Eastern Maghreb Research Articles

The Bougrine Zn-Pb deposit, the largest salt diapir-related Mississippi Valley-type deposit in the Eastern Maghreb salt diapir province, Tunisia

The Bougrine Zn-Pb sulfide deposit (5.5 Mt @ 12 % Zn + 2.5 % Pb) in the Eastern Maghreb salt diapir province of Tunisia is located at the northeastern end of the Lorbeus-Bougrine diapir. The several-km large diapir is composed of Triassic evaporite that intruded about 10 km-thick Jurassic and Cretaceous marine sedimentary rocks during the Cretaceous and Tertiary. Mineralization occurs in breccia zones at the contact with the Triassic diapir and in the peridiapiric Cretaceous strata. Most of the ore is stratabound (F2 orebody) with microsphalerite disseminations in organic-rich laminated black limestone of the Cenomanian-Turonian Bahloul Formation, where sphalerite occurs as infill of Globigerina shells, as massive replacement of Globigerina-rich layers, as banded colloform sphalerite and in voids as euhedral, coarse-grained crystals together with skeletal galena and minor marcasite. Abundant degraded oil seepages in the orebodies and their host rocks testify to the existence of a (former) hydrocarbon reservoir.The various sphalerite generations have highly variable Mn and Fe contents; Cd has an average content of 445 ppm; As and Hg contents are in the 200 ppm range; Tl content is about 40 ppm; Ga content is about 10 ppm; and In content is consistently <1 ppm. Fluid inclusions in celestite, coarse-grained late-stage sphalerite and calcite indicate moderate fluid salinities (16 ± 2 wt% NaCl eq.) and moderate temperatures (85 to 146 °C). The empirical trace-element sphalerite geothermometer gives 184 ± 38 °C for the main-stage microsphalerite and the banded colloform sphalerite which have no measurable fluid inclusions.A wide range of δ13C (-13.6 to 5.6 ‰) and δ18O (-19.8 to 33 ‰) values is observed in ore-stage and post-ore calcite. Ore-stage calcite commonly has negative δ13C values reflecting involvement of carbon from organic matter; its δ18O composition with an average of 22.6 ± 2 ‰ (19.7 to 26.0 ‰) may be explained by a moderate-temperature hydrothermal fluid. There is no systematic δ34S variation between and across the different styles of mineralization. The range of δ34S of 6–19 ‰ with a mean of 13 ‰ is close to the δ34S value of local and regional Triassic marine sulfate (δ34S = 16 ‰). Thermochemical reduction of Triassic sulfate, likely mediated by hydrocarbons, is the most probable source for sulfur in the ores. Lead isotope data on galena are within a narrow field of upper crustal signature (206Pb/204Pb = 18.71–18.77; 207Pb/204Pb = 15.66–15.70; 208Pb/206Pb = 38.81–38.97) indicating a homogeneous ore fluid.The major characteristics of the Bougrine ore deposit are typical of Mississippi Valley type Pb-Zn deposits and indicate large-scale fluid circulation likely related to stacking and uplift in the northern Tunisian Nappes Zone during the Miocene. Both the hydraulic load of meteoric water and over-pressured basinal and basement fluids during compressional movements favored southward fluid migration along basement discontinuities and most permeable strata in the Mesozoic-Cenozoic sedimentary pile. Low-pressure domains around salt diapirs in a rifting environment allowed focused upward fluid flow, and the hydrocarbon reservoir of the Bahloul Formation then provided the geochemical trap for efficient base metal deposition.

The status of minority species and young individuals of the terrestrial malacofauna in Capsian and Neolithic populations in eastern Maghreb

The status of minority species and young individuals of the terrestrial malacofauna in Capsian and Neolithic populations in eastern Maghreb

Re-examination of the Aïn Allegua-Aguiba celestite-Pb-Zn salt diapir-related Mississippi Valley-type deposits (Nappes zone, Northwestern Tunisia): New conceptual model

