DOCSLIB.ORG

The Revised Classification of Eukaryotes

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
The Revised Classification of Eukaryotes The Journal of * Published by the International Society of Eukaryotic Microbiology W? ?ocie,yProtistologists of J. Eukaryot. Microbiol., 59(5), 2012 pp. 429-493 ©2012 The Author(s) Journal o f Eukaryotic Microbiology © 2012 International Society of Protistologists DOI: 10.111 l/j.l550-7408.2012.00644.x The Revised Classification of Eukaryotes SINA M. ADL,ab ALASTAIR G. B. SIMPSON,b CHRISTOPHER E. LANE," JULIUS LUKES,d DAVID BASS," SAMUEL S. BOWSER/ MATTHEW W. BROWN/ FABIEN BURKI,h MICAH DUNTHORN/ VLADIMIR HAMPL,j AARON HEISS,b MONA HOPPENRATH,k ENRIQUE LARA,1 LINE LE GALL,111 DENIS H. LYNN,”’1 HILARY MCMANUS," EDWARD A. D. MITCHELL,1 SHARON E. MOZLEY-STANRIDGE/ LAURA W. PARFREY,“ JAN PAWLOWSKI/ SONJA RUECKERT/ LAURA SHADWICK,* CONRAD L. SCHOCH," ALEXEY SMIRNOV' and FREDERICK W. SPIEGEL* aDepartment of Soil Science, University of Saskatchewan, Saskatoon, SK, S7N 5A8, Canada, and bDepartment of Biology, Dalhousie University, Halifax, NS, B3H 4R2, Canada, and cDepartment of Biological Sciences, University of Rhode Island, Kingston, Rhode Island, 02881, USA, and dBiology Center and Faculty o f Sciences, Institute of Parasitology, University of South Bohemia, Ceské Budêjovice, Czech Republic, and eZoology Department, Natural History Museum, London, SW 7 5BD, United Kingdom, and fWadsworth Center, New York State Department of Health, Albany, New York, 12201, USA, and gDepartment of Biochemistry, Dalhousie University, Halifax, NS, B3H 4R2, Canada, and hDepartment of Botany, University of British Columbia, Vancouver, BC, V6T 1Z4, Canada, and 1Department of Ecology, University of Kaiserslautern, 67663, Kaiserslautern, Germany, and 3Department of Parasitology, Charles University, Prague, 128 43, Predict 2, Czech Republic, and kForschungsinstitut Senckenberg, D ZM B - Deutsches Zentrum für Marine Biodiversitätsforschung, D-26382, Wilhelmshaven, Germany, and 1Institute of Biology, University of Neuchâtel, Neuchâtel, CH-2009, Switzerland, and mMuséum National d’Histoire Naturellem, UMR 7138 Systématique, Adaptation et Evolution, Paris, 75231, Cedex Paris 05, France, and nDepartment of Integrative Biology, University of Guelph, Guelph, ON, NIG 2W1, Canada, and °Department of Biological Sciences, LeMoyne College, Syracuse, New York, 13214, USA, and pDepartment of Biology, Middle Georgia College, Cochran, Georgia, 31014, USA, and qDepartment of Chemistry and Biochemistry, University of Colorado, Boulder, Colorado, 80309, USA, and TDepartment of Genetics and Evolution, University of Geneva, 1211, Geneva 4, Switzerland, and sSchool of Life, Sport and Social Sciences, Edinburgh Napier University, Edinburgh, EH11 4BN, United Kingdom, and department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas, 72701, USA, and uGenBank taxonomy, NIH/NLM /NCBI, Bethesda, Maryland, 20892-6510, USA, and ''Department of Invertebrate Zoology, St.Petersburg State University, St. Petersburg, 199034, Russia ABSTRACT. This revision of the classification of eukaryotes, which updates that of Adi et al.[J. Eukaryot. Microbiol. 52 (2005) 399], retains an emphasis on the protists and incorporates changes since 2005 that have resolved nodes and branches in phyloge­ netic trees. Whereas the previous revision was successful in re-introducing name stability to the classification, this revision provides a classification for lineages that were then still unresolved. The supergroups have withstood phylogenetic hypothesis testing with some modifications, but despite some progress, problematic nodes at the base of the eukaryotic tree still remain to be statistically resolved. Looking forward, subsequent transformations to our understanding of the diversity