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The great majority of continental flood basalts (CFBs) have a marked lithospheric geochemical signature, suggesting derivation from the continental lithosphere, or contamination by it. Here we present new Pb and Os isotopic data and review previously published major element, trace element, mineral chemical, and Sr and Nd isotopic data for geochemically unusual mafic and ultramafic dikes located in the Antarctic segment (Ahlmannryggen, western Dronning Maud Land) of the Karoo CFB province. Some of the dikes show evidence of minor contamination with continental crust, but the least contaminated dikes exhibit depleted mantle –like initial εNd (+9) and 187Os/188Os (0.1244–0.1251) at 180 Ma. In contrast, their initial Sr and Pb isotopic compositions (87Sr/86Sr = 0.7035–0.7062, 206Pb/204Pb = 18.2–18.4, 207Pb/204Pb = 15.49–15.52, 208Pb/204Pb = 37.7–37.9 at 180 Ma) are more enriched than expected for depleted mantle, and the major element and mineral chemical evidence indicate contribution from (recycled) pyroxenite sources. Our Sr, Nd, Pb, and Os isotopic and trace element modeling indicate mixed peridotite-pyroxenite sources that contain ~10–30 % of seawater-altered and subduction-modified MORB with a recycling age of less than 1.0 Ga entrained in a depleted Os-rich peridotite matrix. Such a source would explain the unusual combination of elevated initial 87Sr/86Sr and Pb isotopic ratios and relative depletion in LILE, U, Th, Pb and LREE, high initial εNd, and low initial 187Os/188Os. Although the sources of the dikes probably did not play a major part in the generation of the Karoo CFBs in general, different kind of recycled source components (e.g., sediment-influenced) would be more difficult to distinguish from lithospheric CFB geochemical signatures. In addition to underlying continental lithosphere, the involvement of recycled sources in causing the apparent lithospheric geochemical affinity of CFBs should thus be carefully assessed in every case.

A checklist of 29 species in the smaller carnoid families Acartophthalmidae, Canacidae, Carnidae and Milichiidae (Diptera) recorded from Finland is presented. Tethinidae are also included as a subfamily of Canacidae. Phyllomyza tetragona Hendel is removed from the list as no reliable records of this species within the post-1944 borders of Finland could be found.

A checklist of the Agromyzidae (Diptera) recorded from Finland is presented. 279 (or 280) species are currently known from the country. Phytomyza linguae Lundqvist, 1947 is recorded as new to Finland.

A Finnish checklist of the sphaeroceroid fly families Chyromyidae and Heleomyzidae is provided.

Nearly thirty-five years have passed since Hackman published his “Check list of the Finnish Diptera” (1980). The number of true flies (Diptera) known from Finland has increased by more than two thousand species since then. At the same time, hundreds of erroneous records have been recognized and purged from the checklist. ZooKeys issue 441 provides a new checklist of the Diptera species of the Republic of Finland. This introductory paper presents the rationale behind the project, provides technical documentation on the checklist format and sources used, and summarizes the results. The remaining papers in this issue cover one or more Diptera families in detail. Two electronic appendices are provided: supporting data (additional references to first published records and the previous checklist) and a complete list of Finnish Diptera taxa in Darwin Core compliant format for easy computer access and processing. The new checklist records 6920 fly species from Finland, 2932 belonging to the nematoceran or lower flies and 3989 to the suborder Brachycera. The changes since 1980 are most prominent in the Lower Diptera. For example, more than 400 non-biting midges (Chironomidae) have been added since 1980, and the number of moth flies (Psychodidae) known from Finland has more than tripled. Among the larger families, large increases in known Finnish species are also seen in Cecidomyiidae (161% increase), Pipunculidae (98%), and Chironomidae (90%).

An updated checklist of the Atelestidae, Brachystomatidae, Dolichopodidae, Empididae and Hybotidae (Diptera) recorded from Finland is presented. The genera with uncertain placement within superfamily Empidoidea (= the Iteaphila group) are also included in this paper.

The northern European taxa of genus Melanostoma Schiner, 1860 (Syrphidae, Diptera) are revised. A longstanding question concerning the number of Melanostoma taxa occurring in northern Europe prompted us to contrast and compare their morphological and molecular variability. Particular uncertainty concerned the putative existence of a sibling species of Melanostoma mellinum, and the identity of the taxon Melanostoma dubium in northern Europe due to existence of morphologically similar dark forms of M. mellinum in the northern parts of its distributional range. Partial sequences of two DNA markers, the mitochondrial protein-coding gene cytochrome c oxidase subunit I (COI-3') and the nuclear second internal transcribed spacer (ITS2) were analysed separately under parsimony. The obtained COI-3'gene fragment showed taxon-specific haplotypes and haplotypes that were shared among the taxa. The ITS2 sequences presented genotypes unique to each species, and congruence with our independently established taxonomic entities. Based on congruent signal of the ITS2 sequences and study of morphological characters we establish the presence of four taxa in northern Europe: Melanostoma mellium (= M. dubium nec auctt., syn. n.), M. certum sp. n. (= M. dubium auctt.), M. mellarium stat. n. (= M. mellinum auctt. partim) and M. scalare. Lectotype designations were made for Musca mellina, Syrphus mellarius and Melanostoma mellinum var. melanatus. The following synonymies were established: Melanostoma mellarium=Melanostoma melanatum syn. n.;Melanostoma mellinum = Scaeva dubia syn. n., Melanostoma tschernovi syn. n., and Melanostoma clausseni syn. n. Morphological circumscriptions of the taxa and an identification key are presented.

An updated checklist of the superfamilies Oestroidea and Hippoboscoidea recorded from Finland is presented. The checklist covers the following families: Calliphoridae, Rhiniidae, Sarcophagidae, Rhinophoridae, Tachinidae, Oestridae and Hippoboscidae.

