Évolution des grands Carnivores au Plio Pléistocène en Afrique et en Europe occidentale

This paper provides an overview of current knowledge of Plio Pleistocene Carnivora from Africa and Europe. In Plio Pleistocene times, many genera extend their ranges in Africa and Eurasia but their evolution are quite distinct in the two continents. In Africa, the modern carnivoran guild of sub saharian Africa originated in the early Pleistocene when took place extinction of archaic species. The north African fossil record is far from complete until the middle Pleistocene. In the middle and late Pleistocene, the modern carnivoran guild is associated with other elements: the simian jackal Canis simensis and two species of ursids Ursus arctos and Ursus deningeri which evolved towards Ursus spelaeus. Western European carnivore faunas show due to migration a constant turn-over of the species. Among felids, Panthera schaubi described by Viret (1954) and attributed to the genus Puma by Hemmer et al. (2004), is morphologically close to the snow leopard Panthera (Uncia) uncia. Canis etruscus is the sister group pf the clade including wolf and coyote and Canis arnensis is close to the African jackals. Ursus deningeri appears in the early Pleistocene together with two arctoid forms Ursus rodei and Ursus dolinensis that may be synonymous to Ursus arctos. The genus Hyaena is present in Europe in the middle and late Pleistocene.     

Étude biostratigraphique et paléoenvironnementale des diatomées pyritisées de l'intervalle Paléocène supérieur-Éocène inférieur du sondage de Saint-Josse (France)

