Phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs have been determined by geological processes and climate change in the Late Cenozoic

AIM: Our aims were to assess the phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs and to estimate divergence times using different geological scenarios. We related divergence times to past geological events and discuss the relevance of our data for the systematics of eastern Mediterranean water frogs. LOCATION: The eastern Mediterranean region. METHODS: Genetic diversity and divergence were calculated using sequences of two protein-coding mitochondrial (mt) genes: ND2 (1038 bp, 119 sequences) and ND3 (340 bp, 612 sequences). Divergence times were estimated in a Bayesian framework under four geological scenarios representing alternative possible geological histories for the eastern Mediterranean. We then compared the different scenarios using Bayes factors and additional geological data. RESULTS: Extensive genetic diversity in mtDNA divides eastern Mediterranean water frogs into six main haplogroups (MHG). Three MHGs were identified on the Anatolian mainland; the most widespread MHG with the highest diversity is distributed from western Anatolia to the northern shore of the Caspian Sea, including the type locality of Pelophylax ridibundus. The other two Anatolian MHGs are restricted to south-eastern Turkey, occupying localities west and east of the Amanos mountain range. One of the remaining three MHGs is restricted to Cyprus; a second to the Levant; the third was found in the distribution area of European lake frogs (P. ridibundus group), including the Balkans. MAIN CONCLUSIONS: Based on geological evidence and estimates of genetic divergence we hypothesize that the water frogs of Cyprus have been isolated from the Anatolian mainland populations since the end of the Messinian salinity crisis (MSC), i.e. since c. 5.5-5.3 Ma, while our divergence time estimates indicate that the isolation of Crete from the mainland populations (Peloponnese, Anatolia) most likely pre-dates the MSC. The observed rates of divergence imply a time window of c. 1.6-1.1 million years for diversification of the largest Anatolian MHG; divergence between the two other Anatolian MHGs may have begun about 3.0 Ma, apparently as a result of uplift of the Amanos Mountains. Our mtDNA data suggest that the Anatolian water frogs and frogs from Cyprus represent several undescribed species.     

Mitochondrial HVI sequence variation in Anatolian hares (Lepus europaeus Pallas, 1778)

Variability and phylogenetic relationships of sequences of the hypervariable domain I (HVI) of the mitochondrial DNA was studied in 46 brown hares (Lepus europaeus) from Anatolia, to test the hypotheses that (i) hares from several islands off the Anatolian coast and from Cyprus are phylogenetically close to mainland Anatolian hares, (ii) Anatolian hare sequence variability is higher than that of typical European brown hares, and to iii) infer possible Anatolian source populations of hares from some islands in the eastern Mediterranean. Neighbor joining and Maximum Parsimony analyses revealed reciprocal monophyly for sequences from Anatolia, the considered eastern Mediterranean islands off the Anatolian coast, Cyprus, and those sequences published earlier form NE Greece that were supposed to originate from earlier immigration via the late-Pleistocene/early-Holocene land bridge that connected SE Europe and W Anatolia (Kasapidis et al., 2005. Mol. Phylogenet. Evol. 34, 55–66). A high sequence idiosyncrasy was found among the Anatolian samples. Almost all approaches to compare variability between Anatolian and the downloaded European sequence data indicated higher sequence diversity in Anatolia, in accordance with earlier findings for allozyme loci. Network and principal coordinate analyses of the Anatolian sequences and those from the islands off the Anatolian coast as well as the Anatolian-type NE Greek sequences suggested high mitochondrial gene exchange among local populations in Anatolia with little effect of possible geographic barriers, and did not provide clues for tracing possible origins of island populations.     

