Syntaxonomy and biogeography of dry grasslands on calcareous substrates in the central and southern Balkans

Questions

Which major syntaxa of dry grasslands supported by carbonate bedrock occur in the central and southern Balkans? What is their position along major ecological gradients and in the context of phytogeographic patterns of the region?

Location

Central and southern Balkans, including western Bulgaria, northern Greece, Kosovo, Macedonia (FYROM) and Serbia.

Methods

We compiled a matrix of 660 relevés of dry grasslands over lime‐rich bedrock, previously classified in the Festuco‐Brometea. We applied clustering techniques to classify separately synoptic and relevé data, and applied NMDS with passive projection of indicator values, climatic data and biogeographic geo‐elements onto ordination diagrams to assist interpretation of the syntaxonomic patterns. We constructed elevation distribution profiles for alliances and classes of grasslands of several grassland classes from a broader study area to elucidate the relationship of the elevational sorting of the syntaxa in relation to latitude.

Results

The analysis revealed six major vegetation types, classified into four orders: (1) Stipo pulcherrimae‐Festucetalia pallentis, incl. (sub)montane rocky steppic grasslands of the Saturejion montanae of central Balkans, and the Koelerio‐Festucion dalmaticae – submontane rocky grasslands of southern Serbia and Kosovo; (2) Astragalo onobrychidis‐Potentilletalia represented by the Saturejo‐Thymion (low‐elevation steppic grasslands of southern Balkans); (3) Festucetalia valesiacae represented by grasslands on deep soil and low elevation of northern Greece, and finally (4) high‐elevation rocky grasslands of southern Balkans, classified as a new alliance – Diantho haematocalycis‐Festucion hirtovaginatae, that might belong to a new, yet undescribed, syntaxonomic order. Ordination suggests that the major differentiation of the high‐rank syntaxa follows north–south geographic and low–high elevation gradients.

Conclusions

Because of the transitional biogeographic position of the studied region, as well as considerable large elevation span across latitudes, the diversity of vegetation types is high. The indication a putative new dry grassland order, the mid‐high altitudes of the southern Balkans points to a need to re‐assess the Balkan vegetation occupying the community niche between the low‐elevation dry grasslands (Festuco‐Brometea) and those typical of high elevations (Elyno‐Seslerietea and Daphno‐Festucetea), seeking parallels to patterns described from the western Alps, Pyrenees, and Apennines. This syntaxonomic unit is poised to expand the concept of the Festuco hystricis‐Ononidetea striatae to the Balkans.

     

New insights into the origin and the genetic status of the Balkan donkey from Serbia

The Balkan donkey (Equus asinus L.) is commonly regarded as a large‐sized, unselected, unstructured and traditionally managed donkey breed. We assessed the current genetic status of the three largest E. asinus populations in the central Balkans (Serbia) by analysing the variability of nuclear microsatellites and the mitochondrial (mtDNA) control region of 77 and 49 individuals respectively. We further analysed our mtDNA dataset along with 209 published mtDNA sequences of ancient and modern individuals from 19 European and African populations to provide new insights into the origin and the history of the Balkan donkey. Serbian donkey populations are highly genetically diverse at both the nuclear and mtDNA levels despite severe population decline. Traditional Balkan donkeys in Serbia are rather heterogeneous; we found two groups of individuals with similar phenotypic features, somewhat distinct nuclear backgrounds and different proportions of mtDNA haplotypes belonging to matrilineal Clades 1 and 2. Another group, characterized by larger body size, different coat colour, distinct nuclear gene pool and predominantly Clade 2 haplotypes, was delineated as the Banat donkey breed. The maternal landscape of the large Balkan donkey population is highly heterogeneous and more complex than previously thought. Given the two independent domestication events in donkeys, multiple waves of introductions into the Balkans from Greece are hypothesized. Clade 2 donkeys probably appeared in Greece prior to those belonging to Clade 1, whereas expansion and diversification of Clade 1 donkeys within the Balkans predated that of Clade 2 donkeys.     