of life will be from the discovery of novel lineages in previously under-sampled areas and from environmental genomic information. Key Words.Algae, amoebae, biodiversity, ciliates, flagellates, fungi, parasites, protozoa, systematics, taxonomy. HE classification proposed by Adi et al. (2005) on behalf current revision reflects the need to have a classification of of The Society established name stability as well as a syn­ protistan eukaryotes that incorporates recent advances thesisT of the overall structure of the classification of eukary­ wrought both by the widespread use of phylogenomic-scale otes, based on the information available at that time, and phylogenetic analyses and by massively increased taxon sam­ after the upheaval introduced by molecular phylogenetic stud­ pling in rRNA-based phylogenies, partly due to a renaissance ies over the preceding two decades. Overall, the system pro­ in novel organism discovery. With the current revision, we posed was conservative enough to largely avoid erroneous or have again tried to strike a conservative balance between premature groupings, whilst eliminating wherever possible updating the classification where needed and avoiding formal known polyphyletic groups or groups of convenience, encour­ recognition of uncertain groupings where further investigation aging correction of many of the errors in text books. The would be warranted. One notable advance since 2005 is the consolidation of a classification founded on robust phylogenetic relatedness. The super-groups formalized by Adi et al. (2005) are mostly retained, although some have been assembled into still higher Correponding Author: Sina M. Adi, Department of Soil Science, order groupings (Table 1, see below). One notable exception is University of Saskatchewan, 51 Campus Drive, Saskatoon, SK S7N 5A8, Canada-Telephone number: +306 966 6866; FAX number: the Chromalveolata, which was retained then as useful + 306 966 6881; e-mail:[email protected] although controversial, and with the authors noting concerns 1 Present Address: Department of Zoology, University of British with this grouping of Cryptophyceae, Haptophyta, Strameno- Columbia, Vancouver, BC, V6T 1Z4, Canada. piles, and Alveolata. Since then, evidence has mounted that 429 430 J. EUKARYOT. MICROBIOL., 59, NO. 5, SEPTEMBER-OCTOBER 2012 Table 1. The classification of eukaryotes at the highest ranks. system despite changes; and (iv) separating naming clades from assembling nested hierarchies - in contrast to rank- Super-groups Examples based nomenclature that treats these steps as part of the same process (Adi et al., 2007; Cantino, 2004; Krön, 1997; Pleijel and Amoebozoa Tubulinea Rouse 2003). This approach provides a more stable classifica­ Mycetozoa tion that preserves names, while allowing revisions to reflect our Fungi Amorphea Opisthokonta changing understanding of evolutionary history. We have also Choanomonada Metazoa relied on ideas borrowed from phylogenetic nomenclature, dis­ Apusomonada tinguishing groups with definitions based on apomorphies, Breviata nodes, branches, or combinations of these. These kinds of defi­ Excavata Metamonada nitions are more suited to a classification based on phylogenetic Malawimonas B trees, and can be written as phylogenetic hypotheses that can be o Discoba tested. 