A synopsis of <Emphasis Type="Italic">Bracon</Emphasis> species of Turkey with description of a new species (Hymenoptera: Braconidae: Braconinae)

Our studies on Bracon F. fauna of Turkey started in 1979 and 107 species have been determined so far belonging to the subgenera Habrobracon Ashmead, Asiabracon Tobias, Rostrobracon Tobias, Bracon Fahringer, Cyanopterobracon Tobias, Glabrobracon Fahringer and Lucobracon Fahringer. Thirteen new Bracon species were published from Turkey. With this present study Bracon (Lucobracon) achterbergi Beyarslan sp. n. is described and its diagnostic characters are illustrated. The distribution of Bracon species are discussed according to the Euxin, Subeuxin, Mediterranean, Xeroeuxin, Iran Steppe, Anatolia Steppe, Central Anatolia Steppe and Mesopotamian Steppe phytogeographical provinces of Turkey.     

The importance of Anatolian mountains as the cradle of global diversity in Arabis alpina, a key arctic-alpine species

Background and Aims

Anatolia is a biologically diverse, but phylogeographically under-explored region. It is described as either a centre of origin and long-term Pleistocene refugium, or as a centre for genetic amalgamation, fed from distinct neighbouring refugia. These contrasting hypotheses are tested through a global phylogeographic analysis of the arctic–alpine herb, Arabis alpina.

Methods

Herbarium and field collections were used to sample comprehensively the entire global range, with special focus on Anatolia and Levant. Sequence variation in the chloroplast DNA trnL-trnF region was examined in 483 accessions. A haplotype genealogy was constructed and phylogeographic methods, demographic analysis and divergence time estimations were used to identify the centres of diversity and to infer colonization history.

Key Results

Fifty-seven haplotypes were recovered, belonging to three haplogroups with non-overlapping distributions in (1) North America/Europe/northern Africa, (2) the Caucuses/Iranian Plateau/Arabian Peninsula and (3) Ethiopia–eastern Africa. All haplogroups occur within Anatolia, and all intermediate haplotypes linking the three haplogroups are endemic to central Anatolia and Levant, where haplotypic and nucleotide diversities exceeded all other regions. The local pattern of haplotype distribution strongly resembles the global pattern, and the haplotypes began to diverge approx. 2·7 Mya, coinciding with the climate cooling of the early Middle Pleistocene.

Conclusions

The phylogeographic structure of Arabis alpina is consistent with Anatolia being the cradle of origin for global genetic diversification. The highly structured landscape in combination with the Pleistocene climate fluctuations has created a network of mountain refugia and the accumulation of spatially arranged genotypes. This local Pleistocene population history has subsequently left a genetic imprint at the global scale, through four range expansions from the Anatolian diversity centre into Europe, the Near East, Arabia and Africa. Hence this study also illustrates the importance of sampling and scaling effects when translating global from local diversity patterns during phylogeographic analyses.     

Phylogeography of Troglophilus (Orthoptera: Troglophilinae) based on Anatolian members of the genus: radiation of an old lineage following the Messinian

Active tectonic history of the Eastern Mediterranean, especially around Aegean area, through the Neogene led to interesting radiation patterns of animal lineages, allowing intriguing biogeographical hypotheses to be tested. Descendants of the ancestral stock in the Miocene Aegean Plate presently occur in the Anatolia, Aegean islands and the Balkan Penninsula. Troglophilus (Orthoptera, Rhaphidophoridae, Troglophilinae) is such a genus represented in these areas with approximately 15 species. The present study addresses the phylogeography of the genus, with a special emphasis on its Anatolian members, aiming to test the biogeographical patterns suggested for this area using mitochondrial [cytochome oxidase I (COI)] and nuclear (ITS1‐5.8S–ITS2) markers. Data matrices obtained from sequences of COI and ITS1‐5.8S–ITS2 were used for phylogenetic analyses using Dolichopoda lycia and Dolichopoda sbordonii as outgroups. All sets of the analyses suggested monophyly of the Anatolian haplotypes, although they are not congruent in revealing their relationships. Anatolian haplotypes constituted three main phylogroups in trees calculated from a matrix of short COI sequences: the ECMA (corresponding to the Eastern part of coastal Mediterranean Anatolia); the CWMA (from the Central and Western part of Mediterranean Anatolia); and NA (from Northern Anatolia). Trees obtained using longer sequences resulted in only two phylogroups, namely ECMA and CWMA + NA. The trees based on the ITS1‐5.8S–ITS2 data matrix supported monophyly of Anatolian phylogroups. BEAST analysis of the COI estimated the time to most recent common ancestor for Dolichopoda and Troglophilus as 10.8 Mya, to that for the Anatolian + Balkan Troglophilus as 7.2 Mya, and to that for the Anatolian Troglophilus as 6.3 Mya. BEAST analysis of ITS1–ITS2 intron regions is largely congruent with that of COI. From these results, several conclusions were drawn. First, the divergence of Dolichopoda and Troglophilus possibly started with the opening of the Mid‐Aegean Trench in the Tortonian. Second, Troglophilus possibly originated from an ancestral stock in the old Aegean Plate. It later diverged as Anatolian and Balkan lineages and, possibly, the Cretan population may be regarded the third lineage. Divergence within the Anatolian lineage is estimated to have occurred through the Pliocene and Pleistocene, although before the last four glacial periods in the late Pleistocene. Additionally, the northern Anatolian Troglophilus appears to originate from the dispersal of an ancestral stock from a mountainous lineage through the Taurus Way. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 335–348.     

