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The Eocene Mammalian Fauna of Chambi (Tunisia) in Its Geological Context
Abstract
In the Kasserine area of Tunisia there is a Cenozoic basin containing Paleogene continental deposits. Palustrine limestones located at the base of the Paleogene series have yielded the Chambi mammalian fauna. Representatives of the following mammalian orders and families have been found: Marsupialia, Peradectidae; Placentalia, Protheutheria indet.; ?Lipotyphla, ?Erinaceomorpha; Chiroptera, Rhinolophoidea, Nycterididae and Vespertilionoidea, Philisidae; Primates, Adapidae; Rodentia, Zegdoumyidae; Macroscelidea, Herodotinae; Hyracoidea, Pliohyracidae. The analysis of this local fauna provides insights into its position with regards to other North African mammalian faunas and its possible relationships with Eocene Eurasian faunas.
... In the framework of our paleontological programme in the Early Tertiary of North Africa, since 2008, we have focused our field research in the western part of Central Tunisia (Kasserine region), notably on the geological outcrops exposed in the Natural Park of Djebel Chambi ( Fig. 1A-B). The Djebel Chambi is particularly famous for having yielded a fossiliferous vertebrate locality (Chambi locus 1, CBI-1) dating from the Late Early-to-the Early Middle Eocene (Hartenberger et al. 1985(Hartenberger et al. , 1997(Hartenberger et al. , 2001Coster et al. 2012). The excavations, acid processing and screen washings of several hundred kilograms of sediments (lacustrine limestone; Fig. 1B) of this locality have so far provided a diverse assemblage of aquatic and terrestrial vertebrates including fishes, turtles, crocodiles, squamates (Rage et al. under study), birds (Mourer-Chauvir e et al. 2013) and mammals such as bats, primates, eulipotyphlans, hyracoids, an elephant shrew, a marsupial, a rodent and a sirenian (Hartenberger 1986;Crochet 1986;Sig e 1991;Court & Hartenberger 1992Hartenberger & Marandat 1992;Vianey-Liaud et al. 1994;Hartenberger et al. 1997Hartenberger et al. , 2001Gheerbrant & Hartenberger 1999;Tabuce et al. 2007Tabuce et al. , 2011Ravel et al. 2011Ravel et al. , 2012Benoit et al. 2013a,b,c;Marivaux et al. 2013; see Table S1). ...
... The Djebel Chambi is particularly famous for having yielded a fossiliferous vertebrate locality (Chambi locus 1, CBI-1) dating from the Late Early-to-the Early Middle Eocene (Hartenberger et al. 1985(Hartenberger et al. , 1997(Hartenberger et al. , 2001Coster et al. 2012). The excavations, acid processing and screen washings of several hundred kilograms of sediments (lacustrine limestone; Fig. 1B) of this locality have so far provided a diverse assemblage of aquatic and terrestrial vertebrates including fishes, turtles, crocodiles, squamates (Rage et al. under study), birds (Mourer-Chauvir e et al. 2013) and mammals such as bats, primates, eulipotyphlans, hyracoids, an elephant shrew, a marsupial, a rodent and a sirenian (Hartenberger 1986;Crochet 1986;Sig e 1991;Court & Hartenberger 1992Hartenberger & Marandat 1992;Vianey-Liaud et al. 1994;Hartenberger et al. 1997Hartenberger et al. , 2001Gheerbrant & Hartenberger 1999;Tabuce et al. 2007Tabuce et al. , 2011Ravel et al. 2011Ravel et al. , 2012Benoit et al. 2013a,b,c;Marivaux et al. 2013; see Table S1). Here, we report new dental remains from CBI-1 attributable to Zegdoumys sbeitlai Vianey-Liaud et al. 1994; the unique rodent recorded from that locality. ...
... Late Early-to-Early Middle Eocene (e.g. Hartenberger et al. 1997Hartenberger et al. , 2001Adaci et al. 2007;Coster et al. 2012;Marivaux et al. 2013). ...
The late early to early middle Eocene Zegdoumyidae are the oldest representatives of the order Rodentia to be known in Africa thus far. Despite the fragmentary nature of their fossil record, these early African rodents have been central in discussions surrounding the early evolutionary history of the Anomaluroidea clade (the living forms of which are the ‘scaly-tailed squirrels’, i.e. Anomaluridae). Here, we describe new dental remains attributable to Zegdoumys sbeitlai, a zegdoumyid from Tunisia (Djebel Chambi, CBI-1) dated at ca 50-45 Ma. The original material referred to this taxon was limited to four teeth documenting few dental loci. The new gathered material comprises practically all dental loci (except P4), thereby allowing a better description and characterization of this taxon. This Tunisian species is clearly distinct from its roughly coeval Algerian counterpart (Z. lavocati) or from the younger species recorded from Namibia (Z. namibiensis). We investigated the phylogenetic positions of Z. sbeitlai and the Zegdoumyidae in a high-level rodent phylogeny with a cladistic assessment of the dental evidence. Our results show that zegdoumyids represent the earliest offshoots (pectinately arranged) of a large clade that also encompasses the more advanced Eocene anomaluroids (here named Nementchamyidae), the Miocene Nonanomaluridae, and the stem and crown Anomaluridae. In this phylogenetic context, the species of Zegdoumys are the basalmost members of this large Anomaluroidea clade. Zegdoumyid taxa are therefore the oldest stem anomaluroid candidates, a statement which strengthens support for the hypothesis of the great antiquity of the Anomaluroidea clade in Africa. From a historical biogeography perspective, this assumption suggests that anomaluroids invaded Asia from Afro-Arabia sometime during the middle Eocene, a dispersal event which was likely concomitant but opposite to the dispersals envisaged for the hystricognathous rodents and anthropoid primates from Asia to Afro-Arabia.
... Here, we describe a new fossil of sirenian, which consists of a petrosal (CBI-1-542) (Fig. 1A, B, C). This fossil was recently found in the late Ypresian-early Lutetian fossil mammal-bearing locality of Djebel Chambi in Tunisia [14,15]. Interestingly, this African locality is roughly contemporaneous with the Jamaican locality that yielded Prorastomus. ...
... Other paleontological clues suggest they were probably already adapted to marine environments, but given the greater diversity of aquatic angiosperms in freshwater, one cannot exclude occasional freshwater foraging [1,6,7]. In contrast, the vertebrate locality of Djebel Chambi in Tunisia is a bed of lacustrine limestone, which should correspond to freshwater deposits [14]. The presence of charophytes and amphibian remains in the locality supports this paleoenvironmental reconstruction [14]. ...
... In contrast, the vertebrate locality of Djebel Chambi in Tunisia is a bed of lacustrine limestone, which should correspond to freshwater deposits [14]. The presence of charophytes and amphibian remains in the locality supports this paleoenvironmental reconstruction [14]. Noteworthily, the discovery of a tooth of an electric ray (Chondrichthyes, Torpediniformes) in this fauna suggests occasional connections with sea water (S2). ...
Sea cows (manatees, dugongs) are the only living marine mammals to feed solely on aquatic plants. Unlike whales or dolphins (Cetacea), the earliest evolutionary history of sirenians is poorly documented, and limited to a few fossils including skulls and skeletons of two genera composing the stem family of Prorastomidae (Prorastomus and Pezosiren). Surprisingly, these fossils come from the Eocene of Jamaica, while stem Hyracoidea and Proboscidea - the putative sister-groups to Sirenia - are recorded in Africa as early as the Late Paleocene. So far, the historical biogeography of early Sirenia has remained obscure given this paradox between phylogeny and fossil record. Here we use X-ray microtomography to investigate a newly discovered sirenian petrosal from the Eocene of Tunisia. This fossil represents the oldest occurrence of sirenians in Africa. The morphology of this petrosal is more primitive than the Jamaican prorastomids' one, which emphasizes the basal position of this new African taxon within the Sirenia clade. This discovery testifies to the great antiquity of Sirenia in Africa, and therefore supports their African origin. While isotopic analyses previously suggested sirenians had adapted directly to the marine environment, new paleoenvironmental evidence suggests that basal-most sea cows were likely restricted to fresh waters.
... Mannaï-Tayech 2006Belghithi et al. 2016). The sandstone attributed by Hartenberger et al. (2001) to the Beglia Fm. in fact corresponds to the uppermost Paleogene levels where the Mabrouk locality occurs (Sweydan et al. 2019). The geological study made by Hartenberger et al. (2001) did not include the area where the Neogene sediments outcrop. ...
... The sandstone attributed by Hartenberger et al. (2001) to the Beglia Fm. in fact corresponds to the uppermost Paleogene levels where the Mabrouk locality occurs (Sweydan et al. 2019). The geological study made by Hartenberger et al. (2001) did not include the area where the Neogene sediments outcrop. Furthermore, a recent discussion of the Miocene sandstones capping the Nementcha Fm. in Algeria (Lihoreau et al. 2015b), only 130 km southwest of Sidi Hedri, highlighted the critical role of fossil assemblages in correlating between continental formations, and demonstrates that it is best to avoid using only facies correlations for precise datings. ...
