Ahl Al Oughlam Research Articles

Palaeoecological reconstruction of Plio-Pleistocene herbivores from the Ahl al Oughlam site (Casablanca, Morocco): Insights from dental wear and stable isotopes

In this article, we use a multiproxy approach based on stable isotope analyses (δ18O and δ13C), mesowear, and microwear dental analysis to reconstruct the climate, diet, and habitat of Plio-Pleistocene herbivores at the site of Ahl al Oughlam in western Morocco. This study has been conducted on teeth from several taxa (Sivatherium maurusium, Hipparion pomeli, Metridiochoerus phacochoeroides, Gazella sp., Alcelaphini and Anancus osiris), enabling a robust ecological reconstruction. Our isotopic results suggest a predominance of C3 plant consumption for all analyzed species, while dental wear indicates a varied diet, with evidence for both browsing and grazing behavior being present. The combination of proxies indicates a landscape dominated by open C3 grasslands in an arid climate, as supported by the Mean Annual Precipitation (MAP) results. These findings contribute to the ongoing debate on the ecological context that facilitated the Plio-Pleistocene dispersal, occupation, and evolution of mammals and hominins in northern Africa, for which limited information is available.

Les Tortues Du Pliocène d'Ahl al Oughlam (Casablanca, Maroc) et De Localités Mio-Pliocènes Avoisinantes

Turtles from the Pliocene of Ahl al Oughlam (Casablanca, Morocco) and nearby Mio-Pliocene localities.The Late Pliocene locality Ahl al Oughlam (Casablanca, Morocco) yielded a fauna of cryptodiran testudinoid turtles, all of them related to extant African forms. Two terrestrial and one freshwater species are recognized. First, a new small species of the palaearctic genus Testudo Linnaeus, 1758, T. oughlamensis n. sp., is abundantly represented and is closely related to T. kenitrensis Gmira, 1993, from the Middle Pleistocene of Kenitra, Morocco. It is compared to other new remains of Testudo from nearby localities of similar age, including Dehar Mouak and Ar Rehamna. Together with T. kenitrensis, it constitutes a new clade within Testudo, providing a minimum age for the divergence between this clade and T. graeca Linnaeus, 1758 from Morocco. Secondly, the giant ?Centrochelys marocana n. sp., is probably related to the extant C. sulcata (Miller, 1779). Its description gives the opportunity to examine the earliest member of this genus in Morocco, from the nearby Late Miocene/Early Pliocene locality of Lissasfa. Comparisons with Miocene to extant genera of large size from Europe and northern Africa are made. The freshwater species from Ahl al Oughlam is close to the extant species Mauremys leprosa (Schweigger, 1812). It is the earliest record in Morocco of this second palaearctic genus. Thus, the fossil turtle fauna of Casablanca demonstrates the antiquity in Morocco of the three extant genera. Their geographical origin is considered.

A revision of the fossil Canidae (Mammalia) of north‐western Africa

Abstract: The fossil record of the Canidae in North‐western Africa begins near the Miocene–Pliocene boundary with a form close to Nyctereutes, a genus best known in the late Pliocene of Ahl al Oughlam. This site yields two other canids. Vulpes hassani sp. nov. is a small fox, probably ancestral to the modern V. rueppelli, recorded from the Middle Pleistocene onwards. Lupulella paralius sp. nov. is a primitive jackal that probably belongs to the clade of modern African jackals. In the middle Pleistocene, the most common canid is Lupulella mohibi sp. nov., remarkable by its Nyctereutes‐like dentition and primitive skull‐features. These are all endemic forms, but V. vulpes and C. aureus, of northern origin, appear in the course of the middle Pleistocene. Lycaon has a sparse record in the middle and late Pleistocene.