Spatial patterns of the Iberian and Balearic endemic vascular flora

The Iberian flora has a high degree of originality (1328 endemic species, 24% of endemism), comparable to other regions in the Mediterranean Basin. The richness of Iberian endemic species is unevenly distributed; the greatest diversity is found in the main mountain ranges although the southwestern Atlantic coast and specially the Balearic Islands are rich in range-restricted endemic species. The largest number of endemic genera is found in the northwestern mountains, which might have acted as a refugium area. The Baetic System, which includes nearly half (46%) of the total Iberian endemic species, is by far the richest region of the territory. Its endemic flora is characterized by the great richness of narrow endemics and the high species turnover rate. The k-means partitioning analysis enables us to identify 11 units, generally well defined by the natural geographic features. The clusters including the northwestern mountains, the Cantabrian Mountains, the southwestern coast and especially the Balearic Islands, the Pyrenees and the Baetic System are compact and consist of a high proportion of diagnostic species, and can therefore be considered areas of endemism on a large scale. The regionalization reflects a primary longitudinal division of Iberia between a basic eastern and an acidic western region, but also partly reveals a climatic division between Eurosiberian and Mediterranean regions. Southeastern Iberia seems to be an important center of differentiation for several typically Mediterranean genera (e.g. Centaurea, Linaria, Armeria, Teucrium and Thymus), but other large genera are also highly diversified.     

Historical biogeography and mitogenomics of two endemic Mediterranean gorgonians (Holaxonia,Plexauridae)

Among the Mediterranean plexaurids, four species are endemic and despite their ecological importance, comprehensive studies on the evolution and biogeography of these organisms are lacking. Here, we explore the mitogenomic variability of two endemic, ecologically important Mediterranean Paramuricea species. We assess their phylogenetic relationships and provide first insights into their evolution and biogeography. Complete mitogenome sequences of Paramuricea clavata and Paramuricea macrospina were obtained using long-range PCR, primer-walking and Sanger sequencing. For an enlarged sample of Paramuricea species, maximum likelihood and Bayesian phylogenetic trees of the mitochondrial gene mtMutS were obtained and used to study the biogeographic history of Paramuricea through a statistical Dispersal-Vicariance (S-DIVA) method and a Dispersal Extinction Cladogenesis (DEC) model. Divergence time was estimated under strict and relaxed molecular clock models in BEAST using published octocoral mutation rates. Our results revealed high nucleotide diversity (2.6%) among the two Mediterranean endemics; the highest mutation rates were found in the mtMutS, Nad4 and Nad5. In addition, we found length polymorphisms in several intergenic regions and differences in mitochondrial genome size. The red gorgonian P. clavata was closely related to the Eastern Atlantic Paramuricea grayi rather than its Mediterranean congener, P. macrospina. Our biogeographic results provide evidence for the independent speciation of the Mediterranean species and point to a Miocene origin of the two endemics, highlighting the role played by the Messinian Salinity Crisis in the evolutionary history of Mediterranean organisms.     

