris sp., Cyprinotus? sp., Heterocypris? sp. 1 and sp. 2, Cypris? sp., Ilyocypris sp., Cytheroidea indet. sp. 1 and sp. 2, Limnocytheridae indet. sp. 1, Cypridoidea indet. sp. 1, Cyprididae indet. sp. 1, and Ostracoda indet. sp. 1, 2 and 3. Only Heterocypris sp. 1 occurs in both sections. Although the fauna can as yet not be related to the few other contemporaneous faunas reported from the wider palaeogeographic area, it adds important new information to our poor knowledge on Eocene non-marine ostracods in North Africa and southern Europe. The Méridja sections and area are promising regarding the discovery of more, better preserved material and further studies, and one main limitation to the correlation of the fauna is the hitherto insufficient taxonomic knowledge on many faunal elements of Eocene non-marine ostracods to which our section contributes considerably.     

Late Cretaceous (Santonian-Campanian) sea lilies (Echinodermata,Crinoidea) from the glacial rafts of northwestern Poland

Seven Santonian (Isocrinus? minutus, Tetracrinus jagti, Cyathidium senessei, Bourgueticrinidae indet., Notocrinidae indet., Glenotremites paradoxus, Marsupites testudinarius) and seven Campanian (Nielsenicrinus carinatus, Austinocrinus sp., Bourgueticrinidae indet., Bourgueticrinus cf. constrictus, B. ellipticus, B.? suedicus, Roveacrinina gen. et sp. indet.) crinoid taxa from the Cretaceous glacial rafts of northwestern Poland (near Wolin Island) are described for the first time. The taphonomy and palaeogeographic implications are discussed. The present assemblage differs taxonomically from the crinoid faunules recently described from the glacial rafts of eastern Poland. However, these faunules are similar to crinoids from the autochthonous Upper Cretaceous sediments of southern and eastern Poland (Miechów Trough, Kraków-Cz?stochowa Upland, Middle Vistula River valley). An update of Late Cretaceous crinoids from the Miechów Trough and Kraków-Cz?stochowa Upland is also given.     

Hrabalicrinus zitti gen. et sp. nov., and other Upper Jurassic crinoids (Echinodermata,Crinoidea) from the Brno area (Czech Republic)

Jurassic (Oxfordian) crinoids from the Brno area (Czech Republic) are described in details for the first time. A rich crinoid assemblage consisting of cups, isolated cup elements, brachial plates, columnals, pluricolumnals, and cirrals is assigned to isocrinids Isocrinus amblyscalaris (Thurmann), Balanocrinus subteres (Münster), B. pentagonalis (Goldfuss), Isocrinida indet., comatulid Hrabalicrinus zitti gen. et sp. nov., cyrtocrinids Lonchocrinus sp., Cyrtocrinus cf. nutans (Goldfuss), Pilocrinus moussoni (Desor), Tetracrinus moniliformis Münster, Cyrtocrinida indet., and millericrinids (Millericrinida indet.). Crinoids already mentioned from the Jurassic strata of the Czech Republic are subsequently revised. Apart from the isolated remains of Isocrinida, Millericrinida, and Thiolliericrinida, the presence of any other taxon reported from this area should be treated with extreme caution. A rare example of non-regenerative columnal healing (the so-called callus) in I. amblyscalaris is also described. Based on sedimentology and microfacies, the Jurassic limestones were deposited in various palaeoenvironments of upper carbonate platform and shelf lagoon (0 to > 50 m palaeodepth).     

Early Jurassic (latest Toarcian) brachiopods from the northeastern margin of the Western Tethys (Central Iran) and their paleobiogeographical significance

Brachiopod fauna from central Iran, recorded in the upper part of the Shemshak Group and attributed to the upper Toarcian (Pseudoradiosa-Aalensis zones), are reported for the first time in Iran. The assemblage recognized includes six different taxa: Homoeorhynchia sepahanensis nov. sp., formally described in this paper, Globirhynchia subobsoleta, Pseudogibbirhynchia sp., Tetrarhynchiidae sp. indet., Monsardithyris? aff. haresfieldensis, and Zeilleria cf. leckenbyi. Analysis of faunal affinities with other paleobiogeographical regions shows a free connection of the central Iranian brachiopod fauna and wide areas of the northern shelf margin of the Tethys Ocean. This is due to an apparent disruption of bioprovinciality inferred for the late Toarcian-earliest Aalenian, congruent with a connection through the northern seaway across the peri-Laurasian epicontinental platforms.     