Biochronologie mammalienne du Quaternaire du Maroc atlantique, dans son cadre régional

Recent researches conducted in Morocco in the frame of the “Programme Casablanca” provide a firm ground to set up a biochronology of the Pliocene and Pleistocene based upon the succession of Mammalian faunas, thanks to the occurrence of well-dated landmarks, and despite very scarce absolute datations. The site of Lissasfa, dated to the Miocene-Pliocene boundary by rodents, testifies to important exchanges with Europe. That of Ahl al Oughlam yielded more than 100 vertebrate species. The evolutionary stages of its rodents, carnivores, bovids, Kolpochoerus, as well as the absence of Equus, allow us to refer it to the late Pliocene, at ca. 2.5 Ma. The Thomas and Oulad Hamida quarries in Casablanca cut through a unique succession of marine and continental levels going back to the lower Pleistocene, yielding the earliest evidence of human presence in Morocco. Later stages of this human occupation are documented by the Middle Pleistocene levels of these quarries, and by several other sites, most of which are also located in the Rabat–Casablanca region, thus confirming the importance of this coastal area in human history. However, the first chapter of this history in North-western Africa, at present illustrated by the sites of Aïn Hanech and Mansourah in Algeria, remains poorly documented in Morocco.

The Upper Pliocene avifauna of Ahl al Oughlam, Morocco. Systematics and biogeography. In Proceedings of the VII International Meeting of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy

The Upper Pliocene avifauna of Ahl al Oughlam, Morocco. Systematics and biogeography. In Proceedings of the VII International Meeting of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy

Mammalian faunas from the Pliocene and Pleistocene of Casablanca (Morocco)

In the frame of the Franco–Moroccan ‘Programme Casablanca’, several important new Late Miocene to Late Pleistocene mammalian localities have been excavated in the Casablanca area. The rodent fauna of Lissasfa attests to faunal exchanges with Spain and shows that the ‘Quaternary’ of Casablanca dates back to the Late Miocene. Ahl al Oughlam, with more than 100 vertebrate species, is by far the richest paleontological locality of northwestern Africa, but shows that by Late Pliocene, at 2.5 Ma, faunas of this region were already less diverse than in eastern Africa. The Thomas and Oulad Hamida quarries yield, in several continental levels interstratified with high-marine transgressive ones, a succession of late Early to Late Pleistocene faunas associated with lithic industries and Homo remains, which greatly help calibrating mammalian evolution in this part of Africa. The Late Pleistocene is also documented by several new sites, all of which, unfortunately, were destroyed before they could be properly excavated. All sites are threatened by urbanisation, emphasising the need for preservation of the few remaining ones.

Biogeographic relationships of African carnivoran faunas, 7–1.2 Ma

This study analyses the carnivore component of African fossil faunas from three time slices: 7–5 Ma, 4–3 Ma, and 2.5–1.2 Ma, using cluster analysis and principal coordinate analysis (PCO) of presence/absence data on genera. The faunas mostly cluster by time slice, with the exception of Laetoli (Tanzania) and Ahl al Oughlam (Morocco), which differ from all other faunas. The separation during the Late Miocene of a Chado–Libyan bioprovince from the remainder of Africa is supported. No such distinctions are present in the other time slices. Taxonomic distance is not generally correlated with geographic distance, though if Langebaanweg is removed from the 7–5 Ma time slice, the correlation at that time is significant. Comparison of these paleontological results with phylogeographic studies of modern species leads to some general comments on the analytic power of the fossil record with regard to interregional migrations.

Plio-Pleistocene Carnivora of northwestern Africa: A short review

The fossil record of northwestern African carnivores is very patchy. The richest assemblage is that of the Late Pliocene of Ahl al Oughlam, with more than 20 species belonging to the main modern families. Some additions to its study are made here. The rather poor Early Pleistocene faunas are mainly marked by the arrival of a large Canis. A fauna of modern type, with example, the duo Hyaena– Crocuta, settles in the Earliest Pleistocene site of Tighenif, where some older elements linger on ( Homotherium), beside some taxa of doubtful affinities, like a large Panthera, and a strange canid close to Nyctereutes, dominant at this site as well as at the slightly younger ones of Thomas and Oulad Hamida Quarries in Casablanca. All these faunas consist mostly of African taxa, together with a Palaearctic component whose importance increases towards the end of the Pleistocene.