5* Diaphoretickes Cryptophyceae The most significant changes introduced in this revision are ■a Centrohelida as follows: m Telonemia First, we recognize the grouping of Amoebozoa with Opi- Haptophyta Sar Cercozoa sthokonta. Since 2005, this has become a commonly recog­ Foraminifera nized probable clade, and at present it is usually referred to “Radiolaria” by the informal name “unikont”, sometimes rendered as the Alveolata more formal sounding “Unikonta”. However, the underlying Stramenopiles hypothesis of a monociliated (with only one ciliated basal Archaeplastida Glaucophyta body) ancestry for this cluster of organisms (Cavalier-Smith, Rhodophyceae 2002) is almost certainly incorrect (Kim et al. 2006; Roger Chloroplastida and Simpson, 2009). There is no requirement that names of Incertae sedis Eukaryota Incertae sedis, and taxa reflect the ancestral state of the clade. However, the name table 3 “unikonts” causes confusion because of the apomorphy hypothesized for the ancestral character. To address, this we introduce a new formal name for the probable clade. We have formalized this clade as a new taxon, Amorphea, with a node­ Chromalveolates are probably polyphyletic (Baurain et al. based phylogenetic definition: 2010; Parfrey et al. 2010; Stiller et al. 2009). Instead, multi­ gene phylogenetics and phylogenomic studies generally sup­ port Stramenopiles and Alveolata as specifically related to Amorphea: the least inclusive clade containing H o m o sapi­ Rhizaria (Burki et al. 2009, 2010, 2012; Hampl et al. 2009; ens Linnaeus 1758, Neurospora crassa Shear & Dodge 1927 Parfrey et al. 2010). The two remaining major lineages that (both Opisthokonta), and Dictyostelium discoideum Raper were formerly assigned to the chromalveolates - Haptophyta 1935 (Amoebozoa). This is a node-based definition in and Cryptophyceae/cryptomonads - have been more challeng­ which all of the specifiers are extant; it is intended to apply ing to place phylogenetically (Burki et al. 2010, 2012), and are to a crown clade; qualifying clause - the name does not two examples of several where stable, deep relationships still apply if any of the following fall within the specified clade remain to be established. Analyses with abundant data for - Arabidopsis thaliana (Linnaeus) Heynhold 1842 (Archaep- each taxon are subject to systematic biases that can lead to lastida), Tetrahymena thermophila Nanney & McCoy 1976 high support for incorrect clades (Hedtke et al. 2006; Zwickl (Alveolata), Thalassiosira pseudonana Hasle & Hiemdal and Hillis 2002). Broader taxonomic sampling is
Load more

Recommended publications
  • New Zealand's Genetic Diversity
    1.13 NEW ZEALAND’S GENETIC DIVERSITY NEW ZEALAND’S GENETIC DIVERSITY Dennis P. Gordon National Institute of Water and Atmospheric Research, Private Bag 14901, Kilbirnie, Wellington 6022, New Zealand ABSTRACT: The known genetic diversity represented by the New Zealand biota is reviewed and summarised, largely based on a recently published New Zealand inventory of biodiversity. All kingdoms and eukaryote phyla are covered, updated to refl ect the latest phylogenetic view of Eukaryota. The total known biota comprises a nominal 57 406 species (c. 48 640 described). Subtraction of the 4889 naturalised-alien species gives a biota of 52 517 native species. A minimum (the status of a number of the unnamed species is uncertain) of 27 380 (52%) of these species are endemic (cf. 26% for Fungi, 38% for all marine species, 46% for marine Animalia, 68% for all Animalia, 78% for vascular plants and 91% for terrestrial Animalia). In passing, examples are given both of the roles of the major taxa in providing ecosystem services and of the use of genetic resources in the New Zealand economy. Key words: Animalia, Chromista, freshwater, Fungi, genetic diversity, marine, New Zealand, Prokaryota, Protozoa, terrestrial. INTRODUCTION Article 10b of the CBD calls for signatories to ‘Adopt The original brief for this chapter was to review New Zealand’s measures relating to the use of biological resources [i.e. genetic genetic resources. The OECD defi nition of genetic resources resources] to avoid or minimize adverse impacts on biological is ‘genetic material of plants, animals or micro-organisms of diversity [e.g. genetic diversity]’ (my parentheses).