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.     

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     

A new cytochrome b phylogroup of the common vole (Microtus arvalis) endemic to the Balkans and its implications for the evolutionary history of the species

The phylogeographic architecture of the common vole, Microtus arvalis, has been well‐studied using mitochondrial DNA and used to test hypotheses relating to glacial refugia. The distribution of the five described cytochrome b (cyt b) lineages in Europe west of Russia has been interpreted as a consequence of postglacial expansion from both southern and central European refugia. A recently proposed competing model suggests that the ‘cradle’ of the M. arvalis lineages is in western central Europe from where they dispersed in different directions after the Last Glacial Maximum. In the present study, we report a new cyt b lineage of the common vole from the Balkans that is not closely related to any other lineage and whose presence might help resolve these issues of glacial refugia. The Balkan phylogroup occurs along the southern distributional border of M. arvalis in central and eastern Bosnia and Herzegovina, Montenegro, and eastern Serbia. Further north and west in Slovenia, Bosnia and Herzegovina, and Serbia, common voles belong to the previously‐described Eastern lineage, whereas both lineages are sympatric in one site in Bosnia and Herzegovina. The Balkan phylogroup most reasonably occupied a glacial refugium already known for various Balkan endemic species, in contrast to the recently proposed model. South‐east Europe is an absolutely crucial area for understanding the postglacial colonization history of small mammals in Europe and the present study adds to the very few previous detailed phylogeographic studies of this region. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 788–796.     

Revision of the central Mediterranean xerothermic cliff vegetation

Questions

What are the syntaxonomic and synchorological patterns of the xerothermic chasmophytic vegetation in the central part of the Mediterranean Basin? What are the diagnostic species of the high‐rank syntaxa of Asplenietalia glandulosi, Onosmetalia frutescentis and Centaureo dalmaticae‐Campanuletalia pyramidalis?

Location

Mediterranean coastal and subcoastal areas of southern France, Italy, Malta, Slovenia, Croatia, Bosnia and Herzegovina, Montenegro, Albania and of mainland Greece.

Methods

The data set of 1,261 published relevés was analysed using hierarchical clustering (Flexible Beta method), involving a series of data transformations. Indicator species analysis was used to select the best dendrogram solution and identify diagnostic taxa of the main clusters. The dendrogram was interpreted from a syntaxonomic point of view, using nomenclatural type relevés as a basis. The NMDS ordination was performed in order to visualize the floristic relationships among associations and high‐rank syntaxa. MRPP was used to test for differences among alliances.

Results

The classification revealed four main clusters of relevés representing the chasmophytic vegetation of southern France, Sardinia and the northwestern part of Italy (Asplenienalia glandulosi/Asplenietalia glandulosi), the southwestern part of Italy and Malta (Tinguarrenalia siculae/Asplenietalia glandulosi), the Adriatic Basin area (Centaureo dalmaticae‐Campanuletalia pyramidalis) and the southern Balkans (Onosmetalia frutescentis). The NMDS ordination confirmed the overall pattern, while MRPP showed significant differences among the alliances of the above‐mentioned orders and suborders. The lists of diagnostic taxa of the high‐rank syntaxa were revised according to a supra‐national perspective.

Conclusions

The new syntaxonomic scheme provides a comprehensive overview of the chasmophytic vegetation of the central part of the Mediterranean Basin. This scheme mostly matches the recently published EuroVegChecklist, but also exhibits important novelties concerning the syntaxonomic position of some alliances (Dianthion rupicolae, Centaureion pentadactyli, Arenarion bertolonii and Caro‐Aurinion), and the floristic and chorological relationships among high‐rank syntaxa, with new revised sets of diagnostic taxa. This revision might be useful for further small‐scale phytosociological studies.