The celestite-rich and Pb-Zn-poor Aïn Allegua deposits are located at the northeastern extremity of the Dougas diapir in the Nappes zone of northern Tunisia. The Sr-Ba-Zn-Pb mineral system studied in this paper is enclosed in a Triassic breccia, labeled cap rock, located between the Triassic diapir and the surrounding Cretaceous to Lutetian marine sediments. It consists of evaporite dissolution float breccia where Triassic dolostone clasts are floating in a Triassic matrix of anhydrite and argillites. Five mineralizing stages are recognized, which are from early to late: celestite (stage 1); hydrothermal dolomite (stage 2); crosscutting galena-barite or sphalerite-galena-barite-calcite (stage 3); (4) fracture and karst filled by barite, celestite, and galena (stage 4); and late supergene mineralization (strontianite, smithsonite, and cerussite) (stage 5). The present study shows that the celestite and Pb-Zn mineralization formed from the circulation in the cap rock of at least three fluids: (1) fluid A (100–150 °C; 10 to 25 wt% NaCl eq.) deposited the celestite; (2) fluid B (125 to 185 °C; 29 to 34 wt% NaCl eq.) deposited the hydrothermal dolomite, and (3) fluid C (temperatures around 90 °C; 14 to 22 wt% NaCl eq.) deposited the calcite that accompanied sphalerite and galena precipitation. The δ34S values for sphalerite and galena fall within a narrow range from + 10.4‰ to + 18.9‰ (avr. 13‰) indicating a homogeneous source of sulfur. The δ34S values of celestite and barite are also relatively homogeneous from + 14.7 to + 23‰ (ave. + 16.5‰) and match those of Tunisian Triassic gypsum and Triassic marine sulfates (δ34S = +15‰). The temperature of deposition together with sulfur isotope data indicate that the reduced sulfur in sulfides was derived through thermochemical sulfate reduction of Triassic sulfate. The lead isotopes in galena show that Pb, and probably Zn and Ba were extracted from the upper crustal rocks. Part of the strontium is derived from the Triassic anhydrite and the other part is probably transported to the cap rock in formation waters. The tectonic setting, the carbonated nature of the host rocks, the epigenetic style of the mineralization, and the mineral associations, together with fluid inclusions data and sulfur and lead isotopes show that the Aïn Allegua deposit belongs to the diapir-related Mississippi Valley-type (MVT) deposits of the Eastern Maghreb salt diapirs province, that formed, as the most of the MVT deposits and districts in the world, from basinal brines with no mixing with metamorphic and/or magmatic fluid or involvement of a heat source from a concealed sill or magmatic intrusion.

Ribât in Early Islamic Ifrîqiya: Another Islam from the Edge

After a difficult conquest under the Umayyads, the eastern Maghreb or Ifrîqiya region was turned into the western borderland of the Abbasid Empire in the second half of the 8th century, and its governance was soon delegated to the Aghlabid Emirate (800–909). In this context, the Sahel (or Ifriqîyan coastline) quickly became a major centre of asceticism and pious collective retreat in places dedicated to ribât activities. This practice provided a framework for the life of devout people who kept a watchful eye on the Byzantine enemy while zealously performing their devotions. A genuine frontier society of religious men and devotees, ascetics and traditionalists arose in this burgeoning coastal fringe. Over the last two decades, this topic has given rise to a very rich historiography, notably produced by Tunisian researchers who have profoundly renewed our understanding. Based on these considerable achievements, the present contribution proposes to broaden the analysis in order to show how the rise of this movement of warrior piety, advocating an ideal of jihâd, must be related to a more global phenomenon, considered at the scale of the Abbasid Empire. Remaining in a comparative dimension, this article also proposes several approaches to the specific architecture of ribât sites, especially the place devoted to the community mosque.

Iron age genomic data from Althiburos – Tunisia renew the debate on the origins of African taurine cattle

The Maghreb is a key region for understanding the dynamics of cattle dispersal and admixture with local aurochs following their earliest domestication in the Fertile Crescent more than 10,000 years ago. Here, we present data on autosomal genomes and mitogenomes obtained for four archaeological specimens of Iron Age (∼2,800 cal BP-2,000 cal BP) domestic cattle from the Eastern Maghreb, i.e. Althiburos (El Kef, Tunisia). D-loop sequences were obtained for an additional eight cattle specimens from this site. Maternal lineages were assigned to the elusive R and ubiquitous African-T1 haplogroups found in two and ten Althiburos specimens, respectively. Our results can be explained by post-domestication hybridization of Althiburos cattle with local aurochs. However, we cannot rule out an independent domestication in North Africa considering the shared ancestry of Althiburos cattle with the pre-domestic Moroccan aurochs and present-day African taurine cattle.