    [Show full text]
  • Checklist, Assemblage Composition, and Biogeographic Assessment of Recent Benthic Foraminifera (Protista, Rhizaria) from São Vincente, Cape Verdes
    Zootaxa 4731 (2): 151–192 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4731.2.1 http://zoobank.org/urn:lsid:zoobank.org:pub:560FF002-DB8B-405A-8767-09628AEDBF04 Checklist, assemblage composition, and biogeographic assessment of Recent benthic foraminifera (Protista, Rhizaria) from São Vincente, Cape Verdes JOACHIM SCHÖNFELD1,3 & JULIA LÜBBERS2 1GEOMAR Helmholtz-Centre for Ocean Research Kiel, Wischhofstrasse 1-3, 24148 Kiel, Germany 2Institute of Geosciences, Christian-Albrechts-University, Ludewig-Meyn-Straße 14, 24118 Kiel, Germany 3Corresponding author. E-mail: [email protected] Abstract We describe for the first time subtropical intertidal foraminiferal assemblages from beach sands on São Vincente, Cape Verdes. Sixty-five benthic foraminiferal species were recognised, representing 47 genera, 31 families, and 8 superfamilies. Endemic species were not recognised. The new checklist largely extends an earlier record of nine benthic foraminiferal species from fossil carbonate sands on the island. Bolivina striatula, Rosalina vilardeboana and Millettiana milletti dominated the living (rose Bengal stained) fauna, while Elphidium crispum, Amphistegina gibbosa, Quinqueloculina seminulum, Ammonia tepida, Triloculina rotunda and Glabratella patelliformis dominated the dead assemblages. The living fauna lacks species typical for coarse-grained substrates. Instead, there were species that had a planktonic stage in their life cycle. The living fauna therefore received a substantial contribution of floating species and propagules that may have endured a long transport by surface ocean currents. The dead assemblages largely differed from the living fauna and contained redeposited tests deriving from a rhodolith-mollusc carbonate facies at <20 m water depth.
    [Show full text]
  • Next-Generation Environmental Diversity Surveys of Foraminifera: Preparing the Future Jan Pawlowski, Franck Lejzerowicz, Philippe Esling
    Next-Generation Environmental Diversity Surveys of Foraminifera: Preparing the Future Jan Pawlowski, Franck Lejzerowicz, Philippe Esling To cite this version: Jan Pawlowski, Franck Lejzerowicz, Philippe Esling. Next-Generation Environmental Diversity Sur- veys of Foraminifera: Preparing the Future . Biological Bulletin, Marine Biological Laboratory, 2014, 227 (2), pp.93-106. 10.1086/BBLv227n2p93. hal-01577891 HAL Id: hal-01577891 https://hal.archives-ouvertes.fr/hal-01577891 Submitted on 28 Aug 2017 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/268789818 Next-Generation Environmental Diversity Surveys of Foraminifera: Preparing the Future Article in Biological Bulletin · October 2014 Source: PubMed CITATIONS READS 26 41 3 authors: Jan Pawlowski Franck Lejzerowicz University of Geneva University of Geneva 422 PUBLICATIONS 11,852 CITATIONS 42 PUBLICATIONS 451 CITATIONS SEE PROFILE SEE PROFILE Philippe Esling Institut de Recherche et Coordination Acoust… 24 PUBLICATIONS 551 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: UniEuk View project KuramBio II (Kuril Kamchatka Biodiversity Studies II) View project All content following this page was uploaded by Jan Pawlowski on 30 December 2015.
    [Show full text]
  • The Revised Classification of Eukaryotes
    See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/231610049 The Revised Classification of Eukaryotes Article in Journal of Eukaryotic Microbiology · September 2012 DOI: 10.1111/j.1550-7408.2012.00644.x · Source: PubMed CITATIONS READS 961 2,825 25 authors, including: Sina M Adl Alastair Simpson University of Saskatchewan Dalhousie University 118 PUBLICATIONS 8,522 CITATIONS 264 PUBLICATIONS 10,739 CITATIONS SEE PROFILE SEE PROFILE Christopher E Lane David Bass University of Rhode Island Natural History Museum, London 82 PUBLICATIONS 6,233 CITATIONS 464 PUBLICATIONS 7,765 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Biodiversity and ecology of soil taste amoeba View project Predator control of diversity View project All content following this page was uploaded by Smirnov Alexey on 25 October 2017. The user has requested enhancement of the downloaded file. The Journal of Published by the International Society of Eukaryotic Microbiology Protistologists J. Eukaryot. Microbiol., 59(5), 2012 pp. 429–493 © 2012 The Author(s) Journal of Eukaryotic Microbiology © 2012 International Society of Protistologists DOI: 10.1111/j.1550-7408.2012.00644.x The Revised Classification of Eukaryotes SINA M. ADL,a,b ALASTAIR G. B. SIMPSON,b CHRISTOPHER E. LANE,c JULIUS LUKESˇ,d DAVID BASS,e SAMUEL S. BOWSER,f MATTHEW W. BROWN,g FABIEN BURKI,h MICAH DUNTHORN,i VLADIMIR HAMPL,j AARON HEISS,b MONA HOPPENRATH,k ENRIQUE LARA,l LINE LE GALL,m DENIS H. LYNN,n,1 HILARY MCMANUS,o EDWARD A. D.