     

The Balkans and the colonization of Europe: the post‐glacial range expansion of the wild boar,Sus scrofa

Aim We focus on the biogeographical role of the Balkan Peninsula as a glacial refugium and source of northward post‐glacial dispersal for many European taxa. Specifically, we analysed the genetic structure and variation of wild boar (Sus scrofa) samples primarily from Greece, a region that has repeatedly served as a glacial refugium within the Balkan Peninsula. Location Continental Greece, the Aegean island of Samos and Bulgaria. Methods We analysed wild boar samples from 18 localities. Samples from common domestic breeds were also examined to take into account interactions between wild and domesticated animals. Phylogenetic analyses were carried out on a 637‐bp fragment of the mitochondrial DNA control region in 200 wild boar and 27 domestic pigs. The sequences were also compared with 791 Eurasian wild boar and domestic pig D‐loop sequences obtained from GenBank. Results Ninety‐four haplotypes were identified in the European wild boar data set, of which 68 were found in the Balkan samples and assigned to two previously described clades: the E1 European and Near Eastern clades. All of the continental samples clustered in the E1 clade and the samples from Samos fell into the Near Eastern clade, consistent with the island’s proximity to Asia Minor. Intriguingly, 62 novel haplotypes were identified and are found exclusively in the Balkans. Only six haplotypes were shared between wild boar and domestic pigs. Main conclusions Our data reveal numerous novel and geographically restricted haplotypes in wild boar populations, suggesting the presence of separate refugia in the Balkans. Our analyses support the hypothesis of a post‐glacial wild boar expansion consistent with the leading edge model, north and west from modern day Greece, and suggest little maternal introgression of Near Eastern and domestic haplotypes into wild Balkan populations.     

Fagus sylvatica forest vegetation in Greece: Syntaxonomy and gradient analysis

Abstract. This is the first comprehensive study on Fagus sylvatica s.l. forest communities in Greece. It is based on the multivariate analysis of more than 1100 relevés throughout the distribution area of Fagus in Greece, 60 % of which have been sampled within the last 15 years. The data were compiled from all relevant literature and from unpublished field studies. Prior to data analysis, taxonomic and floristic inaccuracies in older literature have been corrected, and some taxa aggregated. Status and distribution of Fagus in Greece, as well as historical and silvicultural aspects, are outlined. Classification resulted in 16 vegetation types defined by species composition, of which 10 units are ranked as association (of which two were described as new) or community, with six subcommunities included. Synonyms from the literature are given to all types. The floristic structure of the data set is displayed by a synoptic table and by DCA ordination. Community differentiation follows a complex edaphic‐climatic‐phytogeographic pattern. Apart from Fagus sylvatica ssp. orientalis forests of warm and dry habitats and low to medium altitudes which are related to deciduous Quercus forests, four Fagion community groups are distinguished which represent different habitat types: (1) cool, humid, mesotrophic; (2) cool, acidic; (3) high altitude, calcareous; (4) medium altitude, moderately warm and dry. The communities may be assigned to three regional suballiances, viz. (a) Doronico orientalis‐Fagenion at medium altitudes of North‐Central and East‐Central Greece; (b) Doronico columnae‐Fagenion in the far north and at high altitudes southward to East‐Central Greece; (c) Geranio striati‐Fagenion in the Northwest. Various types of thermophilous beech forest are assigned to (a) and (c), acidic forest to (b), and mesic Fagus forest to (b) and (c).     

Anthropogenous areal extension of central European woody species on the British Isles and its significance for the judgement of the present potential natural vegetation

Several central European species have failed to reach the British Isles because of the early separation from the continent. The two tree species Fagus sylvatica and Carpinus betulus reached southern England but were unable to spread much further. Other species, such as Acer pseudoplatanus, were only relatively recently introduced. Recent distribution maps for Fagus and Acer show an almost uninterrupted distribution in the whole British Isles. The beech must be considered to be an important element of the present potential natural vegetation; it has been planted widely and regenerates freely. The woodlands of Brittany may provide a model to enable us to visualize the possible species composition and appearance of these potential Atlantic beech woods.Nomenclature of vascular taxa follows Ehrendorfer (1973).     

