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Comparación métrica y morfológica de la falange del género Homo de Cueva Victoria (Cartagena, Murcia) con las de primates y úrsidos
... par J. Masoliver (Centre de Traitement d'Images de l'Université Autónoma de Barcelona). Des analyses discriminantes (Palmqvist et al., 1996 ;Santamaria et Gibert, 1992) prouvent que CV-0 appartient à un spécimen d'Homo infantile (Fig. 10). ...
... Ce fait démontre l'arrivée d'hominidés au sud-est de la Péninsule Ibérique pendant le début du Pléistocène. (Santamaria et Gibert, 1992). On a obtenu un 100 % de reclassification correcte. ...
... Fig. 10. Discriminating analyses between phalanxes of Homo et Gorilla, Papio et Mandrillas, using conventionnal variables (distance and surface) measured with the analyse of X Ray (Santamaria et Gibert, 1992). 100 % correct reclassification was obtained. ...
This paper describes the human remains and other evidence of human presence (lithic artifacts) located in the early Pleistocene deposits of Orce and Cueva Victoria (SE Spain). We also discuss the age of those sites.
... The CV-0 phalanx was not compared to the fossil phalanges of T. oswaldi or H. erectus or H. habilis because they were not available. However, it was compared to juvenile and adult manual and pedal phalanges of H. sapiens, H. neanderthalensis, Australopithecus , and modern cercopithecoids (Gibert et al., 1985; Gibert and Pérez-Pérez, 1989; see also Pérez-Pérez, 1989; Palmqvist et al., 1995 Palmqvist et al., , 1996 Santamaría and Gibert, 1992). If, as Navarro et al. (2005) argued, CV-0 is a T. oswaldi subadult phalanx, then it should be compared with subadult phalanges of T. oswaldi, not with adult phalanges , which have larger dimensions. ...
... There are morphological differences between the lateral and medial borders of human and Theropithecus phalanges. Generally speaking, in Homo the lateral border is slightly curved, while the medial border is straight (Fig. 5), whereas in all the Theropithecus phalanges illustrated by Navarro et al. (2005) and in other modern cercopithecoids (plates 2 and 3 in Gibert et al., 1985; and plate 1 in Santamaría and Gibert, 1992), both borders are equally curved. As can be seen inFig. ...
... The attribution of CV-0 to Homo is supported by different morphological and internal features. All of the anatomical traits studied (Gibert and Pons-Moyà, 1984; Gibert et al., 1985; Gibert and Pérez-Pérez, 1989; Pérez-Pérez, 1989 ), the distribution of cortical bone (Santamaría and Gibert, 1992), and the discriminant analysis of morphological measurements (Palmqvist et al., 1995Palmqvist et al., , 1996 Pérez-Pérez, 1989) indicate similarity to Homo and not to cercopithecoids. It is unsurprising to find some differences between CV-0 and modern human phalanges, given that Cueva Victoria is a representative of early Pleistocene Homo. ...
... On the other hand, a second phalanx of the fifth finger of the right hand was also found, classified as Homo sp. (CV-0) (Gibert 1985;Gibert and Pons-Moyà, 1985;Pons-Moyà, 1985;Gibert et al., 2008;Ribot et al., 2015). ...
Cueva Victoria is a paleontological site of late Early Pleistocene age (ca. 0.9 Ma) located in the southeast of the Iberian Peninsula (Cartagena, Region de Murcia), which yielded diverse and abundant fossil remains, especially of large mammals. This accumulation is regarded as mainly deriving from the action of the giant hyena Pachycrocuta brevirostris, which used the cave as a den. Here, we present some preliminary taphonomic information deriving from the study of a portion of the large mammal collection from Cueva Victoria, including formerly uncatalogued material collected since the 1980s, which has not been considered in previous research. The study of this material started in 2018, when, owing to a suspension of the excavation activities in the cave, permissions were obtained to examine the uncatalogued specimens recovered during previous campaigns. Alongside a large number of remains of terrestrial mammals such as equids and cervids, Pachycrocuta brevirostris carried into the cave remains of groups uncommon in continental settings, such as marine mammals (cetaceans, seals). Beginning with this case, with then discuss the relevance of hyena-seal interactions in the Pleistocene of Europe in terms of ecology, as well as of seal documentation in the fossil record. From a critical review of the available evidence, Cueva Victoria stands out as a clear example of the consumption of seal by hyenas, with another likely case being represented by the much younger site of Lunel-Viel (late Middle Pleistocene, France). In other known cases of co-occurrence of hyenas and seals, the presence of seals is either due to other factors (e.g., hominins, deposition of marine sediments) or its origin cannot be ascertained for sure.
