Patterns of allozyme variation in western Eurasian Fagus

Fagus sylvatica L. is one of the most widespread broad‐leaved trees in the temperate forests of the northern hemisphere. In addition to two subspecies, F. sylvatica ssp. sylvatica in Europe and F. sylvatica ssp. orientalis in south‐western Asia, two further taxa were described: F. moesiaca (Maly) Czeczott in the south‐western Balkans and F. taurica Popl. in Crimea. The opinions about the number and ranks of taxa within this complex are highly controversial. To assess the degree of genetic differentiation among them, and to reveal geographical patterns of genetic diversity and their relationships to history and biogeography of beech populations, genetic variation at 12 allozyme loci was studied in 279 populations in western Eurasia. A Bayesian analysis of population structure revealed the existence of two clusters, which fairly well coincided with F. sylvatica ssp. sylvatica and F. sylvatica ssp. orientalis, whereby the populations from the south‐western Balkans and Crimea contained a mixture of these two gene pools. On the other hand, a neighbour‐joining tree based on pairwise F_ST failed to separate the subspecies into well‐defined distinct clades. Populations of F. sylvatica ssp. orientalis proved to be incomparably more differentiated than ssp. sylvatica (F_ST = 0.157 and 0.032, respectively). Asian populations also showed higher levels of allelic richness both on population and taxon levels than the European ones (the number of alleles after rarefaction was 3.40 and 4.27 in F. sylvatica ssp. sylvatica and ssp. orientalis, respectively). This indicates that the gene pool of F. sylvatica ssp. orientalis has not been depleted by reduced population sizes during the Pleistocene glaciations, as is the case of F. sylvatica ssp. sylvatica. Genetic similarities between isolated regional populations are explained by shared ancestral polymorphisms and/or range overlaps with subsequent hybridization in the past. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 154 , 165–174.     

The evolutionary history of Fagus in western Eurasia: Evidence from genes, morphology and the fossil record

 Fagus (beech) is among the most abundant and economically important genera of broad-leaved trees in northern hemisphere temperate forests. The number of modern taxa present in Europe and Asia Minor has long been a matter of debate and up to five species have been recognised. To resolve taxonomic and phylogenetic relationships we conducted morphological and molecular genetic analyses in western Eurasiatic taxa and evaluated palaeontological evidence. To place our findings from western Eurasiatic populations in a broader context additional East Asiatic and North American species of the same subgenus Fagus as well as two species of the subgenus Engleriana were included in our study. The morphological features exhibited in western Eurasiatic populations of Fagus show a west-east gradient that is characterised by strongly overlapping variability between geographical races. Fagus populations from Asia Minor exhibit an even higher variability that is also reflected in their genetic variability of nuclear rDNA internal transcribed spacer (ITS) sequences. The intraspecific genetic variability recorded here is in conflict with previous ITS studies in Fagus. The high amount of ITS polymorphism within Fagus from western Eurasia along with the clinal variation observed for morphological characters suggest the presence of only a single species, Fagus sylvatica L., in Europe and Asia Minor. Previously recognised taxa such as F. orientalis Lipsky and Fagus moesiaca (Maly) Czeczott should therefore be treated as synonyms of Fagus sylvatica. Although species belonging to the subgenus Engleriana were genetically distinct from species of the subgenus Fagus, relationships within the subgenus Fagus could not be clearly resolved. A reason for this could be the low rate of diversification in Fagus during the early phase of range expansion of the genus in the Oligocene period as indicated by the uniformity of leaf and cupule/nut fossils. This may account for the low overall ITS divergence and the high degree of polymorphism encountered in the subgenus Fagus and points to a late differentiation of western Eurasiatic and eastern Asiatic species. Area disruptions during the Pleistocene and the post-glacial recolonisation of western Europe appear to have caused the west-east gradient that is apparent in modern Fagus of western Eurasia but absent in Late Tertiary ancestors of Fagus sylvatica. Received June 22, 2001 Accepted February 25, 2002     

Leaf morphological variation in beech (Fagus sylvatica L.) populations in Greece and its relation to their post‐glacial origin