    [Show full text]
  • The Evolution of Early Foraminifera
    The evolution of early Foraminifera Jan Pawlowski†‡, Maria Holzmann†,Ce´ dric Berney†, Jose´ Fahrni†, Andrew J. Gooday§, Tomas Cedhagen¶, Andrea Haburaʈ, and Samuel S. Bowserʈ †Department of Zoology and Animal Biology, University of Geneva, Sciences III, 1211 Geneva 4, Switzerland; §Southampton Oceanography Centre, Empress Dock, European Way, Southampton SO14 3ZH, United Kingdom; ¶Department of Marine Ecology, University of Aarhus, Finlandsgade 14, DK-8200 Aarhus N, Denmark; and ʈWadsworth Center, New York State Department of Health, P.O. Box 509, Albany, NY 12201 Communicated by W. A. Berggren, Woods Hole Oceanographic Institution, Woods Hole, MA, August 11, 2003 (received for review January 30, 2003) Fossil Foraminifera appear in the Early Cambrian, at about the same loculus to become globular or tubular, or by the development of time as the first skeletonized metazoans. However, due to the spiral growth (12). The evolution of spiral tests led to the inadequate preservation of early unilocular (single-chambered) formation of internal septae through the development of con- foraminiferal tests and difficulties in their identification, the evo- strictions in the spiral tubular chamber and hence the appear- lution of early foraminifers is poorly understood. By using molec- ance of multilocular forms. ular data from a wide range of extant naked and testate unilocular Because of their poor preservation and the difficulties in- species, we demonstrate that a large radiation of nonfossilized volved in their identification, the unilocular noncalcareous for- unilocular Foraminifera preceded the diversification of multilocular aminifers are largely ignored in paleontological studies. In a lineages during the Carboniferous. Within this radiation, similar previous study, we used molecular data to reveal the presence of test morphologies and wall types developed several times inde- naked foraminifers, perhaps resembling those that lived before pendently.
    [Show full text]
  • Proposal for Practical Multi-Kingdom Classification of Eukaryotes Based on Monophyly 2 and Comparable Divergence Time Criteria
    bioRxiv preprint doi: https://doi.org/10.1101/240929; this version posted December 29, 2017. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. 1 Proposal for practical multi-kingdom classification of eukaryotes based on monophyly 2 and comparable divergence time criteria 3 Leho Tedersoo 4 Natural History Museum, University of Tartu, 14a Ravila, 50411 Tartu, Estonia 5 Contact: email: [email protected], tel: +372 56654986, twitter: @tedersoo 6 7 Key words: Taxonomy, Eukaryotes, subdomain, phylum, phylogenetic classification, 8 monophyletic groups, divergence time 9 Summary 10 Much of the ecological, taxonomic and biodiversity research relies on understanding of 11 phylogenetic relationships among organisms. There are multiple available classification 12 systems that all suffer from differences in naming, incompleteness, presence of multiple non- 13 monophyletic entities and poor correspondence of divergence times. These issues render 14 taxonomic comparisons across the main groups of eukaryotes and all life in general difficult 15 at best. By using the monophyly criterion, roughly comparable time of divergence and 16 information from multiple phylogenetic reconstructions, I propose an alternative 17 classification system for the domain Eukarya to improve hierarchical taxonomical 18 comparability for animals, plants, fungi and multiple protist groups. Following this rationale, 19 I propose 32 kingdoms of eukaryotes that are treated in 10 subdomains. These kingdoms are 20 further separated into 43, 115, 140 and 353 taxa at the level of subkingdom, phylum, 21 subphylum and class, respectively (http://dx.doi.org/10.15156/BIO/587483).