... CV-0, falange del quinto dedo de la mano derecha, descrita por GIBERT y PONS, 1984;PONS MOYÀ, 1985;GIBERT et alii, 1985, GIBERT y PÉREZ-PÉREZ, 1989, GIBERT et alii, 1992, PALMQVIST y GIBERT et alii, 1996 En los estudios más completos realizados sobre esta pieza (SANTAMARÍA y GIBERT, 1992 y PALMQVIST GIBERT et alii, 1996) se comparó la distribución de la cortical interna de CV-0 con las de H. s. sapiens y con los primates no humanos que en su estudio morfométrico han presentado valores más próximos a los humanos: Macaca sylvana, Papio hamadryas y Gorilla gorilla. ...
Resumen
En este trabajo se describe la historia geológica de la cavidad, la formación de la brecha fosilífera, la edad de la fauna y la naturaleza humana de la falange CV-0 comparándola con las de Theropithecus.
Summary
This work describes the geological history of Cueva Victoria including the formation of the fossiliferous breccia, and the age of the fauna. This work also compares a phalanx classified as Homo with Theropitecus.
... Cavidad kárstica localizada en el término murciano de La Victoria, cuenta con un depósito de notable riqueza fosilífera. Se ha informado de la presencia de restos humanos: una falange, dos fragmentos de húmero, un incisivo y un fragmento de fémur (Gibert, 1999: 230;Gibert, Campillo et alii, 1993;Gibert, Sánchez et alii, 1992;Pons-Moyà, 1985;Santamaría y Gibert, 1992) y un pequeño conjunto lítico y óseo sobre el que se identificó una supuesta acción antrópica (Carbonell, Estévez et alii, 1981). ...
Present paper briefly reviews the mean archaeological record proposed as evidence of human presence during Lower Pleistocene in the Iberian Peninsula. A critical review questions the ancient chronologies assigned to surface lithic collections recorded in high fluvial terraces. Nowadays, archaeological data that most likely traces the Iberian earliest human occupation are those recorded in Barranco León 5, Fuentenueva 3 (Granada), Gran Dolina, and Sima del Elefante (Sierra de Atapuerca, Burgos). Paying attention to this archaeological and palaeontological record, some hominid groups inhabited some Iberian areas between the normal polarity episode Gilsá (OIS 34) and the paleomagnetic boundary Brunhes/Matuyama (OIS 20), Lower/Middle Pleistocene conventional limit. This chronological proposal for the earliest human occupation on the Iberian Peninsula introduces us in the broader view of the earliest human occupation of Eurasia, giving some reflections about this phenomena and the state of the art.
... Esta zona sólo es observable en la radiografía y no se corresponde a ninguna morfología externa de la falange. Esto también queda claro en el pie de figura 2 de Palmqvist et al. (1996), que dice: "Discriminant analysis between Homo and Gorilla, Papio and Mandrillus phalanges, using conventional variates (distances and surface areas) measured by image analysis on radiographs (data from Santamaría & Gibert, 1992)…", y en la imagen, estas zonas se denominan como A d y A i , respectivamente; y a pie de su figura 6 se lee: "Calculation of shape coordinates for triangles of landmarks (redrawn after figure 5.1.2 in Bookstein, 1991), and selected landmarks on radiographs of the phalanges…" (en ambas citas el subrayado es nuestro fig. ...
At Cueva Victoria, two types of primates have been found, the papionin Theropithecus oswaldi and the hominin Homo sp. The former was initially identified by a deciduous molar; recently, the discovery of 4 new teeth has allowed a more accurate classification as T. T. oswaldi leakeyi. Of the second, only one intermediate phalanx of the V finger of the right hand and two humeral fragments have been found up to now, which makes the specific attribution difficult, and they are classified as Homo sp. A claim, published in 2005, stating that the phalanx was of Theropithecus, was followed by a reply and a response in 2008. Here we reply this latter answer and we reaffirm the assignment to Homo sp. The presence of these primates in Cueva Victoria makes this site essential in the discussion of the dispersal of African primates into Europe, and reinforces the hypothesis of the Strait of Gibraltar as a probable route out of Africa.
