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1.
Chloroplast DNA variation was studied in three evergreen Quercus species (Q. suber L., Q. ilex L. and Q. coccifera L.) from the Western Mediterranean Basin using PCR-RFLP. We studied five primer pair/enzyme combinations, four of them previously used in other European Quercus, obtaining a large number of haplotypes (81) grouped in three main types (suber type, ilex-coccifera I type and ilex-coccifera II type). Such level of haplotype diversity is higher than previously reported for the genus. Remarkable differences in haplotype richness between species have been found. Q. ilex and Q. coccifera usually share the same haplotypes, while a number of Q. suber populations possesses variants of the ilex-coccifera I type. This fact is interpreted as a result of genetic introgression between Q. suber and Q. ilex. Reproductive factors that could determine this exchange are discussed, as well as the influence of different species histories on the present structure of evergreen Quercus in the Western Mediterranean Basin.  相似文献   

2.
The leaf trichomes of the evergreen and semi-deciduous species of the genus Quercus living in the Iberian Peninsula were studied by means of SEM and LM. The SEM has been particularly useful because of the high quality of the illustrations and the possibility of recognizing the structures under I,M or the stereomicroscope afterwards. The studied species were: Quercus coccifera, Q ilex , Q rotundifolia, Q suber, Q canariensis, Q faginea, Q brateroi and Q lusitanica . All species show some exclusive particularities that permit their identification using these characters. The abaxial surface of the leaf always has more trichomes than the adaxial and hence provides more characters for identification. Quercus alpestris Boiss. shows the same features in the leaf trichomes as Q faginea Lam. A key for identification using these characters is provided.  相似文献   

3.
The leaf trichomes of the evergreen and semi-deciduous species of the genus Quercus living in the Iberian Peninsula were studied by means of SEM and LM. The SEM has been particularly useful because of the high quality of the illustrations and the possibility of recognizing the structures under LM or the stereomicroscope afterwards. The studied species were-. Quercus coccifera, Q. ilex, Q. rotundifolia, Q suber, Q canariensis, Q. faginea, Q. broteroi and Q. lusitanica. All species show some exclusive particularities that permit their identification using these characters. The abaxial surface of the leaf always has more trichomes than the adaxial and hence provides more characters for identification. Quercus alpestris Boiss. shows the same features in the leaf trichomes as Q. faginea Lam. A key for identification using these characters is provided.  相似文献   

4.
The southwestern Iberian Peninsula is an important biogeographic region, showing high biodiversity levels and hosting several putative glacial refugia for European flora. Here, we study the genetic diversity and structure of the Mediterranean, thermophilous plant Cheirolophus sempervirens (Asteraceae) across its whole distribution range in SW Iberia, as a tool to disentangle some of the general biogeographic patterns shaping this southern refugia hotspot. Null genetic diversity was observed in the cpDNA sequencing screening. Nonetheless, AFLP data revealed high levels of among-population genetic differentiation correlated to their geographic location. Our results suggest longer species persistence in southern Iberian refugia during glacial periods and subsequent founder effects northwards due to colonizations in warmer stages (i.e., the southern richness to northern purity pattern). Additionally, our phylogeographic analyses indicate the presence of two separate genetic lineages within Ch. sempervirens, supporting the hypothesis of multiple minor refugia for SW Iberia in agreement with the refugia within refugia model.  相似文献   

5.
Abstract.  This study presents pollen-analytical data from continental and offshore Iberian Peninsula sites that include pollen curves of Quercus suber , to provide information on the past distribution and ecology of the cork oak ( Q. suber ). Results centre on a new pollen record of Navarrés (Valencia, eastern Spain), which shows that the cork oak survived regionally during the Upper Pleistocene and was important during a mid-Holocene replacement of a local pine forest by Quercus -dominated communities. This phenomenon appears linked to the recurrence of fire and reinforces the value of the cork oak for reforestation programmes in fire-prone areas. In addition to Navarrés, other Late Quaternary pollen sequences (Sobrestany, Casablanca-Almenara, Padul, SU 8103, SU8113, 8057B) suggest last glacial survival of the cork oak in southern and coastal areas of the Peninsula and North Africa. Important developments also occur from the Late Glacial to the middle Holocene, not only in the west but also in the eastern Peninsula. It is suggested that, in the absence of human influence, Q. suber would develop in non-monospecific forests, sharing the arboreal stratum both with other sclerophyllous and deciduous Quercus and Pinus species.  相似文献   