Anthracotheres dispersed from Asia toward Africa at least three times: at the Eocene/Oligocene transition, during the early Miocene and later during the Miocene. Those dispersals are important datum events for African Tertiary biochronology. New fossil remains of early Libycosaurus, the genus implicated in the late Miocene dispersal, are described from a new Tunisian locality of the Kasserine area. The new fossils enhance the hypodigm of Libycosaurus algeriensis and increase the resolution of the phylogenetic position of this species using cladistics analysis. The inclusion of the genus Libycosaurus within the well-described Merycopotamus lineage allows us to constrain its dispersal time. Dispersal of this anthracothere from the Indian sub-continent to Africa was probably facilitated by sea level decrease during the early Tortonian, just preceding the Hipparion dispersal event. This new age estimation refines the resolution of the succession of late Miocene deposits in Maghreb and frames the date of the onset of the Sahara.
... The Djebel Chambi in central Tunisia is famous for having yielded a rich faunal and floral assemblage (Hartenberger et al. 2001), including indeterminate amphibians and squamates, a peradectid marsupial, and several eutherians among which there are chiropterans, a putative erinaceomorph, a strepsirhine primate, a zegdoumyid rodent, a stem macroscelidid, a hyracoid, and charophytes. Hartenberger et al. (2001) have proposed that the fauna is Early Eocene in age, roughly contemporaneous with that of the Glib Zegdou in Algeria. ...
... The Djebel Chambi in central Tunisia is famous for having yielded a rich faunal and floral assemblage (Hartenberger et al. 2001), including indeterminate amphibians and squamates, a peradectid marsupial, and several eutherians among which there are chiropterans, a putative erinaceomorph, a strepsirhine primate, a zegdoumyid rodent, a stem macroscelidid, a hyracoid, and charophytes. Hartenberger et al. (2001) have proposed that the fauna is Early Eocene in age, roughly contemporaneous with that of the Glib Zegdou in Algeria. However, the age of Chambi and Glib Zegdou faunas have long remained a matter of discussion, and current data rather indicate a late Early or early Middle Eocene age, unfortunately without more precision (e. g. adaci et al. 2007;Seiffert 2010). ...
... The Tunisian locality of Chambi (Figure 1.1), dating from the late Ypresian or early Lutetian (Figure 1.2), has yielded mammal fossils since the mid 1980s (e.g., Hartenberger et al., 1985Hartenberger et al., , 1997Hartenberger et al., , 2001. These fossils have strongly contributed to clarify the early evolution of some mammal groups in Africa, notably the elephant-shrews (Hartenberger, 1986;Tabuce et al., 2007;Benoit et al., 2013a), primates (Hartenberger and Marandat, 1992;Marivaux et al., 2013), rodents (Vianey-Liaud et al., 1994;Marivaux et al., 2015), bats (Ravel et al., 2011a(Ravel et al., , 2012(Ravel et al., , 2015 in press), hyracoids (Court and Hartenberger, 1992;, and sirenians (Benoit et al., 2013b). ...
... Based on biostratigraphic data (mammals and charophytes), Hartenberger et al. (1997Hartenberger et al. ( , 2001 envisaged that the locality of Chambi (Tunisia) is Ypresian in age and possibly older than those from the Gour Lazib complex (Algeria). However, based on recent better substantiated biostratigraphic and magnetostratigraphic data, a contemporaneity between both localities have been proposed Coster et al., 2012;Marivaux et al., 2015;Ravel et al., 2015, in press), i.e., dated as late Ypresian or early Lutetian, between 49 and 45 m.y. ...
We present and describe new fossils from the Eocene North African localities of Chambi (Tunisia; late Ypresian or early Lutetian) and Bir el Ater (Algeria; latest Bartonian or earliest Priabonian). The specimens from Chambi allow recognizing two recently described hyainailourines: Furodon crocheti and Parvavorodon gheerbranti; these taxa were previously known from the Gour Lazib area (Algeria; late Ypresian or early Lutetian). The new material from Tunisia includes a fragmentary dentary of Parvavorodon that substantially supports the hyainailourine status of this genus and represents the oldest dentary fragment presently known for a juvenile of Hyaenodonta in Africa. The presence of Furodon and Parvavorodon in Chambi strengthens support for the hypothesis of contemporaneity of the Eocene Gour Lazib and Chambi mammal-bearing localities. In addition, the find of a typical teratodontine fourth premolar in Chambi testifies to the presence of a small representative of this group. The fossil record in Bir el Ater is scarcer than in Chambi. However, we recognize specimens attributable to a hyainailourine and a teratodontine. The latter is referred as Masrasector cf. ligabuei, and is the oldest record for this genus. We tentatively identify a modification of the hyaenodont fauna in the Maghreb after the “Early Eocene Climatic Optimum” (EECO). This faunal change might be related to the decrease of the global temperature after the EECO event. It appears contemporaneous of a drastic replacement in the composition of the mammal faunas in Africa. Finally, the end of the Eocene (Priabonian) shows an increase in the subfamilial richness amongst hyaenodonts.
... Depuis les premiers travaux de Pervinquière en 1903 puis de Flandrin en 1948, la Tunisie centrale et plus particulièrement la région de Kasserine (fig. 1) est considérée, selon les auteurs, comme étant une terre émergée (l'île ou l'archipel de Kasserine), depuis le Coniacien [Burollet, 1956] ou la fin du Maastrichtien [Marie et al., 1984 ; Bishop, 1988 ; Bensalem H., 2002] jusqu'à la base du Miocène moyen [Sassi, 1974 ; Sassi et al., 1984 ; Hartenberger et al., 2001 ; Zouaghi et al., 2005]. De ce fait, et avant les années 1980, aucun dépôt sédimentaire plus vieux que le Miocène moyen n'avait été reconnu en Tunisie centrale. ...
... De ce fait, et avant les années 1980, aucun dépôt sédimentaire plus vieux que le Miocène moyen n'avait été reconnu en Tunisie centrale. A partir de 1984, plusieurs travaux ont alors porté sur des dépôts continentaux constitués de paléosols évolués, de formations conglomératiques ou carbonatées, lacustres ou palustres Truc, 1981 ; Zouari, 1984 ; Abdeljaoued, 1991 ; Sassi et al., 1992 ; Hartenberger et al., 1985 Hartenberger et al., , 2001 Jamoussi et al., 2001 ; Zaaboub et al., 2005]. Ces séries sédimentaires ont livré des faunes de gastéropodes terrestres (Bulimes) ou de riches gisements de vertébrés [Hartenberger et al., 1985], qui ont conduit à les dater du Paléocène et de l'Eocène inférieur. ...
The Kasserine area, in central Tunisia, is for a long time considered by many authors as an emerged island (or islets), since the upper Cretaceous until the middle Miocene. In the eighties, continental deposits were described and dated Eocene. At that time, no marine deposit older than middle Miocene were thus known in central Tunisia, and the only Eocene marine strata, were located in periphery of the emerged zone, in phosphatic basins. It is in the area of Djebel el Kebar, which is situated at the eastern edge of the Kasserine region that a detailed study of the Eocene-middle Miocene deposits was carried out. In the studied series, four major unconformities are observed and correlated on the whole of the site. Between these surfaces, three successive sedimentary units are observed from base to top and are composed of limestone, glauconite and clay deposits (unit I), shell beds limestone (unit II), conglomerate, sandstone and silty-clay (unit III). Paleontological and biostratigraphic results. - An important fossils locality was discovered in Djebel el Kebar, in glauconitic marine deposits of the first sedimentary unit. Fauna, very rich, contains many marine (shark, ray, sirenian, etc.) and continental forms (rodent, hyracoid and primate). The new dating clearly indicates that the deposits of the lower part of the series are Bartonian in age. Sedimentological results. - Thirteen facies were identified and the exceptional conditions of outcrop in Djebel el Kebar have allowed us to map the lateral facies evolution. The sedimentary environments were reconstituted and their succession shows a very shallow water-depth carbonate platform (wave dominated and influenced by storms and tide), passing upward into a sandy fluvial and estuarine environment. Tectono-sedimentary and sequential evolution. - A sequence stratigraphie framework was proposed, in which, on both sides of unconformities primarily of tectonic origin, three very partially preserved sequences (> 3 Ma) are compatible with the second order of Vail et al. [1991]. In these sequences, the major part of the studied series is set up in transgressive system tracts and highstand system tracts. The fossil locality then seems to be associated with several episodes with condensation, formed during the transgressive part of the second order cycle. Conclusions. - The first, new and important result of this work, is the characterization and dating of marine deposits older than middle-Miocene in Djebel el Kebar, in central Tunisia. The studied area is thus not in the emerged part of the zone of Kasserine during the Eocene. Between Bartonian and lower-Miocene, marine facies, evolve within an environment of very shallow marine carbonated platform. A major event (flexure coupled with eustatic fall in Miocene) would cause a change of depositional profile, the arrival of coarse terrigenous material and the installation of a continental fluvial and estuarine sedimentary environment. Finally the whole of the series records large scale tectono-eustatic changes, responsible of several second order sequences, partially preserved.