... A number of new taxa have been defined from Cueva Victoria, including the canid Cuon rosi (Moyà and Moyà-Solà, 1978); the arvicolid Allophaiomys chalinei (Alcalde et al., 1981), later reassigned to a new genus, Victoriamys by Martin (2012) ; and the lagomorph Oryctolagus giberti (de Marfà, 2008). The faunal association also includes Homo sp., based on an intermediate human phalanx (Gibert and Pons-Moyà, 1985; Pons-Moyà, 1985; Gibert et al., , 2002 Santamaría and Gibert, 1992; Palmqvist et al., 1996), which some authors (Martínez-Navarro et al., 2005 ) have suggested may belong to Theropithecus (see discussion in Gibert et al., 2008, and Martínez-Navarro et al., 2008). The finding at Cueva Victoria, a right M 2 , CV-1 (Gibert et al., 1995Gibert et al., , 1999), was the first evidence found for the presence of Theropithecus in Europe. ...
... (Pons Moyá, 1985). A morphological study (Gibert & Pons Moyá, 1984) was undertaken to differentiate the phalanx from other pentadactyl mammals, particularly carnivores and beavers, and a biometric comparison was done with primates and ursids (Gibert, Pons & Ruz, 1985). Using both anatomical data and conventional biometric parameters it was possible to discriminate the Cueva Victoria phalanx form those of cercopithecids (Cercopithecus nictitans, Macaca silvana, Mandrillus sphinx and Papio hamadryas), apes (Gorilla gorilla, Pan troglodytes, Pongo pygmaeus and Hylobates sp.), ursids (Ursus spelaeus and Ursus arctos), terrestrial and marine carnivores (Indarctos vireti, Hyaena sp. and Phoca sp.) and beavers (Stenofiber gaegeri). ...
The presence ofHomoin Europe longer than a million years ago is often discussed. The finding of a Lower Pleistocene phalanx from the karstic site of Cueva Victoria in Murcia (Spain) can contribute important information about this theme if theHomoidentification is confirmed. We have used an array of morphometric analyses to study this phalanx, and to compare it with human, gorilla and cercopithecid phalanges. The morphometric variates included conventional distance and surface area measurements estimated from radiographs of the phalanges, harmonic amplitudes from Fourier series fitted to their contours, and shape coordinates of landmarks. Discriminant analysis was used to compare the groups, and principal and relative warp analyses used to investigate the relative position of landmarks. Results obtained with all morphometric methods consistently indicated a human affinity for the fossil, and thus constitute further evidence of the presence ofHomosp. in the southern Iberian Peninsula during the Lower Pleistocene.
... Deux d'entre eux sont des fragments d'humérus (CV-1 et CV-2 ; qui sont encore moins convaincants anatomiquement que les spécimens VM-1960 et VM-3691 décrits précédemment. Le troisième spécimen (CV-0, Fig. 9) fut identifié comme une deuxième phalange du 5 e doigt de la main (Gibert et Pons-Moyà, 1985 ;Pons-Moyà, 1985). ...
... Deux d'entre eux sont des fragments d'humérus (CV-1 et CV-2 ; ) qui sont encore moins convaincants anatomiquement que les spécimens VM-1960 et VM-3691 décrits précédemment. Le troisième spécimen (CV-0, Fig. 9) fut identifié comme une deuxième phalange du 5 e doigt de la main ( Gibert et Pons-Moyà, 1985 ;Pons-Moyà, 1985). Ce fossile a été comparé au moyen de techniques anatomiques, radiologiques et morphométriques, avec des deuxièmes phalanges d'autres mammifères pentadactyles, tels que des carnivores, des cercopithecoidés, des singes et des humains ( Gibert et Pérez, 1989 ;Santamaría et Gibert, 1992 ;Palmqvist et al., 1996b). ...