    [Show full text]
  • Chamber Arrangement Versus Wall Structure in the High-Rank Phylogenetic Classification of Foraminifera
    Editors' choice Chamber arrangement versus wall structure in the high-rank phylogenetic classification of Foraminifera ZOFIA DUBICKA Dubicka, Z. 2019. Chamber arrangement versus wall structure in the high-rank phylogenetic classification of Fora- minifera. Acta Palaeontologica Polonica 64 (1): 1–18. Foraminiferal wall micro/ultra-structures of Recent and well-preserved Jurassic (Bathonian) foraminifers of distinct for- aminiferal high-rank taxonomic groups, Globothalamea (Rotaliida, Robertinida, and Textulariida), Miliolida, Spirillinata and Lagenata, are presented. Both calcite-cemented agglutinated and entirely calcareous foraminiferal walls have been investigated. Original test ultra-structures of Jurassic foraminifers are given for the first time. “Monocrystalline” wall-type which characterizes the class Spirillinata is documented in high resolution imaging. Globothalamea, Lagenata, porcel- aneous representatives of Tubothalamea and Spirillinata display four different major types of wall-structure which may be related to distinct calcification processes. It confirms that these distinct molecular groups evolved separately, probably from single-chambered monothalamids, and independently developed unique wall types. Studied Jurassic simple bilocular taxa, characterized by undivided spiralling or irregular tubes, are composed of miliolid-type needle-shaped crystallites. In turn, spirillinid “monocrystalline” test structure has only been recorded within more complex, multilocular taxa pos- sessing secondary subdivided chambers: Jurassic
    [Show full text]
  • The Evolution of Early Foraminifera
    The evolution of early Foraminifera Jan Pawlowski†‡, Maria Holzmann†,Ce´ dric Berney†, Jose´ Fahrni†, Andrew J. Gooday§, Tomas Cedhagen¶, Andrea Haburaʈ, and Samuel S. Bowserʈ †Department of Zoology and Animal Biology, University of Geneva, Sciences III, 1211 Geneva 4, Switzerland; §Southampton Oceanography Centre, Empress Dock, European Way, Southampton SO14 3ZH, United Kingdom; ¶Department of Marine Ecology, University of Aarhus, Finlandsgade 14, DK-8200 Aarhus N, Denmark; and ʈWadsworth Center, New York State Department of Health, P.O. Box 509, Albany, NY 12201 Communicated by W. A. Berggren, Woods Hole Oceanographic Institution, Woods Hole, MA, August 11, 2003 (received for review January 30, 2003) Fossil Foraminifera appear in the Early Cambrian, at about the same loculus to become globular or tubular, or by the development of time as the first skeletonized metazoans. However, due to the spiral growth (12). The evolution of spiral tests led to the inadequate preservation of early unilocular (single-chambered) formation of internal septae through the development of con- foraminiferal tests and difficulties in their identification, the evo- strictions in the spiral tubular chamber and hence the appear- lution of early foraminifers is poorly understood. By using molec- ance of multilocular forms. ular data from a wide range of extant naked and testate unilocular Because of their poor preservation and the difficulties in- species, we demonstrate that a large radiation of nonfossilized volved in their identification, the unilocular noncalcareous for- unilocular Foraminifera preceded the diversification of multilocular aminifers are largely ignored in paleontological studies. In a lineages during the Carboniferous. Within this radiation, similar previous study, we used molecular data to reveal the presence of test morphologies and wall types developed several times inde- naked foraminifers, perhaps resembling those that lived before pendently.
