Brachiopodes toarciens de la coupe d’Aït Athmane,Haut Atlas centro-oriental (Maroc)

The Aït Athmane section is located in the eastern part of the central High Atlas, about 20 km north of Errachidia. Its Toarcian series consists of silty marl and silty marly limestone. Its lower part, of early (Levisoni Zone?) to middle Toarcian (Bifrons and Gradata? Zones) age, is rich in brachiopods (rhynchonellides and terebratulides). Its upper part, of late Toarcian age, is rich in terebratulids. Among eighteen reported brachiopod taxa, six are new for the central High Atlas. Their specific assemblages and stratigraphic distributions are similar to those of the Toarcian brachiopods reported from several basins of northern and southern margins of the Tethys and the Subboreal domain, particularly of western France.     

Atlantic paleobathymetry, paleoproductivity and paleocirculation in the late Albian: the benthic foraminiferal record

Spatial distribution patterns of benthic foraminifers in upper Albian sediments from 25 DSDP/ODP sites and 31 onshore sections of the North and South Atlantic Ocean are used to generate paleobathymetric reconstructions and to identify areas of high primary production such as coastal and equatorial upwelling zones. New paleobathymetric estimates are provided for DSDP/ODP sites and onshore locations that are not situated on oceanic crust. Paleobathymetric reconstructions indicate shallow water exchange between the North and South Atlantic but show the existence of a deep-water connection between the western and eastern Tethys (>2500 m) through the Gibraltar Gateway. Strikingly, there is no evidence for a strong latitudinal gradient in deep-water benthic foraminiferal distribution during the late Albian: South Atlantic assemblages show close affinity to North Atlantic and Tethyan assemblages, exhibiting only a minor degree of provincialism. Biogeographic patterns reveal a distinct asymmetry in late Albian paleoproductivity for the North Atlantic. As for the present day, the eastern margins of the Atlantic were generally more productive than the western margins, and a belt of enhanced carbon flux export to the seafloor can be traced around the north African coast, which probably corresponded to a zone of vigorous coastal upwelling. By contrast, assemblage composition in the South Atlantic generally reflects mesotrophic to oligotrophic conditions. Benthic foraminiferal distribution patterns, thus, provide robust proxy data to test predictions from paleocirculation and paleobathymetric models for the mid-Cretaceous Atlantic Ocean and adjacent margins.     

Gradient in seaweed vegetation patterns along the North Icelandic coast,related to hydrographic conditions

The benthic algal vegetation of the North Icelandic coast exhibits particular features, different from those found in southern and western Iceland on the one hand and in eastern Iceland on the other. It is characterised by low-eulittoral belts of Devaleraea ramentacea, Petalonia species and Chordaria flagelliformis as well as by meadows of diverse Acrosiphonia species. In the tide pools Atlantic and typical North Icelandic associations occur side by side. This area thus represents an intermediate region between Atlantic and subarctic growth conditions. A certain gradient in the west-east direction was observed within the North Icelandic vegetation type and is possibly related to the gradual cooling and dilution of the water masses which pass the North Icelandic shelf. The hydrographic discontinuity in the extreme NW creates a sharp floristic and vegetational boundary for benthic algae, whereas in the NE a gradual transition towards the subarctic East Icelandic vegetation was observed. The Atlantic character of the vegetation was more pronounced in the western than in the eastern part of the north coast. An enclave of warm water vegetation was interposed along the western banks of the Tjörnes peninsula.This paper is based on a presentation at the XIII^th International Seaweed Symposium in Vancouver, Canada.     

Stratigraphie et sédimentologie des « Couches rouges » continentales du Jurassique-Crétacé du Haut Atlas central (Maroc) : implications paléogéographiques et géodynamiques

In several synclines of the central High Atlas, the “Redbeds” following the closure of the marine Tethyan Atlasic trough during the Middle Jurassic are constituted by three successive formations or units of continental deposits dated recently with biostratigraphical elements. Some micropaleontological markers, mainly charophytes and ostracods, allow to precise the stratigraphy in agreement with a Bathonian-?Callovian assignment for the lower unit (Guettioua Formation), and in dating the middle and upper units. The Upper Jurassic, mainly the Kimmeridgian, is developed in the lower part of the middle unit (Iouaridene Formation). The Barremian has been recognized in this middle unit and in the upper unit (Jbel Sidal Formation). The Jurassic/Cretaceous boundary is thus delimited for the first time with micropaleontological data. These new data are very significant for the Atlasic history during the Mesozoic. The basaltic flows inserted in the continental Jurassic-Cretaceous deposits of the central High Atlas result from two separate events in the Middle Jurassic and in the Barremian. The tectogenesis in the basins is characterized by a polyphase process including notably a synsedimentary tectonic activity conspicuous in the Barremian. The evidence of marine to brackish intercalations allows moreover to date the first Cretaceous transgressive event on the NW boundary of the High Atlas during the Lower Barremian and to consider an Atlantic paleogeographical interaction. SW margin of the Tethyan trough in the Lower and Middle Jurassic, the central High Atlas is merged with the margin of the central Atlantic Ocean during the Lower Cretaceous.     