6.
Anatomical changes at the cotyledonary node from the embryo to the seedling stage in Quercus coccifera, Q. ilex, and Q. humilis were investigated by light and scanning electron microscopy techniques. Mature embryos of Q. humilis possess 2-3 pairs of leaf primordia and a pair of cotyledonary buds, whereas in Q. coccifera and Q. ilex there are two incipient primordia, and cotyledonary buds are not observed until 1 wk after germination. In all three species the cotyledonary buds multiply, forming bud clusters, and a vascular connection is well established within 5-6 wk after germination. As development proceeds, the cotyledonary region becomes woody, but buds, which are exogenous in origin, never become embedded in the periderm. In comparison with Q. suber, another native Mediterranean Basin oak, the cotyledonary node is short and axillary buds are not present below the insertion of cotyledons. In addition, starch accumulation in the cotyledonary region is not observed from histological analysis in the three oaks. Therefore, in Q. coccifera, Q. ilex, and Q. humilis seedlings the cotyledonary node can be considered to be an important regenerative structure enabling them to resprout after the elimination of the shoot above the cotyledons, despite the absence of a lignotuberous structure.  相似文献   

7.
Gene flow is particularly frequent in the genus Quercus (oaks), especially between closely related species. We focus here on Quercus ilex and the cork-producing Quercus suber , which occasionally hybridize although they are phylogenetically markedly separated. Morphological observations were combined with both allozymic and chloroplastic diagnostic markers to characterize hybridization and introgression and to infer their dynamics in two French regions (French Catalonia and Provence), which are separated by several hundred kilometres. Some hybrids were found in both regions, indicating recent hybridization events. As expected from previous studies, most hybrids resulted from ♀ Q. ilex  × ♂ Q. suber crosses, but our data showed that the reciprocal cross is also possible. Partial independence between nuclear and chloroplastic introgression was observed in the two species. Nuclear introgression was limited in both species and both regions, with no preferred direction. In Provence, chloroplastic introgression was very rare in both species. Conversely, all Q. suber individuals from French Catalonia were introgressed by Q. ilex chlorotypes. This might be explained by introgression in the Iberian Peninsula antedating the first occurrence of the two species in French Catalonia. We also observed a new chlorotype that was created locally, and was exchanged between the two species. However, the two species still remain genetically differentiated. The dynamics and complexity of exchanges and the factors determining them (including human management of Q. suber ) are discussed.  相似文献   

8.
The dynamics of vegetation in the North Iberian Peninsula have been historically established from peaty and lacustrine records obtained in strongly continental mountainous areas. Pollen records located in areas with a more oceanic climate, have allowed a more precise evaluation of the changes and differences in the vegetation development of the various biogeographical areas in the North Iberian Peninsula, during the Late-glacial and Holocene. It was found that: (1) The vegetation of the Cantabrian–Atlantic province in northern Iberian Peninsula responded to climatic changes during the last 16000 years. (2) Tree vegetation declined during the Oldest Dryas, with steppe vegetation in the interior and cryophilous vegetation along the coast. (3) During the time of the Lateglacial Interestadial, lowlands, arboreal vegetation was dominant in the landscape (Pinus, Betula, Quercus, Corylus). (4) During the Younger Dryas the differences between coast and interior are less obvious. (5) During the Holocene oceanic decidious forests of Quercus robur, Corylus avellana, Tilia sp. and Fagus sylvatica were present along the coast. At low altitudes, in the interior, forests of Quercus robur / pyrenaica, enclaves of pine woods and Mediterranean types of forest (Quercus ilex, Olea europea) were present. At higher altitudes Pinus (P. sylvestris and P. uncinata) or mixed forests of Pinus and Betula were dominant.  相似文献   