This paper presents the results of a biostratigraphic analysis of pyritized diatom assemblages in the Saint-Josse core (CC82) located in the northern part of the Paris basin (coordinates: X =555?000, Y =307?324 and Z =56.9 m). During the Paleocene-Eocene this locality was situated in a small bight of the North Sea Basin. The results are obtained from sediments of the Thanet and Mont-Bernon group equivalents belonging to the Paleocene-Eocene interval. Thirteen samples were studied and eleven contain pyritized diatoms. By epigenesis, the siliceous frustule of the diatom is changed into a pyritized skeleton with all the ornamentation very well preserved such as the velum, the areolae, the labiate processes, etc. Dispersive X-Ray analyses show this complete replacement of silica by pyrite. Another form of pyritization is crystallization of pyrite (in cubes, pyritohedrons, octahedrons, framboids) in internal cavities of the diatoms to produce internal moulds, also called steinkerns, which preserve the external morphology of the diatoms and some details such as the girdle. Despite pyritization, diatoms can often be recognized at the species level; brackish and marine species were observed but no fresh water specimens were encountered. Pyrite is easily altered into iron oxides. In outcrops, pyritized diatoms can be obscured or destroyed by oxidation. We have chosen borehole samples for this study to avoid the affects of weathering. Deposits from the Sorrus Unit (of the Thanet Group) have low abundances of pyritized diatoms (3 to 25 specimens per sample). This unit contains allochtonous diatom assemblages comprising Actinoptychussenarius, Coscinodiscus morsianus var. morsianus, and Coscinodiscus morsianus var. moelleri. Sample 34.31-34.45 from this unit is particularly interesting. It contains three specimens of Paralia siberica var. laevis, which is typical of an estuarine or littoral paleoenvironment, and corresponds to the more continental deposit of the Sorrus Unit. Seven samples, taken in the Argile de Saint-Aubin Unit (from the Mont-Bernon Group), contain more than 200 pyritized diatoms and between 9 to 16 different species per sample. A brackish assemblage (Actinoptychus senarius, Coscinodiscus commutatus and Stellarima microtrias) and a marine assemblage (C. morsianus var. morsianus, Coscinodiscus var. moelleri, Fenestrella antiqua, Odontellaheibergii, Rhizosolenia sp. 1 and Trinacria regina) are observed. The brackish assemblage expands in the upper part of the Saint-Aubin Unit indicating more widespread brackish paleoenvironmental conditions. Pyritized diatoms are important fossil markers in the Paris basin mainly when siliceous fossils (silicoflagellates or radiolarians) and others microfossils (foraminifers and dinoflagellates) are very poor or absent. Two diatom assemblages have been identified in the Saint-Josse borehole. The first assemblage D1, comprising C. morsianus var. moelleri (high occurrence), C. morsianus (a few) and T. regina (rare), is recorded in the Sorrus, La Calotterie, Bois Gorguette, Le Goulet and lower Saint-Aubin Units. The second assemblage D2, characterized by great abundance of F. antiqua and decreasing numbers of C. morsianus species, is found in the upper part of the Saint-Aubin Unit. These diatom assemblages can be correlated with other biozonations established in the North Sea basin and based on nannoplankton and diatoms (King, 1983), diatoms (Mitlehner, 1996) and calcareous nannoplankton (Martini, 1971). F. antiqua and C. morsianus var. moelleri are the more important diatom species encountered. F. antiqua (previously named Coscinodiscus sp. 1, King, 1983) is the fossil marker used by King to define his NSP4 biozone. The base of NSP4 corresponds to the lowest occurrence of F. antiqua and the top is marked by the highest occurrence of this species. The Sorrus, La Calotterie, Bois Gorguette, Le Goulet and Saint-Aubin (“Sparnacian” stage) Units, the last one comprising F. antiqua (increasing to acme), correspond to the NSP4 zone of King (1983). The Mont-Hulin and Château de la Bruyère Units correspond probably to the NSP5 zone of King (1983). F. antiqua and C. morsianus var. moelleri were also used by Mitlehner (1996) to differentiate the NSP4a zone (low abundance of F. antiqua and abundance of C. morsianus var. moelleri) from the NSP4b zone (great abundance and acme of F. antiqua). The abundance of C. morsianus var. moelleri is recorded in the lower part of Saint-Aubin Unit and the abundance of F. antiqua (high peak at 46% in the sample 17.87) is found in the upper part of Saint-Aubin Unit. Therefore, Sorrus, La Calotterie, Bois Gorguette, Le Goulet and the lower part of Saint-Aubin Units are correlated with the NSP4a zone and the upper part of Saint-Aubin Unit with the NSP4b zone. By correlating the calcareous nannoplankton biozonation (NP) of Martini (1971) with the biozonation of King (1983), adapted by Mitlehner (1996), the boundary between the NP9 and NP10 zones could be located in the 18.75-17.87 interval and the NP10-NP11 boundary at the top of Saint-Aubin Unit. Lithostratigraphic correlations between the units of the Saint-Josse borehole (Paris basin) and the formations of the North Sea basin can be established. The great abundance of C. morsianus var. moelleri and the great abundance (including acme) of F. antiqua are respectively recorded in the Sele and Balder Formations. Therefore, the interval Sorrus/lower Saint-Aubin Units is correlated with the Sele Formation and upper part of Saint-Aubin Unit with the Balder Formation.     

Groupe de Candona clivosa, nouveau groupe de Candoninae (Crustacea, Ostracoda) et sa diversification dans le Bassin de Turiec (Slovaquie) au Miocène supérieur

The Candona clivosa-group, a new group of Candoninae - is erected on the base of the folded inner lamella observed on the postero-ventral border of the female valve. The elongated sub-triangular and rectangular valves of this group are described from the Miocene and Quaternary deposits of Europe. A folded inner lamella is also observed in 11 species of Candona from the Upper Miocene of the Turiec Basin of which 8 are new - Candona fatrica nov. sp., C. tatrica nov. sp., C. lacustris nov. sp., C. palustris nov. sp., C. stagnosa nov. sp., C. laterisimilis nov. sp., C. jiriceki nov. sp. and C. incurva nov. sp. They show a higher variability of the outline, calcification of the valves and width of the zone of concrescence. The extrinsic and intrinsic aspects of this morphological diversification in the Turiec Basin are discussed.     