Lowland woodland structure and pattern and the distribution of arboreal,phytophagous arthropods

We investigated factors that limited the distribution of phytophagous species within a woodiand system in Midlothian, Scotland. A pattern analysis was conducted of phytophagous species on a total of 45 Fagus sylvatica within 15 woodlands. Species richness counted on collected leaves was tested against within-and between-wood variables. Variables used in a regression with arthropod data from Fagus were used to estimate the phytophage richness on Betula pendula and Quercus robur in the same woods. Convariance in the number of phytophages in sampled woods was found for Fagus over three years and for Fagus, Betula and Quercus in 1992. Association analysis was used to classify the woods into species rich or poor based on presence or absence matrices. The main factors that limit phytophages on Fagus (gaps along the woodland edge, depth and species richness of the field layer. density of leaf litter and the extent of contiguous woodland cover, when including hedgerows and lines of trees) affect phytophages of similar life history strategy on other tree species within the same woods. Eighty-six per cent of species were lost because certain life history stages were vulnerable to factors that prevail in woods of poor structure. The nature conservation value of woodlands could be assessed using the correlated vulnerability of particular phytophages across tree species under specific woodland conditions.     

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.     

Taxonomy and ecology of <Emphasis Type="Italic">Castanea sativa</Emphasis> Mill. forests in Greece

Castanea sativa Mill. is an important species of the Balkan Peninsula with high ecological and economic value. This article contributes to a phytosociological synsystematic approach of the C. sativa plant communities in Greece that covers the southern part of the Balkan Peninsula and describes the ecological parameters involved in their distribution and taxonomy. Phytosociological data have been collected from 14 mountainous areas across continental Greece, which are considered representative of the main forest types of C. sativa in the area. The 14 mountains demonstrate a variety of environmental conditions. Five communities (associations) and seven sub-communities (sub-associations) have been identified and described using TWINSPAN, Corespondence Analysis and the Braun-Blanquet classification scheme. Castanea sativa (chestnut) forests of Greece can be distinguished as three broad groups, defined primarily by their geographical distribution. In northern and central Greece chestnut stands are characterized by their high floristic homogeneity, which is reflected in the identification of only one community per region. Chestnut forests in northern Greece, in particular, exhibit strong floristic similarities with those of the rest of the Balkans. In southern Greece, on the other hand, the southern distribution limit of chestnut in the Balkans, there is a greater floristic diversity between the sampled mountains which is reflected in the presence of three confirmed communities and two sub-associations. Several factors have been identified as important in determining the structure and floristic composition of chestnut forests in Greece. Whilst climate and the grazing regime are influential, the degree of silvicultural management appears to be the most important factor determining the floristic composition of chestnut forests and their long-term sustainability.     

Phylogeographic pattern of the European forest grass species Hordelymus europaeus: cpDNA evidence

The phylogeographical structure of the European forest grass Hordelymus europaeus (Poaceae) was studied by sequencing three non-coding regions of chloroplast DNA. Forty-three populations within the entire natural distribution area were analysed. The greatest haplotype variation and divergence were revealed on Balkan and Apennine Peninsula, suggesting main glacial refugia in these regions. Among southern refugia, probably only the Balkans could have remarkably contributed to postglacial re-colonisation of the northern parts of Europe. A distinctly different haplotype group found on the Crimean Peninsula and in central Italy may represent either relicts of a previously more widespread ancestor or result of long-distance dispersal. The phylogeographical pattern found in H. europaeus is to certain extent similar with that found in Fagus sylvatica. This might imply a partly common postglacial colonisation history of these ecologically narrowly tied species. Nevertheless, unlike in the case of F. sylvatica, we did not found convincing evidence for the existence of Central European glacial refugia for H. europaeus.     