Premise of the study: The locally rare, haploid, lichen-forming fungi Bryoria capillaris , B. fuscescens , and B. implexa are associated with boreal forests and belong to Bryoria sect. Implexae . Recent phylogenetic studies consider them to be conspecific. Microsatellite loci were developed to study population structure in Bryoria sect. Implexae and its response to ecosystem disturbances. • Methods and Results: We developed 18 polymorphic microsatellite markers using 454 pyrosequencing data assessed in 82 individuals. The number of alleles per locus ranged from two to 13 with an average of 4.6. Nei’s unbiased gene diversity, averaged over loci, ranged from 0.38 to 0.52. The markers amplifi ed with all three species, except for markers Bi05, Bi15, and Bi18. • Conclusions: The new markers will allow the study of population subdivision, levels of gene introgression, and levels of clonal spread of Bryoria sect. Implexae . They will also facilitate an understanding of the effects of forest disturbance on genetic diversity of these lichen species.

A checklist of the family Sphaeroceridae (Diptera) recorded from Finland.

Intentional moving of species threatened by climate change is actively being discussed as a conservation approach. The debate, empirical studies, and policy development, however, are impeded by an inconsistent articulation of the idea. The discrepancy is demonstrated by the varying use of terms, such as assisted migration, assisted colonisation, or managed relocation, and their multiple definitions. Since this conservation approach is novel, and may for instance lead to legislative changes, it is important to aim for terminological consistency. The objective of this study is to analyse the suitability of terms and definitions used when discussing the moving of organisms as a response to climate change. An extensive literature search and review of the material (868 scientific publications) was conducted for finding hitherto used terms (N = 40) and definitions (N = 75), and these were analysed for their suitability. Based on the findings, it is argued that an appropriate term for a conservation approach relating to aiding the movement of organisms harmed by climate change is assisted migration defined as follows: Assisted migration means safeguarding biological diversity through the translocation of representatives of a species or population harmed by climate change to an area outside the indigenous range of that unit where it would be predicted to move as climate changes, were it not for anthropogenic dispersal barriers or lack of time. The differences between assisted migration and other conservation translocations are also discussed. A wide adoption of the clear and distinctive term and definition provided would allow more focused research on the topic and enable consistent implementation as practitioners could have the same understanding of the concept.

Systematics, phylogenetics and taxonomy are the scientific fields of species discovery, delimitation, description, classification and evolutionary history. The major task of these fields is to form meaningful groups, such as species and higher taxa, based on certain rules and characters. Species and higher taxa create the units of the Linnean hierarchic classification system, which is needed as the basis of all knowledge on biodiversity. Indeed, there is a great need for a complete classification, covering every species on earth, because all subsequent studies and applications are hindered as long as species remain undescribed. However, the estimated count of nine million species on earth, of which ca. 86% are currently unknown to science, together with the numerous threats to biodiversity pose a real challenge to taxonomy, and efficient tools and procedures are strongly needed. In order to be both effective and high-quality, the taxonomic workflow needs to be divided into different steps in the correct sequence. A sampling scheme, the choice of characters and analytical tools are dependent on the phase performed along the workflow. The main aim of Mari Kekkonen s doctoral thesis is to study and conduct different steps along this taxonomic flowchart by using various gelechioid moths as focal species. Specifically, the thesis includes four chapters: a phylogenetic examination of the superfamily Gelechioidea based on combined data of multi-locus DNA and morphology, and three studies focusing on delineation of putative species (i.e., operational taxonomic units, OTUs) based on single-locus DNA barcodes. The delineation studies cover various topics, from testing different OTU delineation methods (BIN, TCS, ABGD, GMYC) with reference species, developing criteria for discordant results and a protocol for associating short sequences from type specimens to delineated groups, to employing the methods, criteria and protocols in practice. The first chapter presents a phylogene, Laji on luonnon monimuotoisuuden perusyksikkö ja siten kaikki biologinen tutkimus alkaa lajien määrittelystä: niiden rajaamisesta, kuvaamisesta ja nimeämisestä. Maapallon miljoonat lajit tarjoavat lukemattomia hyötyjä ihmiskunnalle esimerkiksi ekosysteemipalvelujen ja uusien lääkkeiden muodossa, mutta näiden resurssien hyödyntäminen on vaikeaa niin kauan kun lajeilta puuttuvat kuvaus ja nimi. Ongelma on huomattava, koska tuoreimpien arvioiden mukaan jopa 86 prosenttia maalla elävistä lajeista on tieteelle tuntemattomia. Lisäksi luonnon monimuotoisuutta uhkaavat monet tekijät, ilmastonmuutos ja elinympäristöjen tuhoaminen muiden muassa, jotka tekevät lajien tutkimuksesta erityisen kiireellistä. Lajien kuvaamiseen ja luokitteluun keskittyvät tieteenalat, systematiikka ja taksonomia, ovat suuren haasteen edessä, ja tarve työn tehostamiseen on ilmeinen. Mari Kekkosen väitöskirja paneutuu erityisesti tähän ongelmaan ja esittelee taksonomisen työjärjestyksen , jossa eri osatutkimukset suoritetaan tietyssä, toisiaan tukevassa järjestyksessä ja jokaiseen vaiheeseen pyritään valitsemaan sopivimmat työkalut. Työkalupakki sisältää perinteisten rakennetuntomerkkien lisäksi eri DNA-jaksojen tutkimista monilla uusilla analyysimenetelmillä. Työjärjestys tehostaa taksonomista tutkimusta ajallisesti, lisää sen objektiivisuutta ja toistettavuutta sekä helpottaa eri tutkimusten vertailua. Väitöskirjassa tutkitaan erityisesti DNA-viivakoodien käyttöä osana taksonomista työtä. DNA-viivakoodit kehitettiin alun perin lajien tunnistamiseen tutkimalla lyhyt pätkä perintöainesta ja vertaamalla sitä laajaan lajitietokantaan. Tätä tekniikkaa on käytetty moniin sovelluksiin kuten ruuan ainesosien alkuperälajin selvittämiseen. DNA-viivakoodit soveltuvat myös tuntemattomien lajien tutkimukseen, koska niillä voidaan tuottaa alustavat lajirajaukset kymmenille tai sadoille lajeille samanaikaisesti. Alustavat lajit varmennetaan muiden tuntomerkkien avulla ennen niiden kuvaamista. Väitöskirjan kohdela

Madeira island presents a unique spider diversity with a high number of endemic species, many of which are still poorly known. A recent biodiversity survey on the terrestrial arthropods of the native forest, Laurisilva, provided a large set of standardized samples from various patches throughout the island. Out of the fifty two species recorded, approximately 33.3% are Madeiran endemics, many of which had not been collected since their original description. Two new species to science are reported – Ceratinopsis n. sp. and Theridion n. sp. – and the first records of Poeciloneta variegata (Blackwall, 1841) and Tetragnatha intermedia Kulczynski, 1891 are reported for the first time for Madeira island. Considerations on species richness and abundance from different Laurisilva locations are presented, together with distribution maps for endemic species. These results contribute to a better understanding of spider diversity patterns and endemic species distribution in the native forest of Madeira island.