... Depuis les premiers travaux de Pervinquière en 1903 puis de Flandrin en 1948, la Tunisie centrale et plus particulièrement la région de Kasserine ( fig. 1) est considérée, selon les auteurs, comme étant une terre émergée (l'île ou l'archipel de Kasserine), depuis le Coniacien [Burollet, 1956] ou la fin du Maastrichtien [Marie et al., 1984 ;Bishop, 1988 ;Bensalem H., 2002] jusqu'à la base du Miocène moyen [Sassi, 1974 ;Sassi et al., 1984 ;Hartenberger et al., 2001 ;Zouaghi et al., 2005]. De ce fait, et avant les années 1980, aucun dépôt sédimentaire plus vieux que le Miocène moyen n'avait été reconnu en Tunisie centrale. ...
... De ce fait, et avant les années 1980, aucun dépôt sédimentaire plus vieux que le Miocène moyen n'avait été reconnu en Tunisie centrale. A partir de 1984, plusieurs travaux ont alors porté sur des dépôts continentaux constitués de paléosols évolués, de formations conglomératiques ou carbonatées, lacustres ou palustres Truc, 1981 ;Zouari, 1984 ;Abdeljaoued, 1991 ;Sassi et al., 1992 ;Hartenberger et al., 1985Hartenberger et al., , 2001Jamoussi et al., 2001 ;Zaaboub et al., 2005]. Ces séries sédimentaires ont livré des faunes de gastéropodes terrestres (Bulimes) ou de riches gisements de vertébrés [Hartenberger et al., 1985], qui ont conduit à les dater du Paléocène et de l'Eocène inférieur. ...
The Kasserine area, in central Tunisia, is for a long time considered by many authors as an emerged island (or islets), since
the upper Cretaceous until the middle Miocene. In the eighties, continental deposits were described and dated Eocene. At that
time, no marine deposit older than middle Miocene were thus known in central Tunisia, and the only Eocene marine strata, were
located in periphery of the emerged zone, in phosphatic basins. It is in the area of Djebel el Kebar, which is situated at
the eastern edge of the Kasserine region that a detailed study of the Eocene-middle Miocene deposits was carried out. In the
studied series, four major unconformities are observed and correlated on the whole of the site. Between these surfaces, three
successive sedimentary units are observed from base to top and are composed of limestone, glauconite and clay deposits (unit
I), shell beds limestone (unit II), conglomerate, sandstone and silty-clay (unit III).
Paleontological and biostratigraphic results An important fossils locality was discovered in Djebel el Kebar, in glauconitic marine deposits of the first sedimentary
unit. Fauna, very rich, contains many marine (shark, ray, sirenian, etc.) and continental forms (rodent, hyracoid and primate).
The new dating clearly indicates that the deposits of the lower part of the series are Bartonian in age.
... The Djebel Chambi in central Tunisia is famous for having yielded a rich faunal and floral assemblage (Hartenberger et al. 2001), including indeterminate amphibians and squamates, a peradectid marsupial, and several eutherians among which there are chiropterans, a putative erinaceomorph, a strepsirhine primate, a zegdoumyid rodent, a stem macroscelidid, a hyracoid, and charophytes. Hartenberger et al. (2001) have proposed that the fauna is Early Eocene in age, roughly contemporaneous with that of the Glib Zegdou in Algeria. ...
... The Djebel Chambi in central Tunisia is famous for having yielded a rich faunal and floral assemblage (Hartenberger et al. 2001), including indeterminate amphibians and squamates, a peradectid marsupial, and several eutherians among which there are chiropterans, a putative erinaceomorph, a strepsirhine primate, a zegdoumyid rodent, a stem macroscelidid, a hyracoid, and charophytes. Hartenberger et al. (2001) have proposed that the fauna is Early Eocene in age, roughly contemporaneous with that of the Glib Zegdou in Algeria. However, the age of Chambi and Glib Zegdou faunas have long remained a matter of discussion, and current data rather indicate a late Early or early Middle Eocene age, unfortunately without more precision (e. g. adaci et al. 2007;Seiffert 2010). ...
A distal tarsometatarsus and a fragment of carpometacarpus of a small galliform, the size of a recent quail, have been found in the late Early or early Middle Eocene of Chambi, in Tunisia. Although a large number of stem group representatives of Galliformes are known from the Eocene of the Northern Hemisphere, and one from the middle Eocene of Namibia, the taxon from Chambi differs from them and is described as a new genus and species. A very small zygodactyl form, represented by three distal tarsometatarsi, is also present in the same locality. This form, described as a new genus and species, is a stem group representative of the recent family Cuculidae. It shows a plesiomorphic morphology compared to the recent members of the Cuculidae, but it is, however, more derived than the younger genus Eocuculus. It is the earliest Cuculidae known so far.
... Situated in the western part of central Tunisia (Kasserine region), the Djebel Chambi ( Fig. 1) is particularly famous for having yielded a fossiliferous locality (Chambi locus 1, CBI-1) dating from the late early to the early middle Eocene (Hartenberger et al., 1985(Hartenberger et al., , 1997(Hartenberger et al., , 2001. This locality has provided a diverse assemblage of aquatic and terrestrial vertebrates (fishes, turtles, crocodiles, squamates), including terrestrial birds (Mourer-Chauviré et al., in press) and mammals such as marsupials, rodents, bats, lipotyphlans, elephant-shrews, hyracoids, and primates (Crochet, 1986;Hartenberger, 1986;Sigé, 1991;Court and Hartenberger, 1992;Hartenberger and Marandat, 1992;Court, 1993;Vianey-Liaud et al., 1994;Hartenberger et al., 1997Hartenberger et al., , 2001Gheerbrant and Hartenberger, 1999;Tabuce et al., 2007;Ravel et al., 2011Ravel et al., , 2012. ...
... Situated in the western part of central Tunisia (Kasserine region), the Djebel Chambi ( Fig. 1) is particularly famous for having yielded a fossiliferous locality (Chambi locus 1, CBI-1) dating from the late early to the early middle Eocene (Hartenberger et al., 1985(Hartenberger et al., , 1997(Hartenberger et al., , 2001. This locality has provided a diverse assemblage of aquatic and terrestrial vertebrates (fishes, turtles, crocodiles, squamates), including terrestrial birds (Mourer-Chauviré et al., in press) and mammals such as marsupials, rodents, bats, lipotyphlans, elephant-shrews, hyracoids, and primates (Crochet, 1986;Hartenberger, 1986;Sigé, 1991;Court and Hartenberger, 1992;Hartenberger and Marandat, 1992;Court, 1993;Vianey-Liaud et al., 1994;Hartenberger et al., 1997Hartenberger et al., , 2001Gheerbrant and Hartenberger, 1999;Tabuce et al., 2007;Ravel et al., 2011Ravel et al., , 2012. The small Euprimates recorded from Chambi CBI-1 consist of Djebelemur martinezi (Chambi being the type-locality of this species; Hartenberger and Marandat, 1992) and another tiny taxon that is similar to and possibly congeneric with Algeripithecus (see Godinot, 1994;Seiffert et al., 2010;Seiffert, 2012). ...
We report the discovery of three isolated primate petrosal fragments from the fossiliferous locality of Chambi (Tunisia), a primate-bearing locality dating from the late early to the early middle Eocene. These fossils display a suite of anatomical characteristics otherwise found only in strepsirhines, and as such might be attributed either to Djebelemur or/and cf. Algeripithecus, the two diminutive stem strepsirhine primates recorded from this locality. Although damaged, the petrosals provide substantial information regarding the ear anatomy of these advanced stem strepsirhines (or pre-tooth-combed primates), notably the patterns of the pathway of the arterial blood supply. Using mu CT-scanning techniques and digital segmentation of the structures, we show that the transpromontorial and stapedial branches of the internal carotid artery (ICA) were present (presence of bony tubes), but seemingly too small to supply enough blood to the cranium alone. This suggests that the ICA was not the main cranial blood supply in stem strepsirhines, but that the pharyngeal or vertebral artery primitively ensured a great part of this role instead, an arterial pattern that is reminiscent of modern cheirogaleid, lepilemurid lemuriforms and lorisiforms. This could explain parallel loss of the ICA functionality among these families. Specific measurements made on the cochlea indicate that the small strepsirhine primate(s) from Chambi was (were) highly sensitive to high frequencies and poorly sensitive to low frequencies. Finally, variance from orthogonality of the plane of the semicircular canals (SCs) calculated on one petrosal (CBI-1-569) suggests that Djebelemur or cf. Algeripithecus likely moved (at least its head) in a way similar to that of modern mouse lemurs.