The spider fauna of the Aladjagiola wetland complex (Nestos Delta,north‐east Greece): a reflection of a unique zoogeographical transition zone in Europe

In order to increase our arachnological knowledge of Greece, and in particular north‐east Greece, an investigation of the arachnofauna of the Aladjagiola wetland complex was conducted from April to July 2008. The study area is part of the Nestos Delta in north‐east Greece, and comprises an area of approximately 20 km². Spiders were collected from 28 sites using pitfall traps. In all, 6694 specimens were sampled, including 206 species from 31 families. Altogether, 30 and 74 species were new to the districts of Greece and Macedonia, respectively. A number of faunistically interesting species were recorded during the study: Brachythele denieri (Simon, 1916), Camillina metellus (Roewer, 1928), Zelotes cingarus (O.P.‐Cambridge, 1874), Zodarion blagoevi Bosmans, 2009, Zodarion epirense Brignoli, 1984, Zodarion hauseri Brignoli, 1985, and Zodarion pirini Drensky, 1921, are Balkan endemics; Tegenaria angustipalpis Levy, 1996 and Titanoeca turkmenia Wunderlich, 1995 were newly discovered in Europe. Widespread elements represented the largest share of the species inventory, although chorological analysis indicated that the study area, as well as the whole north‐eastern part of Greece, is a reflection of a transition zone from the European fauna towards the Mediterranean and eastern (or Ponto–Anatolian) elements. Taken in combination, these affect the local fauna. The Balkan endemics, although small in number of species, further underline the unique identity of the area. Based on comparisons with data sets from the country's southernmost island, i.e. Crete, Greece can be regarded as a zoogeographical gradient. Hereby, Aladjagiola represents the northern edge, with European and Anatolian elements dominating the species inventory, and Crete represents the southern edge, with widespread but also endemic Mediterranean and eastern elements creating a more local and insular zoogeographical character. In conclusion, Greece cannot be viewed as a uniform zoogeographical unit, but rather as a mosaic of various regional zoogeographical patterns. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 217–233.     

Implications of 19th century landscape patterns for the recovery of Fagus crenata forests

Questions: What is the effect of the 19th century (pre‐industrialization) landscape pattern on the recovery of climax forests in cool‐temperate mountain areas dominated by Fagus crenata (Japanese beech)? Location: Secondary forests on Mt. Daisen, western Japan. Methods: Vegetation patterns before and after industrialization were obtained from maps drawn in 1898 and 1979. Tree measurements were made in 12 plots in 1997. Correlation between current Fagus crenata dominance and forest edge in the 19th century was analysed using an S‐shaped regression curve. Fagus juvenile density was counted in the plots, and distances from each plot to the five nearest mother trees were measured to determine the dispersal kernel. Results: Secondary grassland covered a substantial area in 1898, whereas forest covered most of the area in 1997. Fagus was dominant in places in the interior forest 100 years ago, and mature Fagus trees were absent in secondary forests that had been grasslands in 1898. The expected number of juveniles decreased to one individual per 100 m² at 43.5 m from the mother tree. Conclusions: The pre‐industrialization landscape greatly affected recovery of Fagus forest. Forests found on the 1898 vegetation map might have acted as refugia for Fagus. The limited dispersal ability of Fagus suggests that it would take many generations (several hundred years) for Fagus forests to recover at the centre of what had been grasslands in the 19th century.     