First record of Dinofelis (Felidae, Mammalia) from North Africa

Fragmentary dental and cranial remains from the late Pliocene of Ahl al Oughlam in Morocco are referred to Dinofelis ZDANSKY, 1924. The peculiar mastoid process, large conical upper canine and probable retention of P2 set this form aside from other species of the genus, which had not been reported from North Africa previously. However, in other features, it is closer to the Siwalik species D. cristata than to African ones.

Plio-Pleistocene mammalian biostratigraphy of Atlantic Morocco / Biostratigraphie des mammifères Plio-Pléistocènes du Maroc atlantique.

The recent Moroccan-French cooperation programme has brought about the discovery and excavation of several nch sites in the area of Casablanca. The earliest of these is the newly discovered locality of Lissasfa, which is at least 5 Myr old, and shows that the «Quaternary» sequence of Atlantic Morocco is at least twice as long as previously thought. The other localities provide a biochronological frame for the North-African Upper Pliocene and Quaternary Ahl al Oughlam, dated to about 2.5 Myr, is one of the richest faunal localities in the whole African Cenozoic, but man had probably not arrived in North Africa at that time. In Thomas Quarry 1, the earliest archaeological level in Morocco, level L, is probably of lower Pleistocene age. In the same complex of quarries, the faunas of cave fillings, of which the Homo erectus level of Thomas 1 looks the oldest, can be grouped within a single faunal unit.

Theropithecus atlanticus (Thomas, 1884) (Primates: Cercopithecidae) from the late Pliocene of Ahl al Oughlam, Casablanca, Morocco.

The site of Ahl al Oughlam near Casablanca, Morocco, dated to ca. 2.5 Ma, has yielded a good sample of Theropithecus atlanticus (Thomas, 1884), a North African late Pliocene species previously known only by its holotype, a lower molar from Algeria. Theropithecus atlanticus, which can now be much better defined, is clearly distinct from other species of the genus, which is thus more diverse than previously thought. The mandible of T. atlanticus has a very characteristic deep and long post-molar sulcus and a deep and well excavated supra-lateral triangular depression of the ramus, with a sharp postero-inferior ridge. The upper and lower canines are rather large but low. The male P3 is very wide, with well developed posterior crests; the P4 is rounded, with a large talonid and weak notches and clefts. Median lingual notches of the lower molars form an acute angle. Although our incomplete knowledge of T. atlanticus precludes a detailed phylogenetic analysis, we suggest that it arose by cladogenesis from the T. darti-T. oswaldi lineage; it is replaced by the latter species in the Pleistocene.

La faune de Mammifères du Pliocène terminal d'Ahl al Oughlam, Casablanca, Maroc

The Late Pliocene site (ca 2.5 Ma) of Ahl al Oughlam has yielded a complete fauna of macroand micro-mammals, by far the richest of the late Cenozoic era of North Africa; it includes at least 55 species. Carnivora (23 species) are the dominant group. Many of them used the caves and fissures as dens or shelters, bringing in most of the ungulate remains. These remains include mostly medium-size species and juveniles of larger ones. Very few taxa are akin to Palearctic ones, and most of the faunal exchanges were intra-African. However, when compared to East Africa, the poor diversity of these ungulates must be noted, which the karstic nature of the site fails to fully explain, but which could result from a harsh, open and relatively cold environment.

Bovidae (Mammalia) du Pliocène final d’Ahl al Oughlam, Casablanca, Maroc

Only seven bovid species are present at Ahl al Oughlam: aTragelaphus (rare), a Bovine close toPelorovis ? praeafricanus but mostly known by teeth, a new species of kob (perhaps of an endemic lineage), aParmularius slightly more primitive than at Olduvai,Beatragus (mentioned for the first time in North Africa),Gazella thomasi, and a new species of gazelle noticeable by its nasal region. The Ahl al Oughlam bovids are in good agreement with an age of 2.5 Ma. They point towards an open and probably rather unfavourable, perhaps cold, environment.