... Only four other mammal-producing early Eocene (Ypresian) localities have been described from Africa, all from the north of the continent: (1) the Grand Daoui locality in the Ouled Abdoun Basin, Morocco (earliest Eocene, Gheerbrant et al. [2002Gheerbrant et al. [ , 2003Gheerbrant et al. [ , 2005a); (2) N'Tagourt and Adrar Mgorn in the Ouarzazate Basin, Morocco (early Eocene, Sudre et al. [1993]; Hartenberger et al. [2002:fig. 6]; Gheerbrant et al. [2002]); (3) the Chambi locality, Tunisia (late early Eocene, Hartenberger [1986]; Hartenberger et al. [2002]); and (4) the El Kohol locality, Algeria (early Eocene, Mahboubi et al. [1986]; Mehbrouk et al. [1997]; middle Eocene, Hartenberger et al. [2002:fig. 6] and Gheerbrant et al. [2005a:89]). ...
... Recent phylogenetic analyses of Afrotheria (Asher et al., 2003;Zack et al., 2005) have not explicitly included the proboscideans Phosphatherium and Daouitherium, or the macroscelidids and hyracoids from the Chambi locality (Hartenberger et al., 2002). It is conservative to suggest, however, that these taxa, and those from Tamaguélelt, are likely to emerge nested within Afrotheria. ...
Fossils from early Tertiary phosphate deposits of northern Mali include a new diminutive proboscidean taxon intermediate in size between Daouitherium rebouli and Phosphatherium escuilliei. Although a fragmentary specimen, it has two dentary characters previously demonstrated to be diagnostic for basal proboscideans: a labially expanded mandibular corpus in the region of m2 and m3 and a mesiolabially positioned mandibular ramus. The locality that produced the fossils, Tamaguélelt, has historically been hard to date even though it has produced many vertebrates. A section is presented that includes the locality and relates it to surrounding sediments. The locality is Eocene, lies very close to the Paleocene-Eocene boundary, and is most likely early Eocene (Ypresian). The depositional environment suggests substantial reworking of older sediments, and the fossils within them, including the proboscideans, may be even older than early Eocene. The Mali proboscidean is one of the oldest members of Afrotheria that has been found in Africa and this locality is the southernmost African early Eocene locality for mammals.
... Also notable in Europe, bats are the most abundant mammals known in the middle Eocene (MP 11) lagerstätte at Messel, Germany, with four families and four genera (Archaeonycteris in Archaeonycteridae, Palaeo chiropteryx in Palaeochiropterygidae, Hassianycteris in Hassianycteridae, and Tachypteron in Emballonuridae; Russell and Sigé, 1970;Habersetzer et al., 1992Habersetzer et al., , 1994Storch et al., 2002;Morlo et al., 2004). In northern Africa, fossil bats similar in age to the Elderberry Canyon LF include, at Chambi, Tunisia (late early Eocene to early middle Eocene), ?Necromantis (Necromantidae), ?Palaeophyllophora and Hipposideros (Hipposideridae), Vespertiliavus (Emballonuridae), Khoufechia (Nycteridae), Chambinycteris (family indeterminate), Dizzya (Philisidae), and genus indeterminate (Vespertilionidae; Sigé, 1991;Hartenberger et al., 2001;Ravel et al., 2016) and, at Glib Zegdou, Algeria (late early Eocene to early middle Eocene), Pseudovespertilia vus (Emballonuridae) and Drakonycteris (family indeterminate; Ravel et al., 2016). Also noteworthy in eastern Africa is the tanzanycterid Tanzanycteris, from the middle middle Eocene, early Lutetian, from Mahenge, Tanzania (Gunnell et al., 2003;Smith et al., 2012). ...
We report three new taxa of bats from the late early Eocene to earliest middle Eocene (Bridgerian biochrons Br1b–Br2; ca. 50–48 Ma) Elderberry Canyon Quarry, Sheep Pass Formation, in the Egan Mountain Range of eastern Nevada, USA. Volactrix simmonsae gen. et sp. nov., represented by two dentaries, is tentatively referred to the family ?Onychonycteridae and exhibits a p3 that is much smaller than the p2, a semimolariform p4, and lower molars with a postcristid configuration intermediate between necromantodonty and nyctalodonty. The genus Palaeochiropteryx, a palaeochiropterygid otherwise known from the early and middle Eocene of Europe, is the first representative of its genus and of the family Palaeochiropterygidae outside Eurasia and the Indo-Pakistan subcontinent. Palaeochiropteryx is represented at the site by a partial skeleton with a crushed skull and two left dentaries (one unexposed within the same limestone block but revealed by micro-computed tomography scanning) and is formally named Palaeochiropteryx sambuceus sp. nov. A third taxon, Sonor handae gen. et sp. nov., is a probable vespertilionid, evidenced by a dentary fragment with double-rooted p3 and myotodont lower molar morphology, and provides the earliest occurrence of the family Vespertilionidae in the Western Hemisphere. Despite the small number of specimens, these three new taxa comprise the most diverse Eocene assemblage of bats yet found in a single quarry in the Western Hemisphere.
... Neighbouring countries in North Africa (Morocco, Algeria, Libya, Egypt) have all yielded important quantities of Palaeogene mammals, both large and small (Pickford et al. 2008). Tunisia is well known for its Middle Eocene Chambi fauna (Kasserine) (Crochet 1986, Hartenberger 1986, Hartenberger et al. 1985, 1997a, b, 1998, Hartenberger and Marandat 1992, but this locality has yielded few large mammals such as Megalohyrax (Tabuce et al. 2011). ...
Palaeogene large mammals are poorly represented in Tunisia, in contrast to Morocco, Algeria, Libya and Egypt, where abundant and diverse faunas are known. Oligocene proboscideans have been recorded from four localities in Tunisia (Djebel Bou Gobrine, Oued Bazina, Bled Mellaha and Djebel Touila) but little else from this period is known from the country. For this reason it is worth recording the discovery of an arsinoithere tooth fragment from the divide between Oued Cherichera and Oued Grigema, Central Tunisia. This discovery confirms the presence of continental Oligocene strata in the region, and the palaeodistribution of Arsinoitherium 1,300 km to the north-northwest of its previously established range. Arsinoitheres are now known to have been widespread throughout the Afro-Arabian continent. Although palaeoclimatic data for the Oligocene of Tunisia is still scarce, fossil plants suggest that, during the Oligocene, the country enjoyed a tropical to sub-tropical humid climate, in accordance with the presence of Arsinoitherium , Phiomia and an anthracothere, taxa that are also present in the classic Fayum faunas of Egypt and the Ashawq faunas of Oman.
... We present here the 3D digital models of several fossil specimens (Figs 1 and 2; Table 1) documenting a small-bodied primate: Djebelemur martinezi Hartenberger and Marandat, 1992. The fossils were discovered in the western part of central Tunisia (Kassérine region, Djebel Chambi; 2008 and 2009 field campaigns) from lacustrine deposits (CBI-1) dating from the late Early Eocene (Hartenberger and Marandat, 1992;Hartenberger et al., 1997Hartenberger et al., , 2001Tabuce et al., 2011;Marivaux et al., 2015). They were recovered after several rounds of acid processing and wet screening of the indurated calcareous matrix. ...
This contribution contains the 3D models of the fossil remains (maxilla, dentary, and talus) attributed to Djebelemur martinezi, a ca. 50 Ma primate from Tunisia (Djebel Chambi), described and figured in the following publication: Marivaux et al. (2013), Djebelemur, a tiny pre-tooth-combed primate from the Eocene of Tunisia: a glimpse into the origin of crown strepsirhines. PLoS ONE. https://doi.org/10.1371/journal.pone.0080778
... In Pleistocene localities, fossils of bufonids and Discoglossus were found in Filfilla, north-east Algeria (Ginsburg et al., 1968). In Tunisia, fossils of undetermined ranoids were recovered in Eocene deposits (Chambi, Ypresian-Lutetian transition, 48 Ma;Hartenberger et al., 2001). Remains of adult and tadpoles of pipid frogs were also recovered in Early Oligocene deposits (33 Ma) in Libya (Jabal al Hasawinah site), described as a new species, Xenopus (Libycus) hassaunus (Špinar, 1980). ...
Amphibians of North Africa is a comprehensive compilation of available data on the amphibians found in various ecosystems across North West Africa and parts of the Mediterranean region. The book included also the description of a new species of hylid for the northeast Maghreb: Hyla numidica (Holotype-MZB 2019-0924; Terra Typica: Nefza, Tunisia; urn:lsid:zoobank.org:act:154236EB-0CCA-4835-85C7-7516BB6231D1).The first of its kind, this book is essential to identifying and understanding the ecological role of regional herpetofauna and its conservational importance.