The taxonomic and nomenclatural history of Jávorka’s Sorbus bakonyensis is surveyed and its correct authorship and bibliographic citation is provided. The identity of the original material of S. bakonyensis and the history of its interpretations are discussed. The name S. bakonyensis is applicable to an endemic of the eastern Bakony Mountains, restricted to the vicinity of the village of Márkó in Hungary. The illegitimate name S. majeri is a synonym of S. bakonyensis. A new name, S. udvardyana, is introduced here for the species occurring in the western Balaton region of Hungary, which was erroneously named “S. bakonyensis” in recent Hungarian literature.

Two new species of Meropidia Hippa & Thompson, 1983 (Diptera, Syrphidae) are described, Meropidia nitida Morales, sp. n. and M. flavens Hippa & Ståhls sp. n., from Bolivia and Colombia respectively. A key to all described Meropidia species is provided.

Successful conservation plans are conditioned by our ability to detect anthropogenic change in space and time and various statistical analyses have been developed to handle this critical issue. The main objective of this paper is to illustrate a new approach for spatial analysis in conservation biology. Here, we propose a two-step protocol. First, we introduce a new disturbance metric which provides a continuous measure of disturbance for any focal communities on the basis of the surrounding landscape matrix. Second, we use this new gradient to estimate species and community disturbance thresholds by implementing a recently developed method called Threshold Indicator Taxa ANalysis (TITAN). TITAN detects changes in species distributions along environmental gradients using indicators species analysis and assesses synchrony among species change points as evidence for community thresholds. We demonstrate our method with soil arthropod assemblages along a disturbance gradient in Terceira Island (Azores, Portugal). We show that our new disturbance metric realistically reflects disturbance patterns, especially in buffer zones (ecotones) between land use categories. By estimating species disturbance thresholds with TITAN along the disturbance gradient in Terceira, we show that species significantly associated with low disturbance differ from those associated with high disturbance in their biogeographical origin (endemics, non-endemic natives and exotics) and taxonomy (order). Finally, we suggest that mapping the disturbance community thresholds may reveal areas of primary interest for conservation, since these may host indigenous species sensitive to high disturbance levels. This new framework may be useful when: (1) both local and regional processes are to be reflected on single disturbance measures; (2) these are better quantified in a continuous gradient; (3) mapping disturbance of large regions using fine scales is necessary; (4) indicator species for disturbance are se

Individual variability influences the demographic and evolutionary dynamics of spatially structured populations, and conversely ecological and evolutionary dynamics provide the context under which variations at the individual level occur. Therefore, it is essential to identify and characterize the importance of the different factors that may promote or hinder individual variability. Animal signaling is a prime example of a type of behavior that is largely dependent on both the features of individuals and the characteristics of the population to which they belong. After 10 years studying the dynamics of a population of a long-lived species, the eagle owl (Bubo bubo), we investigated the emergence and maintenance of traits that reveal individual identity by focusing on vocal features. We found that individuals inhabiting a high density population characterized by a relative lack of heterogeneity (in terms of prey availability and breeding success) among breeding sites might be selected for reducing the levels of identity. Two non-mutually exclusive hypotheses may explain the structural call patterns we detected: (1) similarity in calls may be principally a consequence of the particular characteristics of the population; and (2) high density may encourage individuals to mimic each other’s vocalizations in a cascade effect, leading to a widespread and unique communication network.

Public genomic databases have provided new directions for molecular marker development and initiated a shift in the types of PCR-based techniques commonly used in plant science. Alongside commonly used arbitrarily amplified DNA markers, other methods have been developed. Targeted fingerprinting marker techniques are based on the well-established practices of arbitrarily amplified DNA methods, but employ novel methodological innovations such as the incorporation of gene or promoter elements in the primers. These markers provide good reproducibility and increased resolution by the concurrent incidence of dominant and co-dominant bands. Despite their promising features, these semi-random markers suffer from possible problems of collision and non-homology analogous to those found with randomly generated fingerprints. Transposable elements, present in abundance in plant genomes, may also be used to generate fingerprints. These markers provide increased genomic coverage by utilizing specific targeted sites and produce bands that mostly seem to be homologous. The biggest drawback with most of these techniques is that prior genomic information about retrotransposons is needed for primer design, prohibiting universal applications. Another class of recently developed methods exploits length polymorphism present in arrays of multi-copy gene families such as cytochrome P450 and β-tubulin genes to provide cross-species amplification and transferability. A specific class of marker makes use of common features of plant resistance genes to generate bands linked to a given phenotype, or to reveal genetic diversity. Conserved DNA-based strategies have limited genome coverage and may fail to reveal genetic diversity, while resistance genes may be under specific evolutionary selection. Markers may also be generated from functional and/or transcribed regions of the genome using different gene-targeting approaches coupled with the use RNA information. Such techniques have the potential t

Background Climatic warming predicts that species move their entire distribution poleward. Poleward movement of the ‘cold’ side of the distribution of species is empirically supported, but evidence of poleward movement at the ‘warm’ distributional side is relatively scarce. Methodology/Principal Finding Finland has, as the first country in the world, completed three national atlas surveys of breeding birds, which we here use to calculate the sizes and weighted mean latitudes of the national range of 114 southern and 34 northern bird species during three periods (1974–1979; 1986–1989; 2006–2010), each denoting species presence in approximately 3 800 10×10 km2 squares. We find strong evidence that southern species (breeding predominantly in central Europe) showed a latitudinal shift of 1.1–1.3 km/year poleward during all three pairwise comparisons between these atlases (covering 11, 20.5 and 31.5 years respectively). We find evidence of a latitudinal shift of 0.7–0.8 km/year poleward of northern boreal and Arctic species, but this shift was not found in all study periods and may have been influenced by increased effort put into the more recent surveys. Species showed no significant correlation in changes in range size and weighted mean latitude between the first (11 year) and second (20.5 year) period covered by consecutive atlases, suggesting weak phylogenetic signal and little scope of species characteristics in explaining latitudinal avian range changes. Conclusions Extinction-driven avian range changes (at the ‘warm’ side) of a species' distribution occur at approximately half the rate of colonisation-driven range changes (at the ‘cold’ side), and its quantification therefore requires long-term monitoring data, possibly explaining why evidence for such changes is currently rare. A clear latitudinal shift in an assemblage of species may still harbour considerable temporal inconsistency in latitudinal movement on the species level. Understanding this inconsistency i