The status of the african fossil suids Kolpochoerus limnetes ( Hopwood, 1926), K. Phacochoeroides ( Thomas 1884) and “K.” Afarensis ( Cooke, 1978)

Kolpochoerus limnetes is the name widely used for an abundant suid in East African Plio-Pleistocene deposits. A recent publication ( Pickford 1994) suggests that the holotype of “ Sus limnetes” from Kaiso could be a Nyanzachoere and proposes that this specific name should be restricted to the Type specimen and also that the other East Africa fossils be given the prior name of Kolpochoerus phacochoeroides. It is shown here that, although the Kaiso Type specimen is morphologically compatible with Kolpochoerus, this single tooth is now an unsatisfactory type specimen in view of the questionable stratigraphic source indicated by the preservation colour. Accordingly the name should be restricted only to that specimen as Pickford suggests. The use of the specific name phacochoeroides is inappropriate as Geraads (1993) has demonstrated that collections from Ahl al Oughlam, Morocco, belong to the North African K. phacochoeroides and are clearly different from the East African species. Accordingly the East African material is best referred in future to K. heseloni. Based on Geraads evaluation of other species and reconsideration of the material, it is proposed here that the Hadar species described as Kolpochoerus afarensis should now be placed in Potamochoerus as P. afarensis.

Carnivores du Pliocène terminalde Ahl al Oughlam (Casablanca, Maroc)

The site of Ahl al Oughlam has yielded the first important carnivore fauna of North Africa, and one of the richest of Africa. It includes 23 taxa, 13 of which are new, belonging to most of the carnivore families, mainly the Felidae (6 species), Hyaenidae and Mustelidae (4 species each). Several genera were not previously known from North Africa, like Pliocrocuta, Chasmaporthetes, Nyctereutes, Prepoecilogale, while the walrus ( Alachtherium) is mentioned for the first time in Africa. Many other genera were unknown in the area earlier than the Middle or late Pleistocene: Herpestes, Viverra, Genetta, Acinonyx, Ursus, Poecilictis, Mellivora, Lutra. The carnivore fauna of Ahl al Oughlam, quite diverse ecologically, is at least as distinct from those of Eurasia as from those of Eastern and Southern Africa; the many similarities with both, however, allow to confirm the age previously proposed for the locality, ca. 2.5 m.y.

Rongeurs et insectivores (Mammalia)du Pliocene final de Ahl Al Oughlam (Casablanca, Maroc)

Late Pliocene Rodents from Ahl al Oughlam (Morocco) are known by several thousands of teeth, but include only 6 species. Hystrix and the Ctenodactylid Irhoudia are very rare; the Murids Paraethomys, Praomys and Mus are slightly more primitive than at Irhoud Ocre. A new species of Gerbillus is similar to G. campestris, despite its early age. Insectivora belong chiefly to the Soricids; the most common one is an “Episoriculus” similar to the one from Irhoud Ocre; a new species of Suncus is the first fossil of this genus in North-Africa; Erinaceus (Atelerix) is known by 2 teeth only. The microfauna is less diverse than in more recent sites, where the predominance of Gerbillids attests a more open and dry environment, and Ahl al Oughlam looks earlier than the late Pliocene climatic event.

Kolpochoerus phacochoeroides (Thomas, 1884)(Suidae, Mammalia), du Pliocène supérieur de Ahl al Oughlam (Casablanca, Maroc)

All suid remains from the upper Pliocene of Ahl al Oughlam (Casablanca, Morocco) are refered to Kolpochoerus phacochoeroides Thomas, 1884 . This species differs from “ K.” afarensis and K. limnetes from East Africa by a shortened muzzle, large canines, even in females, reduced anterior premolars, and P4/ with a postero-lingual fovea. It belonged to a group of grazers which also included K. limnetes, K. paiceae from South Africa and the forest hog, among which K. limnetes split off first. Assuming an uniform rate of M3 lengthening, the age of Ahl al Oughlam can be estimated at 2.4 m.y. This age is in good agreement with the rest of the fauna.