Genetic diversity and structuring of the grey wolf population from the Central Balkans based on mitochondrial DNA variation

The Dinaric-Balkan grey wolf population used to be at a border between the large remaining Eastern European populations and the largely eradicated Western European populations. During the last few decades we have witnessed the Western European wolf population recovery. Substantial genetic variation has previously been reported in the Balkan wolf population, but rigorous genetic characterization has not been done for its central parts. The aims of this research were to determine genetic diversity based on mtDNA sequence variability, to infer possible population structuring, to find genetic signals of population expansions or bottlenecks and to evaluate phylogenetic position of the grey wolf population from the Central Balkans. Six haplotypes were detected, of which three have only been found in the Balkan region. These haplotypes belong to both haplogroups previously determined in Europe. Based on our mtDNA sequence analyses, the Dinaric-Balkan wolf population is vertically differentiated into “western” (Croatia/Bosnia and Herzegovina) and “eastern” (Serbia/Macedonia) subpopulations. None of the results support assumption of population expansion. Instead, significantly positive values for Tajima's D and Fu's Fs may suggest recent population bottleneck. Obtained data may be helpful in observation to which extent gene pool from the Balkans contribute to newly founded populations in Western Europe.     

Phylogeography of the brown hare (Lepus europaeus) in Europe: a legacy of south‐eastern Mediterranean refugia?

Aim We analysed the population genetics of the brown hare (Lepus europaeus) in order to test the hypothesis that this species migrated into central Europe from a number of late glacial refugia, including some in Asia Minor. Location Thirty‐three localities in Greece, Bulgaria, Italy, Croatia, Serbia, Poland, Switzerland, Austria, France, Germany, the Netherlands, Spain, the United Kingdom, Turkey and Israel. Methods In total, 926 brown hares were analysed for mitochondrial DNA (mtDNA) variation by restriction fragment length polymorphism (RFLP) performed on polymerase chain reaction‐amplified products spanning cytochrome b (cyt b)/control region (CR), cytochrome oxidase I (COI) and 12S–16S rRNA. In addition, sequence analysis of the mtDNA CR‐I region was performed on 69 individuals, and the data were compared with 137 mtDNA CR‐I sequences retrieved from GenBank. Results The 112 haplotypes detected were partitioned into five phylogeographically well‐defined major haplogroups, namely the ‘south‐eastern European type haplogroup’ (SEEh), ‘Anatolian/Middle Eastern type haplogroup’ (AMh), ‘European type haplogroup, subgroup A’ (EUh‐A), ‘European type haplogroup, subgroup B’ (EUh‐B) and ‘Intermediate haplogroup’ (INTERh). Sequence data retrieved from GenBank were consistent with the haplogroups determined in this study. In Bulgaria and north‐eastern Greece numerous haplotypes of all five haplogroups were present, forming a large overlap zone. Main conclusions The mtDNA results allow us to infer post‐glacial colonization of large parts of Europe from a late glacial/early Holocene source population in the central or south‐central Balkans. The presence of Anatolian/Middle Eastern haplotypes in the large overlap zone in Bulgaria and north‐eastern Greece reveals gene flow from Anatolia to Europe across the late Pleistocene Bosporus land‐bridge. Although various restocking operations could be partly responsible for the presence of unexpected haplotypes in certain areas, we nevertheless trace a strong phylogeographic signal throughout all regions under study. Throughout Europe, mtDNA results indicate that brown hares are not separated into discernable phyletic groups.     

Short‐term changes affecting regeneration of Fagus crenata after the simultaneous death of Sasa kurilensis

Abstract. Question: The aim of the present study is to determine whether seed/seedling predation will increase and Fagus survival will decline with the recovery of the Sasa cover. Methods: We examined Fagus crenata regeneration for seven years in an old‐growth Fagus‐Sasa forest near Lake Towada, northern Japan, by examining the effects of simultaneous death of Sasa, tree canopy gap formation, mast seeding of Fagus and seed and seedling predation by rodents on the survival of Fagus seeds and current year seedlings. We established four types of sites differing in forest canopy (closed or gap) and Sasa status (dead or alive) following the simultaneous flowering and death of Sasa kurilensis (dwarf bamboo) in 1995. Results: Fallen Fagus seed was abundant in 1997 and 2000 (mast years). In sites with alive Sasa, survival from the first growing season was low due to high seed and seedling predation. In sites with dead Sasa, seed survival under the canopy was high for both mast years, but in gaps it varied between years. Seedling survival was highest in canopy gaps with dead Sasa (gap‐dead) in 1998, because of higher light levels and lower predation by rodents. However, seedling survival in these plots was low in 2001, apparently because rapid Sasa recovery favoured rodent predation. In both mast years, Sasa die‐back had significant positive effects on seed and seedling survival under closed canopies because the seedlings there were more successful in escaping predation. Conclusion: The change in successful sites for the early stage of regeneration of Fagus appears to reflect the combined effects of canopy gap, seed/seedling predation and revegetation of Sasa.     