Phylogeographic analysis of Anatolian bats highlights the importance of the region for preserving the Chiropteran mitochondrial genetic diversity in the Western Palaearctic

Identification of intraspecific conservation units and incorporating the distribution of genetic diversity into management plans are crucial requirements for assessing effective protection strategies. This study investigates the phylogeographic structures of 33 bat species present in the Near East in order to evaluate the conservation implications of their intraspecific genetic diversity both at regional and large-scale levels. To compare Anatolian populations with the European ones, we utilized two commonly used mitochondrial markers, Cytb and ND1, and analysed them together with the available sequences from GenBank. The management requirements of the identified clades and their taxonomical relations were evaluated by analysing their distributions and the levels of their genetic differentiations. In 12 species and the large Myotis complex, we identified a total of 15 genetically distinct populations found in the Near East, some of which might represent biologically distinct taxa. Comparing the phylogeographic patterns of different taxa indicates that three regions, the Balkans, the Caucasus, and the southern Anatolia, harbour genetically divergent populations and should have higher priority in conservation practices. Considering that Turkey has one of the richest bat fauna in the Mediterranean region and the Anatolian populations of various species are genetically distinct, protecting populations in Turkey is critically important for preserving the genetic diversity of the bats in the Western Palaearctic. Both regional and large-scale conservation strategies, which incorporate the distribution of genetic diversity, should be assessed and further ecological studies are needed to clarify the taxonomic relations of the identified clades.     

Genetic diversity in the threatened insular endemic plantAster asa-grayi (Asteraceae)

Genetic diversity in an insular endemic plantAster asa-grayi was examined using enzyme electrophoresis. Distribution ofA. asa-grayi is restricted to only four subtropical islands of Japan, and this species is listed as vulnerable to extinction in the Red Data Book of Japanese wild plants. A total of 161 individuals were sampled from five populations on four islands. Genetic diversity values at the population level were very low, compared to other plant species with a similar life history. Genetic variability at the species level is comparable to the mean value of endemic species. Genetic differentiation among populations is extremely high (G_ST= 0.71), indicating that the gene flow among populations is highly impeded, and pollen and seed dispersal is limited due to the pollinators and the seed morphology. This is because the four islands are geographically isolated. Fixation indices suggested that most populations do not randomly cross. To conserve the genetic diversity of the species, artificial crossings among different island populations are necessary.     

Genetic networking of the Bemisia tabaci cryptic species complex reveals pattern of biological invasions

Background

A challenge within the context of cryptic species is the delimitation of individual species within the complex. Statistical parsimony network analytics offers the opportunity to explore limits in situations where there are insufficient species-specific morphological characters to separate taxa. The results also enable us to explore the spread in taxa that have invaded globally.

Methodology/Principal Findings

Using a 657 bp portion of mitochondrial cytochrome oxidase 1 from 352 unique haplotypes belonging to the Bemisia tabaci cryptic species complex, the analysis revealed 28 networks plus 7 unconnected individual haplotypes. Of the networks, 24 corresponded to the putative species identified using the rule set devised by Dinsdale et al. (2010). Only two species proposed in Dinsdale et al. (2010) departed substantially from the structure suggested by the analysis. The analysis of the two invasive members of the complex, Mediterranean (MED) and Middle East – Asia Minor 1 (MEAM1), showed that in both cases only a small number of haplotypes represent the majority that have spread beyond the home range; one MEAM1 and three MED haplotypes account for >80% of the GenBank records. Israel is a possible source of the globally invasive MEAM1 whereas MED has two possible sources. The first is the eastern Mediterranean which has invaded only the USA, primarily Florida and to a lesser extent California. The second are western Mediterranean haplotypes that have spread to the USA, Asia and South America. The structure for MED supports two home range distributions, a Sub-Saharan range and a Mediterranean range. The MEAM1 network supports the Middle East - Asia Minor region.

