Emmanuelle Stoetzel

Taphonomy and palaeoecology of the late Pleistocene to middle Holocene small mammal succession of El Harhoura 2 cave (Rabat-Témara, Morocco)

The relationship between local and global climatic variations and the origin and dispersal of Homo sapiens in Africa is complex, and North Africa may have played a major role in these events. In Morocco, very few studies are specifically dedicated to small fossil vertebrates, and neither taphonomic nor palaeoecological studies have been undertaken on these taxa, particularly in archaeological contexts. The late Pleistocene to middle Holocene succession of El Harhoura 2 cave, situated in the region of Témara, yields an exceptionally rich small vertebrate assemblage. We present the results of a first systematic, taphonomic, and palaeoecological study of the small mammals from Levels 1 to 8 of El Harhoura 2. The absence of bone sorting and polishing, as well as the presence of significant traces of digestion indicate that the small mammal bones were accumulated in the cave by predators and that no water transport occurred. Other traces observed on the surface of bones consist mainly of root marks and black traces (micro-organisms or more probably manganese) which affected the majority of the material. The percentage of fragmentation is very high in all stratigraphic levels, and the post-depositional breakage (geologic and anthropogenic phenomena) obscure the original breakage patterns of bones by predators. According to the ecology of the different species present in the levels of El Harhoura 2, and by taking into account possible biases highlighted by the taphonomic analysis, we reconstruct the palaeoenvironmental evolution in the region. For quantitative reconstructions we used two indices: the Taxonomic Habitat Index (THI) and the Gerbillinae/Murinae ratio. Late Pleistocene accumulations were characterised by a succession of humid (Levels 3, 4a, 6, and 8) and arid (Levels 2?, 5, and 7) periods, with more or less open landscapes, ending in an ultimate humid and wooded period during the middle Holocene (Level 1). We discuss particular limits of our results and interpretations, due to an important lack of taxonomic, ecological, and taphonomic knowledge in North Africa.

First fossil representative of the salamander crown-group from a Gondwanan continent:Pleurodeles cf. waltl from the Quaternary of Morocco

No fossil belonging to the Caudata crown-group has been hitherto described from a former Gondwanan continent. We report the first known extinct member of the caudate crown-clade from Africa, i.e. a Gondwanan territory. It belongs to the Recent genus Pleurodeles (Salamandridae) and is referred to as Pleurodeles cf. waltl . The fossils come from the Pleistocene of Morocco. They show that Pleurodeles (likely P. waltl ) has been present in Africa for at least two million years and that anthropogenic dispersal is not the cause of its presence in this continent.

Upper Palaeolithic (layer 2) and Middle Palaeolithic (layer 3) large faunas from El Harhoura 2 Cave (Témara, Morocco): paleontological, paleoécological and paleoclimatic data

The El Harhoura 2 cave is situated in the commune of El Harhoura in the Temara region (Morocco). The entrance faces west, toward the ocean, around 200 m away from the current shoreline and about 16 m above sea level. The stratigraphy has currently 12 levels numbered from top to bottom. Level 1 is attributed to the early Neolithic. The archaeological material from level 2 places it within the Upper Palaeolothic (Iberomaurusian). Levels 3 to 11 are assigned to the Middle Palaeolithic (Aterian). Since 1977, a surface of 37 m2 has been excavated, yielding particularly abundant remains of a large mammal fauna. This paper is about paleontological, paleoecological and paleoclimatologic data of layers 2 and 3 from El Harhoura 2.

A cost-effective high-throughput metabarcoding approach powerful enough to genotype ~44 000 year-old rodent remains from Northern Africa.

We present a cost‐effective metabarcoding approach, aMPlex Torrent, which relies on an improved multiplex PCR adapted to highly degraded DNA, combining barcoding and next‐generation sequencing to simultaneously analyse many heterogeneous samples. We demonstrate the strength of these improvements by generating a phylochronology through the genotyping of ancient rodent remains from a Moroccan cave whose stratigraphy covers the last 120 000 years. Rodents are important for epidemiology, agronomy and ecological investigations and can act as bioindicators for human‐ and/or climate‐induced environmental changes. Efficient and reliable genotyping of ancient rodent remains has the potential to deliver valuable phylogenetic and paleoecological information. The analysis of multiple ancient skeletal remains of very small size with poor DNA preservation, however, requires a sensitive high‐throughput method to generate sufficient data. We show this approach to be particularly adapted at accessing this otherwise difficult taxonomic and genetic resource. As a highly scalable, lower cost and less labour‐intensive alternative to targeted sequence capture approaches, we propose the aMPlex Torrent strategy to be a useful tool for the genetic analysis of multiple degraded samples in studies involving ecology, archaeology, conservation and evolutionary biology.