9.
To understand the demographic history of Arabidopsis thaliana within its native geographical range, we have studied its genetic structure in the Iberian Peninsula region. We have analyzed the amount and spatial distribution of A. thaliana genetic variation by genotyping 268 individuals sampled in 100 natural populations from the Iberian Peninsula. Analyses of 175 individuals from 7 of these populations, with 20 chloroplast and nuclear microsatellite loci and 109 common single nucleotide polymorphisms, show significant population differentiation and isolation by distance. In addition, analyses of one genotype from 100 populations detected significant isolation by distance over the entire Iberian Peninsula, as well as among six Iberian subregions. Analyses of these 100 genotypes with different model-based clustering algorithms inferred four genetic clusters, which show a clear-cut geographical differentiation pattern. On the other hand, clustering analysis of a worldwide sample showed a west–east Eurasian longitudinal spatial gradient of the commonest Iberian genetic cluster. These results indicate that A. thaliana genetic variation displays significant regional structure and consistently support the hypothesis that Iberia has been a glacial refugium for A. thaliana. Furthermore, the Iberian geographical structure indicates a complex regional population dynamics, suggesting that this region contained multiple Pleistocene refugia with a different contribution to the postglacial colonization of Europe.  相似文献   

10.
Pleistocene glaciations often resulted in differentiation of taxa in southern European peninsulas, producing the high levels of endemism characteristic of these regions (e.g. the Iberian Peninsula). Despite their small ranges, endemic species often exhibit high levels of intraspecific differentiation as a result of a complex evolutionary history dominated by successive cycles of fragmentation, expansion and subsequent admixture of populations. Most evidence so far has come from the study of species with an Atlantic distribution in northwestern Iberia, and taxa restricted to Mediterranean‐type habitats remain poorly studied. The Iberian Midwife toad (Alytes cisternasii) is a morphologically conserved species endemic to southwestern and central Iberia and a typical inhabitant of Mediterranean habitats. Applying highly variable genetic markers from both mitochondrial and nuclear genomes to samples collected across the species’ range, we found evidence of high population subdivision within A. cisternasii. Mitochondrial haplotypes and microsatellites show geographically concordant patterns of genetic diversity, suggesting population fragmentation into several refugia during Pleistocene glaciations followed by subsequent events of geographical and demographic expansions with secondary contact. In addition, the absence of variation at the nuclear β‐fibint7 and Ppp3caint4 gene fragments suggests that populations of A. cisternasii have been recurrently affected by episodes of extinction and recolonization, and that documented patterns of population subdivision are the outcome of recent and multiple refugia. We discuss the evolutionary history of the species with particular interest in the increasing relevance of Mediterranean refugia for the survival of genetically differentiated populations during the Pleistocene glaciations as revealed by studies in co‐distributed taxa.  相似文献   

11.
The contemporary distribution and genetic composition of biodiversity bear a signature of species’ evolutionary histories and the effects of past climatic oscillations. For many European species, the Mediterranean peninsulas of Iberia, Italy and the Balkans acted as glacial refugia and the source of range recolonization, and as a result, they contain disproportionately high levels of diversity. As these areas are particularly threatened by future climate change, it is important to understand how past climatic changes affected their biodiversity. We use an integrated approach, combining markers with different evolutionary rates and combining phylogenetic analysis with approximate Bayesian computation and species distribution modelling across temporal scales. We relate phylogeographic processes to patterns of genetic variation in Myotis escalerai, a bat species endemic to the Iberian Peninsula. We found a distinct population structure at the mitochondrial level with a strong geographic signature, indicating lineage divergence into separate glacial refugia within the Iberian refugium. However, microsatellite markers suggest higher levels of gene flow resulting in more limited structure at recent time frames. The evolutionary history of M. escalerai was shaped by the effects of climatic oscillations and changes in forest cover and composition, while its future is threatened by climatically induced range contractions and the role of ecological barriers due to competition interactions in restricting its distribution. This study warns that Mediterranean peninsulas, which provided refuge for European biodiversity during past glaciation events, may become a trap for limited dispersal and ecologically limited endemic species under future climate change, resulting in loss of entire lineages.  相似文献   

