Patterns and traits of the endemic plants of Greece

Greece is characterized by high plant diversity (5800 species) and endemism (15.6%). This study attempts a first overall assessment of the taxonomy, distribution, traits and conservation status of the Greek endemic plants. The endemic species belong to 56 families and 242 genera. Most of the endemic plants have a narrow geographical and altitudinal distribution range. The southern floristic regions are richer in endemic species. The species area relationships for endemics (EARs) for island and continental floristic regions explain over 50% of the variation in number of species and are characterized by steep curves. Analysis of the distributional pattern of the endemics by similarity coefficients offers useful insights into the palaeogeography and biogeography of Greece. The endemic species occur at all altitudes, but the altitudinal distribution shows a predominance of local endemics at 0–600 m in the island regions and in higher zones in the continental regions. The life form spectra show a predominance of hemicryptophytes and chamaephytes. This trait seems indicative of their habitat and adaptive strategy and may be related to speciation processes. The overview of the conservation status of the Greek endemics indicates that over 40% of the taxa are threatened or near threatened. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 130–422.     

Species diversity,endemism and conservation of the family Caryophyllaceae in Greece

The family Caryophyllaceae includes world-wide 86 genera and approximately 2100 species. Greece is one of its most important centres of diversity and endemism. A total of 428 Caryophyllaceae taxa are distributed in Greece, 161 of them being endemic to the Greek political territory. The endemic element represents approximately 5% of the global diversity of the family at the species level. The aim of this paper is to discuss the distribution patterns of the Greek endemic Caryophyllaceae, as well as those with a limited distribution range to the neighbouring areas of the Balkans and Anatolia, on the basis of the phytogeographical regions of Greece in order to identify the important regions for their conservation. The majority of the Greek endemic Caryophyllaceae (64.6%) are distributed in only one single phytogeographical region, or even a smaller area indicating the extremely restricted distribution ranges of the endemic plants in Greece. Actually 83 Greek endemic Caryophyllaceae can be grouped on cytotaxonomic criteria. Most of them belong to the category of schizoendemics (91.6%), indicating that the endemism of Caryophyllaceae in Greece has mainly originated in an active way. Cluster analysis has been used to classify the phytogeographical regions according to their floristic similarities. Two iterative complementarity methods were used to evaluate the importance of each phytogeographical region in the conservation of the endemic Caryophyllaceae in Greece. Peloponnisos, Kriti-Karpathos and Sterea Ellas are the most important phytogeographical regions in this respect, followed by North Central and North East. When adding the Balkan-Aegean-Anatolian endemics to the analysis, Peloponnisos, North Central, Kriti-Karpathos, North East and Eastern Aegean result as the most important areas. In every case, an elevated number of sites are required for the conservation of Caryophyllaceae in Greece, reflecting the great dissimilarities in the floristic composition of the various phytogeographical regions. The results provided by the different methods are compared. A catalogue of the Greek endemic Caryophyllaceae is appended.     

Micro-evolution in the weevil genus Larinus: the formation of host biotypes and speciation

Data are presented on allozyme variation between 15 populations of the stenophagous capitulum weevil, Larinus cynarae , and three populations of its congener, L. latus , that had been collected throughout the northern mediterranean range of these species. A phenetic analysis of these data revealed no direct relationship between genetic variation and host-plant association within L. cynarae , but there was a strong geographical structuring of allozyme patterns. Most of the genetic variation was due to differences between geographical regions and variation within these was small. Wright's F _ST values showed that Italian and Greek populations of L. cynarae were most distinct from L. latus , with southern Iberian, northern Spanish and French populations increasingly less so. This pattern was associated with a cline in the frequencies of certain alleles along this geographical arc from France to Greece. A phenogram of Nei's genetic distances indicated the close genetic relationship between the two species of Larinus and separated the populations of L. cynarae into three allopatric groups. These groups have different host-plant spectra - dominated by Cynara cardunculus in Italy and Greece, Cynara humilis/Onopordum in southern Iberia and Onopordum spp. in France/Northern Spain - and can be considered to be host biotypes of L. cynarae. L. latus , which occurs in Greece and further east is also an Onopordum specialist. An analysis of the phylogeny of this group of Larinus indicates a primary separation into eastern ( L. latus ) and western ( L. cynarae ) taxa, with further branching of the L. cynarae lineage into the putative host-biotypes. An hypothesis for the evolution of these taxa is given, based on the evolutionary history of host-plant taxa and geographical constraints.     

Phylogeography and host race differentiation in the European mistletoe (Viscum album L.)