... 1C). Vertebrate fossils (including frogs) were collected from a 30 to 100 cm thick layer of freshwater limestone, representing a channel setting and positioned in the basal part of an unnamed sequence of continental, Palaeogene age sediments (Hartenberger et al. 1997(Hartenberger et al. , 2001. ...
Lissamphibians (frogs, salamanders, caecilians, and the extinct Albanerpetontidae) have a near global distribution. Africa, its associated islands (especially Madagascar and the Seychelles) and the Arabian Plate are home to about 27 families (including 15 endemic) and 1135 species of extant lissamphibians or about 38 and 15 %, respectively, of the global totals. The region also contains an extensive, but patchy and somewhat under-appreciated fossil record. Based on published and unpublished information, we provide here the most comprehensive review to date of the lissamphibian fossil record from the region. We also discuss the insights those occurrences provide into past distributions and diversities of lissamphibians in the region and the establishment of the modern fauna. Our review relies on occurrence data from 93 sets of localities of basal Triassic through Holocene age, distributed across 23 countries. As with the modern lissamphibian fauna of the region, the fossil record is dominated by frogs, but there also are notable occurrences of other lissamphibians, including several genera of enigmatic Cretaceous salamanders, one of two known stem caecilians, and the only Gondwanan records for albanerpetontids. Africa is one of only two continents (the other being North America) to have occurrences for all four lissamphibian clades. Twenty named and currently accepted fossil lissamphibian species are recognised from the region: one stem and 14 crown frogs (11 or possibly 12 of which are pipimorphs, 1 alytid, and 1 neobatrachian possibly referable to the otherwise exclusively South American families Ceratophryidae or Calyptocephalellidae); three salamanders; one stem caecilian; and one albanerpetontid. Additional and as yet unnamed taxa are represented in existing collections, and others undoubtedly remain to be discovered. Of the 27 extant lissamphibian families currently recognised from the region, 12 of 22 frog families (including five endemics: Brevicipitidae, Heleophrynidae, Hyperoliidae, Ptychadenidae, and Pyxicephalidae) and the sole salamander family (Salamandridae) have fossil records; at present, none of the known caecilian fossils can be assigned with confidence to any of the four extant families currently recognised in the region. The biogeographic histories of lissamphibians in Africa, its associated islands and the Arabian Plate are characterised by vicariant and dispersal events related to the complex palaeogeographic history of the region. © 2016 Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg
... Note that Mahenge is the only known locality in Africa that has produced an Eocene mammal from south of the equator. 1 -Egypt (Rasmussen et al. 1992;Simons 1992Simons , 1995. 2 -Libya (Arambourg and Magnier 1961;Savage 1969Savage , 1971). 3 -Algeria (Coiffait et al. 1984;Mahboubi et al. 1986;Godinot and Mahboubi 1994). 4 -Tunisia (Hartenberger et al. 1985(Hartenberger et al. , 2001. 5 -Oman (Thomas et al. 1988). 6 -Angola (Pickford 1986). ...
A partial skeleton of a new fossil microbat, Tanzanycteris mannardi, is the oldest placental mammal found in sub-Saharan Africa. It came from early Lutetian (46 Ma) lake sediments in north-central Tanzania. T. mannardi has enlarged cochleae indicating it was capable of a highly derived form of echolocation. Modern bats sharing similar morphology are capable of precise navigation in dense forest undergrowth. The phylogenetic relationships of T. mannardi are unclear. It shares character states with Eocene Hassianycterididae, with extant Microchiroptera, and with Rhinolophoidea within Microchiroptera. T. mannardi is important in documenting early evolution of sophisticated bat echolocating abilities and demonstrating that Tanzanian crater lakes offer an opportunity for future discoveries of Eocene mammals from the African interior.
... The current picture yielded by the poor fossil record of Ceno− zoic marsupials suggests that the peradectids were widely distrib− uted in the Old World, since they are reported from the early Eocene of Tunisia (Chambi) with Kasserinotherium tunisiense (Crochet 1986;Hartenberger et al. 2001), and from both the early Oligocene of Egypt (Gebel El Qatrani, Fayum) and the Arabic peninsula (Taqah, Sultanate of Oman) with Qatranitherium afri− canum (Simons and Bown 1984;Crochet et al. 1992). The pera− dectids are also known in the early Miocene of China (Songlin− zhuang) with Sinoperadectes clandestinus (Storch and Qiu 2002) and from the middle Miocene of Thailand (Mae Long and Li Bas− ins) with Siamoperadectes minutus (Ducrocq et al. 1992;Mein and Ginsburg 1997). ...
Didelphimorph marsupials were widely distributed in Asia during the Cenozoic, but their occurrence in the Indian subcontinent has not so far been demonstrated. Here, we describe a new herpetotherfid marsupial Asiadidelphis akbarbugtii sp. nov. from the early Oligocene Bugti Member of the Chitarwata Formation, Bugti Hills, Pakistan. The discovery of the herpetotheriids in the Oligocene of Pakistan represents the most southern occurrence of the family, which was thought to have occurred only to the north of the Alpine-Himalayan Suture. Our data suggest that episodic faunal exchanges occurred between the Asian mainland and the Indian subcontinent during the late Paleogene, and that the southern Asian faunas were not as completely isolated by the Himalayan chain as formerly believed.
... The current picture yielded by the poor fossil record of Ceno− zoic marsupials suggests that the peradectids were widely distrib− uted in the Old World, since they are reported from the early Eocene of Tunisia (Chambi) with Kasserinotherium tunisiense (Crochet 1986;Hartenberger et al. 2001), and from both the early Oligocene of Egypt (Gebel El Qatrani, Fayum) and the Arabic peninsula (Taqah, Sultanate of Oman) with Qatranitherium afri− canum (Simons and Bown 1984;Crochet et al. 1992). The pera− dectids are also known in the early Miocene of China (Songlin− zhuang) with Sinoperadectes clandestinus (Storch and Qiu 2002) and from the middle Miocene of Thailand (Mae Long and Li Bas− ins) with Siamoperadectes minutus (Ducrocq et al. 1992;Mein and Ginsburg 1997). ...
Didelphimorph marsupials were widely distributed in Asia during the Cenozoic, but their occurrence in the Indian subcontinent has not so far been demonstrated. Here, we describe a new herpetotheriid marsupial Asiadidelphis akbarbugtii sp. nov. from the early Oligocene Bugti Member of the Chitarwata Formation, Bugti Hills, Pakistan. The discovery of the herpetotheriids in the Oligocene of Pakistan represents the most southern occurrence of the family, which was thought to have occurred only to the north of the Alpine−Himalayan Suture. Our data suggest that episodic faunal exchanges occurred between the Asian mainland and the Indian subcontinent during the late Paleogene, and that the southern Asian faunas were not as completely isolated by the Himalayan chain as formerly believed.
... Dental and postcranial remains of Macroscelididae are common in the locality of Chambi and all are referred to a single species, viz. Chambius kasserinensis (Hartenberger 1986;Hartenberger et al. 1997Hartenberger et al. , 2001Tabuce et al. 2007). The systematic attribution of the isolated petrosals from Chambi to a macroscelid, and thus to Chambius, is based on: (1) overall similarities with extant macroscelids, and (2) results of a cladistic analysis based on 52 petrosal and inner ear characters, coded for a wide sample of taxa including afrotheres and insectivore-grade mammals (see Results and discussion section). ...
Macroscelidea (elephant-shrews or sengis) are small insectivorous mammals restricted to Africa; they belong to the super-cohort Afrotheria along with other insectivorans (aardvarks, tenrecs and golden moles) and ungulates (elephants, sea cows and hyraxes). Though their fossil record extends back to the Eocene, cranial remains of Palaeogene elephant-shrews, including the middle and inner ear structure, remain unknown. Two macroscelid isolated petrosal bones are described from the late Early–early Middle Eocene Djebel Chambi locality in Tunisia. Chambius kasserinensis is the only macroscelid represented in this locality. A cladistic analysis based on petrosal and inner ear characters highlights the crucial interest of the petrosal and inner ear morphology for understanding the evolution of Macroscelididae. It confirms the attribution of these isolated petrosals to C. kasserinensis. This hypothesis is supported by a common pattern of circulatory system, the morphology of the rostral and caudal tympanic processes, and the shape of the cochlea. In addition, Chambius appears to be the basal-most taxon among the macroscelid sample; this position is supported by the lack of some specializations of the middle ear such as inflated ossicular bones and pneumatized bulla. The presence of a secondary common crus in Chambius suggests a convergent loss of this structure, at least in Macroscelidea and Tenrecoidea. The petrosal and inner ear characters support the clade Afroinsectivora, which gathers macroscelids with other endemic African insectivorans (tenrecs and golden moles), reinforcing the hypothesis of an African origin of macroscelids. The petrosal bone and inner ear characters provide further morphological support for the debated clade (Petrodromus, Elephantulus rozeti, Macroscelides). New data underlines the fact that the cranial arterial pattern of the Eocene macroscelid Chambius was already similar to that of modern macroscelid species. It also suggests that early elephant-shrews were probably not as capable of hearing low frequencies as their extant representatives.