Phylogenetics and biogeography of the Balkan ‘sand gobies’ (Teleostei: Gobiidae): vulnerable species in need of taxonomic revision

Within the Atlantic–Mediterranean region, the ‘sand gobies’ are abundant and widespread, and play an important role in marine, brackish, and freshwater ecosystems. They include the smallest European freshwater fish, Economidichthys trichonis, which is threatened by habitat loss and pollution, as are several other sand gobies. Key to good conservation management is an accurate account of the number of evolutionary significant units. Nevertheless, many taxonomic and evolutionary questions remain unresolved within the clade, and molecular studies are lacking, especially in the Balkans. Using partial 12S and 16S mitochondrial ribosomal DNA sequences of 96 specimens of at least eight nominal species (both freshwater and marine populations), we assess species relationships and compare molecular and morphological data. The results obtained do not support the monophyly of Economidichthys, suggesting the perianal organ to be a shared adaptation to hole‐brooding rather than a synapomorphy, and urge for a taxonomic revision of Knipowitschia. The recently described Knipowitschia montenegrina seems to belong to a separate South‐East Adriatic lineage. Knipowitschia milleri, an alleged endemic of the Acheron River, and Knipowitschia cf. panizzae, are shown to be very closely related to other western Greek Knipowitschia populations, and appear conspecific. A distinct Macedonian–Thessalian lineage is formed by Knipowitschia thessala, whereas Knipowitschia caucasica appears as an eastern lineage, with populations in Thrace and the Aegean. The present study combines the phylogeny of a goby radiation with insights on the historical biogeography of the eastern Mediterranean, and identifies evolutionary units meriting conservation attention. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 73–91.     

COMPARATIVE PALEOBIOCHEMISTRY OF SOME FOSSIL AND EXTANT FAGACEAE

A Fagus-like leaf fossil (cuticular compression) with an attached fruit, differing from any known Fagus species (fossil or extant) or other fagoid taxa, has been discovered from the Miocene Clarkia Lake deposits of northern Idaho. Because of its unusual morphology (especially the fruit) the fossil taxon has been described as a new genus and species, Pseudofagus idahoensis Smiley and Huggins. The successful previous use of paleobiochemistry in studies of fossil taxa from the Miocene Succor Creek Flora of Oregon suggested that chemical data might help clarify the taxonomic affinities of Pseudofagus. Indeed, examination of the chemistry of the fossil, Pseudofagus idahoensis, and comparison with extant Fagus species and related fagoid genera indicate that: 1) based on steroid chemistry, Pseudofagus idahonesis does belong in the Fagaceae; 2) like all extant species of Fagus, the fossil lacks the tannin component, ellagic acid, which separates it from other extant fagoid genera, and 3) its simple flavonoid pigment profile places it closest to the extant North American Fagus grandifolia or the European/Eurasian Fagus sylvatica. However, the exclusive presence of an isorhamnetin (3'-methoxyquercetin) 3-0-glycoside, onocerane, and 5α-cholestane imparts a species-specific chemical character to Pseudofagus idahoensis, which also sets it apart from extant species of Fagus. While the chemistry does not decide the taxonomic level to be accorded to the fossil, it certainly supports, along with morphology and anatomy, the distinctness of Pseudofagus and its proposed relationships within the Fagaceae.