Contribution à la connaissance des occupations préhistoriques au Pléistocène supérieur en Tunisie

The Upper Pleistocene in Tunisia is characterized by cyclic climatic alternations between rainy and arid phases. The steppic landscape occupies a large area of the country, in particular in the Centre and the South. Even during the humid phases, the savannah landscape still dominant. The Upper Pleistocene prehistoric sites in Tunisia, such as Oued el Akarit, Aïn Mhrotta, Oued el Bey, El Guettar, Aïn Métherchem and Aïn el Guettar, which are the subject of this study, are all open-air sites, located mainly in Central and southern Tunisia, usually to the south of the springs that are still functioning today, or on the border of the wadis from which their names derive. Prehistoric occupations and human behavior are mainly subject to the climatic shifts occurred during the Upper Pleistocene.     

Palaeoenvironments of vertebrates on the southern shore of Tethys: The nonmarine Early Cretaceous of Tunisia

Through the Late Jurassic and Early Cretaceous, the southern shore of the Tethys Ocean migrated north and south over short distances. These vicissitudes are documented in the ‘continental intercalaire’, a long series of mainly non-marine sediments deposited across North Africa. A combined taxonomic, physical and chemical taphonomic study provides independent lines of evidence for reconstruction of palaeoenvironments within this marginal marine setting.The Douiret, Chenini and Oum ed Diab formations from the Tataouine basin of southern Tunisia span the later part of the Early Cretaceous. Microvertebrates from four sites in these formations show different modes of physical abrasion, time averaging, and mixing, based on a taphonomic analysis using a combination of physical and chemical methods. The taxonomic composition of each assemblage, and trends in rare earth element (REE) compositions of bones, from each locality were used as independent lines of evidence to indicate differences in early depositional environments. The Jebel Boulouha assemblage (Douiret Formation) is interpreted as a terrestrial carbonate-rich environment with relatively little mixing. The Touil el Mra assemblage (Oum ed Diab Formation) suggests a marginal marine environment with some mixing of previously interred bones. The Oued el Khil assemblage (Chenini Formation) and the Oum ed Diab assemblage (Oum Ed Diab Formation) are more equivocal, suggesting mixed freshwater and marine influences. Interpreting salinity in marginal marine settings is difficult, and best attempted from multiple, independent lines of evidence. We suggest that REE geochemistry can contribute to palaeoenvironmental reconstruction when used in combination with other, independent physical, palaeontological and /or geochemical methods.     

A new crocodylomorph tooth assemblage from the Tataouine Basin and comments on the stratigraphic context of the Douiret Formation

A new crocodylomorph tooth assemblage from the Lower Member of the Douiret Formation (uppermost Jurassic–lowermost Cretaceous) in southeastern Tunisia is described. These teeth have been grouped into five morphotypes based on their diagnostic morphological features. The sample includes teeth, which have not been reported until now, such as non-Machimosaurini teleosauroid teeth, and labiolingually compressed tooth crowns with cutting edges devoid of serrations. Certain minor morphometric differences have been interpreted as ontogenetic or being possibly related to the position of the teeth in the jaws. Three trophic entities have subsequently been defined on the basis of the morphological characteristics of the teeth, and a possible niche partitioning between at least three crocodylomorph taxa is highlighted for the first time in the Tataouine Basin. On the other hand, the stratigraphic context of the Douiret Formation is thoroughly discussed in order to show how stratigraphic uncertainties could significantly impact previous attempts at reconstructing the macroevolutionary history of Crocodylomorpha. The latter should be treated with caution when the stratigraphic context of the fossil bearing formation remains a subject of debate.     

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.     

Acoustic Telemetry Validates a Citizen Science Approach for Monitoring Sharks on Coral Reefs

Citizen science is promoted as a simple and cost-effective alternative to traditional approaches for the monitoring of populations of marine megafauna. However, the reliability of datasets collected by these initiatives often remains poorly quantified. We compared datasets of shark counts collected by professional dive guides with acoustic telemetry data from tagged sharks collected at the same coral reef sites over a period of five years. There was a strong correlation between the number of grey reef sharks (Carcharhinus amblyrhynchos) observed by dive guides and the telemetry data at both daily and monthly intervals, suggesting that variation in relative abundance of sharks was detectable in datasets collected by dive guides in a similar manner to data derived from telemetry at these time scales. There was no correlation between the number or mean depth of sharks recorded by telemetry and the presence of tourist divers, suggesting that the behaviour of sharks was not affected by the presence of divers during our study. Data recorded by dive guides showed that current strength and temperature were important drivers of the relative abundance of sharks at monitored sites. Our study validates the use of datasets of shark abundance collected by professional dive guides in frequently-visited dive sites in Palau, and supports the participation of experienced recreational divers as contributors to long-term monitoring programs of shark populations.     