... Given the fragmentary nature of their fossil record, additional evidence is needed to further our understanding of this paleontological scenario. Here, we describe new fossils attributable to Djebelemur martinezi, a<50 Ma primate from the late Early to early Middle Eocene deposits of Djebel Chambi, Tunisia (Fig. 1)414243. This taxon was originally interpreted as a cercamoniine adapid primate (i.e., Adapiformes) based on limited fossil evidence primarily from its lower dentition (a mandible preserving p3-m3 [41]). ...
Background:
Molecular clock estimates of crown strepsirhine origins generally advocate an ancient antiquity for Malagasy lemuriforms and Afro-Asian lorisiforms, near the onset of the Tertiary but most often extending back to the Late Cretaceous. Despite their inferred early origin, the subsequent evolutionary histories of both groups (except for the Malagasy aye-aye lineage) exhibit a vacuum of lineage diversification during most part of the Eocene, followed by a relative acceleration in diversification from the late Middle Eocene. This early evolutionary stasis was tentatively explained by the possibility of unrecorded lineage extinctions during the early Tertiary. However, this prevailing molecular view regarding the ancient origin and early diversification of crown strepsirhines must be viewed with skepticism due to the new but still scarce paleontological evidence gathered in recent years.
Methodological/principal findings:
Here, we describe new fossils attributable to Djebelemur martinezi, a≈50 Ma primate from Tunisia (Djebel Chambi). This taxon was originally interpreted as a cercamoniine adapiform based on limited information from its lower dentition. The new fossils provide anatomical evidence demonstrating that Djebelemur was not an adapiform but clearly a distant relative of lemurs, lorises and galagos. Cranial, dental and postcranial remains indicate that this diminutive primate was likely nocturnal, predatory (primarily insectivorous), and engaged in a form of generalized arboreal quadrupedalism with frequent horizontal leaping. Djebelemur did not have an anterior lower dentition as specialized as that characterizing most crown strepsirhines (i.e., tooth-comb), but it clearly exhibited a transformed antemolar pattern representing an early stage of a crown strepsirhine-like adaptation ("pre-tooth-comb").
Conclusions/significance:
These new fossil data suggest that the differentiation of the tooth-comb must postdate the djebelemurid divergence, a view which hence constrains the timing of crown strepsirhine origins to the Middle Eocene, and then precludes the existence of unrecorded lineage extinctions of tooth-combed primates during the earliest Tertiary.
... Scarce but rich North African Paleogene deposits have provided a great amount of fossil mammals since the last century (Andrews, 1906;Mahboubi et al., 1986;Hartenberger et al., 1998Hartenberger et al., , 2001Rasmussen and Simons, 2000;Gheerbrant et al., 2003;Tabuce et al., 2008;Gheerbrant, 2009;Ravel et al., 2011;see Seiffert, 2010 for an overview). Africa was partly isolated from other continents during the Paleogene (Gheerbrant and Rage, 2006;Tabuce et al., 2008), and given the great scarcity of Paleogene localities in Africa, these fossils are dramatically important to understand the early evolution of the endemic mammalian fauna of this continent. ...
The ear region of mammals has long been considered as morphologically very conservative and accordingly, phylogenetically useful. In this study, the anatomy of the petrosal and bony labyrinth (osseous inner ear) of Numidotherium (Proboscidea) and Arsinoitherium (Embrithopoda) are investigated and compared in order to assess the evolution of ear region characters in proboscideans and embrithopods. Using a cladistic analysis across Paenungulata based on ear region characters only, we found that Arsinoitherium is surprisingly best placed as a crown proboscideans to the exclusion of Numidotherium and Phosphatherium, which results in the paraphyly of proboscidean. The clade Proboscidea is actually well supported by dental and post-cranial characters, and we propose that this result underlines the great amount of morphological convergences in the ear region of Embrithopoda and Proboscidea, possibly due to convergent evolution of capabilities toward infrasonic hearing.
... Djebel Chambi is located at the northern side of Kasserine ''island'', considered by many authors as an emerged area from the Senonian to the lower Miocene (e.g., Burollet, 1956;Sassi et al., 1984;Zouaghi et al., 2005). Djebel Chambi is also famous for having produced one of the most diverse African vertebrate faunas dated to the late early to early middle Eocene (localities of Chambi-1 and Chambi-2) (e.g., Hartenberger et al., 2001;Tabuce et al., 2007a;Ravel et al., 2012;Benoit et al., 2013, in press a, in press b; Mourer-Chauviré et al., in press). The fossiliferous level of Bir Om Ali conformably overlies the fossiliferous strata of Chambi-1, which is located 3 km southwest (Fig. 1). ...
... This material came from deposits that are almost exclusively estuarine or marine; however, Khare (1976) noted that the fossils may have been transported from fresh water. The latter view, that periodic connections existed between Afro-Arabia and Eurasia at times in the Eocene, has been suggested previously based on mammalian fossil material (e.g., Hartenberger et al., 2001). Van Couvering and Van Couvering (1975) noted evidence of a one-way corridor from Eurasia to Africa, which could be explained by the movement of microplates, such as those postulated by McKenzie (1970). ...
Upper Eocene and lower Oligocene deposits of the Jebel Qatrani Formation, Fayum Depression, Egypt, have provided a number of skulls and other bones that belong to a species of snakehead (Channidae). This material is morphologically more similar to species of Parachanna, rather than species of Channa, and is here described as a new species, Parachanna fayumensis. It differs from the other species of the genus in possessing a prominent tooth patch on the posterior end of the parasphenoid. This new species of channid is the oldest member of the family known from Africa. Prior to this record, the oldest African fossil channid material was found in Mio-Pliocene deposits. This lack of information on the African fossil record leads to biogeographic reconstructions in which channids were believed to have arisen in Asia and invaded Africa through fresh waters, only after the two continents were connected in the Miocene. The Egyptian material shows that channids were in the fresh waters of Africa in the latest Eocene. Either a freshwater connection existed between Africa and Asia before or during the late Eocene, or members of the Channidae were able to migrate through marine waters to attain their current distribution.
... These localities share common occurrences of the zegdoumyid Zegdoumys, the macroscelidid Chambius, and the hyracoid Titano hyrax tantulus, all of which are absent from younger localities. Chambi, on the basis of charophytes and mammalian biostratigraphic data, is believed to be late early to early middle Eocene in age (Hartenberger et al., 2001), but the age of this locality is poorly constrained. The El Kohol Formation, dated from middle Ypresian, has yielded abundant remains of early proboscideans. ...
Despite numerous discoveries that have considerably enriched the African-Arabian Tertiary fossil record over the last decades, our knowledge of the evolutionary history of many continental African vertebrate groups during the Paleogene period remains inade-quate, particularly when it is compared with the fossil records of Europe or North Amer-ica. The Eocene Epoch in Africa is especially poorly documented, being restricted to few fossiliferous localities. Our understanding of the early Tertiary emergence, diversifi cation, and paleobiogeographic history of African-Arabian mammals has been further hindered by the lack of a precise temporal framework for these sites. We conducted magnetostratigraphic analy-ses, associated with biostratigraphic studies, in the fossiliferous sequences exposed in the northwestern Hammadas of the Saharan Platform in the Glib Zegdou area and in the Saharan Atlas at the El Kohol locality (Algeria) to further defi ne the age of these Eocene continental deposits. Based on bio-stratigraphic constraints, the six polarity zones identifi ed in the El Kohol section can be correlated with chrons C24n to C22r, pro-viding the fi rst direct age estimates for the El Kohol fossiliferous strata between 52 and 51 Ma. Correlation to the geomagnetic po-larity time scale, using previously published biostratigraphic data for the Glib Zegdou fauna, suggests an age ranging between 49 and 45 Ma for this section. The high-resolution magnetostratigraphic study of the poorly known continental Eo-cene Epoch of Algeria provides new in-sights into the early Tertiary stratigraphy of northwest Africa. The placement of the Algerian localities into a consistent chrono-logical framework constitutes considerable advancement to achieve biostratigraphic cor-relation of the Paleogene African-Arabian mammal localities.
... The possible primitive primate Altiatlasius kouchelli is listed for sake of completeness, but the taxon is not included in analyses, as its systematic position is uncertain. Sources: Andrews, 1901;Andrews and Beadnell, 1902;Matsumoto, 1924;Sudre, 1975Sudre, , 1979Coppens et al., 1978;Crochet, 1984Crochet, , 1986Mahboubi et al., 1984Mahboubi et al., , 1986Sigé, 1985Sigé, , 1991Sigé, , 1988Crochet et al., 1990Crochet et al., , 1992Sigé et al., 1990Sigé et al., , 1994Court and Hartenberger, 1992;Hartenberger and Marandat, 1992;Rasmussen et al., 1992;Sudre et al., 1993;Vianey-Liaud et al., 1994;Court, 1995;Gheerbrant et al., 1996;Gagnon, 1997;Gheerbrant et al., 1993Gheerbrant et al., , 1998Gheerbrant et al., , 2001Gheerbrant et al., , 2002Thomas et al., 1999;Tabuce et al., 2000Tabuce et al., , 2001Hartenberger et al., 2001;Seiffert et al., 2003Seiffert et al., , 2005a. diversity. ...