Le passage Cénomanien–Turonien dans le Nord-Ouest algérien (coupe de Douar Menkouchi,Ouarsenis) : biostratigraphie des radiolaires

The Ouarsenis Massif belongs to the Algerian Tell domain which is considered as the eastern part of the Maghrebian Tethys former margin. The Cenomanian-Turonian Douar Menkouchi section outcropping in the eastern side of the Ouarsenis massif is composed of marls and laminated limestone alternations, including massive limestone and a biosiliceous bed toward the upper part of the succession. It reveals the presence of well-preserved and diversified radiolarians. In this study, we have identified for the first time in the Algerian domain, age-diagnostic radiolarian assemblages characterizing the Cenomanian-Turonian transition (C/T). The organic-rich laminated beds (black shales) confined to Silvae Zone, Cenomanian in age and underlying a biosiliceous interval, have provided radiolarian assemblages dominated by Pseudodictyomitrae. The successions overlying the biosiliceous interval have provided diversified radiolarian assemblages including Hagiastridae (i,e. Crucella cachensis, Alievium cf. superbum) and abundant Dictymitrae taxa (i,e. Dictyomitra cf. napaensis and Dictyomitra undata) indicating early Turonian age. These marl-limestone alternations are assigned to the Alievium superbum Zone. Considering taphonomic and diagenetic implications on assemblages content, we conclude that organic-rich beds and biosiliceous interval outline the latest Cenomanian. The radiolarian assemblage composition exhibits similarities to microfauna recovered from Bonarelli Horizon in Central Italy. Morphological similarities are also deduced from assemblages containing Pseudodictyomitrae tiara morphotypes (synonym of P. nakasekoi) which resemble those recovered from ophiolitic intervals in Cyprus and Oman areas.     

Upper Triassic (Norian) hypercalcified sponges from the Musandam Peninsula (United Arab Emirates and Oman)

Supercalcified sponges, including sphinctozoans, inozoans, chaetetids, spongiomorphids, occurring in Upper Triassic (Norian-Rhaetian) shallow-marine limestones of Musandam Mountains in United Arab Emirates (UAE), are described. The following taxa were determined: sphinctozoans: Hajarispongia osmani Senowbari-Daryan and Yancey, Nevadathalamia arabica n. sp., Nevadathalamia conica n. sp., Fanthalamia milahaensis n. sp., Iranothalamia incrustans (Boiko), Cinnabaria regularis n. sp.; inozoans: Cavsonella triassica n. sp., Molengraaffia regularis Vinassa de Regny, Peronidella? sp., Circopora cf. caucasica Moiseev, Circopora? sp.; spongiomorphids: Spongiomorpha sp.; chaetetids: Lovcenipora chaetetiformis Vinassa de Regny, Lovcenipora musandamensis n. sp., Lovcenipora sp., chaetetid sponge gen. et sp. indet. The most abundant sponge in the studied material is Nevadathalamia arabica n. sp. The described sponge association of the Arabian shelf (Musandam Mountains) shows close affinity to the sponge association known from age-equivalent terranes in the Panthalassa Ocean (Sonora Mountains in Mexico; Pilot Mountains in Nevada, USA), but is remarkably different from sponge associations in carbonates bordering the Tethys. This difference goes along with the biogeography of wallowaconchid bivalves and is most likely attributed to climatic, palaeogeographic or oceanographic factors.     

Upper Cenomanian bivalves from the Guir Basin (southwestern Algeria): Order Veneroida

The Upper Cretaceous rocks are widely distributed and well exposed in south Algeria and consist in beds rich in macrofauna. For the first time, twenty veneroid species (Bivalvia) are systematically described from the upper Cenomanian deposits of the Guir Basin. While three species were reported since long before [Granocardium desvauxi (Coquand), G. productum (J. de C. Sowerby) and Glossus aquilinus (Coquand)]; Lucina fallax Forbes, Crassatella (Rochella) tenuicostata (Seguenza), Protocardia hillana (J. Sowerby), G. productum (J. de C. Sowerby) var. byzacenica (Pervinquière), Arctica cordata (Sharpe), A. humei (Cox), A. inornata (d’Orbigny), A. picteti (Coquand), Tenea delettrei (Coquand), Paraesa faba (J. de C. Sowerby), Meretrix desvauxi (Coquand) were previously unknown from the Cretaceous of Algerian Sahara. Because of reduced degree of preservation, Sphaera cf. corrugata J. Sowerby, Maghrebella cf. forgemoli (Coquand), Maghrebella sp., Granocardium cf. carolinum (d’Orbigny), Protocardia sp. and Meretrix sp. are tentatively determined. The studied material, found in the lower part of the “Calcaires de Sidi Mohamed Ben Bouziane” Formation, evidences palaeobiogeographic affinities occurring over a wide geographical area: from North Africa, southern Europe to Middle East and India. The present study provides new information to the knowledge of the upper Cenomanian palaeobiology of the studied region.     