    [Show full text]
  • Biocenosis De Foraminíferos Y Su Relación Con Variables Ambientales En La Plataforma Española
    Biocenosis de foraminíferos y su relación con variables ambientales en la plataforma española. Alumno: Uxue Muñoz Berruezo Tutora: Ana María Blázquez Morilla Curso académico: 2019-2020 AGRADECIMIENTOS En primer lugar, quiero dar las gracias a mis padres. Siempre me han apoyado y me han alentado a estudiar lo que quería, costase lo que costase, fuese en la ciudad que fuese. Gracias a mi madre, que ha pasado horas al teléfono conmigo, ya sea para desahogarme o simplemente para cubrir un trayecto caminando hacia la universidad. Y por pasar la última semana antes de la entrega del TFG conmigo impidiéndome que trabajase 13 horas seguidas y que se me comiesen los nervios. A mi padre, porque siempre tiene las palabras adecuadas en el momento adecuado, por enseñarme a no ser tan dura conmigo misma, por hacerme ver que lo que hago es suficiente e incluso bueno, aunque yo nunca esté satisfecha del todo. A mi hermana, Andrea. Por aguantarme siempre, por tu paciencia infinita, por todo lo que me aportas y lo poco que te lo agradezco. Muchos de mis logros son gracias a ti. Pronto tendrás tus grandes logros también. A Rubén. Por ser mi segunda familia, por enseñarme a querer Valencia, por estar todos los días (menos cuando te embarcas) al pie del cañón. A la COCSABO y la UTM por permitir la campaña que ha dado pie a mi TFG. Y a todas las personas de la universidad que pasaron años organizándola. A la tripulación y todos los técnicos del Sarmiento de Gamboa, por enseñarnos todo lo necesario a bordo y estar siempre dispuestos para lo que necesitásemos.
    [Show full text]
  • Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes
    University of Rhode Island DigitalCommons@URI Biological Sciences Faculty Publications Biological Sciences 9-26-2018 Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes Christopher E. Lane Et Al Follow this and additional works at: https://digitalcommons.uri.edu/bio_facpubs Journal of Eukaryotic Microbiology ISSN 1066-5234 ORIGINAL ARTICLE Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes Sina M. Adla,* , David Bassb,c , Christopher E. Laned, Julius Lukese,f , Conrad L. Schochg, Alexey Smirnovh, Sabine Agathai, Cedric Berneyj , Matthew W. Brownk,l, Fabien Burkim,PacoCardenas n , Ivan Cepi cka o, Lyudmila Chistyakovap, Javier del Campoq, Micah Dunthornr,s , Bente Edvardsent , Yana Eglitu, Laure Guillouv, Vladimır Hamplw, Aaron A. Heissx, Mona Hoppenrathy, Timothy Y. Jamesz, Anna Karn- kowskaaa, Sergey Karpovh,ab, Eunsoo Kimx, Martin Koliskoe, Alexander Kudryavtsevh,ab, Daniel J.G. Lahrac, Enrique Laraad,ae , Line Le Gallaf , Denis H. Lynnag,ah , David G. Mannai,aj, Ramon Massanaq, Edward A.D. Mitchellad,ak , Christine Morrowal, Jong Soo Parkam , Jan W. Pawlowskian, Martha J. Powellao, Daniel J. Richterap, Sonja Rueckertaq, Lora Shadwickar, Satoshi Shimanoas, Frederick W. Spiegelar, Guifre Torruellaat , Noha Youssefau, Vasily Zlatogurskyh,av & Qianqian Zhangaw a Department of Soil Sciences, College of Agriculture and Bioresources, University of Saskatchewan, Saskatoon, S7N 5A8, SK, Canada b Department of Life Sciences, The Natural History Museum, Cromwell Road, London, SW7 5BD, United Kingdom
    [Show full text]
  • Protista (PDF)