Recent fossil discoveries, phylogenetic analyses, revised reconstructions of continental drift, and accumulating molecular evidence have all yielded new information relating to anthropoid origins within the broader context of primate evolution. There is an emerging consensus among molecular studies that four superorders of eutherian mammals can be recognized: Afrotheria, Euarchontoglires (to which primates belong), Laurasiatheria, and Xenarthra. Overall, molecular phylogenies for mammals agree with some statistical analyses of the primate fossil record in indicating an early origin for primates around 85 Ma ago, and the divergence of haplorhines and strepsirrhines at ca. 77 Ma. Such an ancient date for the origin of haplorhines is some 17 Ma prior to the first known possible primate, and some 22 Ma before the earliest fossil evidence of undoubted euprimates. Because anthropoid fossils date back at least to the late Eocene and perhaps to the middle Eocene, and given indications of an early origin for primates, it is unlikely that ancestral anthropoids arose within any other currently known clade of fossil primates (adapiforms, omomyiforms, strepsirrhines, or tarsiiforms). Implications of new molecular, morphological, and biogeographic lines of evidence are explored with respect to the likely time and place of the origin of anthropoids. Four competing, testable hypotheses are reviewed in detail: 1) the Paratethyan hypothesis, 2) the continental Asian hypothesis, 3) the Indo-Madagascar hypothesis, and 4) the African hypothesis. A case is made that current evidence best supports a relatively ancient Gondwanan origin for primates, as well as a Gondwanan (African or Indo-Madagascan) origin for anthropoids at least as old as that of any other currently documented major primate clade. Available fossil evidence at present seems to be most compatible with the African hypothesis, but it is noteworthy that primates are included not in Afrotheria but in Euarchontoglires.
... Intensive palaeontological work in Early to Middle Eocene North African localities in Algeria (Gour Lazib) and Tunisia (Chambi) have led to the recovery of several new fossils of mammals. Based on biostratigraphic studies, the Gour Lazib formation has been assigned either a late Ypresian or Early Lutetian age (Tabuce et al. 2004) and the Chambi locality an Ypresian age (Hartenberger et al. 2001). The newly discovered Algerian fossils document rodents, primates (particularly assigned to Algeripithecus, the oldest alleged anthropoid), creodonts, chiropterans, insectivores and hyraxes. ...
The phylogenetic pattern and timing of the radiation of mammals, especially the geographical origins of major crown clades, are areas of controversy among molecular biologists, morphologists and palaeontologists. Molecular phylogeneticists have identified an Afrotheria clade, which includes several taxa as different as tenrecs (Tenrecidae), golden moles (Chrysochloridae), elephant-shrews (Macroscelididae), aardvarks (Tubulidentata) and paenungulates (elephants, sea cows and hyracoids). Molecular data also suggest a Cretaceous African origin for Afrotheria within Placentalia followed by a long period of endemic evolution on the Afro-Arabian continent after the mid-Cretaceous Gondwanan breakup (approx. 105-25 Myr ago). However, there was no morphological support for such a natural grouping so far. Here, we report new dental and postcranial evidence of Eocene stem hyrax and macroscelidid from North Africa that, for the first time, provides a congruent phylogenetic view with the molecular Afrotheria clade. These new fossils imply, however, substantial changes regarding the historical biogeography of afrotheres. Their long period of isolation in Africa, as assumed by molecular inferences, is now to be reconsidered inasmuch as Eocene paenungulates and elephant-shrews are here found to be related to some Early Tertiary Euramerican 'hyopsodontid condylarths' (archaic hoofed mammals). As a result, stem members of afrotherian clades are not strictly African but also include some Early Paleogene Holarctic mammals.
We here describe Terastiodontosaurus marcelosanchezi, a new amphisbaenian genus and species from the Eocene of Chambi, Tunisia. Using micro-computed tomography (μCT), we document the peculiar anatomy of the new taxon, which is characterized by extreme dental morphology, including one massive tooth on the maxilla and dentary, flat cheek teeth, and an array of other diagnostic features that readily differentiate it from all other amphisbaenians. We also redescribe the oldest named African amphisbaenian, Todrasaurus gheerbranti, from the late Palaeocene of Morocco, using μCT. Phylogenetic analysis recovers Terastiodontosaurus and Todrasaurus as sister taxa and provides strong support for a sister-group relationship of those two large-toothed amphisbaenians with extant Trogonophis. Accordingly, Todrasaurus shows that the divergence of crown Trogonophidae occurred much earlier than currently thought. Our survey of μCT scans reveals that Terastiodontosaurus, Todrasaurus, and Trogonophis are characterized by a great enamel thickness on their teeth, a feature that is absent in other examined amphisbaenians. Size estimates show that Terastiodontosaurus was the largest known amphisbaenian ever to have lived, with an estimated skull length of >5 cm. Based on new muscle data of Trogonophis, we estimate very high bite forces for Terastiodontosaurus, which would allow it to crush a wide variety of snails.
Pan‐trionychids (soft‐shelled turtles) are known from the Neogene and Quaternary fossil record of Africa. They have so far been totally absent in Palaeogene localities of the continent although they are commonly found in Palaeogene localities across Europe, North America, and Australasia; as such, their absence from the Palaeogene of Africa has been envisaged to be genuine. Here, I describe a large costal from the Eocene of Mali, which documents for the first time the presence of pan‐trionychid turtles in the Palaeogene of the then isolated Afro‐Arabia. The affinities of the new Malian pan‐trionychid with Neogene and Quaternary forms from Africa are discussed, as well as its biogeographic origins and potential dispersal scenarios from other landmasses to Afro‐Arabia during the Palaeogene.
We define 17 African land mammal ages, or AFLMAs, covering the Cenozoic record of the Afro-arabian continent, the planet’s second largest land mass. While fossiliferous deposits are absent on the eroded plateau of the continent’s interior, almost 800 fossil genera from over 350 locations have now been identified in coastal deposits, karst caves, and in the Neogene rift valleys. Given a well-developed geochronologic framework, together with continuing revision to the fossil record—both stimulated by the story of human evolution in Africa—and also to compensate for the variation in fossil ecosystems across such great distances, the AFLMAs are biochronological units defined by type localities, and not biozones to be recognized by the occurrence of certain genera. Disparities are notable: Africa is the highest of all continents, but almost every Paleogene locality was formed at sea level; the fossil record of its great rainforest ecosystem remains virtually unknown; and the Paleogene fauna is relatively isolated, whereas the Neogene begins with open exchange with Laurasia following the Tauride collision, with a simultaneous opening of the East African rift valleys in which the newly revolutionized fauna is abundantly preserved. Notably, the continent-wide and comprehensive documentation of the African mammalian record reveals an unparalleled rate of transformation in the hominin lineage, unmatched by any other group, in response to the Neogene expansion of the open-country ecosystem.
A large collection of lizard vertebrae from northern Africa represents the oldest unambiguous occurrence of the genus Varanus. The fossils come from late Eocene and early Oligocene freshwater deposits of the Fayum, Egypt, an area noted for many significant primate finds. The recovery and identification of this material indicate that the genus Varanus arose in Africa, before dispersing to Australia and Asia. This dispersal occurred prior to the early to mid-Miocene, by which time fossil Varanus are known from Australia and Eurasia. Although the dispersal route remains unknown, the lizard material reported here supports the hypothesis that a corridor existed allowing freshwater and terrestrial organisms to cross from Africa to Asia.
Among the new dental remains from the late Early Eocene of Chambi (Kasserine area, Tunisia) is a large-sized upper molar of a new bat species, Witwatia sigei nov. sp. (Chi-roptera, Vespertilionoidea, Philisidae), described herein. The locality of Chambi has revealed evidence for an early appear-ance of two modern microchiropteran superfamilies in Africa: Dizzya exsultans, a Philisidae, which is considered to be an archaic Vespertilionoidea, and an indeterminate Rhinolophoi-dea. In addition to D. exsultans, the new species, W. sigei, is the second representative of the Philisidae in this locality. W. sigei extends back to the late Early Eocene the occurrence of the genus Witwatia, which was previously only reported from the early Late Eocene of the Fayum (BQ-2, Egypt). By analogy with the largest extant microbats, the large size of Witwatia suggests a tendency to the opportunistic diet of this taxon, thereby con-trasting with the strict insectivory characterizing primitive bats found in other continents in the same epoch.