Evolution of the Madrean–Tethyan disjunctions and the North and South American amphitropical disjunctions in plants

Abstract The present paper reviews advances in the study of two major intercontinental disjunct biogeographic patterns: (i) between Eurasian and western North American deserts with the Mediterranean climate (the Madrean–Tethyan disjunctions); and (ii) between the temperate regions of North and South America (the amphitropical disjunctions). Both disjunct patterns have multiple times of origin. The amphitropical disjunctions have largely resulted from long‐distance dispersal, primarily from the Miocene to the Holocene, with available data indicating that most lineages dispersed from North to South America. Results of recent studies on the Mediterranean disjuncts between the deserts of Eurasia and western North America support the multiple modes of origin and are mostly consistent with hypotheses of long‐distance dispersal and the North Atlantic migration. Axelrod's Madrean–Tethyan hypothesis, which implies vicariance between the two regions in the early Tertiary, has been favored by a few studies. The Beringian migration corridor for semiarid taxa is also supported in some cases.     

Paleobiogeography of Mesozoic brachiopod faunas from Andean–Patagonian areas in a global context

During the Mesozoic, the Andean region has played a hinging role between high- and low-latitude faunas, which are, respectively, characterized by stocks that display long-term fidelity. This paper is aimed at providing an updated review of Late Triassic to Late Cretaceous South American articulated brachiopods in the light of previous knowledge at worldwide scale. Late Triassic brachiopods from the Argentine–Chilean Andes show unmistakable Maorian (or Notal) faunal elements alongside some more cosmopolitan genera, with certain influence of Eastern Pacific taxa. By Early Jurassic times, differentiation of Tethyan and Boreal Realms became progressively evident in Europe. In South America, Hettangian–Sinemurian brachiopod faunules from the Argentinian Andes are somewhat impoverished, with mostly cosmopolitan genera showing certain affinities to Maorian species, and with the addition of some endemics later. Increasingly, diverse Pliensbachian Andean brachiopods denote close relationships to Celto-Swabian taxa, then by Domerian times, a certain degree of endemism was developed, though somewhat delayed Tethyan influences, and persistent links with New Zealand are subordinately recognizable, too; most Toarcian assemblages reveal basically Celto-Swabian and Iberian affinities as well. East-west austral links across the Pacific may have been favored by migratory routes fringing the Gondwana margin, whereas faunal exchange with the western end of the Tethys appears to reflect an intermittent shallow-marine connection through the Hispanic Corridor. During the Middle Jurassic, distinction of Tethyan and Boreal Realms was maintained in the northern Hemisphere, and the differentiation of an Ethiopian or Southern Tethyan fauna became better characterized. Aalenian and Bajocian brachiopods of the Andes display generic affinities mainly with those from western Europe, with some minor endemic developments; brachiopods recorded from the Bathonian–Callovian of Argentina (and Chile) also occur along the northern Tethyan margin, yet with some genera extending into Indo-Ethiopian areas. During the Late Jurassic, Boreal faunas from high-latitudes became even more strongly differentiated from low-latitude, Tethyan ones. Oxfordian and Tithonian brachiopods from the Andes apparently belong to genera of cosmopolitan or northern Tethyan affiliation, yet there are few elements in common with other eastern Pacific areas, such as Mexico. Early Cretaceous brachiopods, in addition to Andean basins of Chile and western Argentina, are known also from Patagonia and Tierra del Fuego. They belong mostly to widely distributed, mainly Tethyan genera, with some quasi-cosmopolitan and circum-Pacific components (some shared with Antarctica become noticeable). Late Cretaceous brachiopods from northern Patagonia show significant affinities to Maastrichtian ones of northwest Europe and central Asia, which calls for further assessing the potential role that may have played the trans-Saharan passageway in such dispersal. Broad aspects of Mesozoic brachiopod paleobiogeography are fairly well understood, yet details of ranking and naming of certain units are still in need of more agreement.     