The influence of gummy sharks, Mustelus antarcticus, on observed fish assemblage structure

Large predatory fish that alter the behaviour of smaller species may affect visual estimates of abundance by making organisms more or less difficult to observe and quantify. To evaluate the non-consumptive influence of large predators on rocky reef fishes observed with Baited Remote Underwater Video (BRUV), we tested the hypothesis that fish assemblages monitored when large (ca 1.5 m) piscivorous sharks, Mustelus antarcticus, were present differ from those observed when sharks were absent. We did this in two ways using recordings from rocky reefs in Batemans Marine Park, NSW, Australia. First, we examined 6 min of each 30 min tape; 3 min when sharks were present and 3 min when they were absent, in a paired sample design from 17 sites. Second, we compared fish assemblages for complete tapes (30 min) at sites with sharks present compared to sites where they were absent. The diversity and total abundance of fishes was consistently lower in the presence of sharks; we detected significant assemblage-wide change (PERMANOVA, P?<?0.05). Importantly, the diversity and total abundance of fishes for complete tapes (30 min) decreased in the presence of sharks by 18 % and 36 %, respectively, underscoring the likelihood that sharks influence observed fish assemblages. Individual species responses were variable and while the abundance of many fish species decreased in response to sharks, others increased 6-fold. Overall, these results highlight the need to consider systematic differences in the abundance of large predators to avoid biases when testing hypotheses about fish assemblages using non-destructive visual methodologies.     

Occurrence of an in situ fern grove in the Aptian Douiret Formation, Tataouine area, South-Tunisia

A level with fern stumps was discovered in the Aptian Douiret Formation, South-Tunisia. These stumps are preserved as external moulds or casts, without any preservation of anatomical structures. These stumps are considered to be affiliated with the numerous fossil plants assigned to the fern genera Alstaettia and Piazopteris that are widely distributed in coeval strata from the same region, either as leaf imprints or as permineralized remains. The record of in situ fossil forests for the Southern Hemisphere reveals that their systematic components are different, i.e. mainly corystosperms and/or conifers, and rarely under tidal influence. The way this fern grove settled in a margino-littoral environment is discussed.     

Nouveaux gisements à rongeurs dans le Miocène moyen continental du Bassin d'Aït Kandoula (Maroc)

The Azdal section in the Aït Kandoula basin show a succession of four fossiliferous layers delivering micromammals particularly rodents are very abundant than the great mammals. Mellalomys kandoulati nov. sp. is described in the oldest layer of the section. Compared to the Middle Miocene fauna of North Africa all the species from different layers are not yet reach the evolutionary stage of those of Pataniak 6. The old level has an age similar to those of Beni Mellal. The magnetostratigraphic study allows to situating these fore layers in chron C5An.2n and C5AAn.     

La limite Cénomanien-Turonien dans le Haut Atlas central, Maroc

On the northern side of the Central High Atlas, new lithological and biostratigraphical data, especially realized in the Naour syncline, allow us to precise the Cenomanian-Turonian boundary. We recognized in that syncline an apparently complete series, the Ben Cherrou series, that represent the Upper Cenomanian-Turonian interval. We recommend it as the referring section of the Ben Cherrou Formation, that takes the place of the incomplete and not representative Aït Attab Formation. More, the sedimentological and paleontological analysis, suggest a Tethyan origin for the Cenomanian-Turonian transgressions in the studied area.     

Phu Din Daeng,a new Early Cretaceous vertebrate locality on the Khorat Plateau,NE Thailand

In this paper, we report on a new Early Cretaceous vertebrate locality, Phu Din Daeng, in Nakhon Phanom Province, NE Thailand. The Phu Din Daeng site has yielded a diverse vertebrate assemblage, including sharks (Heteroptychodus steinmanni), bony fishes (Pycnodontiformes; Sinamiidae cf. Siamamia and ?Vidalamiinae, and Ginglymodi), adocid turtles, indeterminate neosuchian crocodiles, pterosaurs and dinosaurs (spinosaurids and indeterminate theropods). A new adocid turtle, Protoshachemys rubra n. g. n. sp. is described on the basis of shell material. Field investigations on the geology and comparisons with other vertebrate faunas place Phu Din Daeng in the Sao Khua Formation (Barremian) of the Khorat Group.     