The fossils reported here were collected from the sites of Thaytiniti and Taqah, Oman. They comprise the first known Palaeogene hyracoids from the Arabian Peninsula. Three (or four) species are represented in the collections, cf. Saghatherium bowni, Thyrohyrax meyeri and two unidentified medium sized brachyodont species (? Megalohyrax sp.,) similar to undescribed fossils from quarry L 41 Fayum, Egypt. Comparisons of hyracoids from Oman and Egypt indicate that the Omani sites correlate closely with Faunal Zone 1 of the Fayum. This in turn indicates that all four faunal zones of the Fayum are Oligocene in age. -Authors
21 species have been recovered from six Tertiary localities of Algeria. Their stratigraphical indications allow to determine the age of non-marine deposits: those of Atlas Saharian and of Hauts Plateaux are attributed to the Ilerdian and those of the north-western Saharian Hammada, possibly more recent, to the Upper Ilerdian up to the Lower Lutetian. We place the El Biod locality (Tinh'ert Plateau, Central Sahara) in the Priabonian-Stampian interval. The floristic relationships with northern regions allow correlations with the biozonation established in Europe.
Twelve years ago, just after the publication of the volume dedicated to platyrrhine origins (Ciochon and Chiarelli, 1980), Hoffstetter made a remarkable attempt to synthesize information and debates about simiiform (= anthropoid) evolution. What he wrote then holds true today: “We are still far away from a consensus about phylogeny and biogeographical history, that is to say about the evolution, of these animals [primates]” (Hoffstetter, 1982, p. 242, translated). In this paper, Hoffstetter drew attention to the inconveniences of the vernacular term “anthropoid,” ambiguous in many languages. In all these languages, anthropoid means apes, often great apes only, which is quite different from “Anthropoidea.” Furthermore, the Code of Zoological Nomenclature recommends that the suffix -oidea be reserved for super-families, an almost universal use in mammalogy. Hoffstetter added: “As Szalay and Delson (1979) revealed the existence of a genus Anthropus, synonym of Homo, Anthropoidea is a synonym of Hominoidea and must be definitely rejected” (Hoffstetter, 1982, translated). I follow the authors who, since this time, use Simiiformes Hoffstetter, 1974 instead of Anthropoidea Mivart, 1864 (vernacular: simiiform or simian).
Dental remains of a new hyracoid mammal, Titanohyrax tantulus sp. nov., are described from Eocene deposits at Chambi in Tunisia. This new form is one of very few Palaeogene hyracoid occurrences outside the Early Oligocene Jebel Qatrani Formation of Egypt. Very small size and low crown height reflect the primitive nature of the new species relative to other members of the genus. The Chambi rodent fauna and data from charophytes provisionally indicate an early Eocene age for the locality. T. tantulus is thus potentially the oldest known representative of the order Hyracoidea. -Authors
We detail here the study of Phosphatherium escuilliei from the Paleocene of the Ouled Abdoun Basin, 040 Morocco. This fossil is both the oldest proboscidean and the oldest modern ungulate. The species, known by two specimens, is dated as Thanetian on the basis of the foraminifera and the selachians identified in the matrix. P. escuilliei shows several derived proboscidean features, especially the true lophodont pattern of the molars, but also remarkable primitive features such as a vestigial dilambdodonty and a very small size, the smallest reported in the order. The vestigial dilambdodonty is interpreted here as a reminiscence of the pantomesaxonian super-group. Primitive tethythere features are also noticed, especially the orbit in anterior position and the occurrence of a postentoconule. Numidotherium koholense from El Kohol (late Early Eocene of Algeria) is the most closely related proboscidean to P. escuilliei, and the two species are included in the same family Numidotheriidae. However, the available material shows only one possible synapomorphy: the absence of lingual cingulum in the upper molars. P. escuilliei is the most primitive known lophodont proboscidean. The dP4/ of P. escuilliei strongly recalls the single known tooth of the enigmatic species Khamsaconus bulbosus from the Ypresian site of N'Tagourt 2, Morocco. Shared features such as the occurrence of a postentoconule and the incipient bilophodonty indicate affinities of K. bulbosus with tethytheres, and others such as the absence of conules are even possibly suggestive of affinities with lophodont proboscideans. The K. bulbosus morphology remains however clearly more primitive than that of P. escuilliei, especially in being more bunodont. Phosphatherium, Numidotherium, Barytherium and even Khamsaconus raise the problem of a lophodont ancestral proboscidean morphotype, which is in accordance with the recent hypothesis of Court (1995) regarding Moeritherium as more derived than Numidotherium. P. escuilliei supports the early age of the radiation of modern orders of mammals, which was earlier than the late Paleocene, as well as its “explosive” pattern if it is closely connected to the extinctions of the K/T boundary. It shows also, more clearly than the taxa from the Adrar Mgorn fauna (late Paleocene of Morocco), the old age of the eutherian presence in the Arabo-African domain. It also supports the African origin of true proboscideans.
The carbonates studied include several types 1.(1) Lacustrine dolomites which have resulted from the concentration of products leached from neighbouring Paleozoic and/or Mesozoic rocks. Certain well-crystallised limestones contain phantoms of rhombohedra, and are the result of dedolomitisation.2.(2) Limestones deposited in fluviatile channels, or at the outflow of rivers into lakes, which are always sandy (with corroded quartz) and often contain calcitic intraclasts, bioclasts and oncolites. (Cyanophyceae, Rivulariaceae.)3.(3) True lacustrine limestones, with an entirely subaqueous diagenetic evolution, which are homogenous, or show evidence of reworking: breccias, gravels, coated gravels. Burrowing animals have left traces of their activity (in particular striotubules∗) in sediments deposited on the bottom lakes.4.(4) “Paludine” limestones which have resulted from emersion of lacustrine muds, have been affected by soil processes and show various signs of reworking: penetration of roots, burrows, nodulisation∗, marmorisation, etc. All these phenomena occur together giving facies and microfacies which may be very complicated.5.(5) Silicification, which may occur in all continental limestones, including lacustrine or paludine types. It exhibits various morphological aspects; on a microscopic scale, it is an accumulation of opal and/or thin veins of chalcedony.These different facies are not distributed haphazardly, but constitute sequences which are repeated, with some variations, in stratigraphical series. Lataral variation in the style of vertical sequence in a basin enables the reconstitution of the paleo-environment at a given period.
A new Adapiform Primate from the locality of Chambi (Tunisia) is described and assigned toDjebelemur martinezi gen.nov., sp. nov. Holotype CBI 33, fragment of left mandible with P/3-M/3. Other material includes a lower and two upper
isolated molars and one probable lower canine. This new material is assigned to the Adapidae.
Thèse. Sc. Nat. Paris. 1948. No 3074.
Sea level sequence stratigraphy is especially valuable for correlating marine stages on passively subsiding continental margins. The continental Gebel Qatrani Formation in Egypt is separated from the underlying marine Qasr el-Sagha Formation by a major unconformity with a minimum of 76 m of section missing due to erosion and/or non-deposition. This unconformity is constrained by Priabonian planktonic foraminifera in the Gehannam Formation to be younger than early late Eocene and it is constrained by radiometric ages and a great thickness of Gebel Qatrani Formation to be older than late Oligocene. The only "type-1" sequence boundary within these age constraints that involved a low enough sea stand to explain the unconformity was at the Priabonian-Rupelian (Eocene-Oligocene) boundary, which means that the Gebel Qatrani Formation is entirely Oligocene in age. This corroborates earlier age assignments based on invertebrate and vertebrate faunal succession and it is consistent with new paleomagnetic evidence. The Gebel Qatrani Formation has yielded the earliest primates of anthropoid grade and the evolutionary emergence of higher primates may be related to profound environmental change during the Eocene-Oligocene transition. Peer Reviewed http://deepblue.lib.umich.edu/bitstream/2027.42/30939/1/0000609.pdf
Nouveou mammifèze insedivore (?Lipotypla ?Erinaceomorpha) de 1’ Eocène
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Un gisement de la zone de Mammifères d’Issel (Éocène moyen)
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Kasserinotherium tunisiense nov. gen., nov. sp., troisième marsupial découvert en Afrique
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Les remplissages karstiques polyphasés (Eocène, Oligocène, Pliocène) de St-Maximin (phosphorites du Gard) et leur apport à la connaissance des faunes européennes
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Charophytes du Tertaire continental de l’Algérie
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Oligocene age for the Gebel Qatrani Formation
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Kasserinotherium tunisiense nov. gen., nov. sp., troisième marsupial découvert en Afrique (Eocène inférieur de Tunisie)
- J.-Y Crochet
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Un gisement de la zone de Mammifères d’Issel (Éocène moyen) dans les calcaires lacustres d’Aumelas (Hérault), Comptes-rendus som
- J.-L Hartenberger
- J-L Hartenberger
Découverte de Mammifères d’âge Eocène inférieur en Tunisie Centrale, Comptes-rendus de l’Académie des Sciences
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