Hinterland and foreland structures of the eastern Maghreb Tell and Atlas thrust belts: tectonic controlling factors, pending questions, and oil/gas exploration potential of the Pre-Triassic traps

Hinterland and foreland structures of the eastern Maghreb Tell and Atlas thrust belts: tectonic controlling factors, pending questions, and oil/gas exploration potential of the Pre-Triassic traps

Borrowings and exogenous contact in the Eastern Maghreb in the holocene: an examination of the bone industry

This article considers the bone industries of the Capsian and the Neolithic of Capsian Tradition. It place these industries in the broader context of the Near-East and Europe in order to identify their unique characteristics and the elements they share with other industries. The criteria considered are at once stylistic, morphological, and technical. Both the Capsian and the Neolithic of Capsian Tradition (NCT), present commonalities with the industries of the Natufian and the PPNB. Is this the result of exogenous borrowings, internal dynamics, or documentation biases? The question of possible contact between cultures and populations is examined here.

Morphological variability in Lophyra flexuosa (Fabricius, 1787) (Coleoptera, Cicindelidae) in desert countries is affected by sexual dimorphism and geographic aspect.

Lophyra flexuosa, a eurytopic tiger beetle characterized by long phenological activity, wide geographic and altitudinal distribution, and occurring in the highest number of habitats among all Cicindelidae known from North Africa, was chosen to study its geographic variation in morphology and sexual dimorphism. Here, we found significant sexual dimorphism exhibited in larger body size of females and longer mandibles in males, which can be explained by different roles of particular sexes in courtship. Moreover, we recorded significant differences in body sizes between western and eastern Maghreb populations which could suggest genetic isolation between these populations. As the species is related to habitats placed close to the water reservoirs, which in the desert countries are under significant human pressure (including climate change), we expect a reduction of habitat occupied by this taxon. Therefore, the geographic morphological variability that we observe today in the tiger beetle Lophyra flexuosa in the future could lead to speciation.

Biogeographic and demographic history of the Mediterranean snakes Malpolon monspessulanus and Hemorrhois hippocrepis across the Strait of Gibraltar

BackgroundThe contribution of North Africa to the assembly of biodiversity within the Western Palaearctic is still poorly documented. Since the Miocene, multiple biotic exchanges occurred across the Strait of Gibraltar, underlying the high biogeographic affinity between the western European and African sides of the Mediterranean basin. We investigated the biogeographic and demographic dynamics of two large Mediterranean-adapted snakes across the Strait and assess their relevance to the origin and diversity patterns of current European and North African populations.ResultsWe inferred phylogeographic patterns and demographic history of M. monspessulanus and H. hippocrepis, based on range-wide multilocus data, combined with fossil data and species distribution modelling, under present and past bioclimatic envelopes. For both species we identified endemic lineages in the High Atlas Mountains (Morocco) and in eastern Iberia, suggesting their persistence in Europe during the Pleistocene. One lineage is shared between North Africa and southern Iberia and likely spread from the former to the latter during the sea-level low stand of the last glacial stage. During this period M. monspessulanus shows a sudden demographic expansion, associated with increased habitat suitability in North Africa. Lower habitat suitability is predicted for both species during interglacial stages, with suitable areas restricted to coastal and mountain ranges of Iberia and Morocco. Compiled fossil data for M. monspessulanus show a continuous fossil record in Iberia at least since the Pliocene and throughout the Pleistocene.ConclusionsThe previously proposed hypothesis of Pleistocene glacial extinction of both species in Europe is not supported based on genetic data, bioclimatic envelopes models, and the available fossil record. A model of range retraction to mountain refugia during arid periods and of glacial expansion (demographic and spatial) associated to an increase of Mediterranean habitats during glacial epochs emerges as a general pattern for mesic vertebrates in North Africa. Moreover, the phylogeographic pattern of H. hippocrepis conforms to a well-established biogeographic partition between western and eastern Maghreb.

The Birth of a Successful Prayer Book

Abstract The vast project to reconstruct a history and geography of the spread of the Dalāʾil al-Khayrāt necessarily involves looking into the beginnings of the prayerbook’s manuscript transmission. Composed in Morocco before 869/1465, the prayerbook was already known in the Eastern Maghreb from the mid-11th/17th century. It then reached Turkey and the rest of the Mashriq. After that it found its way to Central, South and Southeast Asia. Returning to the core of the book’s diffusion, this article questions the existence of an autograph copy of Dalāʾil al-Khayrāt. How was the manuscript tradition of one of the most copied religious books in pre-modern times established? This article also poses essential questions about the work of the actors (copyists, illuminators) responsible for the diffusion of the book in its early days.