12.
Chloroplast (cp) and mitochondrial (mt) DNA variation were studied in 97 populations of cork oak (Quercus suber) in Morocco; in 31 of these populations, holm oak (Quercus ilex), a clearly distinct species, also occurred and was compared with Q. suber. Three cpDNA and one mtDNA primer pairs were used in the survey, each in combination with one restriction enzyme. Six haplotypes belonging to two very divergent lineages were detected; one lineage predominates in each species, and is probably ancestral, as inferred from comparisons with other oak species. In the mixed-species populations, cytoplasmic genomes were frequently shared across species, as indicated by an introgression ratio of 0.63. This index is a new measure of the propensity of species to share locally genetic markers, varying from zero (complete differentiation) to one (no differentiation). By contrast, more closely related deciduous oak species (Q. robur, Q. petraea and Q. pubescens) have introgression ratios varying from 0.82 to 0.97. The introgression events appear to have been more frequent in the direction Q. ilex (female) x Q. suber (male), a finding which seems attributable to the flowering phenology of these two species. This asymmetry may have favoured immigration of Q. suber beyond its main range, in regions already colonized by Q. ilex. There, rare hybridization and further introgression through long distance pollen flow have established populations that are morphologically indistinguishable from Q. suber but that have cytoplasmic genomes originating from the local Q. ilex populations.  相似文献   

13.
The role of glacial refugia in shaping contemporary species distribution is a long-standing question in phylogeography and evolutionary ecology. Recent studies are questioning previous paradigms on glacial refugia and postglacial recolonization pathways in Europe, and more flexible phylogeographic scenarios have been proposed. We used the widespread common vole Microtus arvalis as a model to investigate the origin, locations of glacial refugia, and dispersal pathways, in the group of “Continental” species in Europe. We used a Bayesian spatiotemporal diffusion analysis (relaxed random walk model) of cytochrome b sequences across the species range, including newly collected individuals from 10 Iberian localities and published sequences from 68 localities across 22 European countries. Our data suggest that the species originated in Central Europe, and we revealed the location of multiple refugia (in both southern peninsulas and continental regions) for this continental model species. Our results confirm the monophyly of Iberian voles and the pre-LGM divergence between Iberian and European voles. We found evidence of restricted postglacial dispersal from refugia in Mediterranean peninsulas. We inferred a complex evolutionary and demographic history of M. arvalis in Europe over the last 50,000 years that does not adequately fit previous glacial refugial scenarios. The phylogeography of M. arvalis provides a paradigm of ice-age survival of a temperate continental species in western and eastern Mediterranean peninsulas (sources of endemism) and multiple continental regions (sources of postglacial spread). Our findings also provide support for a major role of large European river systems in shaping geographic boundaries of M. arvalis in Europe.  相似文献   

14.
Lumaret R  Mir C  Michaud H  Raynal V 《Molecular ecology》2002,11(11):2327-2336
Variation in the lengths of restriction fragments (RFLPs) of the whole chloroplast DNA molecule was studied in 174 populations of Quercus ilex L. sampled over the entire distribution of this evergreen and mainly Mediterranean oak species. By using five endonucleases, 323 distinct fragments were obtained. From the 29 and 17 cpDNA changes identified as site and length mutations, respectively, 25 distinct chlorotypes were distinguished, mapped and treated cladistically with a parsimony analysis, using as an outgroup Q. alnifolia Poech, a closely related evergreen oak species endemic to Cyprus where Q. ilex does not grow. The predominant role of Q. ilex as maternal parent in hybridization with other species was reflected by the occurrence of a single very specific lineage of related chlorotypes, the most ancestral and recent ones being located in the southeastern and in the northwestern parts of the species’ geographical distribution, respectively. The lineage was constituted of two clusters of chlorotypes observed in the ‘ilex’ morphotyped populations of the Balkan and Italian Peninsulas (including the contiguous French Riviera), respectively. A third cluster was divided into two subclusters identified in the ‘rotundifolia’ morphotyped populations of North Africa, and of Iberia and the adjacent French regions, respectively. Postglacial colonization probably started from three distinct southerly refugia located in each of the three European peninsulas, and a contact area between the Italian and the Iberian migration routes was identified in the Rhône valley (France). Chlorotypes identical or related to those of the Iberian cluster were identified in the populations from Catalonia and the French Languedoc region, which showed intermediate morphotypes, and in the French Atlantic populations which possessed the ‘ilex’ morphotype, suggesting the occurrence of adaptive morphological changes in the northern part of the species’ distribution.  相似文献   