Phylogeography and population dynamics of the white-sided dolphin (Lagenorhynchus acutus) in the North Atlantic

Highly mobile species in the marine environment may be expected to show little differentiation at the population level, but this is often not the case. Instead cryptic population structure is common, and effective conservation will require an understanding of how these patterns evolve. Here we present an assessment from both sides of the North Atlantic of differentiation among populations of a dolphin species that inhabits mainly pelagic waters, the Atlantic white-sided dolphin. We compare eleven putative populations in the western and eastern North Atlantic at mtDNA and microsatellite DNA loci and find reduced nucleotide diversity and signals for historical bottlenecks and post-bottleneck expansions in all regions. We calculate expansion times to have occurred during the early Holocene, following the last glacial maximum (LGM). We find evidence for connectivity among populations from either side of the North Atlantic, and differentiation between putative populations in the far northeast compared with all other areas sampled. Some data suggest the possibility of separate refugia during the LGM explaining this pattern, although ongoing ecological processes may also be a factor. We discuss the implications for developing effective programs of conservation and management in the context of ongoing anthropogenic impact.     

Impact of mountain chains, sea straits and peripheral populations on genetic and taxonomic structure of a freshwater turtle, Mauremys leprosa (Reptilia, Testudines, Geoemydidae)

Mauremys leprosa, distributed in Iberia and North‐west Africa, contains two major clades of mtDNA haplotypes. Clade A occurs in Portugal, Spain and Morocco north of the Atlas Mountains. Clade B occurs south of the Atlas Mountains in Morocco and north of the Atlas Mountains in eastern Algeria and Tunisia. However, we recorded a single individual containing a clade B haplotype in Morocco from north of the Atlas Mountains. This could indicate gene flow between both clades. The phylogenetically most distinct clade A haplotypes are confined to Morocco, suggesting both clades originated in North Africa. Extensive diversity within clade A in south‐western Iberia argues for a glacial refuge located there. Other regions of the Iberian Peninsula, displaying distinctly lower haplotype diversities, were recolonized from within south‐western Iberia. Most populations in Portugal, Spain and northern Morocco contain the most common clade A haplotype, indicating dispersal from the south‐western Iberian refuge, gene flow across the Strait of Gibraltar, and reinvasion of Morocco by terrapins originating in south‐western Iberia. This hypothesis is consistent with demographic analyses, suggesting rapid clade A population increase while clade B is represented by stationary, fragmented populations. We recommend the eight, morphologically weakly diagnosable, subspecies of M. leprosa be reduced to two, reflecting major mtDNA clades: Mauremys l. leprosa (Iberian Peninsula and northern Morocco) and M. l. saharica (southern Morocco, eastern Algeria and Tunisia). Peripheral populations could play an important role in evolution of M. leprosa because we found endemic haplotypes in populations along the northern and southern range borders. Previous investigations in another western Palearctic freshwater turtle (Emys orbicularis) discovered similar differentiation of peripheral populations, and phylogeographies of Emys orbicularis and Mauremys rivulata underline the barrier status of mountain chains, in contrast to sea straits, suggesting common patterns for western Palearctic freshwater turtles.     

Relicts of the Tethyan fauna in the Recent oceans

In tropical, two subtropical, low boreal, and notal zones of the Recent oceans, the distribution of benthic relict animal taxa is clarified for brachiopods, mollusks, echinoderms, and some other taxonomic groups. The richest places for relict taxa are situated at the depths of the bathyal zone (intermediate zone between shallow water and the abyss). Preservation of relicts is a result of oceanic asymmetry and the disposition of faunal centers in the western parts of the oceans. This asymmetry in hydrological and biological structure of the oceans depends on the global peculiarities of the Earth. The Atlantic and Indian ocean centers show faunal exchanges and their origin from West Tethyan ancestors.     