Diversity in leaf morphological traits was assessed in 38 Greek Fagus sylvatica populations, covering the distribution of the species in the country. The relationship between the post‐glacial origin of these populations and leaf morphology was investigated. The results showed a complex and geographically continuous morphological diversity pattern, influenced mainly by traits expressing leaf size, leaf shape and petiole length. Two simultaneous trends appeared to be responsible for the existing diversity pattern. One was geographical, with leaf types resembling F. sylvatica subsp. sylvatica occurring in the western part of the distribution of beech in Greece and types resembling F. sylvatica subsp. orientalis being dominant in the eastern part. A second trend seemed to be connected with the post‐glacial origin of the populations, as described by previous plastid DNA haplotype studies of the same trees. The genetic background and the possible adaptation of beech populations to different environmental conditions have resulted in a complex morphological pattern, especially in areas in which different post‐glacial lineages appear to meet. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 422–436.     

Unexpected presence of <Emphasis Type="Italic">Fagus orientalis</Emphasis> complex in Italy as inferred from 45,000-year-old DNA pollen samples from Venice lagoon

Background

Phylogeographic analyses on the Western Euroasiatic Fagus taxa (F. orientalis, F. sylvatica, F. taurica and F. moesiaca) is available, however, the subdivision of Fagus spp. is unresolved and there is no consensus on the phylogeny and on the identification (both with morphological than molecular markers) of Fagus Eurasiatic taxa.For the first time molecular analyses of ancient pollen, dated at least 45,000 years ago, were used in combination with the phylogeny analysis on current species, to identify the Fagus spp. present during the Last Interglacial period in Italy.In this work we aim at testing if the trn L-trn F chloroplast DNA (cpDNA) region, that has been previously proved efficient in discriminating different Quercus taxa, can be employed in distinguishing the Fagus species and in identifying the ancient pollen.

Results

86 populations from 4 Western Euroasistic taxa were sampled, and sequenced for the trn L-trn F region to verify the efficiency of this cpDNA region in identifying the Fagus spp.. Furthermore, Fagus crenata (2 populations), Fagus grandifolia (2 populations), Fagus japonica, Fagus hayatae, Quercus species and Castanea species were analysed to better resolve the phylogenetic inference.Our results show that this cpDNA region harbour some informative sites that allow to infer relationships among the species within the Fagaceae family. In particular, few specific and fixed mutations were able to discriminate and identify all the different Fagus species.Considering a short fragment of 176 base pairs within the trn L intron, 2 transversions were found able in distinguishing the F. orientalis complex taxa (F. orientalis, F. taurica and F. moesiaca) from the remaining Fagus spp. (F. sylvatica, F. japonica, F. hayataea, F. crenata and F. grandifolia). This permits to analyse this fragment also in ancient samples, where DNA is usually highly degraded.The sequences data indicate that the DNA recovered from ancient pollen belongs to the F. orientalis complex since it displays the informative sites characteristic of this complex.

Conclusion

The ancient DNA sequences demonstrate for the first time that, in contrast to current knowledge based on palynological and macrofossil data, the F. orientalis complex was already present during the Tyrrhenian period in what is now the Venice lagoon (Italy).This is a new and important insight considering that nowadays West Europe is not the natural area of Fagus orientalis complex, and up to now nobody has hypothesized the presence during the Last Interglacial period of F. orientalis complex in Italy.

     

Small genome size variation across the range of European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>)

Interpopulation variation of relative and absolute genome size was studied in Fagus sylvatica subsp. sylvatica and subsp. orientalis. The study included 18 populations of beech planted in a common-garden experiment in central Slovakia and three additional populations from the Caucasus. Nuclear DNA content was determined by means of flow cytometry using the AT-specific fluorochrome 4′,6-diamidino-2-phenylindole and non-specific propidium iodide, and its associations with climate, growth, phenology and physiological traits were assessed. The approximate average nuclear DNA content (2C) across all samples was 1.178 ± 0.020 pg. The lowest mean relative genome sizes were observed in the Alpine range, whereas they increased toward the range margins; no clear trend was observed for 2C values. Temperature seasonality and temperature annual range were found to be negatively associated with genome size. Among phenotypic traits, the maximum chlorophyll a fluorescence yield (Fv/Fm) was found to be negatively correlated with relative genome size, whereas phenology and some photosynthetic parameters were correlated with the 2C values.     