Conclusion/Significance

The network analyses show a high level of congruence with the species identified in a previous phylogenetic analysis. The analysis of the two globally invasive members of the complex support the view that global invasion often involve very small portions of the available genetic diversity.     

Threatened fishes of the world: <Emphasis Type="Italic">Pseudophoxinus anatolicus</Emphasis> (Hanko 1924) (Cyprinidae), Central Anatolia,Turkey

Being an endemic and endangered fish species in Anatolia, P. anatolicus restricted to Lakes Akgöl, Bey?ehir, Çavu?cu, Akkaya Dam and Ere?li marshes as well as some other small marshes in Central Anatolia region, Turkey. Current population of P. anatolicus tends to decrease in the habitats. A detailed study of current population status, biology, ecology and life history of P. anatolicus is required. It should also be included into national threatened fish category.     

Molecular phylogeography of Troglophilus cave crickets (Orthoptera,Rhaphidophoridae): A combination of vicariance and dispersal drove diversification in the East Mediterranean region

In this study, we investigated the molecular phylogenetic divergence and historical biogeography of cave crickets belonging to the genus Troglophilus (Orthoptera, Rhaphidophoridae) from caves in eastern Mediterranean and Anatolia regions. Three mitochondrial DNA genes (COI, 12S rDNA, and 16S rDNA) and two nuclear ones (18S rDNA and 28S rDNA) were amplified and partially sequenced to reconstruct phylogenetic relationships among most of the known Troglophilus species. Results showed a well‐resolved phylogeny with three main clades representing the Balkan, the Anatolian, and the Cycladian–Cretan lineages. Based on Bayesian analyses, we applied a relaxed molecular clock model to estimate the divergence times between these three lineages. Dating estimates indicate that radiation of the ingroup might have been triggered by the opening of the Mid‐Aegean trench, while the uplift of the Anatolian Plateau in Turkey and the changes of relief, emergence, and disappearance of orographic and hydrographical barriers in the Balkan Peninsula are potential paleogeographic events responsible for the initial diversification of the genus Troglophilus. A possible biogeographic scenario, reconstructed using S‐DIVA with RASP software, suggested that the current distribution of Troglophilus species can be explained by a combination of both dispersal and vicariance events that occurred in particular in the ancestral populations. The radiation of Troglophilus species likely started from the Aegean and proceeded eastward to Anatolia and westward to the Balkan region. Results are additionally compared to those available for Dolichopoda, the only other representative genus of Rhaphidophoridae present in the Mediterranean area.     

Y DNA and mitochondrial lineages in European and Asian populations of the brown hare (Lepus europaeus)

Both the Cytb gene of mtDNA and Y chromosome markers were studied in a relatively large sample of brown hares (L. europaeus) from Europe and Anatolia (Turkey and Israel), together with other seven Lepus species, in order to enable comparative analysis of possible sex-specific gene flow. Furthermore, Y chromosome markers were compared with data from biparentally inherited markers in an attempt to understand whether or not their pattern of distribution was congruent with that of allozymes or whether they rather matched mtDNA phylogenies, with which they share uniparental inheritance. Consistent with the general observation, levels of interspecific genetic variability were very low for the Y chromosome markers compared with mtDNA. Moreover, lack of interspecific variation for the Y-DNA studied within Lepus genus rendered these markers improper for any further phylogenetic analysis. With the highest nucleotide diversity in Anatolia compared with Europe, both marker systems confirmed an unbroken species history in Anatolia, corroborated the hypothesis of continuous gene flow from Anatolia's neighbouring regions, and supported the idea of a quick postglacial colonization followed by expansion of the species in large parts of Europe. Phylogenetic analysis under mtDNA revealed the existence of four different haplogroups with a well defined distribution across Europe and Anatolia. Both genetic systems supported the deep separation of Anatolian and European lineages of L. europaeus. Nevertheless, Anatolian Y-DNA lineages extended across a longer geographic distance in south-eastern Europe than Anatolian mtDNA haplotypes, probably as a result of higher female philopatry that makes mtDNA introgression more difficult in brown hares.     