Bacteriophage sensitivity patterns among bacteria isolated from marine waters

Phage-host cross-reaction tests were performed with 774 bacterial strains and 298 bacteriophages. The bacteria (bacteriophages) were isolated at different times from water samples collected in the Atlantic Ocean between the European continental shelf and the Sargasso Sea: 733 (258) strains; in the North Sea near Helgoland: 31 (31) strains; and in the Bay of Biscay: 10 (9) strains. Of the Atlantic Ocean bacteria 326 were found to be susceptible to one or more Atlantic Ocean bacteriophage(s). The bacteriophage sensitivity patterns of these bacteria vary considerably, placing 225 of them in two large clusters of bacteriophage-host systems. Taking all into account, 250 of the 326 Atlantic Ocean bacteria are different from each other. This high degree of variation among the bacteria distinguishes microbial populations derived from widely separated eastern and western regions of the Atlantic Ocean. It also sets apart from each other the populations derived from samples collected at successive stations some 200 miles apart, although to a lesser degree. With bacterial populations found from samples collected on the way back and forth between Europe and the Sargasso Sea a gradual change was observed from "western" phage sensitivity patterns to "eastern" ones. Sixty-nine Atlantic Ocean bacteria are sensitive to bacteriophages isolated from the North Sea and the Bay of Biscay; of these only 26 strains are also susceptible to Atlantic Ocean phages. The interpretation of the results is based on the hydrographical conditions prevailing in the northern Atlantic Ocean including the North Sea, and on the assumption that the microbial populations investigated have undergone genetic changes while being transported within water masses from west to east.     

Multicolony tracking reveals the winter distribution of a pelagic seabird on an ocean basin scale

Aim An understanding of the non‐breeding distribution and ecology of migratory species is necessary for successful conservation. Many seabirds spend the non‐breeding season far from land, and information on their distribution during this time is very limited. The black‐legged kittiwake, Rissa tridactyla, is a widespread and numerous seabird in the North Atlantic and Pacific, but breeding populations throughout the Atlantic range have declined recently. To help understand the reasons for the declines, we tracked adults from colonies throughout the Atlantic range over the non‐breeding season using light‐based geolocation. Location North Atlantic. Methods Geolocation data loggers were deployed on breeding kittiwakes from 19 colonies in 2008 and 2009 and retrieved in 2009 and 2010. Data from 236 loggers were processed and plotted using GIS. Size and composition of wintering populations were estimated using information on breeding population size. Results Most tracked birds spent the winter in the West Atlantic, between Newfoundland and the Mid‐Atlantic Ridge, including in offshore, deep‐water areas. Some birds (mainly local breeders) wintered in the North Sea and west of the British Isles. There was a large overlap in winter distributions of birds from different colonies, and colonies closer to each other showed larger overlap. We estimated that 80% of the 4.5 million adult kittiwakes in the Atlantic wintered west of the Mid‐Atlantic Ridge, with only birds from Ireland and western Britain staying mainly on the European side. Main conclusions The high degree of mixing in winter of kittiwakes breeding in various parts of the Atlantic range implies that the overall population could be sensitive to potentially deteriorating environmental conditions in the West Atlantic, e.g. owing to lack of food or pollution. Our approach to estimating the size and composition of wintering populations should contribute to improved management of birds faced with such challenges.     

Genetic differentiation between parapatric 'nearshore' and 'offshore' populations of the bottlenose dolphin.

The existence of nearshore and offshore populations of the bottlenose dolphin has been documented throughout its range. In several cases the two regional forms have been shown to be morphologically distinct, although there is considerable overlap for most characters. The populations off the eastern coast of North America have been the subject of a long-term programme of research on their distribution and movements. In this study, we compare mitochondrial and nuclear genetic markers between dolphins classified as either nearshore or offshore type. These putative populations were found to be distinct at both nuclear and mitochondrial genetic markers. Further, the level of variation among the nearshore dolphins was reduced compared with the offshore population. A broader geographical comparison suggests a shared lineage between offshore dolphins from the western North Atlantic and both offshore and nearshore dolphins from the eastern Atlantic. These results are consistent with local differentiation based on habitat or resource specialization in the western North Atlantic, and suggest differences in the character of the nearshore/offshore distinction in different parts of the world.     