New data on crinoid assemblages from the oldest sedimentary rocks of the Polish Outer Carpathians (Jurassic-Cretaceous)

Crinoids (Crinoidea) preserved in the so-called Cieszyn Beds (Tithonian-Hauterivian; southern Poland) are relatively numerous, especially in some levels, but not highly diversified. They are absent in the Lower Cieszyn Shale (Tithonian) and the Upper Cieszyn Shale (Valanginian-Hauterivian). The Lower Cieszyn Limestone (Tithonian) yielded crinoid remains classified as Isocrinus cf. amblyscalaris (Thurmann) and Isocrinida indet. In the Upper Cieszyn Limestone (Berriasian) crinoids are much more numerous and diversified. The following taxa are recorded: Isocrinus? annulatus (Roemer), Balanocrinus subteres (Münster), B. cf. smithi Hess and Gale, and Hemicrinus sp. The presence of specimens resembling Isocrinus amblyscalaris (Thurmann) in the Tithonian strata is surprising since the last occurrence of this taxon was reported from the Kimmeridgian. The Lower Cieszyn Limestone crinoidal assemblage is dominated by isocrinids (Isocrinida). This probably reflects the much shallower sedimentary environment of these sediments compared to that of the Upper Cieszyn Limestone, where excepting isocrinids, hemicrinids (Hemicrinidae, Cyrtocrinida) typical for deep marine environments were found. It is consistent with the analytical results from the foraminiferal assemblages, among which the benthic forms known from shelfal and paralic epicontinental environments first dominated, and then were supplanted by deep-sea taxa at the turn of the Jurassic and Cretaceous.     

Le genre leptaenopyxis (brachiopoda,leptaenidae) dans le Devonien du Massif Armoricain (France)

The study of genus LeptaenopyxisHavlicek, 1963 in the Lower Devonian from Armorican Massif allows to recognize four species: Leptaenopyxis kerfornei n. sp., Leptaenopyxis cf. bouei (Barrande), Leptaenopyxis sp. 1 and Leptaenopyxis sp. 2. These species were described, until now, under specific name boueiBarrande, 1848. From this study it follows some biostratigraphical and palaeobiogeographical results for the Armorican Massif and its relations with surrounding areas during Lower Devonian time.     

Upper Pliocene bivalve shell concentrations from the Lower Chelif basin (NW Algeria): Systematics,sedimentologic and taphonomic framework

Sedimentologic and palaeontological investigation of the Upper Pliocene Slama Formation in the Lower Chelif Basin (NW Algeria) led us to collect important bivalve assemblages for taxonomic and taphonomic purposes. A rather comprehensive inventory list of Upper Pliocene bivalves from northwestern Algeria is now available and consists of 30 species, 17 of which are extinct ones. Four principal taphonomic attributes were analysed: bioerosion, encrustation, fragmentation, and abrasion. Physical and biogenic sedimentary structures are used for palaeoenvironmental interpretations. The taphonomic, sedimentologic and ichnological characteristics of most of the deposits suggests they originated from discontinuous processes of winnowing and bypassing of sediments, probably due to the action of storms in shallow waters, mainly in the shoreface depositional environment. The bivalve assemblage is dominated by disarticulated valves and displays significant taphonomic alteration in the shells. Sclerobionts traces in shells particulary affect the oyster shells. Bioerosion traces are predominately those of clionid sponges (Entobia isp.), polychaetes (Maeandropolydora isp. and Caulostrepsis isp.), bivalves (Gastrochaenolites isp.), and of predatory gastropods (Oichnus isp.). Among the sclerobionts, the identified encrusters were juvenile oyster recruits, barnacles, polychaetes (serpulid tubeworms), bryozoans (Microporella sp. and Acanthodesia sp.), and vermetid gastropods (Petaloconchus intortus).     

Plicathyridine brachiopods (Athyridida) from the Frasnian (Late Devonian) of Western Europe and Middle East

Plicathyridine brachiopods (Athyridida) from the early–middle Frasnian of southern Belgium and northern France (Dinant Synclinorium) are systematically described for the first time. They include two species: Anathyris (Anathyris) calestiennensis nov. sp., and A. (A.) sp. indet. 1. They are uncommon in the mainly shally La Prée (Nismes Formation) and Ermitage (Moulin Liénaux Formation) members and are absent from the mixed argillaceous–carbonate late Frasnian succession in this area contrary to what is observed in Russia, notably in the East-European Platform and Siberia. In the Middle East, two species of Anathyris are recognized on the basis of a limited material from the Dascht-e-Nawar area in Afghanistan (A. (A.) sp. indet. 2) and the Kuh-e Kaftar mountains in Central Iran (A. (A.) sp. indet. 3). Anathyris (A.) calestiennensis nov. sp. is sometimes encrusted by epizoans (tabulate and rugose corals, and bryozoans) and rarely displays single, small circular drill holes. The past and current epizoan–brachiopod interactions are also discussed (Anathyris vs. Lingula, respectively).     