The Early Pleistocene locality at Venta Micena (Orce, Guadix-Baza basin, province of Granada, Spain) has provided four fossil remains – skull fragment VM-0, and long bone diaphyses VM-1960, VM-3691, and VM-12000 – which have been tentatively attributed to the hominids. Although several methodologies have been used to ascertain the human affinities of these specimens – including anatomical, morphometric and immunological analyses – the results obtained have not been conclusive, instigating a persistent debate. A taphonomic approach is used here for estimating the probability that a taxon the size of Homo sp. (~50 kg) could be represented in the fossil assemblage by four bone fragments and no tooth remain. A least-squares regression analysis between the percentage of teeth and the body mass estimated for each taxon of large mammals (N=20) predicts a raw abundance of six teeth for Homo sp. in the assemblage. Given that up to the present moment no tooth remains attributable to the hominids has been unearthed during systematic excavations in the Venta Micena quarry, which has provided more than 15,000 fossils of large mammals, this argues strongly against the possibility that the three bone specimens could belong to Homo sp. The phalanx CV-0 from the Early Pleistocene site of Cueva Victoria (Cartagena, Spain) has also been attributed to the genus Homo. The taxonomic assignment of this specimen is biased, however, because it was not compared with Theropithecus oswaldi, the only primate species actually recorded from this karstic locality. A comparative anatomical and morphometric analysis of fossil and modern specimens of Theropithecus suggests that CV-0 can be attributed to T. oswaldi. As a result, Cueva Victoria does not contribute additional information concerning the first human settlements in Europe. By these reasons, apart from the paleoanthropological and archaeological findings from Atapuerca (TD lower levels and Sima del Elefante), the rich archaeological assemblages from Barranco León and Fuente Nueva-3 in Orce, dated 1.3-1.2 Myrs, which include fourteen hundred stone tools of Oldowan technology, constitute at present the only unequivocal evidence of human presence in Southeast Spain during Early Pleistocene times.
... Deux d'entre eux sont des fragments d'humérus (CV-1 et CV-2 ; qui sont encore moins convaincants anatomiquement que les spécimens VM-1960 et VM-3691 décrits précédemment. Le troisième spécimen (CV-0, Fig. 9) fut identifié comme une deuxième phalange du 5 e doigt de la main (Gibert et Pons-Moyà, 1985 ;Pons-Moyà, 1985). ...
... Based on published photographs and on the recent publications of Gibert and coworkers (Santamaría and Gibert, 1992;Palmqvist et al., 1996;Gibert et al., 2002), one question raised by us (Martínez-Navarro et al., 2005: 519) was whether the Cueva Victoria phalanx ''corresponds to a juvenile or subadult individual, as it does not preserve the proximal epiphysis, which had not fused at the time of death.'' Now, after direct study of the specimen, we know that the proximal facets are present, but they are partially eroded, especially on the palmar/plantar aspect, where it is possible to see traces of the suture between the epiphysis and the diaphysis of the phalanx. ...
... CV-0, falange del quinto dedo de la mano derecha, descrita por GIBERT y PONS, 1984;PONS MOYÀ, 1985;GIBERT et alii, 1985, GIBERT y PÉREZ-PÉREZ, 1989, GIBERT et alii, 1992, PALMQVIST y GIBERT et alii, 1996 En los estudios más completos realizados sobre esta pieza (SANTAMARÍA y GIBERT, 1992 y PALMQVIST GIBERT et alii, 1996) se comparó la distribución de la cortical interna de CV-0 con las de H. s. sapiens y con los primates no humanos que en su estudio morfométrico han presentado valores más próximos a los humanos: Macaca sylvana, Papio hamadryas y Gorilla gorilla. ...
Resumen En Cueva Victoria se han encontrado dos tipos de primates, el papionino Theropithecus oswa/di y el hominino Horno sp. El primero se definió por un molar inferior deciduo; recientemente, el descubrimiento de 4 nuevos dientes ha per-mitido una clasificación más ajustada como Theropithecus T oswa/di /eakeyi. Los restos de homininos hallados hasta ahora son una falange intermedia del V dedo de la mano derecha (CV-O) y dos fragmentos humerales (CV-1 y CV-2), lo que hace muy difícil su atribución específica, clasificándose como Horno sp. Respecto a la hipótesis publicada en 2005 de que la falange pertenezca en realidad a Theropithecus, se publicaron en 2008 una réplica y una contra réplica y en este trabajo se responde a la contrarréplica, ratificando la asignación a Horno sp. La presencia de estos primates en Cueva Victoria hace que este yacimiento sea fundamental para la discusión de las dispersiones de primates africanos a Europa, y refuerza la hipótesis de que probablemente la vía de entrada al continente europeo se produjo a través del Estrecho de Gibraltar. Palabras clave Theropithecus oswaldi, Horno sp., Dispersión, Falange, Pleistoceno inferior. Summary At Cueva Victoria, two types of primates have been found, the papionin Theropithecus oswaldi and the hominin Horno sp. The former was initially identified by a deciduous molar; recently, the discovery of 4 new teeth has allowed a more accurate classification as T T oswaldi leakeyi. Of the second, only one intermediate phalanx of the V finger of the right hand and two humeral fragments have been found up to now, which makes the specific attribution difficult, and they are classified as Horno sp. A claim, published in 2005, stating that the phalanx was of Theropithecus, was followed by a reply and a response in 2008. Here we reply this latter answer and we reaffirm the assignment to Homo sp. The presence of these primates in Cueva Victoria makes this site essential in the discussion of the dispersal of African primates into Europe, and reinforces the hypothesis of the Strait of Gibraltar as a probable route out of Africa.