Assessing metabolic constraints on the maximum body size of actinopterygians: locomotion energetics of Leedsichthys problematicus (Actinopterygii,Pachycormiformes)

Maximum sizes attained by living actinopterygians are much smaller than those reached by chondrichthyans. Several factors, including the high metabolic requirements of bony fishes, have been proposed as possible body‐size constraints but no empirical approaches exist. Remarkably, fossil evidence has rarely been considered despite some extinct actinopterygians reaching sizes comparable to those of the largest living sharks. Here, we have assessed the locomotion energetics of Leedsichthys problematicus, an extinct gigantic suspension‐feeder and the largest actinopterygian ever known, shedding light on the metabolic limits of body size in actinopterygians and the possible underlying factors that drove the gigantism in pachycormiforms. Phylogenetic generalized least squares analyses and power performance curves established in living fishes were used to infer the metabolic budget and locomotion cost of L. problematicus in a wide range of scenarios. Our approach predicts that specimens weighing up to 44.9 tonnes would have been energetically viable and suggests that similar body sizes could also be possible among living taxa, discarding metabolic factors as likely body size constraints in actinopterygians. Other aspects, such as the high degree of endoskeletal ossification, oviparity, indirect development or the establishment of other large suspension‐feeders, could have hindered the evolution of gigantism among post‐Mesozoic ray‐finned fish groups. From this perspective, the evolution of anatomical innovations that allowed the transition towards a suspension‐feeding lifestyle in medium‐sized pachycormiforms and the emergence of ecological opportunity during the Mesozoic are proposed as the most likely factors for promoting the acquisition of gigantism in this successful lineage of actinopterygians.     

New chondrichthyan fauna from the Palaeogene deposits of Barmer district,Rajasthan, western India: Age,palaeoenvironment and intercontinental affinities

Field investigations of the Palaeogene sections exposed in Barmer district, Rajasthan (India) led to the discovery of a new fossiliferous horizon in the Padma Rao Open Cast Bentonitic Clay Quarry. The bentonitic clay sequence of this quarry is considered as representing the upper part of the shale, carbonaceous shale, lignite and bentonitic clay succession of the Akli Formation exposed in the Giral Lignite Mine, 2.5 km to its north. Screen-washing of the sediments from a fossiliferous level within the bentonitic clay of the new site has yielded nine taxa of sharks and batoids: Squatiscyllium nigeriensis White, 1934, Ginglymostoma sokotoense White, 1934, Ginglymostomatidae gen. et sp. indet., Brachycarcharias sp., Abdounia recticona Winkler, 1873, Premontreia (Oxyscyllium) peypouqueti Noubhani et Cappetta, 1997, Eotorpedo hilgendorfi Jaekel, 1904, Coupatezia sp. cf. C. danica, and Myliobatidae gen. et sp. indet. The fauna recovered is mainly represented by nearshore shallow marine forms. This fauna is quite different from that of Palaeocene Fatehgarh Formation, the Lower Eocene Kapurdi Formation (Barmer Basin), the Khuiala Formation (Jaisalmer Basin) and the Cambay Shale (Cambay Basin), and favours a late Palaeocene (Thanetian) age for the upper part of the Akli Formation. A nearshore shallow marine palaeoenvironment is favored for the investigated stratigraphic horizon on the basis of the selachian fauna. Majority of the fossil shark species described from the Padma Rao quarry section are known from a number of sites in Africa, western Europe, Asia and southeastern USA, suggesting that free faunal interchanges were possible between the western Tethys and Indo-Madagascan faunal provinces during the late Palaeocene.     

Taxonomical and palaeoecological investigations of the chondrichthyan and osteichthyan fish remains from the Middle-Late Triassic deposits of the Villány Hills (Southern Hungary)

Two sites in the Villány Hills, Hungary, have yielded rich fish assemblages from Middle to Late Triassic shallow marine deposits. The collected material comes from the Ladinian Templomhegy Dolomite Member and from the Carnian Mészhegy Sandstone Formation. The ichthyofauna is composed of both chondrichthyans (Hybodontidae indet., Palaeobates angustissimus, ‘Polyacrodus’ sp., ?Lissodus sp.) and osteichthyans (Gyrolepis sp., Birgeria sp., and further indeterminate actinopterygians). Despite the large sample size, no remains of neoselachians have been found. The Ladinian Templomhegy Dolomite is dominated by durophagous hybodontiforms (Palaeobates angustissimus, ?Lissodus sp.), but the piscivorous hybodontid and the generalist ‘Polyacrodus’ sp. are missing, while in the fish fauna collected from the Carnian Mészhegy Formation indeterminate piscivorous hybodontids are the most common elements and durophagous forms are much less abundant. The dominance of piscivorous hybodontids in the Carnian Mészhegy Sandstone could be related to the global decrease of diversity of marine fish-eating reptiles (e.g., nothosaurs) or to a change of paleoenvironmental conditions. The present study improves our knowledge on the poorly known Triassic vertebrate faunas of the Tisza Mega-unit, which formed a segment of the passive Neotethys margin of the European Plate and shows an important example of a potential vertebrate faunal shift during the Middle to Late Triassic.     