Échinodermes de l'Ordovicien supérieur (Ashgill) de Sardaigne et d'Algérie

Research on the influence of sea level variations on the benthic faunas have been carried out in the Upper Ordovician of Sardinia. Study of the depositional facies and sequence analysis of the upper part (Lower Ashgill) of the Portixeddu Formation led to the identification of the sedimentary environments. Cystoids and crinoids are associated to bryozoans and brachiopods in most levels. The numerical analysis of associations and megaguilds shows that crinoids and cystoids have a higher frequency in the proximal and median facies of the upper offshore. The columnal association characterized by Conspectocrinus celticus and the coronoid Mespilocystites tregarvanicus has been discovered in the upper part of the formation. This material and complementary samples from Upper Ordovician of Sardinia and Kabylia (Algeria) bring additional data on the systematic and show the wide distribution of this fauna outside of the Ibero-armorican domain. The distribution of this echinoderm association supports a palaeogeographical position of the Ibero-armorican domain and Sardinia within the north gondwanan margin during the Lower Palaeozoic.     

Microvertebrates from unit G/layer 17 of the archaeological site of Hummal (El Kowm,Central Syria): Preliminary results

We present a first look at the microvertebrate fauna of the Middle Pleistocene site of Hummal in Central Syria. Some 2000 microvertebrate remains (1200 mammalian; 230 reptilian; 600 unidentified) were found in unit G/layer 17 by screen-washing sediments in an area of 4 m². The following taxa have been identified: Reptilia: Agaminae indet., Gekkota indet., Lacertidae indet. (2–3 taxa), Eryx sp., Natricinae indet.; and Mammalia: Crocidurinae indet., Chiroptera indet., Lepus sp., Arvicolinae indet., Ellobius sp., Microtus sp., Murinae indet. (large form), Mus sp., Meriones sp., Gerbillus sp. The presence of Ellobius indicates a Middle Pleistocene age of the fauna. This genus does not occur in Syria today, but is recorded in Israel in Late Acheulean to Early Mousterian sites. In North Africa, the occurrence of this genus is restricted to the early part of the Middle Pleistocene. The ecological requirements of the nearest living relatives of the recorded taxa indicate mainly open habitats, but also the presence of vegetation and wet conditions, at least close to the site. Lepus, Ellobius, Meriones, Gerbillus, and Eryx live to various extents in steppes, semi-deserts, and deserts. Various extant species of large murids that come into consideration occupy a variety of wooded habitats, grassland and savannah and require the presence of water. Hummal offers the most detailed Pleistocene portrait yet of hominin palaeoenvironment in an area that, at the end of the Pleistocene, became known as part of the Fertile Crescent.     

Rodents, insectivores and chiropterans (Mammalia) from the late Aragonian of Can Missert (Middle Miocene, Vallès-Penedès Basin, Spain)

In this paper, the insectivores, chiropterans and rodents from the middle Miocene site of Can Missert are described. The faunal list of this locality includes the following species: Miosorex grivensis, Desmanella sp., Talpidae indet., Vespertilionidae indet., Hispanomys daamsi, Megacricetodon minor, Megacricetodon ibericus, Fahlbuschia crusafonti, Democricetodon brevis nemoralis, Eumyarion medium, Muscardinus hispanicus, Eomuscardinus sp., Paraglirulus werenfelsi and Spermophilinus bredai. H. daamsi is a new Cricetodontine species which is characterized by complete ectolophs and relatively long third lower molars. The rodent association of Can Missert enable one to assign this locality to the late Aragonian, MN 8, being close in age to other localities in the Vallès-Penedès Basin such as Castell de Barberà. However, the proportion in which each species is represented is very different in the two cases. This evidence points to the existence climatic pulses at the end of the Aragonian Mammal-Stage.     