Mistletoes are bird dispersed, hemi-parasitic shrubs infecting a large number of woody host plants. Chloroplast fragment length polymorphisms were used to study genetic differentiation among presumed host races, population genetic structure, and to elucidate the postglacial migration history of mistletoe ( Viscum album ) across the entire natural distribution range in Europe. The populations sampled belong to four closely related taxa, three of which are widely distributed and differ in their host trees, whereas a fourth taxon is rare and endemic to the Island of Crete. The molecular analysis of chloroplast DNA variation supported the distinction of these four taxa. We further found evidence for phylogeographical structure in each of the three widely distributed host races. Independent of host race, mistletoe haplotypes from Turkey were distinct and distant from those found elsewhere in Europe, suggesting that highly differentiated populations, and possibly new taxa, exist at the range limit of the species.     

Molecular Systematics of Threatened Seed Plant Species Endemic in the Caribbean Islands

A review of available Caribbean Island red-lists species (CR and EN categories based on the IUCN guidelines from 2001, and E category established according to the IUCN guidelines from 1980) is presented. A database of over 1,300 endemic species that are either Critically Endangered or Endangered sensu IUCN was created. There are molecular systematic studies available for 112 of them. Six of these species (in six genera) are the only members of early divergent lineages that are sister to groups composed of a large number of clades. Seven of the species (in seven genera) belong to clades that have a small number of taxa but are sister to species/genus-rich clades. Ten of the species (in six genera) are sister to taxa restricted to South America or nested in clades endemic to this region. Fifty-seven of the species (in 35 genera) are sister to Caribbean Island endemic species. Erigeron belliastroides, an Endangered (EN) Cuban endemic, is sister to the Galapagos genus Darwiniothamnus. The phylogenetic placement of four of the threatened species resulted in changes in their taxonomic placement; they belong to polyphyletic or paraphyletic genera.     

The North Mediterranean Barbus lineage: phylogenetic hypotheses and taxonomic implications based on allozyme data

On the basis of distinct ecological and morphological characters, the European Barbus taxa have been clustered in two groups: a fluvio-lacustrine and a rheophilic or strictly riverine one. These two groups (or ecophenotypes) were recognized in different parts of Europe, and formed either a species assemblage ( Barbus barbus group) or a polytypic species ( Barbus meridionalis ). The hypothesis was that species of the same group belong to the same phylogenetic lineage (clade) and are the result of the same transcontinental colonization event. The analysis, using allozyme markers, of 10 taxa of the genus Barbus from France, Italy, Greece, the Czech Republic and Slovakia, showed that the taxa thought to belong to the fluvio-lacustrine and the rheophilic groups are not monophyletic. The results suggest that probably in each sub-region, the founding taxon has diverged independently to form species of two different ecophenotypes, one occupying the upstream rivers and the other the lowland rivers. Accordingly, Barbus species groups represent clusters of morphologically convergent taxa living in equivalent biotopes.     

Genetic differentiation and natural hybridization between the Sardinian endemic Maniola nurag and the European Maniola jurtina

The Mediterranean island of Sardinia is known for its multitude of unique genetic lineages. We view one of them in a larger phylogeographic context. The endemic Sardinian Meadow Brown butterfly, Maniola nurag, is restricted to the mountainous areas of the island, whereas its widespread close relative, Maniola jurtina, also occurs on the coast. At intermediate altitudes the species' distributions overlap. There, a number of individuals exhibit phenotypic characteristics intermediate between the two species. We examined patterns of intra- and interpopulation variation in 10 M. nurag populations from Sardinia and 16 M. jurtina populations from Sardinia and continental Europe, as well as 17 intermediate individuals, sampled in 1999-2002, by means of allozyme markers, combining it with a morphometric analysis based on 18 wing-characters of 52 males. At the 15 loci studied (aldolase, aat-1, aat-2, g6pdh, gpd, idh-1, idh-2, mdh-1, mdh-2, mpi, me, leu-ala, pgi, pgm, and 6pgdh), 76 different alleles were detected, 63 of which were shared by M. nurag and M. jurtina. None of the loci was found to be alternatively fixed between the two species. In that respect, this study testifies to the difficulties that may arise when trying to identify hybrids from genotypic data. Levels of genetic variation in island populations (M. jurtina: H(O) = 0.137-0.189; M. nurag: H(O) = 0.141-0.270) were comparable to those of mainland M. jurtina (H(O) = 0.141-0.236). A Bayesian admixture analysis supported the hypothesis of mixed (hybrid) ancestry of individuals occurring at intermediate altitudes. Similarly, neighbour-joining and unweighted pair-group method with arithmetic averaging (UPGMA) analyses, as well as morphometrics hinted at the existence of a Maniola-hybrid zone in Sardinia at intermediate altitudes. We discuss the results in the light of the phylogeography of other Sardinian taxa with the aim to reach a general understanding of the biogeographic history of this island's endemic species.     