Succession des peuplements d’échinides du Crétacé inférieur dans le Haut Atlas atlantique (Maroc) ; révision systématique et intérêt stratigraphique

The rich echinoid fauna from the Lower Cretaceous of the Agadir, Haha and Essaouira basins has been revised taxonomically, based on new and abundant collections. Forty-seven species have been identified, of which 20 are reported for the first time from this region and a further three represent new species left in open nomenclature. Their stratigraphic ranges have been determined accurately by reference to the regional ammonite zonation. However, it is also possible to use a local biostratigraphical scheme based on the echinoid succession, where ammonites are rare. The Toxasteridae, which are very abundant and underwent rapid evolution, are the most useful biostratigraphically. The genus Toxaster appears in the Lower Berriasian and continues until the Gargasian passing through the following succession: T. africanus, T. granosus, T. kiliani, T. lorioli, T. obtusus, T. exilis, T. retusus, T. maurus, T. peroni, T. collegnoi. The presence of Eoholaster, Macraster, Douvillaster, Palhemiaster, Heteraster and Pygopyrina is signaled for the first time in the Atlantic High Atlas region.     

Révision des rudistes crétacés (Bivalvia) de la Paléontologiefrançaise d’Alcide d’Orbigny

In the volume IV of the “Paléontologiefrançaise”, A. d’Orbigny deals in particular with Cretaceous rudists. Numerous type specimens of Hippuritidae, Caprinidae, Radiolitidae, Caprotinidae, Requieniidae belonging to his collection and deposited in the Muséumnationald’histoirenaturelle, Paris. Among the 78 species that he has studied, A. d’Orbigny introduced 41 as new ones. This review presents a picture of all types with a comment on the original definition, and set the statute, and the precise stratigraphical level of illustrated forms. The present work gives rise to the designation of 40 lectotypes, two syntypes and three neotypes.     

Interactions between root and shoot competition among four plant species in an alpine meadow on the Tibetan Plateau

At low altitudes, the interactions between root and shoot competition on plant growth have been extensively studied, and such interaction effects were found to be positive, negative, or neutral. However, little is known about such effects at high altitudes where the environmental conditions are harsher than those at low altitudes. We carried out a field experiment in an alpine meadow in the northeast Tibetan Plateau to test the hypothesis that a negative interaction between root and shoot competition exists for alpine plant species. Root and shoot competition were experimentally manipulated in the four grass species (Kobresia humilis, Saussurea superba, Stipa aliena and Elymus nutans). We found that K. humilis and S. aliena grew better without competition, whereas S. superba grew better with shoot competition and E. nutans grew better with root competition. The interactions between root and shoot competition were negative in K. humilis, positive in S. superba, but neutral in S. aliena and E. nutans, suggesting that the interaction effects are species-specific. This study also suggested that alpine plants may trade off both plant–plant interactions and competition shift between root and shoot to adapt to stressful environments.     

Composition des faunes de mammifères quaternaires en Chine selon un gradient Nord-Sud

During Pleistocene Epoch, quite a number of meridionale mammals inhabited north China, such as Stegodon, Macaca, Hystrix, Dicerorhinus, Bubalus and several kinds of other probscideans. On the contrary, some boreal mammals once appeared in south China, such as Megaloceros and Ursus arctos etc. The frequent appearance of warm-adapted mammals in North China can be explained by two alternative assumptions: one is that those mammals were of palearctic origin; the other is the climate-driven northward dispersal. According to the fossil evidences, it seems that the northward dispersal events were much more frequent than the southward ones during Pleistocene in China. It maybe means that during Pleistocene, the temperature fluctuated very frequently and the boundary between the Paleartic and Oriental Regions were always keeping on shifting. But it's still not clear how many episodes or phases of such dispersal events had occurred. The Qinling Mountains weren't responsible for the faunal differentiation between the North and the South, neither was the Yangtze River; because the former is not big enough, and the latter cannot be regarded as an efficient barrier for those animals, which are capable of flying and swimming. Maybe the climate zonation is the root cause of zoogeographical changes.     