Traditionally, genomic or transcriptomic data have been restricted to a few model or emerging model organisms, and to a handful of species of medical and/or environmental importance. Next-generation sequencing techniques have the capability of yielding massive amounts of gene sequence data for virtually any species at a modest cost. Here we provide a comparative analysis of de novo assembled transcriptomic data for ten non-model species of previously understudied animal taxa.

Predation affects life history traits of nearly all organisms and the population consequences of predator avoidance are often larger than predation itself. Climate change has been shown to cause phenological changes. These changes are not necessarily similar between species and may cause mismatches between prey and predator. Eurasian sparrowhawk Accipiter nisus, the main predator of passerines, has advanced its autumn phenology by about ten days in 30 years due to climate change. However, we do not know if sparrowhawk migrate earlier in response to earlier migration by its prey or if earlier sparrowhawk migration results in changes to predation risk on its prey. By using the median departure date of 41 passerine species I was able to show that early migrating passerines tend to advance, and late migrating species delay their departure, but none of the species have advanced their departure times as much as the sparrowhawk. This has lead to a situation of increased predation risk on early migrating long-distance migrants (LDM) and decreased the overlap of migration season with later departing short-distance migrants (SDM). Findings highlight the growing list of problems of declining LDM populations caused by climate change. On the other hand it seems that the autumn migration may become safer for SDM whose populations are growing. Results demonstrate that passerines show very conservative response in autumn phenology to climate change, and thus phenological mismatches caused by global warming are not necessarily increasing towards the higher trophic levels.

Reproductive, phenotypic and life-history traits in many animal and plant taxa show geographic variation, indicating spatial variation in selection regimes. Maternal deposition to avian eggs, such as hormones, antibodies and antioxidants, critically affect development of the offspring, with long-lasting effects on the phenotype and fitness. Little is however known about large-scale geographical patterns of variation in maternal deposition to eggs. We studied geographical variation in egg components of a passerine bird, the pied flycatcher (Ficedula hypoleuca), by collecting samples from 16 populations and measuring egg and yolk mass, albumen lysozyme activity, yolk immunoglobulins, yolk androgens and yolk total carotenoids. We found significant variation among populations in most egg components, but ca. 90% of the variation was among individuals within populations. Population however explained 40% of the variation in carotenoid levels. In contrast to our hypothesis, we found geographical trends only in carotenoids, but not in any of the other egg components. Our results thus suggest high within-population variation and leave little scope for local adaptation and genetic differentiation in deposition of different egg components. The role of these maternally-derived resources in evolutionary change should be further investigated.

A new classification and linear sequence of the gymnosperms based on previous molecular and morphological phylogenetic and other studies is presented. Currently accepted genera are listed for each family and arranged according to their (probable) phylogenetic position. A full synonymy is provided, and types are listed for accepted genera. An index to genera assists in easy access to synonymy and family placement of genera.

The Caucasus region is a hotspot of biodiversity and is one of the few areas in the Northern Hemisphere which harbor Pleistocene glacial refugia. The region encompasses Armenia, Azerbaijan, Georgia, the southernmost European Russia, NE Turkey, and northern Iran. The study on fungal composition of the Caucasus region and its connection and possible contribution to the present mycota of Europe has largely escaped empirical scrutiny. Using taxonomic surveys, phylogenetic reconstruction methods, haplotype analysis, and similarity tests, this study has aimed to, 1) summarize the knowledge on the occurrence of corticioids and polypores in the Caucasus region, 2) resolve the phylogenetic relationships of selected, resupinate wood-inhabiting basidiomycetes for which the Caucasus region is currently the mere, or one of the noteworthy areas of distribution, and, 3) assess the similarity of Caucasian corticioid fungi to those of Europe and important areas in the Northern Hemisphere, and to examine the significance of the Caucasus region as a glacial refugium for these fungi. This study provides the first catalogue of corticioids and polypores (635 species) occurring in the Caucasus region. The phylogeny and systematics of the Caucasian resupinate taxa in focus has been resolved and the usefulness of some morphological characters has been re-evaluated. In this context, four new genera and two new species were described and five new combinations were proposed, two of which were supplemented with modern descriptions. The species composition of corticioids in the Caucasus region is found to be distinctly more similar to Europe and North America than to East Asia and India. The highest molecular diversity and within population pairwise distance for Peniophorella praetermissa has been detected in the Caucasus and East Asia, with the isolates of the latter area being highly divergent from the European ones. This, and the assignment of root haplotype to the Caucasian isolates in a hap, Kaukasia, joka perinteisesti tunnetaan Euroopan ja Aasian kohtauspaikkana, käsittää Georgian, Armenian ja Azerbaidjanin sekä Euroopan puoleisen Venäjän eteläisimmät osat, koillisen Turkin ja Iranin pohjoisosat. Alueen elollinen monimuotoisuus on huomattava ja se on eräs pohjoisen pallonpuoliskon ns. refugioista, eli alueista joilla viimeisen jääkauden aikana säilyi muualta kadonnutta lajistoa. Kaukasian sienilajistoa ja sen roolia Euroopan nykyisen lajiston lähteenä ei ole tarkasti tutkittu. Tietoja alueen puilla kasvavista kantasienistä (käävistä ja orvakoista) ei ole aiemmin koottu yhteen ja tällainen perusselvitys oli tarpeen, jotta lajiston taksonomiaa ja eliömaantiedettä voitaisiin tutkia. Toisaalta sellaisten puilla kasvavien kantasienten, joilla on kasvualustan myötäinen eli resupinaattinen itiöemä, lajit tunnetaan vielä hyvin huonosti ja siksi niiden systematiikan selvittäminen oli tässä työssä etusijalla. Tässä työssä pyrittiin 1) kokoamaan yhteen tiedot Kaukasian käävistä ja orvakoista, 2) selvittämään sellaisten resupinaattisten lajien fylogenia, joiden ainoa tai pääasiallinen levinneisyysalue on Kaukasia, ja 3) tutkimaan Kaukasian merkitystä Euroopassa ja eräillä muilla pohjoisen pallonpuoliskon alueilla esiintyvien puilla kasvavien kantasienten jääkauden aikaisena vetäytymisalueena vertailemalla näiden alueiden ja Kaukasian nykyistä lajistoa. Tutkimuksessa esitetään ensimmäinen yhteenveto alueen käävistä ja orvakoista (noin 626 lajia). Useiden alueen resupinaattisten taksonien asema selvitettiin ja eräiden morfologisten ominaisuuksien käyttökelpoisuus arvioitiin uudelleen. Tieteelle uusina kuvattiin neljä uutta sukua ja kaksi uutta lajia morfologisten, sekä silloin kun tietoa oli saatavilla, myös ekologisten piirteiden ja sekvenssitason aineiston fylogeneettisten analyysien perusteella. Tämän lisäksi työ sisältää viisi nimistöllistä kombinaatiota ja kahteen niistä sisältyvät myös uudet taksonomiset kuvaukset. Kaikkien taksonomisten uutuuksien kuvaus per