New Species of <Emphasis Type="Italic">Inoceramus</Emphasis>-like Bivalves of the Subfamily Kolymiinae from the Middle Permian of Northeastern Asia

Middle Permian Inoceramus-like bivalves of the genera Kolymia Licharew and Cyrtokolymia Astafieva endemic to the East Boreal Biogeographic Realm are considered. Cyrtokolymia, previously regarded by the author as endemics of the Verkhoyansk–Okhotsk Province and including only the type species, are also recorded in the Kolyma–Omolon Province, where they are represented by the endemic species C. bobini sp. nov. An emended diagnosis of the genus Cyrtokolymia is provided. The genus Kolymia comprises about 30 species. The greatest diversity of Kolymia (24 species, 13 of which are endemic) is known from the Verkhoyansk–Okhotsk Province, which is the center of diversification of this genus. The Kolyma–Omolon Province is characterized by 12 species, only three of which are endemic. In other provinces of the East Boreal Realm, only individual members of Kolymia are known. From the Middle Permian of the Omolon Massif, northern Verkhoyansk Region, and Penzhinsky Ridge, the following new species are described: Kolymia posneri Muromzeva, Kusnezov et Biakov, sp. nov., K. pontoneica Biakov, sp. nov., K. simkiniformis Biakov, sp. nov., and Cyrtokolymia bobini Biakov, sp. nov.     

Geographical structuring of genetic diversity across the whole distribution range of Narcissus longispathus, a habitat-specialist, Mediterranean narrow endemic

Background and Aims

High mountain ranges of the Mediterranean Basin harbour a large number of narrowly endemic plants. In this study an investigation is made of the levels and partitioning of genetic diversity in Narcissus longispathus, a narrow endemic of south-eastern Spanish mountains characterized by a naturally fragmented distribution due to extreme specialization on a rare habitat type. By using dense sampling of populations across the species'' whole geographical range, genetic structuring at different geographical scales is also examined.

Methods

Using horizontal starch-gel electrophoresis, allozyme variability was screened at 19 loci for a total of 858 individuals from 27 populations. The data were analysed by means of standard statistical approaches in order to estimate gene diversity and the genetic structure of the populations.

Key Results

Narcissus longispathus displayed high levels of genetic diversity and extensive diversification among populations. At the species level, the percentage of polymorphic loci was 68 %, with average values of 2·1, 0·11 and 0·14 for the number of alleles per locus, observed heterozygosity and expected heterozygosity, respectively. Southern and more isolated populations tended to have less genetic variability than northern and less-isolated populations. A strong spatial patterning of genetic diversity was found at the various spatial scales. Gene flow/drift equilibrium occurred over distances <4 km. Beyond that distance divergence was relatively more influenced by drift. The populations studied seem to derive from three panmictic units or ‘gene pools’, with levels of admixture being greatest in the central and south-eastern portions of the species'' range.

Conclusions

In addition to documenting a case of high genetic diversity in a narrow endemic plant with naturally fragmented populations, the results emphasize the need for dense population sampling and examination of different geographical scales for understanding population genetic structure in habitat specialists restricted to ecological islands.Key words: Allozymes, genetic diversity, geographical scale, habitat isolation, Narcissus longispathus, Mediterranean endemism, mountain range, natural fragmented distribution     

Plant biodiversity in the Saint Catherine area of the Sinai peninsula, Egypt

This study focuses on plant species richness, taxonomic diversity, and endemism of the Sinai peninsula as a whole and Saint Catherine area in particular. Beta diversity was also measured using _T to represent the biotic change between different landforms in the St. Catherine area. The peninsula supports about 1285 species including the infraspecific taxa of which about 800 species (also including the infraspecific taxa) are recorded in the southern Sinai south of El-Tih Desert. The present study reveals that southern Sinai is more diverse as compared with the Sinai peninsula as a whole. Southern Sinai also supports more than the expected numbers of rare and very rare species (about 62%) and about 4.3% endemic species. Beta diversity between different landforms in the St. Catherine area reflect a large biotic change between slopes and terraces on the one hand and between terraces and ridges on the other. Finally, the study recommends appropriate management of the peninsula and more research for the management requirements of the numerous endemic and rare species in the Sinai peninsula.     