A new elephant fossil from the late Pleistocene of Alghero: The puzzling question of Sardinian dwarf elephants

Endemic elephants, variously reduced in size, have been reported from a number of Mediterranean islands. Most of these originated from the mainland species Palaeoloxodon antiquus. A few dwarf mammoth remains are recorded from Crete and Sardinia. In Sardinia, a largely incomplete skeleton and a few mammoth teeth have been reported from localities believed to range in age from the late middle to the late Pleistocene. The chronology of colonisation by the ancestral species, the actual persistence through time of Mammuthus lamarmorai on the island, and the morphological and dimensional range of the species are, however, poorly known. This research aims to describe a distal portion of a left tibia of a dwarf elephant found in the Alghero area (NW Sardinia), showing some morphological traits and dimensions consistent with those of the endemic Sardinian mammoth (Mammuthus lamarmorai). The main unanswered questions about chronology, colonisation and population dynamics of endemic Sardinian elephants are highlighted and briefly discussed.     

The endemic bovids from Sardinia and the Balearic Islands: State of the art

Bovids are not so common in endemic insular faunas and are mainly recorded in Southeast Asia, Japan and some Mediterranean islands. In the Western Mediterranean, endemic bovids have been recorded during the late Miocene in the Tusco-Sardinian palaeobioprovince (Baccinello-Cinigiano basin, South Tuscany, and Fiume Santo, north-western Sardinia). In the latest Neogene and Quaternary, bovids showing highly endemic features were restricted to the Balearic Islands and Sardinia, while Bovini only slightly reduced in size were present on Pianosa, Malta and Sicily. On Sardinia, the richest bovid sample comes from Monte Tuttavista (Orosei), where at least three species have been identified: Asoletragus genthry, Nesogoral aff. N. melonii, and Nesogoral sp. 2. On Mallorca (Balearic Islands) six chronospecies belonging to the Myotragus endemic phylogenetic lineage have been described, spreading in age from the Early Pliocene to the Holocene. For decades, a close phylogenetic relationship between Nesogoral and Myotragus has been widely accepted by scholars. Morphological and biometrical differences shown by Balearic and Sardinian bovids have generally been regarded as the result of the evolution into two different island ecological systems, characterized by different inter and intra-guild selection pressures. Indeed, the more diversified environment of Sardinia, as well as the presence of other large mammals (similar-sized competitors belonging to the same guild and a running predator), increased the interspecific competition, forcing Sardinian bovids to exploit different resources and to occupy different niches, while Myotragus exploited under a monopoly regime the supply of resources available for large herbivores on the Eastern Balearic Islands. Nonetheless, new data suggest that Nesogoral and Myotragus possibly originated from different taxa.     

无量山种子植物区系的特有现象

地处滇中南的无量山地区迄今计有种子植物２０７科,１０２６属,２５４０种,其中东亚特有科１０个,中国特有属２７个,无量山特有种子６７个,这一地区不仅植物种类丰富,而且特有现象较为显著,在对科级持有现象进行较细致的研究后,本文认国无量山具有较多的分类系统上隔离的东亚特有科反映了该地作为东亚古老植物区系的一部分,其地质历史与整个东亚是一致的,且与东亚植物区系的发端密切相关,属的特有现象表明,地处中国特有     

The endemic Cladophorales (Ulvophyceae) of ancient Lake Baikal represent a monophyletic group of very closely related but morphologically diverse species

Lake Baikal, the oldest lake in the world, is home to spectacular biodiversity and extraordinary levels of endemism. While many of the animal species flocks from Lake Baikal are famous examples of evolutionary radiations, the lake also includes a wide diversity of endemic algae that are not well investigated with regards to molecular‐biological taxonomy and phylogeny. The endemic taxa of the green algal order Cladophorales show a range of divergent morphologies that led to their classification in four genera in two families. We sequenced partial large‐ and small‐subunit rDNA as well as the internal transcribed spacer region of 14 of the 16 described endemic taxa to clarify their phylogenetic relationships. One endemic morphospecies, Cladophora kusnetzowii, was shown to be conspecific with the widespread Aegagropila linnaei. All other endemic morphospecies formed a monophyletic group nested within the genus Rhizoclonium (Cladophoraceae), a very surprising result, in stark contrast to their morphological affinities. The Baikal clade represents a species flock of closely related taxa with very low genetic differentiation. Some of the morphospecies were congruent with lineages recovered in the phylogenies, but due to the low phylogenetic signal in the rDNA sequences the relationships within the Baikal clade were not all well resolved. The Baikal clade appears to represent a recent radiation, based on the low molecular divergence within the group, and it is hypothesized that the large morphological variation results from diversification in sympatry from a common ancestor in Lake Baikal.     