Tutkimus käsittelee kääpien sukulaisuussuhteita. Käävät ovat kantasienten (Basidiomycota) muotoryhmä, joiden itiöemien alapinta muodostuu yhteensulautuneista pilleistä. Muotoryhmänä kääpiä voi verrata vaikka puihin siinä mielessä, että käävät kuten puutkaan eivät ole samankaltaisuudestaan huolimatta kaikki sukua toisilleen. DNA:n käyttö sukulaisuussuhteiden selvittämisessä on aloittanut mullistuksen kääpien luokittelussa. Aiemmin käytetty, itiöemien ominaisuuksiin perustunut luokittelu on osoittautunut keinotekoiseksi sukulaisuussuhteiden kannalta. Tutkimuksessani syvennyttiin useamman kääpäsuvun polveutumishistoriaan hyödyntäen DNA:ta ja perinteisiä menetelmiä. Tutkimuksen keskeisimmät tulokset liittyvät sitkokääpien sukuun (Antrodiella). Tämä noin 70 lajia sisältävä suku osoittautui rikkonaiseksi - sitkokääpiin luetut lajit kuuluvat kahteen sienilahkoon ja oikesti vähintään 13 sukuun. Tutkimuksessa löytyi kaksi Suomelle uutta sitkokääpää, leppikääpä (A. ichnusana) ja nipukkakääpä (A. leucoxantha). Uudet suvut kuvattiin Suomessa esiintyville sirppikääville (Sidera) ja talikääville (Obba). Uusi kääpäsuku ja -laji kuvattiin myös Indonesiasta (Sebipora aquosa). Valtaosa sitkokääpiin luetuista lajeista kuuluu orakarakoiden heimoon (Steccherinaceae), joka rajattiin tässä tutkimuksessa uudelleen. Heimoon kuuluvat mm. karakäävät (Junghuhnia) ja orakasmaiset orakarakat (Steccherinum). Sen sisällä selvitettiin kääpien ja orakkaiden sukulaisuussuhteita. Perinteisesti käävät ja orakkaat on viety eri sukuihin riippumatta niiden mikroskooppisesta samankaltaisuudesta. Tulosten valossa orakarakoiden heimossa käävät ja orakkaat pysyvät pääosin erillisissä suvuissa, mutta tästä on myös poikkeuksia (Antrodiella, Metuloidea ja Steccherinum). Lähes kaikki DNA:n perusteella määriteltävissä olevat suvut ovat tunnistettavissa itiöemien ominaisuuksiensa perusteella. Tulokset antavat eväitä kääpien luokitteluun laajemminkin osoittamalla, mitkä ominaisuudet ovat luokittelun kannalta merkity, This thesis deals with genealogy of polypores. Polypores are a form group of basidiomycete fungi (Basidiomycota). Their underside is formed of fused tubes. As a form group polypores can be compared to trees in that neither all polypores nor all trees are related despite their similar appearance. The use of DNA is revolutionising our understanding of how polypores are related. The classification formerly in use was based on fruiting body characters, and has turned out to be highly unnatural from the perspective of genealogy. In the thesis work I studied evolutionary history of several polypore genera using DNA and traditional methods. Main results of the study are related to the polypore genus Antrodiella, which includes about 70 species. Antrodiella turned out to be a heterogeneous assemblage of species. Species currently included in the genus belong to two orders and to a minimum of 13 genera. Two Antrodiella species new to Finland were found during the study (A. ichnusana and A. leucoxantha). Three new polypore genera were also described (Obba, Sidera and Sebipora). Majority of Antrodiella belong to the family Steccherinaceae, which was redefined in this study. In addition to polypore genera, Steccherinaceae includes genera with fruiting bodies that have a spinose (hydnoid) underside. Typically poroid and hydnoid fungi are classified in separate genera regardless of their microscopic similarity. This study focused on the evolutionary relationship between hydnoid and poroid species within Steccherinaceae. The results show that poroid and hydnoid fungi mostly seem to belong to separate genera, but there are also exceptions (Antrodiella, Metuloidea and Steccherinum). Almost all genera that were defined with the aid of DNA could be characterised with fruiting body and microscopic characters. The results have wider implications in polypore classification by showing which characters are significant in classifying polypores. Increased information about species diversity