Genetic differentiation and phylogeny of beech on the Balkan peninsula

Genetic differentiation of 57 beech populations originating from a wide geographical space from the foothills of the Eastern Alps over the Balkan peninsula was investigated employing isozyme markers. Pooled datasets of European beech populations originating from four adjacent regions and Eastern beech populations originating from Thrace and Western Asia Minor were compared. Considerable differences of allele frequencies among regions were found in several marker loci. The highest level of genetic multiplicity and differentiation was found in the populations from the southern Balkans; however, the north-western populations showed higher genetic diversity. The pattern of genetic differentiation based on multilocus genetic distances is a clinal one. The populations belonging to the putative taxon Fagus moesiaca Czeczott seem to form an independent group. Three hypotheses of the evolutionary origin of this taxon are discussed: selection, introgressive hybridization and continuous evolution.     

Culturability and Persistence of <Emphasis Type="Italic">Francisella noatunensis</Emphasis> subsp. <Emphasis Type="Italic">orientalis</Emphasis> (syn. <Emphasis Type="Italic">Francisella asiatica</Emphasis>) in Sea- and Freshwater Microcosms

Francisella noatunensis subsp. orientalis (syn. Francisella asiatica), the causative agent of franciselliosis in warm-water fish, is a Gram-negative facultative intracellular bacterium. Although it has been characterized as one of the most pathogenic bacteria in fish, the water conditions that allow for its survival and infectious capacities outside the fish host are not known. Data obtained in this project indicate that both temperature and salinity are important factors in the culturability and persistence of F. noatunensis subsp. orientalis in both sea- and freshwater microcosms. These results indicate that culturable F. noatunensis subsp. orientalis persist for longer periods of time and at higher numbers in seawater, and its persistence is inversely related to water temperature. Moreover, the pathogenic properties of the bacteria suspended in water microcosms appear to decrease after only 24 h and become non-infective after 2 days in the absence of the fish host.     

Evolutionary melting pots: a biodiversity hotspot shaped by ring diversifications around the Black Sea in the Eastern tree frog (Hyla orientalis)

Hotspots of intraspecific genetic diversity, which are of primary importance for the conservation of species, have been associated with glacial refugia, that is areas where species survived the Quaternary climatic oscillations. However, the proximate mechanisms generating these hotspots remain an open issue. Hotspots may reflect the long‐term persistence of large refugial populations; alternatively, they may result from allopatric differentiation between small and isolated populations, that later admixed. Here, we test these two scenarios in a widely distributed species of tree frog, Hyla orientalis, which inhabits Asia Minor and southeastern Europe. We apply a fine‐scale phylogeographic survey, combining fast‐evolving mitochondrial and nuclear markers, with a dense sampling throughout the range, as well as ecological niche modelling, to understand what shaped the genetic variation of this species. We documented an important diversity centre around the Black Sea, composed of multiple allopatric and/or parapatric diversifications, likely driven by a combination of Pleistocene climatic fluctuations and complex regional topography. Remarkably, this diversification forms a ring around the Black Sea, from the Caucasus through Anatolia and eastern Europe, with terminal forms coming into contact and partially admixing in Crimea. Our results support the view that glacial refugia generate rather than host genetic diversity and can also function as evolutionary melting pots of biodiversity. Moreover, we report a new case of ring diversification, triggered by a large, yet cohesive dispersal barrier, a very rare situation in nature. Finally, we emphasize the Black Sea region as an important centre of intraspecific diversity in the Palearctic with implications for conservation.     

Galega orientalis is more diverse than Galega officinalis in Caucasus--whole-genome AFLP analysis and phylogenetics of symbiosis-related genes

Legume plants can obtain combined nitrogen for their growth in an efficient way through symbiosis with specific bacteria. The symbiosis between Rhizobium galegae and its host plant Galega is an interesting case where the plant species G. orientalis and G. officinalis form effective, nitrogen‐fixing, symbioses only with the appropriate rhizobial counterpart, R. galegae bv. orientalis and R. galegae bv. officinalis, respectively. The symbiotic properties of nitrogen‐fixing rhizobia are well studied, but more information is needed on the properties of the host plants. The Caucasus region in Eurasia has been identified as the gene centre (centre of origin) of G. orientalis, although both G. orientalis and G. officinalis can be found in this region. In this study, the diversity of these two Galega species in Caucasus was investigated to test the hypothesis that in this region G. orientalis is more diverse than G. officinalis. The amplified fragment length polymorphism fingerprinting performed here showed that the populations of G. orientalis and R. galegae bv. orientalis are more diverse than those of G. officinalis and R. galegae bv. officinalis, respectively. These results support the centre of origin status of Caucasus for G. orientalis at a genetic level. Analysis of the symbiosis‐related plant genes NORK and Nfr5 reveals remarkable diversity within the Nfr5 sequence, although no evidence of adaptive evolution could be found.     