Limpet Shells from the Aterian Level 8 of El Harhoura 2 Cave (Témara, Morocco): Preservation State of Crossed-Foliated Layers

The exploitation of mollusks by the first anatomically modern humans is a central question for archaeologists. This paper focuses on level 8 (dated around ∼ 100 ka BP) of El Harhoura 2 Cave, located along the coastline in the Rabat-Témara region (Morocco). The large quantity of Patella sp. shells found in this level highlights questions regarding their origin and preservation. This study presents an estimation of the preservation status of these shells. We focus here on the diagenetic evolution of both the microstructural patterns and organic components of crossed-foliated shell layers, in order to assess the viability of further investigations based on shell layer minor elements, isotopic or biochemical compositions. The results show that the shells seem to be well conserved, with microstructural patterns preserved down to sub-micrometric scales, and that some organic components are still present in situ. But faint taphonomic degradations affecting both mineral and organic components are nonetheless evidenced, such as the disappearance of organic envelopes surrounding crossed-foliated lamellae, combined with a partial recrystallization of the lamellae. Our results provide a solid case-study of the early stages of the diagenetic evolution of crossed-foliated shell layers. Moreover, they highlight the fact that extreme caution must be taken before using fossil shells for palaeoenvironmental or geochronological reconstructions. Without thorough investigation, the alteration patterns illustrated here would easily have gone unnoticed. However, these degradations are liable to bias any proxy based on the elemental, isotopic or biochemical composition of the shells. This study also provides significant data concerning human subsistence behavior: the presence of notches and the good preservation state of limpet shells (no dissolution/recrystallization, no bioerosion and no abrasion/fragmentation aspects) would attest that limpets were gathered alive with tools by Middle Palaeolithic (Aterian) populations in North Africa for consumption.

Identifying predation on rodent teeth through structure and composition: A case from Morocco.

Predation by nocturnal birds of prey is one of the most frequent modes leading to the concentration of rodents in fossil assemblages. This mode of accumulation leaves characteristic surface alterations on bones and teeth. In order to evaluate and characterize the effects of these pre-diagenesis alterations on rodent fossil samples, we have carried out microstructural and chemical analyses on incisors collected from present day Moroccan wild animals and owl pellets. The microstructure of both dentine and enamel was well preserved, but chemical changes were evident in pellet samples and depended on the particular tissue and the nature of the predator. The comparison of compositional data obtained from electron microprobe chemical analyses and infrared spectrometry has allowed us to assign a possible predator to an incisor extracted from a pellet of an unknown origin. This method has further implications for the understanding of taphonomy and palaeoecology of archaeological and fossil sites.

Taphonomy of Small Predators multi-taxa accumulations: palaeoecological implications

Abstract Most small terrestrial vertebrate accumulations in archaeological and palaeontological sites result from predation but we are far from having an exhaustive knowledge of modern predators’ diet, ecological niches and bone modification patterns especially in North African sites. The few neotaphonomic referentials available result from taxon-specialized palaeontologists’ initiatives. A survey of the literature on North Africa predators shows that their prey diversity is high and not only include rodent and shrews but also amphibians, squamates, bats and insects. We performed here a pilot taphonomic study of a Moroccan Tyto alba nest pellets including the whole taxa consumed (birds, rodents, shrews, amphibians, insects). We analyzed bone representation, fragmentation and digestion for each taxa and then compared the results. We observe differences between the taxa but on the whole find higher modification levels for this assemblage compared to roost sites of barn owls from other sites. We used for the comparisons homologous bones like the femur and the humerus and also found differences between the taxa. For example, digestion was lower on amphibians and birds than on micromammals. We discuss here some methodological issues as well as archaeological and palaeoenviromental ones by comparison with the Pleisto-Holocene site of El Harhoura II (Morocco).

Diet of the Lesser Antillean barn owl Tyto insularis (Aves: Strigiformes) in Dominica, Lesser Antilles

Abstract. This study presents new information on the diet of Tyto insularis in Dominica, Lesser Antilles. The study of 57 pellets and bulk material collected in 1999 (23) and 2000 (34) contained 517 prey items of relatively high diversity. These included squamates (2 species), rodents (2 species), bats (7 species), birds (17 identified taxa and several unidentified Passeriformes) and insects. Although our inferences stem a few owls, results suggested that the diet of T. insularis in Dominica is similar to the diet of T. glaucops in Hispaniola.

Out of Africa: demographic and colonization history of the Algerian mouse ( Mus spretus Lataste)

North Africa is now recognized as a major area for the emergence and dispersal of anatomically modern humans from at least 315 kya. The Mediterranean Basin is thus particularly suited to study the role of climate versus human-mediated changes on the evolutionary history of species. The Algerian mouse (Mus spretus Lataste) is an endemic species from this basin, with its distribution restricted to North Africa (from Libya to Morocco), Iberian Peninsula and South of France. A rich paleontological record of M. spretus exists in North Africa, suggesting hypotheses concerning colonization pathways, and the demographic and morphologic history of this species. Here we combined genetic (3 mitochondrial DNA loci and 18 microsatellites) and climatic niche modeling data to infer the evolutionary history of the Algerian mouse. We collected 646 new individuals in 51 localities. Our results are consistent with an anthropogenic translocation of the Algerian mouse from North Africa to the Iberian Peninsula via Neolithic navigators, probably from the Tingitane Peninsula. Once arrived in Spain, suitable climatic conditions would then have favored the dispersion of the Algerian mice to France. The morphological differentiation observed between Spanish, French and North African populations could be explained by a founder effect and possibly local adaptation. This article helps to better understand the role of climate versus human-mediated changes on the evolutionary history of mammal species in the Mediterranean Basin.