This work examines stable isotope ratios of carbon, oxygen and hydrogen in annual growth rings of trees. Isotopic composition in wood cellulose is used as a tool to study past climate. The method benefits from the accurate and precise dating provided by dendrochronology. In this study the origin, nature and the strength of climatic correlations are studied on different temporal scales and at different sites in Finland. The origin of carbon isotopic signal is in photosynthetic fractionation. The basic physical and chemical fractionations involved are reasonably well understood. This was confirmed by measuring instantaneous photosynthetic discrimination on Scots pine (Pinus sylvestris L.). The internal conductance of CO2 was recognized to have a significant impact on the observed fractionation, and further investigations are suggested to quantify its role in controlling the isotopic signal of photosynthates. Isotopic composition of the produced biomass can potentially be affected by variety of external factors that induce physiological changes in trees. Response of carbon isotopic signal in tree ring cellulose to changes in resource availability was assessed in a manipulation experiment. It showed that the signal was relatively stable despite of changes in water and nitrogen availability to the tree. Palaeoclimatic reconstructions are typically based on functions describing empirical relationship between isotopic and climatic parameters. These empirical relationships may change depending on the site conditions, species and timeframe studied. Annual variation in Scots pine tree ring carbon and oxygen isotopic composition was studied in northern and in central eastern Finland and annual variation in tree ring latewood carbon, oxygen and hydrogen isotopic ratio in Oak (Quercus robur L.) was studied in southern Finland. In all of the studied sites at least one of the studied isotope ratios was shown to record climate strongly enough to be used in climatic reconstructions., Tässä väitöskirjassa tutkittiin hiilen, hapen ja vedyn stabiilien eli pysyvien isotooppien suhdetta puiden vuosilustoissa ja niiden käyttöä tiedonlähteenä menneistä ilmastonvaihteluista. Kemiallisissa, fysikaalisissa ja biologisissa prosesseissa saman alkuaineen isotooppien suhde hieman muuttuu, eli tapahtuu fraktioitumista. Puuaineksessa isotooppisuhteet muuttuvat riippuen siitä millaisissa kasvuoloissa ja ilmastossa puu on kasvanut. Kun isotooppisuhde mitataan puun vuosilustosta, lusto voidaan dendrokronologisten menetelmien avulla ajoittaa ja isotooppisuhde yhdistää kyseisen vuoden kasvuoloihin. Aikaisemmissa tutkimuksissa Suomesta on mitattu lähinnä hiilen isotooppisuhteita puiden vuosilustoista. Tässä työssä perehdyttiin tarkemmin niihin prosesseihin jotka puussa muokkaavat hiilen isotooppisuhdetta sekä mitattiin hiilen lisäksi myös hapen ja vedyn pysyvien isotooppien suhdetta puiden vuosilustoissa. Kasvimateriaalin hiilen isotooppikoostumus määräytyy ensisijaisesti kasvin sitoessa hiiltä ilmakehästä yhteyttämällä. Tätä tutkittiin männyllä (Pinus sylvestris L.) kammiomittauksilla ja mallintamalla Etelä-Suomessa. Lisäksi tutkittiin männyn kasvuresurssien muutoksen vaikutusta vuosiluston hiilen isotooppikoostumukseen ja puun kasvuun Lapissa. Isotooppisuhteessa ei havaittu merkittävää muutosta, huolimatta kasvussa tapahtuneista muutoksista. Lapista ja Itä-Suomesta mitattiin 400 vuotta pitkät hiili- ja happi-isotooppi sekä lustonleveys aikasarjat mäntyjen vuosilustoista. Vuosittaista vaihtelua isotooppisuhteissa verrattiin säähavaintoihin viimeisen sadan vuoden ajalta. Hiilen isotooppien havaittiin korreloivan vahvasti kesälämpötilojen kanssa molemmilla alueilla. Itä-Suomessa havaittiin hapen ja myös hiilen isotooppien korreloivan kesän sademäärän kanssa. Pitkiä aikasarjoja käyttäen tehtiin 400 vuoden lämpötilarekonstruktio Lappiin ja lämpötila- ja sademäärärekonstruktiot Itä-Suomeen. Pitkiä aikasarjoja myös vertailtiin keskenään ja havaittiin niiden välisessä korr

Rimbachia leucobryi, a small pleurotoid agaric, is described as new to science. The species grew on living stems of Leucobryum sanctum in lowland rainforest. It is characterised by lamellate hymenophore, hymenial cystidia and small, ellipsoid spores. Pterygellus armeniacus, a bright-coloured stipitate stereoid species, is reported as new to Indonesia. The two species are illustrated.

For studies on the fern family Tectariaceae for the Flora of China, four species previously treated as Ctenitopsis need new combinations in Tectaria. The new combinations are proposed here and information on their types is provided.

Background: Malaria was prevalent in Finland in the 18th century. It declined slowly without deliberate counter-measures and the last indigenous case was reported in 1954. In the present analysis of indigenous malaria in Finland, an effort was made to construct a data set on annual malaria cases of maximum temporal length to be able to evaluate the significance of different factors assumed to affect malaria trends. Methods: To analyse the long-term trend malaria statistics were collected from 1750–2008. During that time, malaria frequency decreased from about 20,000 – 50,000 per 1,000,000 people to less than 1 per 1,000,000 people. To assess the cause of the decline, a correlation analysis was performed between malaria frequency per million people and temperature data, animal husbandry, consolidation of land by redistribution and household size. Results: Anopheles messeae and Anopheles beklemishevi exist only as larvae in June and most of July. The females seek an overwintering place in August. Those that overwinter together with humans may act as vectors. They have to stay in their overwintering place from September to May because of the cold climate. The temperatures between June and July determine the number of malaria cases during the following transmission season. This did not, however, have an impact on the longterm trend of malaria. The change in animal husbandry and reclamation of wetlands may also be excluded as a possible cause for the decline of malaria. The long-term social changes, such as land consolidation and decreasing household size, showed a strong correlation with the decline of Plasmodium. Conclusion: The indigenous malaria in Finland faded out evenly in the whole country during 200 years with limited or no counter-measures or medication. It appears that malaria in Finland was basically a social disease and that malaria trends were strongly linked to changes in human behaviour. Decreasing household size caused fewer interactions between families