Acoustic telemetry reveals cryptic residency of whale sharks

Although whale sharks (Rhincodon typus) have been documented to move thousands of kilometres, they are most frequently observed at a few predictable seasonal aggregation sites. The absence of sharks at the surface during visual surveys has led to the assumption that sharks disperse to places unknown during the long ‘off-seasons’ at most of these locations. Here we compare 2 years of R. typus visual sighting records from Mafia Island in Tanzania to concurrent acoustic telemetry of tagged individuals. Sightings revealed a clear seasonal pattern with a peak between October and February and no sharks observed at other times. By contrast, acoustic telemetry demonstrated year-round residency of R. typus. The sharks use a different habitat in the off-season, swimming deeper and further away from shore, presumably in response to prey distributions. This behavioural change reduces the sharks'' visibility, giving the false impression that they have left the area. We demonstrate, for the first time to our knowledge, year-round residency of unprovisioned, individual R. typus at an aggregation site, and highlight the importance of using multiple techniques to study the movement ecology of marine megafauna.     

A missing piece in the Arctic food web puzzle? Stomach contents of Greenland sharks sampled in Svalbard, Norway

Harbour seals in Svalbard have short longevity, despite being protected from human hunting and having limited terrestrial predation at their haulout sites, low contaminant burdens and no fishery by-catch issues. This led us to explore the diet of Greenland sharks (Somniosus microcephalus) in this region as a potential seal predator. We examined gastrointestinal tracts (GITs) from 45 Greenland sharks in this study. These sharks ranged from 229 to 381?cm in fork length and 136–700?kg in body mass; all were sexually immature. Seal and whale tissues were found in 36.4 and 18.2%, respectively, of the GITs that had contents (n?=?33). Based on genetic analyses, the dominant seal prey species was the ringed seal (Pusa hispida); bearded seal (Erignathus barbatus) and hooded seal (Cystophora cristata) tissues were each found in a single shark. The sharks had eaten ringed seal pups and adults based on the presence of lanugo-covered prey (pups) and age determinations based on growth rings on claws (≤1?year and adults). All of the whale tissue was from minke whale (Balenoptera acutorostrata) offal, from animals that had been harvested in the whale fishery near Svalbard. Fish dominated the sharks’ diet, with Atlantic cod (Gadus morhua), Atlantic wolffish (Anarhichas lupus) and haddock (Melanogrammus aeglefinus) being the most important fish species. Circumstantial evidence suggests that these sharks actively prey on seals and fishes, in addition to eating carrion such as the whale tissue. Our study suggests that Greenland sharks may play a significant predatory role in Arctic food webs.     

Nouveaux sites à vertébrés paléogènes dans la région des Gour Lazib (Sahara nord-occidental,Algérie)

Recent fieldworks in the Palaeogene of the Gour Lazib area (Hammada du Dra, Algeria) led us to reassess the stratigraphic allocations of the Glib Zegdou formation and to complement the lithostratigraphy of this formation. This continental series unconformably overlies a Palaeozoic substratum. The prospecting–exploration works in the middle member of the formation allowed us to discover several new vertebrate sites (with lungfish, actinopterygians, chelonians, and notably mammals), associated with Early to Middle Eocene charophyte oogones. We describe a hyracoid (Titanohyrax tantulus) and a macroscelidid (?Chambius sp.). We here also report the discovery of two rich mammalian sites.     