Reconstitution de la végétation et du paléoclimat à partir de charbons de bois prélevés sur les sites du Paléolithique dans la région de Jungwon en Corée

Hundreds samples of the charcoals and waterlogged woods excavated from three late Paleolithic sites (Suyanggae, Gunang-gul and Sorori) in the Jungwon Region in Korea were analyzed to reconstruct the vegetation and paleoclimate in the region. Analyzed charcoal samples from the Suyanggae site show that the climate was cold in the lower part and it became more warm and dry in the upper part. From the Gunang-gul site, only Picea spp. was identified, indicating cool climates. From the Sorori site, two species were found; Alnus spp. (95%) and Ulmus spp. (5%). No conifers were discovered. The species composition indicates that the climate condition of central Korean peninsula around 8,800 BP was little cooler and wetter condition.     

Size analyses of the coccolith species Biscutum constans and Watznaueria barnesiae from the Late Albian “Niveau Breistroffer” (SE France): taxonomic and palaeoecological implications

The size variability of the coccolith species Biscutum constans and Watznaueria barnesiae has been studied in 50 samples of the Late Albian “Niveau Breistroffer” black shales (Col de Palluel section, SE France). For each species, length and width of the total coccolith and the central unit have been measured in 60 specimens per sample. In addition, ellipticity and central unit length/total coccolith length ratio have been calculated. This study aims to improve our understanding of the taxonomic concepts of B. constans and W. barnesiae, and to test whether the size changes correspond to palaeoceanographic changes interpreted from other proxies. Two morphotypes (varieties) were differentiated for each of the two taxa studied. B. constans includes morphotypes with a narrow (B. constans var. constans) and a wide central unit (B. constans var. ellipticum). Between these two morphotypes no significant size differentiation has been observed. They seem to represent end members of a size continuum, relating to both the total coccolith- and central unit size distribution. The two morphotypes of W. barnesiae differ only in the absence (W. barnesiae var. barnesiae) and the presence of a narrow central opening (W. barnesiae var. fossacincta). No significant size differences have been observed between these two morphotypes, and both show similar distributions of the measured characteristics. We recommend that the studied morphotypes of both B. constans and W. barnesiae should not be assigned to different morphospecies. Throughout the studied black shale succession the measured parameters show mostly statistically insignificant, short-term fluctuations, but significant long-term trends have been observed for B. constans. These show a trend to smaller forms with a narrower central unit in the upper part of the succession. This change coincides with a cooling trend, as indicated by a nannofossil based temperature index. Therefore the two morphotypes of B. constans are interpreted to represent ecophenotypic varieties rather than two different morphospecies. No clear relationship has been recognized between the size of W. barnesiae and the palaeoenvironmental conditions.     

Trace and soft body fossils from the Pedroche Formation (Ovetian, Lower Cambrian of the Sierra de Córdoba, S Spain) and their relation to the Pedroche event

The low Lower Cambrian rocks from the Sierra de Córdoba contain one of the best successions in Europe, which consists of well exposed mixed facies with abundant fossil assemblages showing long stratigraphical ranges throughout the Pedroche Formation. These assemblages include diverse Ovetian archaeocyaths, trilobites, small shelly fossils, calcimicrobes, trace fossils and stromatolites. Trace fossils are still poorly known, and thus they are the main objective of this work. Ichnological data are obtained from the Arroyo de Pedroche 1, Arroyo de Pedroche 2 and Puente de Hierro sections. Trace fossils include the ichnogenera Bergaueria, aff. Bilinichnus, Cochlichnus, aff. Cosmorhaphe?, Cylindrichnus, Dactyloidites, Dimorphichnus, Diplichnites, Monocraterion, Palaeophycus, aff. Phycodes, Planolites, Psammichnites, Rusophycus, Skolithos, Torrowangea and Treptichnus, as well as faecal pellets, meniscate trace fossils and others. They are abundant in shales and sandstones, and indicate important changes in the benthic conditions with respect to the underlying Torreárboles Formation. Changes in fossil assemblages within Member I of the Pedroche Formation indicate palaeoecological disruptions, which led to the disappearance of numerous archaeocyath species and the decrease of stromatolite biodiversity. This was followed by dominance of trilobite and brachiopod assemblages, accompanied by trace fossils of the Psammichnites ichnosp. A ichnoassociation. This biotic turnover (Pedroche event) occurred at the lower part of the archaeocyath Zone III, within the Bigotina bivallata biozone. The diagnoses of the ichnospecies Cochlichnus anguineus and Dactyloidites cabanasi are emended.     