Geographic patterns in morphometric and genetic variation for coyote populations with emphasis on southeastern coyotes

Prior to 1900, coyotes (Canis latrans) were restricted to the western and central regions of North America, but by the early 2000s, coyotes became ubiquitous throughout the eastern United States. Information regarding morphological and genetic structure of coyote populations in the southeastern United States is limited, and where data exist, they are rarely compared to those from other regions of North America. We assessed geographic patterns in morphology and genetics of coyotes with special consideration of coyotes in the southeastern United States. Mean body mass of coyote populations increased along a west‐to‐east gradient, with southeastern coyotes being intermediate to western and northeastern coyotes. Similarly, principal component analysis of body mass and linear body measurements suggested that southeastern coyotes were intermediate to western and northeastern coyotes in body size but exhibited shorter tails and ears from other populations. Genetic analyses indicated that southeastern coyotes represented a distinct genetic cluster that differentiated strongly from western and northeastern coyotes. We postulate that southeastern coyotes experienced lower immigration from western populations than did northeastern coyotes, and over time, genetically diverged from both western and northeastern populations. Coyotes colonizing eastern North America experienced different selective pressures than did stable populations in the core range, and we offer that the larger body size of eastern coyotes reflects an adaptation that improved dispersal capabilities of individuals in the expanding range.     

The occurrence of Acanthurus monroviae(Perciformes: Acanthuridae) in the south‐western Atlantic, with comments on other eastern Atlantic reef fishes occurring in Brazil

The presence of 'vagrants' of the eastern Atlantic surgeonfish Acanthurus monrovia e is confirmed for the south‐eastern coast of Brazil. Three other species, Aulostomus strigosus (Aulostomidae), Parablennius pilicornis (Blenniidae) and Epinephelus marginatus (Serranidae) have apparently also crossed the Atlantic from east to west, whereas the great majority of 'amphi‐Atlantic' species appears to have their origin in the western Atlantic.     

Population structure in the pan-oceanic wreckfish, Polyprion americanus (Teleostei: Polyprionidae), as indicated by mtDNA variation

The wreckfish Polyprion americanus , a large [>1 m total length ( L _T)] demersal teleost, is distributed globally in temperate waters, including both sides of the North and South Atlantic Oceans, the Mediterranean, the western South Pacific, and the southern Indian Ocean. Wreckfish spawn off the south-eastern U.S. on an area of the Blake Plateau (the Charleston Bump) characterized by an extensive ridge having approximately 100 m relief, in 450–600 m depths. Juvenile wreckfish (<60 cm L _T) are pelagic and, in the North Atlantic, are not reported from the Blake Plateau fishing area, but occur in by-catch and fishery landings in the eastern Atlantic. Analysis of nine restriction fragment length profiles from a PCR-amplified fragment (∼1.5 kb) of the ND1 mitochondrial gene indicated no stock separation between eastern North Atlantic (Azores, Majorca, Madeira), and western North Atlantic (Blake Plateau) wreckfish. Restriction site differences separate western South Atlantic wreckfish from the North Atlantic; however, South Atlantic wreckfish share restriction-site similarities with western Pacific wreckfish that are not shared with North Atlantic wreckfish. North Atlantic circulation provides a mechanism for a long-lived pelagic stage to be dispersed from Blake Plateau spawning grounds to the eastern North Atlantic. Global circulation patterns may explain both the dispersal of mtDNA haplotypes and the disjunct distribution of wreckfish body lengths in a temperate, deep-water vagile species with an extended pelagic juvenile stage such as wreckfish.     

贵州关岭上三叠统的楯齿龙类化石

记述的新铺中国豆齿龙（Sinocyamodus xinpuensis gen. et sp. nov.）代表了齿龙目的一新属。这是齿龙类化石在我国的首次发现,也是这类特殊的海生爬行动物在欧洲、北非和中东以外地区的首次记录。标本产自贵州关岭晚三叠世瓦窑组一段的泥晶灰岩中,与鱼龙类、幻龙类及海龙类等多种海生爬行类同时产出。这一化石组合表明,贵州关岭三叠纪海生爬行动物群的性质与西特提斯动物群（wstern-Tethyan fauna）的性质极为相似,这两个地区在晚三叠世早期有广泛的动物交流。     

Corals,molluscs, and paleogeography of late Jurassic strata of the Cerro Pozo Serna,Sonora, Mexico

Jurassic strata of late Oxfordian to early Kimmeridgian age are reported from the Cerro Pozo Serna, northwest-central Sonora, Mexico. The presence of these marine strata greatly alters previous paleogeographic reconstructions of the Tethyan embayment in this area. Approximately 60% of the Sonorian molluscan fauna has been previously reported from Jurassic horizons in the Gulf of Mexico and west Texas regions. The remainder of the molluscs have previously been reported from either the western interior United States or Canada. The Pozo Serna faunas seem to be zoogeographically transitional between communities present in the southern North American Tethyan realm and regions within the central North American Tethyan realm. Additionally, the Pozo Serna fauna greatly resembles contemporaneous communities reported from the Jurassic of Morocco. Corals from the Sonorian Jurassic prove to be, in large part, conspecific with well-known central European and Algerian reef-building species. The scleractinian Lepidophylliopsis gen. nov. and Macgeopsis sonorensis sp. nov. are described from central Sonora.     