Swimming against the flow—Environmental DNA can detect bull sharks (Carcharhinus leucas) across a dynamic deltaic interface

Human activities in coastal areas are accelerating ecosystem changes at an unprecedented pace, resulting in habitat loss, hydrological modifications, and predatory species declines. Understanding how these changes potentially cascade across marine and freshwater ecosystems requires knowing how mobile euryhaline species link these seemingly disparate systems. As upper trophic level predators, bull sharks (Carcharhinus leucas) play a crucial role in marine and freshwater ecosystem health. Telemetry studies in Mobile Bay, Alabama, suggest that bull sharks extensively use the northern portions of the bay, an estuarine–freshwater interface known as the Mobile‐Tensaw Delta. To assess whether bull sharks use freshwater habitats in this region, environmental DNA surveys were conducted during the dry summer and wet winter seasons in 2018. In each season, 5 × 1 L water samples were collected at each of 21 sites: five sites in Mobile Bay, six sites in the Mobile‐Tensaw Delta, and ten sites throughout the Mobile‐Tombigbee and Tensaw‐Alabama Rivers. Water samples were vacuum‐filtered, DNA extractions were performed on the particulate, and DNA extracts were analyzed with Droplet Digital™ Polymerase Chain Reaction using species‐specific primers and an internal probe to amplify a 237‐base pair fragment of the mitochondrial NADH dehydrogenase subunit 2 gene in bull sharks. One water sample collected during the summer in the Alabama River met the criteria for a positive detection, thereby confirming the presence of bull shark DNA. While preliminary, this finding suggests that bull sharks use less‐urbanized, riverine habitats up to 120 km upriver during Alabama''s dry summer season.     

Juvenile Greenland sharks <Emphasis Type="Italic">Somniosus microcephalus</Emphasis> (Bloch &amp; Schneider, 1801) in the Canadian Arctic

Life-stage-based management of marine fishes requires information on juvenile habitat preferences to ensure sustainable population demographics. This is especially important in the Arctic region given very little is known about the life histories of many native species, yet exploitation by developing commercial and artisanal fisheries is increasing as the ice extent decreases. Through scientific surveys and bycatch data from gillnet fisheries, we document captures of rarely reported juvenile Greenland sharks (Somniosus microcephalus; ≤200 cm total length [TL]) during the ice-free period in the Canadian Arctic. A total of 22 juvenile animals (42 % of total catch; n = 54), including the smallest reliably measured individual of 117 cm TL, were caught on scientific longlines and bottom trawls in Scott Inlet and Sam Ford Trough over three consecutive years. Molecular genetic nuclear markers confirmed species identity for 44 of these sharks sampled; however, two sharks including a juvenile of 150 cm TL were identified as carrying a Pacific sleeper shark (Somniosus pacificus) mitochondrial cytochrome b (cyt b) haplotype. This represents the first record of a Pacific sleeper shark genetic signature in Greenland sharks in Eastern Arctic waters. Juvenile sharks caught as bycatch in gillnet fisheries were only observed offshore in Baffin Bay surrounding a fishery closure area, while larger subadult and mature Greenland sharks (>200 cm TL) were caught in all fishing locations, including areas where juveniles were observed. The repeatable occurrence of juvenile Greenland sharks in a fjord and their presence at two offshore sites indicates that these smaller animals either reside in nurseries or have defined home ranges in both coastal and offshore regions or undertake large-scale inshore–offshore movements.     

An exceptionally preserved Late Devonian actinopterygian provides a new model for primitive cranial anatomy in ray-finned fishes

Actinopterygians (ray-finned fishes) are the most diverse living osteichthyan (bony vertebrate) group, with a rich fossil record. However, details of their earliest history during the middle Palaeozoic (Devonian) ‘Age of Fishes'' remains sketchy. This stems from an uneven understanding of anatomy in early actinopterygians, with a few well-known species dominating perceptions of primitive conditions. Here we present an exceptionally preserved ray-finned fish from the Late Devonian (Middle Frasnian, ca 373 Ma) of Pas-de-Calais, northern France. This new genus is represented by a single, three-dimensionally preserved skull. CT scanning reveals the presence of an almost complete braincase along with near-fully articulated mandibular, hyoid and gill arches. The neurocranium differs from the coeval Mimipiscis in displaying a short aortic canal with a distinct posterior notch, long grooves for the lateral dorsal aortae, large vestibular fontanelles and a broad postorbital process. Identification of similar but previously unrecognized features in other Devonian actinopterygians suggests that aspects of braincase anatomy in Mimipiscis are apomorphic, questioning its ubiquity as stand-in for generalized actinopterygian conditions. However, the gill skeleton of the new form broadly corresponds to that of Mimipiscis, and adds to an emerging picture of primitive branchial architecture in crown gnathostomes. The new genus is recovered in a polytomy with Mimiidae and a subset of Devonian and stratigraphically younger actinopterygians, with no support found for a monophyletic grouping of Moythomasia with Mimiidae.