15.
The forests in the Iberian Peninsula have been strongly influenced by past climatic changes, but reconstructing their historical distributions and dynamics is very difficult due to the complex climatic characteristics and relief of the region. Research disciplines such as phylogeography and species distribution modelling can describe the past range dynamics of individual tree species in relatively great detail and help elucidate how these species have reacted to climatic changes. Here we review phylogeographical and modelling studies from species representative of the major Iberian forest types and attempt to extract general trends from the diversity of individual species histories in the Peninsula. To date most studies focus on geographical range dynamics during the Pleistocene, but an increasing body of evidence shows that some species have also retained the genetic imprints of much more ancient processes. Many widespread Iberian species show a deep and often remarkably clear-cut divide between populations from the Mediterranean and from the Atlantic regions of the Iberian Peninsula, suggesting that both areas have independently sustained viable populations over extended periods. In fact, phylogeographical studies commonly find that species had several glacial refugia across the Iberian Peninsula. On the other hand, distribution models help identifying further suitable areas that could have sustained so far undetected refugia. Such studies are of interest for species conservation, because refugium populations are high-priority targets due to their long-term persistence and unique evolutionary trajectory. Overall, we conclude that palaeoecology, phylogeography and species distribution modelling have a great potential to inform each other because of their complementary perspectives and results. A true integration of these approaches is therefore fundamental for further progress in our understanding of past Iberian environments and the organisms they harboured.  相似文献   

16.
Aim To analyse the role of the Balearic Islands as a refuge area for evergreen Quercus (cork oak: Quercus suber L., holm oak: Q. ilex L., kermes oak: Q. coccifera L.), by using molecular, historical and palaeobotanical data. Location The Western Mediterranean Basin (Balearic Islands, eastern Iberia, Provence, Sardinia, Corsica, Sicily, Malta, Italy, Northern Africa). Methods We sampled 108 populations and used the PCR‐RFLP technique with five universal cpDNA primers to define haplotypes in the sampled populations. Diversity, differentiation parameters and spatial analysis of the populations, using a spatial version of amova , were linked to the geological history of the Western Mediterranean Basin in order to explain the present spatial pattern of the evergreen Quercus populations in the Balearics. Results Evergreen Quercus cpDNA shows a complex structure, with remnants of ancient diversity in the Balearics. Balearic populations of holm oak are related to Iberian populations, while for cork and kermes oaks, we found both Tyrrhenian and Iberian haplotypes. Main conclusions The complex spatial patterns of cpDNA in Balearic evergreen Quercus appears explicable in terms of a combination of physical (vicariance and long distance dispersal) and biological (introgressive hybridization) factors. The Balearics constitute a glacial refuge area and a reservoir of genetic variation with traces of ancient diversity from Messinian–Pliocene stages.  相似文献   

17.
Hybridisation between species of the genus Quercus is a common phenomenon as a result of weak reproductive isolation mechanisms between phylogenetically close species that frequently co-occur in mixed stands. In this study, we use microsatellite markers to analyse introgression between kermes ( Quercus coccifera L.) and holm ( Q. ilex L.) oak, two closely related taxa that frequently dominate the landscape in extensive areas in the Mediterranean region. All tested microsatellites amplified and were polymorphic in both kermes and holm oaks. Bayesian admixture analyses showed a good correspondence between each species and one of the two inferred genetic clusters. Five sampled individuals were a priori tentatively identified as hybrids on the basis of intermediate morphological characteristics, and it was confirmed that they also presented mixed genotypes. However, we also detected different levels of genetic introgression among morphologically pure individuals, suggesting that successful backcrossing and/or reduced phenotypic expression of genetic variance in certain individuals may have resulted in strong convergence towards a single species phenotype.  相似文献   