Karyotype variation, evolution and phylogeny in Borago (Boraginaceae), with emphasis on subgenus Buglossites in the Corso-Sardinian system

BACKGROUND AND AIMS: Karyological variation in the Mediterranean genus Borago and cytogeography of subgenus Buglossites in Corsica, Sardinia and the Tuscan Archipelago were investigated in combination with a molecular phylogenetic analysis aimed at elucidating relationships between subgenera and taxa with different chromosome features. METHODS: Karyotype analysis was performed on population samples of B. pygmaea, B. morisiana, B. trabutii and B. officinalis. Phylogenetic analyses were based on ITS1 nrDNA and matK cpDNA sequences. KEY RESULTS: Four base numbers were found, x = 6, 8, 9 and 15, and three ploidy levels based on x = 8. In subgenus Buglossites the Sardinian endemic B. morisiana is diploid with 2n = 18, while B. pygmaea includes three allopatric cytotypes with 2n = 30 (Sardinia), 2n = 32 (southern Corsica) and 2n = 48 (central northern Corsica and Capraia). In subgenus Borago, the Moroccan endemic B. trabutii and the widespread B. officinalis have 2n = 12 and 2n = 16, respectively. Molecular data support the monophyly of Borago, while relationships in subgenus Borago remain unclear. Borago trabutii appears as the earliest divergent lineage and is sister to a clade with B. officinalis, B. morisiana and B. pygmaea. Subgenus Buglossites is also monophyletic, but no correspondence between ITS1 phylogeny and B. pygmaea cytotypes occurs. CONCLUSIONS: Chromosome variation in Borago is wider than previously known. Two base numbers may represent the ancestral condition in this small genus, x = 6 or x = 8. An increase in chromosome number and karyotype asymmetry, a decrease in chromosome size and heterochromatin content, and the appearance of polyploidy are the most significant karyological changes associated with the divergence of the Buglossites clade. High ITS1 variation in the tetra- and hypotetraploid races of B. pygmaea suggests a multiple origin, while the lower polymorphism of the hexaploid race and its allopatric distribution in the northernmost part of the range is better explained with a single origin via union of unreduced and reduced gametes.     

Levels of endemism are not necessarily biased by the co-presence of species with different range sizes: a case study of Vilenkin and Chikatunov's models

Aim  To illustrate problems in the methods proposed by B. Vilenkin and V. Chikatunov to study levels of endemism and species–area relationships.
Location  The study used data on the distribution of tenebrionid beetles (Coleoptera, Tenebrionidae) on the Aegean Islands (Greece).
Methods  A total of 32 islands and 170 taxa (species and subspecies) were included in this study. Levels of endemism were evaluated both as the proportion of endemic taxa, and according to the methods proposed by Vilenkin and Chikatunov, which are based on the number of non-endemic taxa and various relationships with area. A model of the species–area relationship proposed by these authors was also analysed.
Results  The number of endemic taxa was positively correlated with the number of taxa with different distribution types, but this positive correlation did not influence the estimation of the level of endemism. In fact, the commonly used estimate of endemicity as a percentage was strongly correlated with the endemism values calculated according to the method of Vilenkin and Chikatunov. The usual power function fitted the species–area relationship as well as the most complicated method of Vilenkin and Chikatunov.
Main conclusions  As hypothesized by Vilenkin and Chikatunov, the number of endemic taxa was influenced both by the number of taxa of other biogeographical ranks, and by an island's area. However, explanations for the positive relationship between the number of endemic taxa and taxa of different biogeographical ranks are equivocal. Importantly, this relationship did not necessarily influence the level of endemism, which could be expressed adequately by percentages. The method proposed by Vilenkin and Chikatunov to estimate the species–area relationship cannot be clearly justified on theoretical grounds and is of questionable practical utility.     