Die Fagus-Sippen Europas und ihre geographisch-soziologische Korrelation zur Verbreitung der Assoziationen des Fagion s.l.

Studies of the beeches and beech woods of eastern central Europe revealed, that in the postglacial period not only Fagus sylvatica (L.) emend. reimmigrated from the refugial territories in the SE-as has generally been accepted, but also the transitional taxa, which originated from hybrids with F. orientalis Lipsky. The NW area limit of these intermediate taxa must be revised. The presence in central Europe of these taxa-which form own Fagion alliances and associations in the SE (Fagus intermedia ssp. moesiaca and ssp. taurica) as well as the presence of other southeastern species in central European beach woods shows, that their postglacial development is parallel to, but different from other areas. The taxon Fagus intermedia (ssp. neglecta and ssp. transitus) became differential taxa of a central European region of the Fagion medioeuropaeum. Plant sociologists are therefore requested to make new and critical analyses of the beech woods in Europe, with special attention to the Fagus taxa, in order to establish in detail the geographical distribution and phytosociological significance of Fagus intermedia.The author offers to determine or revise Fagus material (herbarium collections).

Danksagung. Ich möchte meinen tiefsten und allzeitigen Dank besonders den Herren Prof. Dr. A. O. Horvat (Pécs), Prof. Dr. Ch. Moulopoulos, Prof. Dr. B. Jovanovié, Prof. Dr. I. Dumitriu-Tataranu, Prof. Dr. R. Bornkamm, Prof. Dr. M. A. Kotschkin, Prof. Dr. P. Fukarek, Prof. Dr. K. Browicz für gewährte Unterstützung aussprechen und ganz besonders unserem unvergeßlichen verstorbenen Prof. Dr. Drs. h.c. R. Tüxen, der mir zur Fortführung dieser Studien Mut machte.     

The Holocene history of low altitude Mediterranean Fagus sylvatica forests in southern France

Questions

As the dominant tree in many European forests, Fagus sylvatica functions as an ecosystem engineer, yet its istory remains little understood. Here we ask: (a) are there indications for its presence in southeast France during the last Glacial period; (b) what was the timing of the expansion and decline of F. sylvatica dominated forests; (c) which factors influenced their dynamics and in particular to what extent did past precipitation changes impact upon them; and (d) at which altitudes did these beech forests occur within the region?

Location

Languedoc, the French Mediterranean area.

Method

This article presents a well dated and high‐resolution pollen sequence covering the last 7,800 years from the Palavas Lagoon in the Languedoc together with a review of Fagus charcoal occurrences in the Languedoc and the lower Rhône Valley, and a review of pollen data from a compilation of 69 sites in southeast France.

Results

The Palavas pollen sequence provides a regional summary of F. sylvatica abundance changes near the Mediterranean coast. Around 6,000 years cal BP , an abrupt transition from small beech populations to well‐developed forests is recorded. The maximum development of beech forests occurred between 4,000 and 3,000 years cal BP , while F. sylvatica started to regress after 3,000 years cal BP .

Conclusion

Scattered F. sylvatica populations probably survived throughout southern France during the last Glacial period. F. sylvatica started to spread around 8,000 years cal BP while beech forests never expanded before 6,000 years cal BP . The complex patterns of F. sylvatica expansion in southern France after 6,000 years cal BP suggests that a combination of global (climate change) and local (human impact) factors were responsible for this major change. Recurrent abrupt climate changes, the aridity trend and human deforestation caused beech forests to decline after 3,000 years cal BP .

     

Local differentiation of Albanian provenances and chorology of beech species in balcan area

Abstract

Some quantitative characters of different Albanian beech provenances are analysed in order to study their local differentiation. The results obtained by discriminant analysis showed the formation of two principal groups. Biza provenance was discriminated from both groups. Principal components and cluster analysis give a good evidence on the similarities among Albanian and Bulgarian beech populations. The results suggest that all area of Balcan is the contact zone of Fagus sylvatica L. and Fagus orientalis Lypski.     

