Guelb el Ahmar (Bathonian, Anoual Syncline, eastern Morocco): First continental flora and fauna including mammals from the Middle Jurassic of Africa

Abstract We report the discovery in Mesozoiccontinental “ red beds” of Anoual Syncline, Morocco, of the new Guelb el Ahmar (GEA) fossiliferous sites in the Bathonian Anoual Formation. They produced one of the richest continental biotic assemblages from the Jurassic of Gondwana, including plants, invertebrates and vertebrates. Both the sedimentological facies and the biotic assemblage indicate a lacustrine depositional environment. The flora is represented by tree trunks (three families), pollen (13 species, five major clades) and charophytes. It suggests local forests and humid (non-arid) conditions. The vertebrate fauna is dominated by microvertebrates recovered by screening–washing. It is rich and diverse, with at least 29 species of all major groups (osteichthyans, lissamphibians, chelonians, diapsids, mammals), except chondrichthyans. It includes the first mammals discovered in the Middle Jurassic of Arabo-Africa. The GEA sites yielded some of the earliest known representatives of osteoglossiformfishes, albanerpetontid and caudate amphibians, squamates (scincomorphans, anguimorphan), cladotherian mammals, and likely choristoderes. The choristoderes, if confirmed, are the first found in Gondwana, the albanerpetontid and caudatan amphibians are among the very few known in Gondwana, and the anguimorph lizard is the first known from the Mesozoicof Gondwana. Mammals ( Amphitheriida, cf. Dryolestida) remain poorly known, but are the earliest cladotherians known in Gondwana. The GEA biotic assemblage is characterized by the presence of Pangean and Laurasian (especially European) taxa, and quasi absence of Gondwanan taxa. The paleobiogeographical analysis suggests either a major fossil bias in Gondwanaduring the Middle Jurassic, and an overall vicariantPangean context for the GEA assemblage, or alternatively, noticeable Laurasian (European) affinities and North-South dispersals. The close resemblance between the Bathonian faunas of GEA and Britain is remarkable, even in a Pangean context. The similarity between the local Anoual SynclineGuelb el Ahmar and Ksar Metlili faunas raises questions on the ?Berriasian age of the latter.