Changing Plant-based Subsistence Practices among Early and Middle Holocene Communities in Eastern Maghreb

ABSTRACT The eastern Maghreb is a key area for understanding environmental and cultural dynamics during the early and middle Holocene. Capsian populations from around 10000–7500 cal BP were among the last foragers in the region. Capsian sites are known as escargotières (land shell middens), and locally called rammadiyat (meaning ashy mound). As taphonomic conditions in Capsian open-air sites generally favour the preservation of resistant materials such as shells and bones rather than fragile plant remains, this study integrates macro-botanical and microfossil evidence from phytoliths, calcitic wood ash pseudomorphs and dung spherulites, since each is influenced by different formation and post-depositional processes. Archaeobotanical results from recent excavations in four Tunisian sites located across the lowland steppe, the Dorsale Mountains and the eastern coast display a wide range of wild plant resources, such as pine, oak, wild legumes, and grasses, in particular Alfa grass. These findings suggest that these plants could have played an important role in Capsian diet, whereas the presence of Alfa grass suggests the use of fibre sources for basketry, matting, building and fuel. Integrated macro-botanical and microfossil records contribute to a better understanding of changing subsistence practices on the threshold of early food-producing systems in north-west Africa.

Common Cultural Markers in the Bone and Lithic Production of the Upper Capsian: A Comparative Approach

This article discusses two major types of the Upper Capsian cultural materials in the Eastern Maghreb (ca. 8.2–6.0 cal BP): worked bones and knapped stones. These materials represent the main “toolset” of Capsian groups. The comparative observation of the two systems of production—bone and lithic—has allowed the recognition of certain analogies in their methods of exploitation and fabrication. These similarities, which represent a significant cultural marker of the Upper Capsian period, have not previously been explored. In this paper, two particular aspects of the system of production will be considered: multi-structured matrix partition in the bone industry and pressure knapping in the lithic industry.

Lophyra flexuosa (Fabricius, 1787) (Coleoptera: Cicindelidae) in desert countries: morphological variability in geographic aspect as potential beginning of speciation?

Lophyra flexuosa is one of only several eurytopic tiger beetles species known from Palearctic realm. Its geographical distribution shows several populations that are spread from the Iberian Peninsula and Morocco, across some regions of south-western Europe and northern Africa to Israel and Syria. The species is characterized by long phenological activity, wide altitudinal distribution, and occurs in the highest number of habitats among all Cicindelidae known from Maghreb region. In the present study the geographical variation in morphology and sexual dimorphism in north African populations of L. flexuosa was studied. In total 52 samples with over 700 specimens were collected including 20 samples in Morocco and 32 in Tunisia. To test the variation in morphometric traits measurements of eight body parameters were taken from all males (383) and females (352) including right mandible length, length of head, width of head, length of pronotum, maximum pronotum width, length of elytra, maximum elytra width, and total body length. We discovered significant sexual dimorphism expressed by larger body size of females and longer mandibles in males, what can be explained by different roles of particular sexes in courtship. Moreover, we recorded significant differences in body sizes between western and eastern Maghreb populations which can suggest genetic isolation between these populations. As the species is related to habitats placed close to water reservoirs, which in the desert countries are under significant human pressure (including climate change), we expect an reduction of habitats occupied by this taxon. Therefore, the geographic morphological variability that we observe today in the tiger beetle Lophyra flexuosa may lead to speciation and creation of separate species in the future.