Some evidence for infrageneric classification in Ziziphora L. (Lamiaceae:Mentheae)

The genus Ziziphora L. (Lamiaceae) consists of annual herbs, except Ziziphora clinopodioides, and is represented by six taxa in the flora of Turkey: Z. clinopodioides, Z. capitata, Z. persica, Z. tenuior, Z. taurica subsp. taurica and Z. taurica subsp. cleonioides. They are strong aromatic herbs and important medicinal plants. This study has been conducted to determine the palynological and karyological features of these taxa. The pollen morphology and exine structure of Ziziphora species were investigated by light microscope and scanning electron microscope. Pollen grains were determined as hexacolpate, medium in size and a circular or elliptic amb. The shape of pollen grains varies from oblate to prolate spheroidal. Ornamentation types of pollen grains are bireticulate, microreticulate or perforate. Somatic chromosome number has been determined and counted as 2n = 16, 18. Results indicate that the pollen characters and chromosome numbers of the genus are valuable for taxonomic applications and may be useful for infrageneric classification.     

Elevation of Francisella philomiragia subsp. noatunensisMikalsen et al. (2007) to Francisella noatunensis comb. nov. [syn. Francisella piscicida Ottem et al. (2008) syn. nov.] and characterization of Francisella noatunensis subsp. orientalis subsp. nov., two important fish pathogens

Aims: This study was conducted to clarify the taxonomic status of Francisella sp. strain Ehime‐1, a fish pathogen, in relation to the fish pathogens F. piscicida and F. philomiragia subsp. noatunensis and to F. philomiragia subsp. philomiragia. Methods and Results: Francisella sp. Ehime‐1 was compared to F. piscicida, F. philomiragia subsp. noatunensis and several F. philomiragia subsp. philomiragia isolates through sequencing of the 16S rRNA‐gene and several house‐keeping genes and determination of biochemical and phenotypic properties. Results show that F. piscicida is indistinguishable from F. philomiragia subsp. noatunensis by sequence and phenotypic traits. Francisella sp. Ehime‐1 and F. philomiragia subsp. noatunensis are clearly separated from F. philomiragia. Francisella sp. Ehime‐1 is biochemically, phenotypically and genetically different from F. philomiragia subsp. noatunensis (=F. piscicida), but DNA–DNA hybridization does not clearly support establishment as a separate species (level of relatedness 64% and 73·4%, mean 68·7%). Conclusions: We propose to elevate F. philomiragia subsp. noatunensis to species rank as F. noatunensis comb. nov., while F. piscicida is considered a heterotypic synonym of F. noatunensis comb. nov. Evidence suggests that Francisella sp. Ehime‐1 represents a novel subspecies of F. noatunensis, for which the name F. noatunensis subsp. orientalis subsp. nov. is proposed (=DSM21254^T, = LMG24544^T). Significance and Impact of the Study: This study contributes to the taxonomy and characteristics of fish‐pathogenic Francisella spp.     

Phylogeographical structure and conservation genetics of wild grapevine

The distribution of Vitis vinifera subsp. silvestris, the wild grapevine subspecies of Vitis vinifera L., has been dramatically reduced in its major sites of diffusion, at first by the spread, over the last 150 years, of pathogens from North America and, more recently, with fragmentation of habitat and disbranching by humans. In this work, 418 wild grapevine samples, belonging to 78 populations, were collected in their main Mediterranean distribution areas, including the Caucasus area, and the extent of their genetic variability evaluated by analysing plastid microsatellite DNA polymorphism. Results show low haplotype diversity value, with five haplotypes detected within the analysed populations. The highest within-population haplotypic diversity, with the presence of all five detected haplotypes, was found in the Caucasus regions and in the central regions of Italy. The distribution of all detected haplotypes suggests the Caucasian region as the possible center of origin of Vitis vinifera subsp. silvestris. A principal plastid lineage was found to be fixed in several locations, in the Northernmost European countries and in the Southern island of Sardinia. These results draw attention to two different refugium sites in the Mediterranean basin and suggest that conservation priority should be given to grapevine populations still preserved in hotspots of these regions.     