18.
The Moroccan distribution of Testudo graeca graeca L. is defined by locality records (109) and related to data from nearby or bioclimatically equivalent meteorological stations (86) to produce a rain-temperature climagram. Tortoises attain 1900 m in the High Atlas range. Tortoises occur where mean annual rainfall (P) is up to 1062 mm, and even 1112 mm where mean minimum monthly temperature for the coolest month (m) is below 0oC and they are usually absent unless associated (90.0%) with Quercus ilex L. and Quercus suber L. woodland. Tortoises are absent from the Arctic-Alpine and Arid or Saharan bioclimates and where, without riverine habitat, Emberger's Quotient (Q) is below 19.5 (P = 139 mm) in the Steppe bioclimate. Below Q_= 39.1, they occur (88.9%) where there is Argania spinosa (L.) Maire woodland in the south-west and other broad-leaved and mixed wood- and scrubland species elsewhere. The Moroccan climagram includes and is extended by further stations (36) in the North African and southern European ranges generally ( T. g. graeca is reported in NW Egypt). The Moroccan range potentially occupies 167 000 m2. Of a total of 45 stations below 550 m in NW Europe, the Western Mediterranean climagram only overlaps 12 in more oceanic Britain and Ireland; 11 are coastal or insular and 11 where Quercus woodland extends the natural range with m below 0oC in montane areas. Tortoises do not occur where mean annual sunshine much exceeds 3250 h in northern Africa or below c. 2400 h in NW Europe. Tortoises introduced from the Mediterranean region by the pet trade are unable to reproduce if they survive in NW Europe for insolation is insufficient for egg incubation and hatchling survival.  相似文献   

19.
This study investigated the phylogeographic structure of Cistus ladanifer, in order to locate its Quaternary refugia, reconstruct its recolonisation patterns and assess the role of geographical features (mountain ranges, rivers and the Strait of Gibraltar) as barriers to its seed flow and expansion through the Western Mediterranean. Thirty-eight populations were screened for length variation of polymorphic chloroplast simple sequence repeats (cpSSRs). Statistical analyses included estimation of haplotypic diversity, hierarchical analysis of molecular variation (amova) and fixation indices. Mantel tests, SAMOVA and BARRIER analyses were applied to evaluate the geographical partitioning of genetic diversity across the entire species range. Pollen data from bibliography were used to complement molecular inferences. Chlorotype diversity within populations was similar throughout the natural range of C. ladanifer (mean haplotypic diversity=0.32). High differentiation among populations was estimated (G(ST)=0.60). Our data suggest that the barriers of the Strait of Gibraltar and the Betic ranges may have favoured the divergence during glacial periods of four different lineages of populations inferred with SAMOVA. The main northward colonisation of in the Iberian Peninsula occurred from refugia in southwest Iberia. This process may have been influenced by human activities (forest clearance, livestock grazing and even commerce) in the Iberian Peninsula. In contrast, populations in the Betic area have conserved a specific haplotype.  相似文献   

20.
In Europe, southern peninsulas served as major refugia during Pleistocene cold periods. However, growing evidence has revealed complex patterns of glacial survival within these southern regions, with multiple glacial refugia within each larger refugial area. We investigated the extent to which patterns of endemism and phylogeographic are concordant across animal species in the Iberian Peninsula, one of the most important unglaciated areas in Europe during the Pleistocene, can be explained in terms of climatic stability. We found that historical climatic stability (notably climate velocity measures integrating macroclimatic shifts with local spatial topoclimate gradients) was often among the most important predictors of endemic species richness for different taxonomic groups using models that also incorporated measures of modern climate. Furthermore, for some taxonomic groups, climatic stability was also correlated with patterns of spatial concordance in interpopulation genetic divergence across multiple taxa, and private haplotypes were more frequently found in relatively stable areas. Overall, our results suggest that both endemism patterns and cross‐taxa concordant phylogeographic patterns across the Iberian Peninsula to some extent are linked to spatial variation in Late Quaternary climate stability, in agreement with the proposed ‘refugia‐within‐refugia’ scenario. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 13–28.  相似文献   

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