A review of the crustal architecture and related pre-salt oil/gas objectives of the eastern Maghreb Atlas and Tell: Need for deep seismic reflection profiling

The Atlas and Tell domains constitute a major segment of the Circum-Mediterranean Alpine fold and thrust belts in the Maghreban part of North Africa. South of the Tell, the Atlas results from a progressive Late Cretaceous to present foreland inversion of former Jurassic to Cretaceous extensional tectonic features inherited from the opening of the Tethys along the northern African margin. In contrast, the Tell is made up of allochthonous Cretaceous to Neogene basinal units detached along a tectonic sole made up of remobilized Triassic salt and tectonic mélanges.Since the last decade, the major tectonic steps accounting for the development of the eastern Maghreb structures are becoming more precisely constrained by seismic data and associated structural analyses. However their overall configuration at depth, as well as the deep architecture of the underlying basement remain poorly understood because only few wells yet penetrated deeper than the Triassic series in the Atlas and adjacent foreland basin. Therefore, this paper is intended to give an overview on the deep structural features affecting the Pan-African basement in the eastern Maghreb by means of tentative regional cross-sections. Our purpose is to stimulate exploration of deep hydrocarbon plays and give some insights to help both the industry, universities and other public research institutes to undertake jointly a major research program on the deep crustal configuration in the eastern Maghreb with particular focus on the pre-Triassic series beneath the Tellian allochthon in the north, the parautochthonous inverted Atlasic structures in the center, and in the adjacent autochthonous foreland domain extending southeastward in the offshore.

Diversification and speciation in tree frogs from the Maghreb (Hyla meridionalis sensu lato), with description of a new African endemic

Comparative molecular studies emphasized a new biogeographic paradigm for the terrestrial fauna of North Africa, one of the last uncharted ecoregions of the Western Palearctic: two independent east-west divisions across the Maghreb. Through a comprehensive phylogeography, we assessed how this model suits the genetic diversification documented for the tree frog Hyla meridionalis sensu lato. Analyses of mtDNA variation and thousands of nuclear loci confirmed the old split (low-Pliocene) between Tunisian and Algerian populations. These lineages meet but barely admix in the eastern Maghreb (Algerian-Tunisian border), a sign of putatively advanced reproductive isolation. In the western Maghreb, we report a Pleistocene divergence between Moroccan and Algerian populations. Tree frogs thus follow both predictions: a double east-west break that gave rise to two suture zones characteristic of North-African phylogeography. Moreover, some intraspecific mtDNA variation is not mirrored by the nuclear data, emphasizing that evolutionary units should always be designated by multilocus approaches. Last but not least, we describe the Tunisian lineage as a new species endemic to Africa.

Phylogeography of the Ibero-Maghrebian red-eyed grass snake (Natrix astreptophora)

We examined phylogeographic differentiation of the red-eyed grass snake (Natrix astreptophora) using 1984 bp of mtDNA and 13 microsatellite loci from specimens collected across its distribution range in southwestern Europe and northwestern Africa. Based on phylogenetic analyses of mtDNA, European N. astreptophora constituted the sister clade to a weakly supported North African clade comprised of two deeply divergent and well-supported clades, one corresponding to Moroccan snakes and the other to snakes from Algeria and Tunisia. This tripartite differentiation was confirmed by analyses of microsatellite loci. According to a fossil-calibrated molecular clock, European and North African N. astreptophora diverged 5.44 million years ago (mya), and the two Maghrebian clades split 4.64 mya. These dates suggest that the radiation of the three clades was initiated by the environmental changes related to the Messinian Salinity Crisis and the reflooding of the Mediterranean Basin. The differentiation of N. astreptophora, with distinct clades in the Iberian Peninsula and in the western and eastern Maghreb, corresponds to a general phylogeographic paradigm and resembles the differentiation found in another co-distributed Natrix species, the viperine snake (N. maura). Despite both species being good swimmers, the Strait of Gibraltar constitutes a significant biogeographic barrier for them. The discovery that North Africa harbours two endemic lineages of N. astreptophora necessitates more conservation efforts for these imperilled snakes.

Underground cryptic speciation within the Maghreb: Multilocus phylogeography sheds light on the diversification of the checkerboard worm lizard Trogonophis wiegmanni

Biogeographic and evolutionary patterns in the North African portion of the Western Palaearctic are poorly known. A high fraction of undescribed diversity is expected in this region, especially in groups such as reptiles. Here we used mitochondrial (12S, 16S, cytb) and nuclear (pomc, rag2, cmos) markers and morphological data to investigate phyletic diversification and phylogeographical structure in the amphisbaenian Trogonophis wiegmanni endemic to the Maghreb. Phylogenetic and molecular dating analyses based on gene trees and species trees support three deeply divergent lineages of Pliocene origin, two in Morocco and one in central Algeria and Tunisia. Parapatry, reciprocal monophyly, high genetic divergence and limited morphological differentiation between them suggest that these lineages represent independent cryptic taxonomic units. Emerging lines of evidence from this study and from available literature on Maghreb taxa support (i) a major biogeographic break between western and eastern Maghreb and (ii) a role of the Atlas as a biogeographic divide within the western Maghreb (Morocco). The origin of these biogeographic units is probably associated with the evolutionary events prompted by the Late Miocene palaeogeographic setting and later by Plio-Pleistocene climatic changes and their interplay with prominent orographic barriers within North Africa.