The plankton community of Lake Matano: factors regulating plankton composition and relative abundance in an ancient,tropical lake of Indonesia

Recent evidence reveals that food webs within the Malili Lakes, Sulawesi, Indonesia, support community assemblages that are made up primarily of endemic species. It has been suggested that many of the species radiations, as well as the paucity of cosmopolitan species in the lakes, are related to resource limitation. In order to substantiate the possibility that resource limitation is playing such an important role, a study of the phytoplankton and zooplankton communities of Lake Matano was implemented between 2000 and 2004. We determined species diversity, relative abundances, size ranges, and total biomass for the phytoplankton and zooplankton, including the distribution of ovigerous individuals throughout the epilimnion of Lake Matano in three field seasons. The phytoplankton community exhibited very low biomass (<15 μg l^?1) and species richness was depressed. The zooplankton assemblage was also limited in biomass (2.5 mg l^?1) and consisted only of three taxa including the endemic calanoid Eodiaptomus wolterecki var. matanensis, the endemic cyclopoid, Tropocyclops matanensis and the rotifer Horaella brehmi. Zooplankton were very small (<600 μm body length), and spatial habitat partitioning was observed, with Tropocylops being confined to below 80 m, while rotifer and calanoid species were consistently observed above 80 m. Less than 0.1% of the calanoid copepods in each year were egg-bearing, suggesting very low population turnover rates. It was concluded that chemical factors as opposed to physical or biological processes were regulating the observed very low standing crops of phytoplankton which in turn supports a very minimal zooplankton community restricted in both species composition and abundance. As chemical factors are a function of the catchment basin of Lake Matano, it is predicted that resource limitation has long played an important role in shaping the unique endemic assemblages currently observed in the food web of the lake.     

Multilocus phylogenetic analysis of true morels (Morchella) reveals high levels of endemics in Turkey relative to other regions of Europe

The present study was conducted to better understand how the phylogenetic diversity of true morels (Morchella) in Turkey compares with species found in other regions of the world. The current research builds on our recently published surveys of 10 Turkish provinces and the northern hemisphere in which DNA sequence data from 247 and 562 collections respectively were analyzed phylogenetically. Herein we report on phylogenetic analyses of 243 additional collections made in spring 2009 and 2010 from eight additional provinces in the Aegean, Black Sea, central Anatolia, eastern Anatolia and Marmara regions of Turkey. Our analysis revealed that five species within the Esculenta clade (yellow morels) and 15 species within the Elata clade (black morels) were present in Turkey. Our preliminary results also indicate that M. anatolica, recently described from a collection in Mu?la province in the Aegean region of Turkey, is a closely related sister of M. rufobrunnea; these two species comprise a separate evolutionary lineage from the Esculenta and Elata clades. Nine species of Morchella currently are known only from Turkey, four species were present in Turkey and other European countries and seven species might have been introduced to Turkey anthropogenically. Three of the putatively exotic species in Turkey appear to be endemic to western North America; they are nested within a clade of fire-adapted morels that dates to the late Oligocene, 25 000 000 y ago. Our results indicate that there are roughly twice as many Morchella species in Turkey compared with the other regions of Europe sampled. Knowledge of Morchella species diversity and their biogeographic distribution are crucial for formulating informed conservation policies directed at preventing species loss and ensuring that annual morel harvests are sustainable and ecologically sound.     

Glacial refuges for three‐spined stickleback in the Iberian Peninsula: mitochondrial DNA phylogeography

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