Mutation and recombination are the fundamental processes leading to genetic variation in natural populations. This variation forms the raw material for evolution through natural selection and drift. Therefore, studying mutation rates may reveal information about evolutionary histories as well as phylogenetic interrelationships of organisms. In this thesis two molecular tools, DNA barcoding and the molecular clock were examined. In the first part, the efficiency of mutations to delineate closely related species was tested and the implications for conservation practices were assessed. The second part investigated the proposition that a constant mutation rate exists within invertebrates, in form of a metabolic-rate dependent molecular clock, which can be applied to accurately date speciation events. DNA barcoding aspires to be an efficient technique to not only distinguish between species but also reveal population-level variation solely relying on mutations found on a short stretch of a single gene. In this thesis barcoding was applied to discriminate between Hylochares populations from Russian Karelia and new Hylochares findings from the greater Helsinki region in Finland. Although barcoding failed to delineate the two reproductively isolated groups, their distinct morphological features and differing life-history traits led to their classification as two closely related, although separate species. The lack of genetic differentiation appears to be due to a recent divergence event not yet reflected in the beetles molecular make-up. Thus, the Russian Hylochares was described as a new species. The Finnish species, previously considered as locally extinct, was recognized as endangered. Even if, due to their identical genetic make-up, the populations had been regarded as conspecific, conservation strategies based on prior knowledge from Russia would not have guaranteed the survival of the Finnish beetle. Therefore, new conservation actions based on detailed studies of the, Mutaatiot ja geneettinen rekombinaatio ovat luonnonpopulaatioiden perinnöllisen muuntelun perusta. Evoluutio muovaa tätä ainesta luonnonvalinnan ja ajautumisen kautta. On ilmeistä, että mutaatiofrekvenssin tutkiminen valaisee niin evoluution kulkua kuin eliöiden sukulaisuussuhteita. Tässä väitöskirjassa tutkittiin kahta molekyylibiologien työkalua, DNA-viivakoodia ja molekyylikelloa. Työn ensimmäisessä osassa tarkasteltiin eri geenialueiden kykyä luonnehtia lähisukuisia lajeja ja tämän kysymyksen merkitystä luonnonsuojelubiologiassa. Työn toisessa osassa tutkittiin hypoteesia, jonka mukaan hyönteisten mutaatiofrekvenssi noudattaa molekyylikelloa. Kohteena oli erityisesti hypoteesi, jonka mukaan molekyylikellon tarkkuutta voi parantaa huomioimalla eri lajien lämpötila. DNA-viivakoodauksella otaksutaan voitavan erottaa lajeja toisistaan tietyn geenialueen avulla. Tutkimuksessa verrattiin Suomesta hiljattain uudelleen löydetyn kovakuoriaisen, haapasepikän ja lajin venäläisten yksilöiden geenialueita keskenään. Kaikki yksilöt osoittautuivat identtisiksi jokaisen tutkitun geenialueen suhteen. Toisaalta suomalaiset ja venälaiset yksilöt erosivat toisistaan niin elintapojensa kuin rakenteensa suhteen. Suomessa laji kehittyi kovapuisissa halavissa ja mustuvapajuissa, Venäjällä suurissa lahoavissa haavoissa. Suomalaiset yksilöt olivat myös hiukan hoikkarakenteisempia kuin venäläiset ja molempien muotojen koiraiden sukupuolielimet olivat selvästi toisistaan eroavat. Näillä perusteilla Suomesta kuvattu laji piti vanhan tieteellisen nimensä, Hylochares cruentatus (Gyllenhal), mutta suomeksi se ristittiin uudelleen halavasepikäksi. Haapasepikän nimi jätettiin maamme rajojen ulkopuolella elävälle lajille, joka kuvattiin tieteelle uutena nimellä Hylochares populi Brüstle -- Muona. Haapasepikän biologiaa ei tunnettu Suomesta, koska siitä tiedettiin vain kaksi hyvin vanhaa löytöä. Lajin katsottiin hävinneen maastamme suurten haapojen myötä, elihän se tällaisissa puissa Venäjän Karja

A new subgenus [Allograpta (Costarica Mengual & Thompson), type Allograpta zumbadoi Thompson], and one new species [Allograpta (Costarica) nishida Mengual & Thompson; type-locality: Costa Rica, type-depository: Instituto Nacional de Biodiversidad de Costa Rica] of flower flies (Diptera: Syrphidae) are described from the Neotropical biotic region. A checklist of the world species of Allograpta including synonyms is provided, and a key to and diagnoses of the subgenera are also supplied. The phylogenetic relationships among Allograpta species, representing all hitherto detected morphological diversity of the genus, and related genera were studied under parsimony based on morphological characters.

This study addressed the large-scale molecular zoogeography in two brackish water bivalve molluscs, Macoma balthica and Cerastoderma glaucum, and genetic signatures of the postglacial colonization of Northern Europe by them. The traditional view poses that M. balthica in the Baltic, White and Barents seas (i.e. marginal seas) represent direct postglacial descendants of the adjacent Northeast Atlantic populations, but this has recently been challenged by observations of close genetic affinities between these marginal populations and those of the Northeast Pacific. The primary aim of the thesis was to verify, quantify and characterize the Pacific genetic contribution across North European populations of M. balthica and to resolve the phylogeographic histories of the two bivalve taxa in range-wide studies using information from mitochondrial DNA (mtDNA) and nuclear allozyme polymorphisms. The presence of recent Pacific genetic influence in M. balthica of the Baltic, White and Barents seas, along with an Atlantic element, was confirmed by mtDNA sequence data. On a broader temporal and geographical scale, altogether four independent trans-Arctic invasions of Macoma from the Pacific since the Miocene seem to have been involved in generating the current North Atlantic lineage diversity. The latest trans-Arctic invasion that affected the current Baltic, White and Barents Sea populations probably took place in the early post-glacial. The nuclear genetic compositions of these marginal sea populations are intermediate between those of pure Pacific and Atlantic subspecies. In the marginal sea populations of mixed ancestry (Barents, White and Northern Baltic seas), the Pacific and Atlantic components are now randomly associated in the genomes of individual clams, which indicates both pervasive historical interbreeding between the previously long-isolated lineages (subspecies), and current isolation of these populations from the adjacent pure Atlantic populations. These mixed pop, Pohjolaa reunustavien merialueiden eliöyhteisöt ovat muotoutuneet vasta jääkauden jälkeen, viimeisten 10 000 vuoden aikana. Tätä ennen aluetta peitti paksu mannerjää, jonka väistyttyä vapautuneet reunameret, etenkin Itämeri, ovat olleet valtamerilajeille haasteellisia elinympäristöjä. Niissä menestyvät vain harvat, vähäsuolaisuutta sietävät lajit. Näitä ovat liejusimpukka (Macoma balthica) ja idänsydänsimpukka (Cerastoderma glaucum). Ennen oletettiin, että reunamerten simpukat polveutuvat suoraan läheisen Koillis-Atlantin populaatioista. Uudempi tutkimus on kuitenkin viitannut täysin erilaiseen historiaan, kun eräiden Itämeren simpukoiden lähimmät sukulaiset ovatkin löytyneet Tyynestämerestä. Väitöstutkimuksessa jäljitettiin simpukoiden historiaa Pohjois-Euroopan reunamerissä ja laajemmin pohjoisissa merissä erilaisten geenituntomerkkien sisältämän informaation perusteella. Mitokondrio-DNA:n polveutumistutkimus osoitti, että liejusimpukat käsittävät useita eri kehityslinjoja, jotka historian saatossa ovat eri aikoina itsenäisesti saapuneet Tyynestämerestä Jäämeren kautta Atlantin altaaseen. Euroopassa näitä linjoja on kaksi: jo ennen jääkausiaikaa saapunut Pohjanmeren linja (alalaji) ja toinen, ilmeisesti vasta pian jääkauden jälkeen Tyynestämerestä saapunut, Itämeressä ja Vienanmeressä nyt vallitseva linja. Idänsydänsimpukan osalta, joka puolestaan on pelkästään eurooppalainen laji, tutkimus kumosi oletuksen perusjaosta Välimeren ja Atlantin populaatioiden välillä; vain Mustanmeren ja itäisimmän Välimeren sydänsimpukat eroavat toisesta, yleiseurooppalaisesta kehityslinjasta, jonka levinneisyys ulottuu Kreikasta Itämereen. Erityisesti liejusimpukka-analyysin tulokset muuttavat perusteellisesti perinteisiä käsityksiä Itämeren ja Pohjois-Euroopan muiden reunamerten eliöstön historiasta ja omaperäisyydestä. Ensiksi, hiljattain mutta kuitenkin ennen ihmisen väliintuloa Tyynestämerestä saapunut simpukkakanta edustaa täällä ennen tuntematonta eläinmaantieteellistä element