Differentiation in the marbled white butterfly species complex driven by multiple evolutionary forces

AbstractAimGenetic and phenotypic data may show convergent or contrasting spatial patterns. Discrepancies between markers may develop in response to different evolutionary forces. In this study we analyse inter‐ and intraspecific differentiation of closely related taxa in the marbled white butterfly species group. Based on genetic and phenotypic characters we test for potential evolutionary drivers and propose a taxonomic revision.LocationWestern Palaearctic (including north‐western Africa).MethodsWe compared distributions of mitochondrial cytochrome c oxidase subunit I gene (COI) sequences, of several allozyme loci, and of the shape of wings and genitalia obtained by applying landmark‐based techniques for the three butterfly species Melanargia galathea (central and eastern Europe), M. lachesis (Iberia) and M. lucasi (North Africa).ResultsAll studied markers showed a strong spatial structure, although discordance among their patterns was detected. COI sequences, wing shape and genitalia indicated a main split between M. galathea and M. lucasi. A lower differentiation between M. galathea and M. lachesis was found in wing shape and reflected in two mutations of the COI gene, while allozymes indicated a strong divergence. Within M. galathea, allozyme data and COI, but not morphology, revealed the existence of a slightly differentiated lineage in the Italian Peninsula, France and Switzerland. Based on COI, Melanargia lucasi was split into two subgroups, a western and an eastern Maghreb lineage.Main conclusionsLong‐term isolation of Melanargia populations between North Africa and Europe led to divergence between M. galathea and M. lucasi. This was followed by a recent differentiation among populations isolated during the cold periods of the Pleistocene, such as M. lachesis in Iberia. These lineages are characterized by a tendency not to overlap in secondary sympatry. The different patterns of the four markers may arise from divergent evolutionary processes and pressures: wings may be mainly affected by natural selection, genital structures by sexual selection, whereas long‐term isolation and drift may have driven divergence of mitochondrial DNA and allozymes.

The Late Eocene and Late Miocene fronts of the Atlas Belt in eastern Maghreb: integration in the geodynamic evolution of the Mediterranean Domain

Regional cross sections at the scale of the eastern Maghreb based on subsurface and field data allow presenting the structural styles related to the Middle-Late Eocene compressional events. The structural cross sections depict that the Late Eocene front of the Atlas Belt extends far through the Northern Africa plate margin comparatively to the Late Miocene front cropping out in the Eastern Tunisian Atlas. The sections allow proposing a new subsurface front for the Atlas belt encompassing a large part of the Pelagian-Sirt basins. The consequences of this particular configuration are discussed at the scale of the south Tethys margin and replaced in the frame of the geodynamic evolution of the Mediterranean Domain.

Genetic differentiation of the pine processionary moth at the southern edge of its range: contrasting patterns between mitochondrial and nuclear markers.

The pine processionary moth (Thaumetopoea pityocampa) is an important pest of coniferous forests at the southern edge of its range in Maghreb. Based on mitochondrial markers, a strong genetic differentiation was previously found in this species between western (pityocampa clade) and eastern Maghreb populations (ENA clade), with the contact zone between the clades located in Algeria. We focused on the moth range in Algeria, using both mitochondrial (a 648 bp fragment of the tRNA‐cox2) and nuclear (11 microsatellite loci) markers. A further analysis using a shorter mtDNA fragment and the same microsatellite loci was carried out on a transect in the contact zone between the mitochondrial clades. Mitochondrial diversity showed a strong geographical structure and a well‐defined contact zone between the two clades. In particular, in the pityocampa clade, two inner subclades were found whereas ENA did not show any further structure. Microsatellite analysis outlined a different pattern of differentiation, with two main groups not overlapping with the mitochondrial clades. The inconsistency between mitochondrial and nuclear markers is probably explained by sex‐biased dispersal and recent afforestation efforts that have bridged isolated populations.