    1 = Astasiopsis distortum (Dujardin,1841) Bütschli,1885 South Scandinavian Marine Protoctista ? Dingensia Patterson & Zölffel,1992, in Patterson & Larsen (™ Heteromita angusta Dujardin,1841) Provisional Check-list compiled at the Tjärnö Marine Biological * Taxon incertae sedis. Very similar to Cryptaulax Skuja Laboratory by: Dinomonas Kent,1880 TJÄRNÖLAB. / Hans G. Hansson - 1991-07 - 1997-04-02 * Taxon incertae sedis. Species found in South Scandinavia, as well as from neighbouring areas, chiefly the British Isles, have been considered, as some of them may show to have a slightly more northern distribution, than what is known today. However, species with a typical Lusitanian distribution, with their northern Diphylleia Massart,1920 distribution limit around France or Southern British Isles, have as a rule been omitted here, albeit a few species with probable norhern limits around * Marine? Incertae sedis. the British Isles are listed here until distribution patterns are better known. The compiler would be very grateful for every correction of presumptive lapses and omittances an initiated reader could make. Diplocalium Grassé & Deflandre,1952 (™ Bicosoeca inopinatum ??,1???) * Marine? Incertae sedis. Denotations: (™) = Genotype @ = Associated to * = General note Diplomita Fromentel,1874 (™ Diplomita insignis Fromentel,1874) P.S. This list is a very unfinished manuscript. Chiefly flagellated organisms have yet been considered. This * Marine? Incertae sedis. provisional PDF-file is so far only published as an Intranet file within TMBL:s domain. Diplonema Griessmann,1913, non Berendt,1845 (Diptera), nec Greene,1857 (Coel.) = Isonema ??,1???, non Meek & Worthen,1865 (Mollusca), nec Maas,1909 (Coel.) PROTOCTISTA = Flagellamonas Skvortzow,19?? = Lackeymonas Skvortzow,19?? = Lowymonas Skvortzow,19?? = Milaneziamonas Skvortzow,19?? = Spira Skvortzow,19?? = Teixeiromonas Skvortzow,19?? = PROTISTA = Kolbeana Skvortzow,19?? * Genus incertae sedis.
    [Show full text]
  • New Freshwater Species of Centrohelids Acanthocystis Lyra Sp. Nov. and Acanthocystis Siemensmae Sp. Nov.(Haptista, Heliozoa, Centrohelea) from the South Urals, Russia
    Acta Protozool. (2016) 55: 231–237 www.ejournals.eu/Acta-Protozoologica ACTA doi:10.4467/16890027AP.16.024.6011 PROTOZOOLOGICA New Freshwater Species of Centrohelids Acanthocystis lyra sp. nov. and Acanthocystis siemensmae sp. nov. (Haptista, Heliozoa, Centrohelea) from the South Urals, Russia Elena A. GERASIMOVA1,2, Andrey O. PLOTNIKOV1,3 1 Center of Shared Scientific Equipment “Persistence of microorganisms”, Institute for Cellular and Intracellular Symbiosis UB RAS, Orenburg, Russia; 2 Laboratory of Water Microbiology, I.D. Papanin Institute for Biology of Inland Waters RAS, Borok, Russia; 3 Department of Hygiene and Epidemiology, Orenburg State Medical University, Orenburg, Russia Abstract. Two new species of centrohelids Acanthocystis lyra sp. nov. and A. siemensmae sp. nov. from the Pismenka River in the South Urals, Russia, have been studied with scanning electron microscopy. Cells of these species have both long and short spine scales with hollow shafts and circular basal plates. A. lyra has the long spine scales divided into two curved S-shaped branches possessing small teeth on their inner surface. The short spine scales have primary and secondary bifurcations. Every secondary branch ends with two teeth. A. siemensmae has both long and short scales with funnel-like apices, which possess small teeth. Based on the scale morphology A. lyra has been attributed to the A. turfacea species group, whereas A. siemensmae has been attributed to the A. pectinata species group, both according to classifica- tion proposed by Mikrjukov, 1997. Similarities and differences of the new species with other members of the genus Acanthocystis have been discussed. Key words: Heliozoa, Centrohelids, Acanthocystis, protists, SEM, taxonomy.
    [Show full text]