Early Permian (Asselian) brachiopods from Karakorum (Pakistan) and their palaeobiogeographical significance

Abstract: Early Permian (Asselian) brachiopods collected from the Gircha Formation of western Karakorum (Pakistan) are described. They include Bandoproductus girchensis sp. nov., Kiangsiella sp. indet., Trigonotreta lyonsensis Archbold and Thomas, Trigonotreta larghii sp. nov., Spirelytha petaliformis (Pavlova), Punctospirifer afghanus Termier, Termier, de Lapparent and Marin, and ?Dielasma sp. indet. and belong to the Trigonotreta lyonsensis–Punctospirifer afghanus Assemblage Biozone, the oldest so far recovered from the Permian succession of Karakorum. The faunal succession of Karakorum records a significant biotic change from the Asselian to the Sakmarian, a shift in diversity and composition that is also recorded along most of the Gondwanan margin and Peri‐Gondwanan regions and that should be related to a major climatic change: the end of the Gondwanan glaciation. A palaeobiogeographical analysis has been performed by means of multivariate methods applying cluster and ordination analyses based on the Jaccard Coefficient and Simpson Index to a matrix consisting of the presence/absence of 23 brachiopod genera from seven geographical operational units from central Afghanistan to eastern Australia. The results suggest the occurrence of a single biotic province during the Asselian, the Indoralian Province, embracing all the faunal stations examined, as a consequence of the global cold phase related to the last pulse of the Gondwanan glaciation.     

The latest Early Pleistocene amphibians and reptiles from Kaiafas (Greece) and the first record of fossil Ophiomorus (Squamata,Scincidae)

Together with the rest of the Balkan Peninsula, Greece acted as a refuge for thermophilic amphibians and reptiles during the Quaternary Glaciations. In this work we provide new herpetological data on the latest Early Pleistocene locality of Kaiafas, in western Peloponnese. The site yielded one indeterminate salamandrid urodelan (Salamandridae indet.), one tree frog (Hyla gr. H. arborea), one indeterminate testudinid (Testudinidae indet.), at least two lacertid lizards (Lacertidae indet. A and B), one scincid lizard (Ophiomorus sp.), one anguid lizard (Pseudopus sp.), and one indeterminate “colubrine” snake (“Colubrinae” indet.). This assemblage is indicative of a surrounding environment with water bodies, wooded areas, and possibly loose soils with rocks and stones. In spite of being based on few remains, the study of the amphibians and reptiles from Kaiafas resulted in the identification of the first fossil of Ophiomorus known worldwide. Moreover, the two lacertid taxa displayed very peculiar morphological features which are either previously unknown or rare in that clade.     

Biostratigraphy and Paleobiogeographic implications of the Cenomanian–Early Turonian Ostracods of Egypt

The present study deals with the Cenomanian–early Turonian ostracods in the southern Sinai, Egypt. The investigated sequence includes the Raha and Abu Qada formations. The Raha Formation consists of distinctive Cenomanian taxa such as Cytherella aegyptiensis, Veeniacythereis maghrebensis, V. jezzineensis, V. streblolophata streblolophata, Monoceratina trituberculata, and Glenocythere reticulata. Based on the ostracod species, four local biozones are recognized, including Veeniacythereis jezzineensis – Metacythereopteron berbericum, Cytherella eosulcata – Xestoleberis derorimensis, Amphicytherura ziregensis and Perissocytheridea istriana – Cythereis fahrioni. The paleobiogeography of the considered ostracods suggests that there are two bioprovinces, the first of which is the North African province and includes Egypt, Tunisia and Algeria. The second bioprovince represents the Middle East province and includes Lebanon, Oman and Iran. The strong resemblance between the two bioprovince indicates a relatively good communication along the margin of Southern Tethys during the Cenomanian and improved marine ostracod exchange.     

Famennian (Late Devonian) ammonoids from the Ouarourout section (Saoura Valley,Algeria)

In North Africa, the ammonoids constitute an important part of the Devonian marine macro-invertebrates. New material has been recently collected in the ‘Argiles de Marhouma’ Formation from several sections located near Béni-Abbès in the Saoura Valley (Ougarta Mountains, Algerian Sahara). Red nodular limestones (i.e., ‘griotte’ facies) characterize this formation rich in ammonoids. The assemblages from the Ouarourout section are composed of Goniatitida and Clymeniida constituting 30 species and 19 genera belonging to 11 families, with Cyrtoclymenia, Erfoudites, Kosmoclymenia, Planitornoceras, Platyclymenia, and Prionoceras, as the most abundant genera. The taxonomic comparison and the biostratigraphic correlations are made by considering the assemblages recorded in the Anti-Atlas of Morocco, which are very close to those observed in Algeria. In the Ouarourout area, the ‘Argiles de Marhouma’ Formation contains middle and late Famennian ammonoid assemblages and three biostratigraphic intervals can be identified: Planitornoceras, Platyclymenia and Medioclymenia.