Cueva Victoria has provided remains of more than 90 species of fossil vertebrates, including a hominin phalanx, and the only specimens of the African cercopithecid Theropithecus oswaldi in Europe. To constrain the age of the vertebrate remains we used paleomagnetism, vertebrate biostratigraphy and (230)Th/U dating. Normal polarity was identified in the non-fossiliferous lowest and highest stratigraphic units (red clay and capping flowstones) while reverse polarity was found in the intermediate stratigraphic unit (fossiliferous breccia). A lower polarity change occurred during the deposition of the decalcification clay, when the cave was closed and karstification was active. A second polarity change occurred during the capping flowstone formation, when the upper galleries were filled with breccia. The mammal association indicates a post-Jaramillo age, which allows us to correlate this upper reversal with the Brunhes-Matuyama boundary (0.78 Ma). Consequently, the lower reversal (N-R) is interpreted as the end of the Jaramillo magnetochron (0.99 Ma). These ages bracket the age of the fossiliferous breccia between 0.99 and 0.78 Ma, suggesting that the capping flowstone was formed during the wet Marine Isotopic Stage 19, which includes the Brunhes-Matuyama boundary. Fossil remains of Theropithecus have been only found in situ ∼1 m below the B/M boundary, which allows us to place the arrival of Theropithecus to Cueva Victoria at ∼0.9-0.85 Ma. The fauna of Cueva Victoria lived during a period of important climatic change, known as the Early-Middle Pleistocene Climatic Transition. The occurrence of the oldest European Acheulean tools at the contemporaneous nearby site of Cueva Negra suggest an African dispersal into SE Iberia through the Strait of Gibraltar during MIS 22, when sea-level was ∼100 m below its present position, allowing the passage into Europe of, at least, Theropithecus and Homo bearing Acheulean technology.
The vertebrate fossils from Cueva Victoria are found in the sediment that infilled karstic cavities during the Early Pleistocene. The sediment, part of an alluvial fan developed on the slopes of San Ginés Hill, penetrated through at least three subvertical entrances. The infilling was relatively rapid and continuous until it filled up the cave. Later, the sediment underwent differential compactation and percolation of saturated CaCO3 waters cemented the upper part of the breccia and formed a capping flowstone. The taphonomic processes affecting the fossils are discussed. The macromammal remains were accumulated by hienas (Pachycrocuta brevirostris) using the cave as a den, as shown by fractures and tooth marks in the bone remains, including pitting, the presence of digested bone fragments, the number of hiena juvenile individuals, the abundance of hiena coprolites, the presence of marine mammal remains in the breccia, and the ratio of skeletal parts of macromammal remains.
Cueva Victoria is a karst-infilling site with early Pleistocene vertebrate fossil remains. It was discovered in the first half of the XXth century by mining explotation. The fossil record of Cueva Victoria is exceptional because of its large biodiversity and because it is the only locality in Europe where the African primate Theropithecus oswaldi occurs. Here, we summarize the history of paleontological research and excavation campaigns. We provide a catalogue of the different locations in the site, and a reference list of the publications on this site.
Recently Toro-Moyano et al. (2013) reported a deciduous tooth from Barranco Leo´n (Spain; BL02-J54-100) and claimed it to be the oldest human fossil in Europe. In that paper, the authors suggest that a previously reported human molar fragmentfrom the same site (BL5-0) was not human but a deciduous molar of Hippopotamus found out of stratigraphic context. Here, we show the stratigraphic and spatial position of BL5-0, and we separate it from deciduous teeth of Hippopotamus. We conclude that two human deciduous molars have been discovered at the Barranco Leo´n site. Both teeth were found 9 meters apart, have a similar size, are heavily worn on the occlusal surface, have a nearly identical interstitial contact facet, and in both cases the roots are practically missing due to resorption. These similarities and the proximity of the finds suggest that both molars probably belonged to the same individual.