Biostratigraphie,paléoécologie et zones d’acmé des foraminifères planctoniques au passage Crétacé-Paléogène dans la Téthys (Tunisie et Espagne) et l’Atlantique (France)

Biostratigraphical high-resolution analyses and quantitative data confirm that deposition is continuous across the K-Pg transition in several sections in Tunisia (El Kef, stratotype section) and Spain (Agost and Caravaca sections) located in the Tethyan realm and the Bidart sections in the Atlantic realm, without any relevant hiatus. The Upper Maastrichtian assemblages of planktic foraminifera from these sections are largely dominated by small biserial heterohelicids. They are associated to common species having planispiral test (i.e. globigerinelloids), trochospiral test (i.e. hedbergellids, rugoglobigerinids globotruncanids), to rare triserial heterohelicids (i.e. guembelitriids) and trochospiral species showing tubulospines (i.e. schackoinids). Stratigraphical ranges of these diverse taxa through the late Maastrichtian in the Tethyan and Atlantic realms show very few changes in the planktonic foraminiferal assemblages and most of the species are present in the Abathomphalus mayaroensis biozone. By our high-resolution sampling and the intensive research for the A. mayaroensis index species in the uppermost Maastrichtian samples, we confirm that this species is omnipresent up to the top of the Maastrichtian. Therefore, A. mayaroensis is present in almost all samples which are late Maastrichtian in age, but this species became very scarce in the uppermost Maastrichtian samples. This scarceness could be due to a climate cooling. A sharp decrease in relative abundance of the deep dwellers species, like as Abathomphalus intermedius and A. mayaroensis as well as in other keeled globotruncanids is observed at the studied sections from the Tethyan realm (indicative of low latitude) across the latest Maastrichtian. At the K/Pg boundary, all the globotruncanids disappeared. They are considered specialists living in tropical-subtropical deep seawater habitat. At this boundary, large and ornate heterohelicids also disappeared. Therefore, all the studied sections show that about 90 % of the Maastrichtian species became extinct according to a catastrophic mass extinction pattern. Only about 10 % crossed the K/Pg boundary and survived during the earliest Danian. The minor difference in the number of disappeared taxa is related to their latitude location or environment paleodepth. The changes in the species relative abundance, observed in the successive planktic foraminiferal assemblages, make it possible to recognize the Acme-stage 0 typical of the upper Maastrichtian interval. It is characterized by the highest species richness of Globotruncanids and heterohelicids specialists of tropical to subtropical marine conditions, the Acme-stage 1 typical of the Guembelitria cretacea Zone, and in particular of the Hedbergella holmdelensis Subzone dominated by “opportunists” species belonging to Guembelitria, the Acme-stage 2 which corresponds to the Pv. eugubina Zone dominated mainly by specimens belonging to Palaeoglobigerina and Parvularugoglobigerina and the Acme-stage 3 which characterizes mainly the Ps. pseudobulloides Zone dominated by biserial species belonging to Chiloguembelina and Woodringina.     

Crustacés décapodes du Kimméridgien de Bure (Lorraine, France)

A new fauna of crustacean decapods has been discovered on the occasion of the digging of two large wells in Kimmeridgian marls and limestones at Bure (Lorraine, France). Among the stratigraphically well repaired identified fossils Thalassinoidea are abundant with the species Etallonia isochela (Woodward, 1876). Other fossils are represented by a Paguridae (Palaeopagurus sp.) and two Erymidae: Eryma cf. babeaui (Etallon, 1861) and E. ventrosa (Von Meyer, 1835) this last one has provided fragments of carapace and fragments of chelipeds. Propodes of E. isochela sometime display remains of coloration patterns that indicate a low oxygen level of depositional and diagenetic environments.     