A new species of Nezumia (Gadiformes: Macrouridae) from Fieberling Guyot, eastern North Pacific Ocean

A new species of the macrourine genus Nezumia is described from specimens collected from the crest of Fieberling Guyot in the eastern North Pacific Ocean. The new species is distinguishable from all other Nezumia species chiefly on the basis of a high number of pelvic fin rays, a relatively high number of first dorsal fin soft rays, a steep blunt snout lacking scales ventrally, a relatively weak suborbital ridge, and a moderately long, thin barbel. Although possibly endemic to the Baja California Seamount Province, the new species appears to lack close relatives among the other Nezumia species of the eastern North Pacific Ocean. It might have arrived there either from a disjunct Tethyan distribution, thus retaining close relatives among the Atlantic Ocean species of Nezumia, or by way of stepping-stone dispersal from the Indo-west Pacific Ocean.     

小叶、中间和柠条三种锦鸡儿的分布式样及其生态适应

确定了小叶锦鸡儿、中间锦鸡儿和柠条锦鸡儿3种植物的分布式样:小叶锦鸡儿为蒙古高原东部-F松辽平原西部-华北山地分布种,中间锦鸡儿为东戈壁-鄂尔多斯高原-黄土高原北部分布种,柠条锦鸡儿为南阿拉善-西鄂尔多斯分布种。小叶锦鸡儿适应分布于蒙古高原典型草原带和森林草原带以及华北山地落叶阔叶林带,在草原带高平原上可形成灌丛化草原的景观,在草原带的沙地上可形成以小叶锦鸡儿为建群种的沙地灌丛植被;中间锦鸡儿适应分布于蒙古高原的荒漠化草原及草原化荒漠带、鄂尔多斯高原的典型草原和荒漠化草原带的沙地及梁地上、黄土高原北部的黄土丘坡上,常形成以中间锦鸡儿为建群种的沙地灌丛植被;柠条锦鸡儿则适应分布于草原化荒漠和典型荒漠带的固定和半固定沙地上。近缘的3种锦鸡儿的地带性分布从东至西或从北向南形成明显的有规律的地理替代分布格局。     

Evolution and radiation in the scorpion Buthus elmoutaouakili Lourenco and Qi 2006 (Scorpiones: Buthidae) at the foothills of the Atlas Mountains (North Africa)

When low dispersal ability of an organism meets geographical barriers, the evolution of inter- and intraspecific differentiation is often facilitated. In the Atlas massif of North Africa, the genus Buthus splits into several species and diverges into numerous genetic lineages, often following the orographic structures of mountain systems. Such high mountain ranges often act as barriers for species with reduced mobility even on small spatial scales. To study the effect of orographic structures on organisms with low dispersal ability, we collected 61 individuals of the scorpion species Buthus elmoutaouakili at 18 locations around the southwestern foothills of the High Atlas and Antiatlas and in the Sousse valley (western Morocco). We analyzed intraspecific differentiation patterns within this geographically restricted area of about 100 × 50 km using 452 bp of the cytochrome oxidase I mitochondrial gene. We detected 5 distinct genetic lineages. In a second analysis, we added 61 previously published sequences from Buthus species from Europe and North Africa. Using a molecular clock approach, we detected old splits (4-5 Ma) separating the samples from 1) the western High Atlas and north of these mountains, 2) the Sousse valley and adjoining mountain areas, and 3) the southwestern Antiatlas. Further differentiation happened in the first 2 geographical groups about 3 Ma. Thus, the divergence time estimates based on a Bayesian approach support the onset of differentiation into these main clades along the Pliocene (5-2.3 Ma) when climatic oscillations started and a constant global cooling preceded the glacial-interglacial cycles of the Pleistocene. Further genetic splits into parapatric groups are detectable for the Sousse valley main group in the early Pleistocene. The climatic oscillations of the Pliocene and early Pleistocene might have caused repeated range shifts, expansions, and retractions leading to repeated vicariance, hereby producing the hierarchical structure of genetic differentiation in B. elmoutaouakili. A taxonomic revision, including morphological and molecular data, is needed to assess the status of each of these Buthus scorpion lineages.