Echium anchusoides (Boraginaceae), a new species from Sardinia (Italy)

The new species Echium anchusoides , endemic to the main siliceous massifs of Sardinia, is described and illustrated. It was formerly confused with E. pustulatum , a Mediterranean form of E. vulgare , but differs in characters regarding habit, leaves, inflorescence, corolla, stamens and mericarps. Contrary to the taxa of the E. vulgare group, E. anchusoides is also characterized by a higher ecological specificity, being linked to natural habitats with siliceous growth substratum. Micromorphological analyses provide additional information on the indumentum and reproductive structures, such as pollen, stigmatic papillae, nectar ducts and mericarp coat surface.     

The endemic vascular flora of Peloritani Mountains (NE Sicily): Plant functional traits and phytogeographical relationships in the most isolated and fragmentary micro-plate of the Alpine orogeny

This study is aimed at (1) producing a complete and updated inventory of the endemic vascular flora of Peloritani Mountains, (2) defining the geographical limits of Peloritani, regarded here as a biogeographical district and (3) highlighting possible paleogeographic connections with other Mediterranean lands. The heterogeneity analysis of the endemic flora was performed by means of contingency tables, through the χ² test. The endemic flora of this area consists of 129 specific and infraspecific taxa, of which 15 are restricted to the Peloritani Mountains. The analysis of habitats revealed that endemic taxa are most abundant on cliffs, rangelands, woods and garrigues. A large number of surveyed taxa are endemic to central-southern Italy and Sicily, while the number of endemic taxa in common with Calabria, Etna and Aeolian Islands turned out to be rather low, in spite of the geographical proximity. The endemic flora of Peloritani allows to emphasize palaeogeographical relationships not only with the neighbouring Mediterranean territories, but also with currently remote ones, such as southern Spain, Sardinia and Corsica. The phytogeographical framework substantiates the hypothesis that the Peloritani floristic district coincides with the limit given to Peloritani Mountains by structural geologists.     

Morphological and molecular characterization of two novel species of Agaricus section Xanthodermatei

Agaricus specimens collected in France belong to two novel entities resembling small forms of A. moelleri and A. xanthodermus, two common species in section Xanthodermatei. Molecular (IT1+ITS2 DNA sequence) and morphological comparisons between eight presumed similar taxa of the section support the elevation of both entities to species rank. The new entities are described as A. parvitigrinus and A. xanthodermulus. They form a group with A. laskibarii, a rare species also recently described from France, and A. californicus, a North-American species. The well known A. moelleri and A. xanthodermus are the most related species among the studied sample. Like other species of the section, both new species have a phenolic odor and are probably toxic.     

Evolutionary patterns and genetic structure in localized and widespread species in the Stylidium caricifolium complex (Stylidiaceae)

The Stylidium caricifolium (Stylidiaceae) complex consists of seven currently recognized species and a taxon of putative hybrid origin endemic to southwest Western Australia. These taxa vary in geographical distribution from widespread, extending over a range of 500 km, to extremely localized, covering a range of only 0.5 km. Patterns of allozyme variation were investigated in 61 populations covering all taxa and two closely related species. Measures of genetic diversity were consistently lower and in some cases significantly lower in four rare and geographically restricted taxa compared with their widespread relatives. In contrast, genetic diversity in two other localized taxa was comparable or higher than in the widespread taxa. The level of divergence among populations was moderate to high, with a significant trend of higher F(ST) values for the widespread species to lower values for the geographically restricted and rare taxa. Phylogenetic relationships and levels of divergence indicate that most taxa are probably relictual rather than recently evolved. Geographical localization and rarity in this complex can be attributed to a range of factors associated with habitat specificity, historical and ecological processes that characterize the southwest region, and mode of origin.     

Taxonomy of the Houbara Bustard Chlamydotis undulata subspecies considered on the basis of sexual display and genetic divergence

The Houbara Bustard Chlamydotis undulata is a polytypic species with three subspecies classified according size, colour and distribution: C. u. undulata, C. u. fuertaventurae and C. u. macqueenii. A comparative study of the male display behaviour of these three subspecies has shown similarities between C. u. undulata and C. u. fuertaventurae , whereas appreciable differences exist in the visual and auditory aspects of the display between C. u. undulata and C. u. macqueenii. These observations make the current taxonomy of Houbara Bustard questionable, particularly in view of the major importance of the sexual display in bird speciation processes. Furthermore, DNA analysis of the cytochrome b gene indicates the existence of a significant differentiation between the C. u. undulata and C. u. macqueenii populations. These findings indicate the need for a revision of the taxonomy of genus Chlamydotis which should separate the macqueenii and undulata populations at species rank. Until further investigations are completed, C. u. fuertaventurae should be kept as a subspecies of undulata since the courtship display is alike in both populations.