Taxonomie du grand canidé de la grotte du Vallonnet (Roquebrune-Cap-Martin, Alpes-Maritimes, France)

The large canid of Vallonnet cave belongs to Xenocyon lycaonoides. Its remains are similar to the Xenocyon from Untermassfeld in Germany. X. Lycaonoides, and evoluated form of the Xenocyon genus, is characteristic of the second half of the Lower Pleistocene. The genus Xenocyon may be the ancestor of both Cuon and Lycaon.     

New material of Ouranopithecus macedoniensis from late Miocene of Macedonia (Greece) and study of its dental attrition

During the last five years our continued excavations in the known late Miocene mammal localities of Macedonia (Greece) provided several new specimens of the hominoid primate Ouranopithecus macedoniensis. This new material includes maxillary and mandibular remains and it is described and compared to the old material of Ouranopithecus in the present article. The material of Ouranopithecus from the three known localities “Ravin de la Pluie” (RPl), “Xirochori 1” (XIR) and “Nikiti 1” (NKT) includes a complete series of tooth rows representing all wearing stages. Thus, the study of the dental wear of Ouranopithecus upper and lower teeth is studied and compared to that of the recent hominoids Gorilla and Pan, as well as to Australopithecus afarensis. The latter species is well known by a series of tooth rows of different wearing stages. The canine’s attrition of Ouranopithecus has a more derived pattern than that of the recent hominoids (Gorilla and Pan) and less derived than A. afarensis. The p3 of Ouranopithecus has similar attrition to that of A. afarensis, the attrition of the molars in Ouranopithecus, A. afarensis and Pan follows a similar pattern, while in Gorilla it is different.     

Evidence for the longest stratigraphic range of a post-Triassic Ichthyosaur: a Leptonectes tenuirostris from the Pliensbachian (Lower Jurassic) of Switzerland

A three-dimensional well-preserved ichthyosaur skull and parts of the postcranial skeleton are attributed to the species Leptonectes tenuirostris (Conybeare, 1822). It was found vertically embedded in Pliensbachian deposits representing three successive biozones (ibex to margaritatus Zone). The find is dated as early Late Pliensbachian (margaritatus Zone) by a rich ammonite and ostracod fauna. It is the first record of the genus Leptonectes from Switzerland and from the Late Pliensbachian. It is so far the best preserved and most complete ichthyosaur from this time interval worldwide. With diagnostic specimens known from the Rhaetian (Late Keuper) up to the early Late Pliensbachian (Middle Liassic), L. tenuirostris (Conybeare, 1822) has the most extensive stratigraphic range documented for any post-Triassic ichthyosaur so far.     

Les foraminifères du Paléocène et de l'Éocène basal du sillon nord-pyrénéen (Aquitaine, France)

This paper presents for the first time the inventory of the Paleocene and Lower Eocene foraminifers located in the North Pyrenean trough, between the Atlantic Ocean and the neighbourhood of the town Pau. They have been studied from three outcrops. The Bidart Beach section shows the Lasseube Formation from the Cretaceous/Paleogene boundary to the base of the P 3a zone. The Loubieng Quarry section, near Orthez, represents the upper P 3a zone and the lower P 3b zone as well as the Lasseube Formation / Pont Labau Formation boundary. The interval between the upper part of the P 3b zone and the upperest part of the P 5 zone crops out along the Gan - Rébénacq road with a hiatus located at the Paleocene / Eocene boundary, the whole interval belonging to the Pont Labau Formation. 394 taxons of foraminifers are present in this formations: 349 benthic and 45 planktonic species. The Velasco type benthic foraminifers show a middle bathyal depositional environment, with a paleobathymetry included between 500-600 m and 1000 m: Nuttallides truempyi, Osangularia velascoensis, Bulimina trinitatensis. The Midway type species which were transported by the turbidite currents from the lower to middle neritic environments are frequent as well as the Cretaceous reworked species. The species number is low: 29 in the iridium layer of the Cretaceous/Paleogene boundary (P 0 zone). Fauna grows rich quickly in the Pα zone reaching 129 species. The diversity grows up progressively from P α to the P 4a zone (NP 8). The disappearances are rare until his horizon, but their number is bigger than the number of appearances from the P 4b zone. It reaches their maximum between the P 4c zone and the P 5 zone. It shows that the decline of the Paleocene fauna begins around 2 million years before the thermal event of the Paleocene / Eocene boundary in the Aquitaine sections. The disappearances stay important in the Lower Eocene - Ypresian, but the appearance of lots of Eocene species show that the fauna renewal is located in this stage. The Cretaceous taxa dominate in the Paleocene benthonic fauna. The appearance or the disappearance of some species has a stratigraphic value in the Aquitaine region: the disappearance of Coryphostoma incrassata in the P 1b zone, the appearances of Plectina dalmatina, Elongobula grata (P α); Bulimina tuxpamensis (P 2); Tritaxilina cubensis, Thalmannita madrugaensis (P 3a); Svenia bulbosa (P 3b); Discorbis perovalis (P 4a/NP 7); Elongobula pulchra, E. pupa, Asterigerina bartoniana, Neorotalia gr. tuberculata (P 4a/NP 8); Bigenerina pannonica, Pentellina pseudosaxorum (P 5/NP 9-10).     