Cortinarius is the largest genus of Agaricales with a worldwide distribution. So far, over 4000 Cortinarius names and combinations have been published. Cortinarius spp. form ectomycorrhizae with different trees and shrubs. A majority of the Cortinarius species have narrow ecological preferences and many form ectomycorrhiza with only one or few host species. The subgenus Telamonia sensu lato (s. lat.), comprising the greatest number of species, is the most poorly known of the subgenera of Cortinarius. The centre of diversity is in the northern hemisphere, although some species of the group are also recognized in the southern hemisphere. The aim of this thesis was to study the taxonomy of Cortinarius subgenus Telamonia p.p. species based on morphological and molecular data, as well as to study the ecology and distribution of the species in North Europe. The taxonomical problems encountered and the difficulty in finding and studying all the relevant names and types slowed down the study. The diversity of the subgenus Telamonia s. lat. in North Europe (excluding sect. Hydrocybe, Icrustati and Anomali) was found to be far greater than previously thought. Even many of the common species have not yet been described. So far, ca. 200 species have been recognised from the Nordic countries, but the sampling in most groups does not cover the whole diversity and especially the southern deciduous forest species are underrepresented in our study. In most cases phylogenetic (only based on ITS data) and morphological species recognition were in concordance, but in a few cases morphologically delimited species had almost identical ITS sequences, raising the question as to whether ITS is always variable enough for species recognition. The opposite situation, in which a morphologically uniform species included two phylogenetically distinct lineages, however, was also encountered, suggesting the possibility of cryptic species in Cortinarius. In our studies no taxa below species level we, Seitikit ovat helttasienten suurin suku. Ne ovat ektomykorritsasieniä ja niillä on huomattava merkitys metsäpuiden ravinnetaloudelle. Monet seitikit ovat herkkiä ympäristönmuutosten ilmentäjiä ja niitä on käytetty arvokkaiden luontokohteiden indikaattorilajeina mm. Ruotsissa ja Tanskassa. Huopaseitikit on seitikkien alasuvuista kaikkein suurin ja huonoimmin tunnettu. Sen levinneisyyden pääpaino on pohjoisella pallonpuoliskolla. Tämän tutkimuksen tarkoituksena oli selvittää huopaseitikkien taksonomiaa, ekologiaa ja levinneisyyttä Pohjois-Euroopassa. Huopaseitikkejä kasvaa Pohjois-Euroopassa paljon enemmän kuin aiemmin oletettiin. Tähän mennessä olemme tunnistaneet jo yli 200 lajia, mutta kaikkia lajiryhmiä ei ole tutkittu yhtä tarkasti ja etenkin eteläisempien lehtimetsien lajit olivat tutkimuksessamme aliedustettuina. Lajit on rajattu morfogeneettisen lajikonseptin perusteella eli ne voidaan tunnistaa myös morfologisesti. Osa lajeista sisältää kuitenkin geneettisesti eriytyneitä yksiköitä, jotka saattavat olla kryptisiä lajeja. Kryptisiä lajeja on löydetty myös muista sienisuvuista ja voikin olla, että huopaseitikkien todellinen lajimäärä on arvioimaamme vieläkin suurempi. Uudet molekyylisystemaattiset tutkimusmenetelmät ovat haastaneet myös käsityksemme seitikkien suvun sisäisestä luokittelusta. Osa aiemmin käytetyistä huopaseitikkien sisäisistä ryhmittelyistä osoittautui tutkimuksissamme epäluonnollisiksi. Tutkimuksen tuottama aineisto lisää olennaisesti tietämystä pohjoismaiden seitikkilajistosta sekä sen levinneisyydestä ja ekologiasta. Pohjois-Euroopasta löytyy vielä runsaasti meillä ennestään tuntemattomia ja myös kokonaan tieteelle uusia seitikkilajeja. Monet aiemmin tuntemattomista lajeista ovat hyvin yleisiä ja osa on myös melko helposti maastossa tunnistettavia. Tutkimuksemme ovat keskittyneet Pohjois-Eurooppaan, mutta löysimme myös joitakin yhtäläisyyksiä Pohjois-Amerikan lajiston kanssa. Vielä on kuitenkin liian aikaista arvioida seitikkien monimuotoisuu