The phylogeny of Lens (Leguminosae): new insight from ITS sequence analysis

 Lens includes L. culinaris subsp. culinaris (the cultivated lentil) and several wild species distributed from the Mediterranean region to western Asia. We compared sequence variation in the ITS region among species of Lens in an effort to end persisting uncertainty regarding the phylogeny of the genus. The parsimony analysis revealed a single minimum-length tree with a topology congruent with patterns derived by previous studies of nuclear and chloroplast DNA RFLPs. The basal and highly divergent status of the L. nigricans clade is depicted, and the progenitor-derivative relationship between L. culinaris subsp. orientalis and L. culinaris subsp. culinaris is reaffirmed. Resolution in the tree was improved by combining the ITS data set with a pre-existing set of chloroplast DNA restriction site data obtained from the same group of samples. Received May 8, 2000 Accepted October 26, 2001     

Morphology and Phylogeny of Two Species of Loxodes (Ciliophora,Karyorelictea), with Description of a New Subspecies,Loxodes striatus orientalis subsp. n.

The morphology and phylogeny of Loxodes vorax and L. striatus orientalis subsp. n. were investigated based on infraciliature and small subunit (SSU) rRNA gene sequence data. Loxodes striatus orientalis subsp. n. was separated from L. striatus striatus stat. n. by having fewer dikinetids in the intrabuccal kinety (35–55 vs. 50–70) and a variable number of macronuclei (2–4 vs. 2). In addition, the SSU rRNA gene sequence of the new subspecies differs in 13 and 11 nucleotides from that of two populations of the nominotypic subspecies. We also summarized the morphological differences between Loxodes and Remanella based on the data available. Phylogenetic analyses revealed that the genus Loxodes was monophyletic and nested within Remanella species. This study might, therefore, support the hypothesis that the freshwater genus Loxodes evolved from the marine genus Remanella.     

Rapid climate change-related growth decline at the southern range edge of Fagus sylvatica

Studies on Fagus sylvatica show that growth in populations toward the southern limit of this species' distribution is limited strongly by drought. Warming temperatures in the Mediterranean region are expected to exacerbate drought where they are not accompanied by increases in precipitation. We studied levels of annual growth in mature F. sylvatica trees over the last half‐century in the Montseny Mountains in Catalonia (northeast Spain). Our results show significantly lower growth of mature trees at the lower limit of this species' distribution when compared with trees at higher altitudes. Growth at the lower Fagus limit is characterized by a rapid recent decline starting in approximately 1975. By 2003, growth of mature trees had fallen by 49% when compared with predecline levels. This is not an age‐related phenomenon, nor is it seen in comparable populations at higher altitudes. Analysis of climate‐growth relationships suggests that the observed decline in growth is a result of warming temperatures and that, as precipitation in the region has not increased, precipitation is now insufficient to ameliorate the negative effects of increased temperatures on tree growth. As the climate‐response of the studied forest is comparable with that of F. sylvatica forests in other southern European regions, it is possible that this growth decline is a more widespread phenomenon. Warming temperatures may lead to a rapid decline in the growth of range‐edge populations and a consequent retreat of the species distribution in southern Europe. Assessment of long‐term growth trends across the southern range edge of F. sylvatica therefore merits further attention.     

Genetic diversity and structure among Iranian tall fescue populations based on genomic-SSR and EST-SSR marker analysis

Tall fescue (Festuca arundinacea Schreb. subsp. arundinacea) is one of the most economically important forage grasses in cold and temperate regions of the world. In this study, we have assessed the genetic diversity and structure of wild Iranian tall fescue populations. Thirty-seven individuals from nine natural populations from northern, western, and southern Iranian mountain ranges were analyzed using six genomic-SSRs and five EST-SSRs primer pairs. Our analysis has demonstrated that transcribed EST-SSR regions showed levels of polymorphism similar to genomic-SSR regions. UPGMA, repeated bisection, and model-based Bayesian STRUCTURE clustering methods coupled with neighbor-net network were used to establish six divergent groups of individuals. F _ST estimates among clusters showed moderate to low genetic structure. Within-group genetic diversity estimate H and partial correlations between genetic and geographic distances among populations suggested that western Zagros population was related to the rest of the Iranian individuals. The isolation-by-distance hypothesis was not supported by SSR data and the present geographical sampling.