Histoire plio-pléistocène des écosystèmes végétaux de Méditerranée sud-occidentale : apport de l’analyse pollinique de deux sondages en mer d’Alboran

The pollen analysis of sediments of two boreholes located in the northeastern Morocco (Nador 1) and in the southeastern Spain (Andalucia G1) has allowed shedding a new light on the vegetation and climate of the North African littoral plains, the Rif Massif and the Betic Range during the Pliocene. The vegetation around the Alboran Sea was open and xeric during the Zanclean, dominated by herbs including subdesertic elements as Calligonum, Lygeum, Nitraria and Neurada. This type of vegetation indicates a dry and hot climate. The southwestern Mediterranean steppes have therefore a climatic character; they existed before the presence and the heavy pressure of Man on the environment. From the Piacenzian, the development, at Andalucia as well as at Nador, of Artemisia and the appearance of some altitude trees such as Cedrus and Cathaya indicate a vegetation change linked to a climatic change. Modifications in the vegetation observed during the same period in northwestern Mediterranean seem to indicate that the vegetation changes observed at Andalucia and Nador are controlled by the appearance of the first arctic glacial-interglacial cycles. The cedar tree appears at Nador only at the end of the Piacenzian, at time of the first cooling, while it has been found punctually in Messinian and early Pliocene sediments of Habibas in Algeria and in Messinian sediments of the Bou Regreg section at Salé in Morocco. The regular presence of Cedar at Bou Regreg during the Messinian, allows supposing that it was present in the Middle Atlas Mountains and that its development was favoured by colder conditions.     

Analysis of the factors that affect the distribution and abundance of three Neobuxbaumia species (Cactaceae) that differ in their degree of rarity

We studied three species of columnar cacti in the genus Neobuxbaumia which differ in their degree of rarity: Neobuxbaumia macrocephala (the rarest), Neobuxbaumia tetetzo (intermediate), and Neobuxbaumia mezcalaensis (the most common). To investigate the ecological factors that limit their distribution and abundance, we surveyed 80 localities within the region of Tehuacan-Cuicatlán, in Central Mexico. At each locality we measured several environmental variables, and the density of the Neobuxbaumia populations present. We used a principal component analysis (PCA) to identify the factors that are associated to the presence/absence of each species. Additionally, we carried out multiple regressions between environmental variables and population density to test whether the variation in these variables was related to changes in abundance. The results show that factors significantly affecting the distribution of these species are mean annual temperature, altitude, rainfall, and soil properties such as texture and organic matter content. N. mezcalaensis reaches maximum population densities of 14,740 plants per ha (average density = 3943 plants per ha) and is associated with localities with relatively abundant rainfall. N. tetetzo shows maximum population densities of 14,060 plants per ha (average = 3070 plants per ha), and is associated with sites located at high latitudes and with high phosphorous content in the soil. The rarest species, N. macrocephala, shows maximum densities of 1180 plants per ha (average = 607 plants per ha) and is associated with localities with high soil calcium content. The distribution of this species is limited to sites with specific values of the environmental variables recorded, conferring it a high habitat specificity which accounts for its rarity.