During the last decade, new discoveries in several Iberian basins, together with the description of previously unpublished finds, have significantly increased the recorded paleodiversity of fossil Primates (Mammalia: Euarchonta) in the Iberian Peninsula. Here we provide an updated compendium of the primate fossil record in Iberia during the Cenozoic and further summarize the changes in primate paleodiversity through time, which are then analyzed in the light of changing climatic conditions. Thanks to favorable climatic conditions, the highest diversity of Iberian primates was reached during the Eocene, thus reflecting the radiation of both adapoids and omomyoids; only a single plesiadapiform genus is in contrast recorded in the Iberian Peninsula. Near the Eocene-Oligocene boundary, paleoclimatic changes led to a primate diversity crisis and other faunal changes, although two Iberian omomyoids survived the Grande Coupure. From the Middle Miocene onwards, catarrhine primates are recorded in the Iberian Peninsula. During the Middle and Late Miocene, they are represented by pliopithecoids and hominoids, restricted to NE Iberia. The Miocene hominoids from Iberia are of utmost significance for understanding the Eurasian hominoid radiation and its role in the origins of the great-ape-and-human clade. Following the local extinction of these taxa during the early Late Miocene, due to progressively increased seasonality and concomitant changes in plant communities, cercopithecoids are also recorded in the Iberian Peninsula from the latest Miocene through the Plio-Pleistocene, although they finally became locally extinct, whereas hominoids are again represented by fossil humans during the Pleistocene.
This paper describes the human remains and other evidence of human presence (lithic artifacts) located in the early Pleistocene deposits of Orce and Cueva Victoria (SE Spain). We also discuss the age of those sites.
Well-defined human anatomical characteristics are present on humeral fragments of a child (VM-1960) and an adult (VM-3691) from early Lower Pleistocene sediments at Venta Micena: both have narrower medullary cavities than in AfricanHomo erectus/ergaster (KNM-ER 1808), and the child’s humeral shaft is longer than in recent 8-to-9-year-olds even though its muscle markings are less pronounced
than theirs. We infer that exposure of growing children to high mechanical loading favoured Plio-Pleistocene skeletal evolution
inHomo of humeral robusticity and elongation. Precocious childhood arm-bone development, occurring before pubertal growth-spurt
increments in shoulder and arm muscularity, implies a different balance from today between prepubertal hormonal influences
exerted on ossification (growth hormone and somatomedin C) and the adolescent gonadal hormones of our modern growth spurt
which may have still been in the process of evolution by natural selection.
A molar tooth fragment from the Plio-Pleistocene Barranco León site 5 at Orce is shown to belong toHomo by analyses of its enamel in terms of the arrangement of the striae of Retzius and Hunter-Shreger bands, presence of perikymata,
and of the thickness of the enamel, when compared with teeth of similarsized mammals of other taxa.
In SE Spain, recent excavations in the Orce basin and at Cueva Victoria indicate presence of both hominids and hominid activity
from the Plio-Pleistocene boundary and early Lower Pleistocene.
The karstic filling of Cueva Victoria in southeastern Spain, dated from the latest early Pleistocene (ca. 1.1 Ma), is famous for providing primate fossil remains (Theropithecus) of typical African origin, in the general controversy on the antiquity of the first hominid settlements in Western Europe and their possible entrance into Europe through the Strait of Gibraltar. Cueva Victoria has also furnished the following fauna of anurans and squamate reptiles: cf. Pelodytes sp. (Pelodytidae), Bufo cf. B. bufo (Bufonidae), Blanus cinereus (Blanidae), Tarentola sp. (Geckonidae), Chalcides cf. Ch. bedriagai (Scincidae), Timon cf. T. lepidus and indeterminate small lacertids (Lacertidae), Natrix maura, Coronella girondica, Rhinechis scalaris and Malpolon cf. M. monspessulanus (Colubridae). This faunal association seems to suggest a mean annual temperature slightly fresher than nowadays (approximately 1°C less than at present in the area), with cooler winters but warmer summers and above all higher mean annual precipitations (+ 400 mm). The landscape may correspond to an open forest environment of a Mediterranean type, with some still water points.
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